Book of Abstracts - 18th International Congress of Arachnology 2010

University of Podlasie
&
International Society of Arachnology
Book of Abstracts
Editor: Marek Żabka
Siedlce 2010
Congress Scientific Committee
Dr. Leon Baert, Brussels, Belgium; Dr. Jonathan Coddington, Washington DC,
USA; Prof. Ansie Dippenaar-Schoeman, Pretoria, South Africa; Dr. Charles
Griswold, San Francisco, USA; Dr. Jason Dunlop – ISA Secretary, Berlin,
Germany; Dr. Mark Harvey, Welshpool, Australia; Prof. Daiqin Li, Singapore;
Dr. Kirill G. Mikhailov, Moscow, Russia; Dr. Norman Platnick, New York,
USA; Prof. Jerzy Prószyński, Warsaw, Poland; Dr. Ferenc Samu, Budapest,
Hungary; Dr. Nikolaj Scharff – ISA President, Copenhagen, Denmark; Dr. Cor
Vink, Lincoln, New Zealand; Dr. Richardo Pinto-da-Rocha, São Paulo, Brazil
Congress Honorary Committee

Prof. Barbara Kudrycka – Minister for Science and Higher Education
Prof. Antoni Jówko – Rector of the University of Podlasie
Mr. Wojciech Kudelski – Mayor of the City of Siedlce
Congress Organizing Committee

Prof. Marek Żabka – chairman, Dr. Barbara Patoleta – secretary, Dr. Izabela
Hajdamowicz, Dr. Joanna Gardzińska, Dr. Piotr Jastrzębski, Dr. Marzena
Stańska, Dr. Maciej Bartos
Technical assistance: Mrs. Małgorzata Kozłowska, Mr. Łukasz Nicewicz
Congress logo: Joanna Gardzińska
ISBN: 978-83-7051-575-1
The publication co-financed by the Polish Ministry for Science and Higher
Education (grant for the University of Podlasie), the International Society of
Arachnology and the City of Siedlce
Akademia Podlaska
ul. Konarskiego 2, 08-110 Siedlce
Format B-5
Druk: ELPIL, Siedlce
In Memoriam
To Colleagues and Friends – Arachnologists,

who have passed away over the last 3 years
Joachim Adis
4 March 1950 - 29 August 2007
Ekaterina Mikhailovna Andreeva
(Katarzyna Andrejewa Prószyńska)
16 November 1941 - 18 September 2008
Nina Sergeevna Azsheganova
8 March 1914 - 8 March 2008
James Carico
20 March 1937 - 31 March 2009
Song Daxiang
9 May 1935 - 21 January 2008
Lyndsay McLaren Forster
19 September 1925 - 20 January 2009
Jean Kekenbosch
17 March 1921 - 13 January 2009
April Kinchloe
16 March 2008
Nancy Kreiter
30 December 2007
Erich Kritscher
22 February 1927 - 27 February 2010
Gershom Levy
1937 - 2009
(Photo G. Simon)
Jean-Pierre Maelfait
1 June 1951 - 6 February 2009
Robert Wetsel Mitchell
25 April 1933 - 18 March 2010
(Photo: William R. Elliott)
Pavel Iustinovitch Marikovski
1912 - 2008
Sergei Vladimirovitch Ovtchinnikov
5 January 1958 - 18 December 2007
Michael H. Robinson
1929 - 20 March 2008
Michael Saaristo
1 September 1938 - 27 April 2008
Manuel Ángel González Sponga
30 April 1929 - 1 March 2009
Changmin Yin
4 October 1923 - 4 October 2009
Abstracts
Congress Plenary Speakers
Prof. Friedrich Barth, Universität Wien, Austria
Dr. William Eberhard, Smithsonian Tropical Research Institute, Costa Rica
Prof. Mark Elgar, University of Melbourne, Australia
Prof. Gonzalo Giribet, Harvard University, Cambridge Mass., USA
Dr. Rudy Jocqué, Royal Museum for Central Africa, Tervuren, Belgium
Prof. Wayne Maddison, University of British Columbia, Vancouver, Canada
Dr. Robert Raven, Queensland Museum, Brisbane, Australia
Prof. Paul Selden, University of Kansas, Lawrence, USA
Prof. Gabriele Uhl, Universität Greifswald, Germany
Dr. Samuel Zschokke, University of Basel, Switzerland
Book of Abstracts, 18th International Congress of Arachnology 2010, Siedlce, Poland
Wstęp
Pajęczaki to fascynujące zwierzęta. Pierwsze kopalne ślady ich istnienia
pochodzą sprzed blisko 400 milionów lat. Dziś znamy kilkadziesiąt tysięcy
gatunków (oprócz roztoczy), zasiedlających różne strefy klimatyczne i ekosystemy:
od skrajnie nieprzyjaznych pustyń, wysokich gór i podbiegunowej tundry, po
tropikalne lasy, sawanny i stepy. Wiele gatunków bytuje w miejscach wilgotnych i
okresowo zalewanych a nieliczne żyją w wodzie na stałe.
Rysunki i ryty naskalne dowodzą, że niektóre pajęczaki, zwłaszcza pająki i
skorpiony, budzą naszą fascynację i strach od tysięcy lat. Także współcześnie
zwierzęta te obecne są w mitach i wierzeniach, a jako bohaterowie filmów
fabularnych, komiksów i stron internetowych, zyskują wręcz rangę zwierzęcych
celebrytów.
Miarą ewolucyjnego sukcesu pajęczaków jest nie tylko ich długa historia,
różnorodność zajmowanych środowisk i liczba gatunków, ale także mnogość
strategii życiowych, zachowań związanych z rozrodem, polowaniem, życiem
społecznym i dyspersją. Pajęczaki są ważnym elementem sieci troficznych: jako
drapieżcy regulują liczebność swych ofiar, ale i same mają wielu wrogów.
Wszystkie te zagadnienia są przedmiotem zainteresowania współczesnej
arachnologii. Wśród jej najważniejszych nurtów kilka rozwija się szczególnie
dynamicznie. Domeną systematyki i biogeografii jest m. in. badanie pokrewieństw
między taksonami i rekonstruowanie historii faun kontynentalnych (regionalnych).
Prace z zakresu ekologii i bioróżnorodności dotyczą funkcjonowania zgrupowań
pajęczaków w różnych ekosystemach i ich roli w ocenie stanu środowiska. Badania
etologiczne i neurobiologiczne pozwalają zrozumieć uniwersalne mechanizmy
zachowań rozrodczych, łowieckich i społecznych a także poznać wyrafinowane
sposoby komunikowania się. Efektem badań nad wykorzystaniem jadu jest
wyodrębnienie substancji biologicznie czynnych o znaczeniu medycznym, zaś prace
nad własnościami przędzy i procesem jej produkcji zmierzają do uzyskania
supermateriałów o niezwykłych własnościach mechanicznych.
Nieodłączną cechą współczesnej arachnologii jest upowszechnienie się metod
biologii molekularnej i technik mikroskopii elektronowej. Zastosowanie
modelowania komputerowego pozwala na prognozowanie procesów ekologicznych
i zjawisk ewolucyjnych, statystyczne analizy wyników osiągają nieznany dotąd
stopień zaawansowania zaś elektroniczne katalogi, bazy danych i publikacje
ogromnie ułatwiają obieg informacji naukowej.
Prezentowany tu zbiór abstraktów daje jedynie wyobrażenie o różnorodności
tematyki poruszanej w trakcie Kongresu. Pełne teksty prac ukażą się w Journal of
Arachnology i innych prestiżowych wydawnictwach, do których odsyłamy
czytelników.
Organizacja Kongresu to efekt zbiorowego wysiłku i życzliwości wielu osób,

instytucji i międzynarodowej społeczności arachnologicznej.
Patronat honorowy ze strony Pani Minister Nauki i Szkolnictwa Wyższego,
Rektora Akademii Podlaskiej i Prezydenta Miasta Siedlce nadał wydarzeniu
wyjątkową rangę.
47
Władze Akademii Podlaskiej, Dziekani Wydziałów Przyrodniczego i Nauk

Ścisłych, Dyrekcja Biblioteki Głównej, Koleżanki i Koledzy z Instytutu Biologii,
studenci oraz pracownicy administracji i obsługi pomagali na miarę swych
możliwości i kompetencji.
Międzynarodowe Towarzystwo Arachnologiczne (ISA) - współorganizator
Kongresu, udzieliło finansowego wsparcia a Prezydent ISA - dr Nikolaj Scharff i
Sekretarz - dr Jason Dunlop współpracowali z Komitetem Organizacyjnym na
wszystkich etapach przygotowań.
Dziękujemy Sponsorom i Partnerom, którzy, rozumiejąc rolę nauki w
cywilizacyjnym rozwoju, udzielili pomocy finansowej, rzeczowej i logistycznej.
Wyrazy wdzięczności kierujemy pod adresem uczonych, którzy przyjęli
zaproszenie do Komitetu Naukowego i tych, którzy zgodzili się wygłosić wykłady
plenarne.
Dziękujemy wszystkim uczestnikom za prezentację ich dorobku i dzielenie
się wiedzą i doświadczeniem.
Witamy osoby towarzyszące, których obecność cenimy sobie na równi z
głównymi uczestnikami.
W bieżącym roku obchodzimy pięćdziesiątą rocznicę Pierwszego

Międzynarodowego Kongresu w Bonn (1960). Zapoczątkował on regularne
spotkania arachnologów z całego świata; jubileusz półwiecza jest więc dobrą okazją,
aby wyrazić szacunek i wdzięczność tym wszystkim, którzy przez minione dekady
kierowali pracami ISA i nadawali ton badaniom arachnologicznym. Wielu z nich
uczestniczy w niniejszym Kongresie; wśród nich - pierwszy Prezydent, Prof. Otto
Kraus, jeden z „ojców założycieli” naszej organizacji.
Spotykamy się w Uczelni, w której przed 40 laty Prof. Jerzy Prószyński

zainicjował aktywność grupy badawczej, specjalizującej się w systematyce,
biogeografii i ewolucji Salticidae. Kilka lat później do zespołu dołączył Prof.
Wojciech Starga; poszerzyło to zakres naszych zainteresowań o faunistykę i
ekologię pająków oraz systematykę kosarzy. Dziś pragniemy podziękować obu
Profesorom - zarówno w imieniu grupy siedleckiej, jak i tych wszystkich Koleżanek
i Kolegów, którzy czują się uczniami i współpracownikami naszych Mistrzów.
Wielu gości Kongresu jest w Polsce po raz pierwszy, a większość nigdy

przedtem nie odwiedziła Siedlec. Mamy nadzieję, że program Kongresu zadowoli
wszystkich, a atmosfera pobytu w Polsce pozostanie w pamięci na długo.
W imieniu Komitetu Organizacyjnego,

Marek Żabka
48
Preface
Arachnids are fascinating creatures, inhabiting the Earth for the last 400
million years. Today (excluding Acari) some 60,000 species are known to exist
in a variety of climatic zones and ecosystems: from hostile deserts, high
mountains and tundra, through tropical forests, savannahs and steppes. Many
species can survive periodically submerged and a few are permanent water
dwellers.
For thousands of years arachnids, especially spiders and scorpions, have
generated lively interest, fascination and fear - well evidenced by the rock
engravings, cave paintings and today’s myths and beliefs. At present, sensational
reports, movies, comics and web pages have raised arachnids to the role of
animal celebrities and pop-culture heroes.
The most deadly spiders such as black widow, recluse spider, Sydney
funnel web spider or many scorpion species attract the attention of general
public, stimulating (or limiting) nature tourism and becoming research subject
for biotechnology and medicine. Chemical and mechanical properties of spider
webs and silk are extensively studied in order to produce silk-like super
materials.
The long history of arachnids, their great species diversity, various habitat
preferences, living strategies, behaviours and dispersal abilities are the best
evidence of their evolutionary success. As top predators and other animals’ prey
they are extremely important in the ecological webs.
Arachnology today is a multidisciplinary science, with a wide range of
research areas such as palaeontology, systematics, biogeography, ecology,
biodiversity and conservation, physiology, genetics, neurobiology, behaviour,
reproduction or social life. In most cases the methods of molecular biology,
electron microscopy and sophisticated statistics have become a necessity in the
profound analyses.
All those aspects are presented and discussed during the Congress and are
covered in the Book of Abstracts. Full texts of the papers will be presented in the
Proceedings published by the Journal of Arachnology and other prestigious
periodicals.
The Congress has been a joined venture of many people, institutions and
the entire arachnological community.
The members of the Honorary Committee supported us mentally and were
“good spirits” of the Congress.
We wish to thank the members of the Scientific Committee and the
Plenary Speakers for accepting our invitation.
The Rector and Chancellor, Deans of the Faculties of Science and Natural
Sciences, Colleagues and Friends from the Institute of Biology, Director of the
Library, administration and students of the University of Podlasie - all are
acknowledged for their commitment and help.
The International Society of Arachnology, the Congress co-organiser, has
provided the financial support, and the ISA President - Dr. Nikolaj Scharff and
Secretary - Dr. Jason Dunlop have been especially co-operative.
49
We are grateful to the Arachnologists from 6 continents and almost 50

countries for presenting their research results and sharing their knowledge and
experience.
We extend our warm welcome to all the accompanying persons, who are
here with us today.
We thank the sponsors for understanding the role of science in modern
world.
The 18th International Congress takes place on the 50th anniversary of the
first meeting in Bonn (1960). Therefore, it gives us an excellent opportunity to
pay our respect to all those, whose efforts and hard work have contributed to the
development of arachnology over the last decades. Our special gratitude goes to
the former ISA officials, including Professor Otto Kraus - the first President and
one of the “founding fathers” of our organization.
The Congress is being held at the University of Podlasie, where 40 years

ago Professor Jerzy Prószyński started to organise the research team working on
salticid taxonomy, biogeography and evolution. In the years to follow, the group
was joined by Prof. Wojciech Staręga to expand the research to such areas as
faunistics, ecology, and harvestmen systematics. On behalf of all those who have
had a privilege to co-operate with both Professors, we would like to express our
gratefulness.
Many Congress participants and accompanying persons have not been in

Poland before. We hope that the programme will meet your expectations and
your stay in Siedlce will leave you only with good memories of the people and
the place itself.
On behalf of the Organizing Committee,
Marek Żabka
50
Phylogeny and sociogeography of Anelosimus spiders

Ingi Agnarsson
Department of Biology, University of Puerto Rico, San Juan, PR, USA,
iagnarsson@gmail.com
The diversity and distribution of species across habitats and landmasses

results from interplay between various factors, such as dispersal ability,
phylogenetical and other historical constraints, as well as the ecology and
behaviour of species. Here I present new phylogenetic and ecological data and use
these to speculate about biogeographical, ecological, and behavioural determinants
of the geographical distribution and diversity of behaviourally diverse Anelosimus
spiders across the globe. In particular, I focus on the potential role of social
behaviour in explaining global geographical distribution and diversity patterns in
the genus, what might be referred to as sociogeography. Solitary species form a
clade of apparently good dispersers, evidenced by their colonization of multiple
landmasses worldwide. Allopatric speciation following colonization of novel areas
appears to have been the primary force of diversification in solitary species.
Solitary species are restricted to habitats where subsocial and social species are
rare or absent: narrowly restricted to coastal habitats in the tropics, but widely
distributed at high latitudes. Social species do not form a clade-sociality evolved
repeatedly, but once social, they fail to diversify, presumably due to various
consequences of their population structure and mating system, including
inbreeding. Social species are mostly restricted to low to mid elevation forest the
Americas. Ecological factors, such as prey size, seem to explain the absence of
social species at high latitudes and altitudes, while the absence of social species
from lowland tropical forests in other parts of the world is best explained in a
biogeographical context; (1) one of the consequences of Anelosimus becoming
social is reduced dispersal, thus social species are unlikely to colonize new
landmasses, and (2) perhaps merely by chance, lineages that have given rise to all
the social species in the Americas have not colonized other parts of the world.
Subsociality is ancestral for the genus as a whole, subsocial species are most
diverse, and subsocial lineages have colonized most continents. Their relatively
high diversity apparently results from good dispersal abilities, and an interplay
between allopatric and, possibly, sympatric speciation. In sum, I propose
speculative hypotheses regarding the influence of social structure on geographical
distribution and diversity patterns. My findings suggest that the diversity and
distribution of Anelosimus spiders worldwide may be influenced by the level of
social behaviour displayed by species and lineages, and thus proposes variation in
social behaviour as an important biogeographical force in this, and perhaps other
lineages.
51
Behavioural and biomaterial coevolution in spider orb webs
Ingi Agnarsson1, Todd A. Blackledge2, Andrew Sensenig3,

Nikolaj Scharff4 & Jonathan Coddington5
1
Department of Biology, University of Puerto Rico, San Juan, PR, USA,
iagnarsson@gmail.com
2
University of Akron, Akron, OH, USA blackledge@uakron.edu
3
University of Akron, Akron, OH, USA
4
Zoological Museum, University of Copenhagen, Copenhagen, Denmark
5
National Museum of Natural History, Washington DC, USA
Mechanical performance of biological structures, such as tendon, byssal

threads, muscles, and spiders webs, is determined by a complex interplay
between material quality (intrinsic material properties, larger scale morphology)
and proximate behaviour. Spider orb webs are a system in which fibrous
biomaterials-silks are arranged in a complex design to produce effective energy
absorbing traps for flying prey. Orb webs show an impressive range of designs,
some effective at capturing tiny and flimsy insects, others that can stop even
small birds in mid flight. Here, we test whether material quality and the
behaviours used to spin webs co-evolve to fine-tune web function, by
quantifying both the intrinsic material properties of silk and web architectures
across diverse species of orb weaving spiders. We find a dominant pattern of
material and behavioural coevolution. Prominently, evolutionary increases in
body size, a common result of fecundity selection in spiders, are repeatedly
accompanied by improved overall web performance, which results from both
material and behavioural changes. After controlling for spider size, spiders that
evolve higher quality silk use it more sparsely in webs. This implies that
improvements in silk quality can relax selection on web architectural designs, or
alternatively that spiders producing lower quality silk must compensate
architecturally for the inferior performance their silk. In summary, spider silk
material properties are fine-tuned to the architectures of webs, resulting in a
coevolutionary pattern that may underlie many biological systems such as
tendon, mollusk byssal threads, and keratin.
52
Costs of burrow-digging and sexual dimorphismin immune

ability and fat reserves in the sex role reversed spider
Allocosa brasiliensis (Lycosidae)
Anita Aisenberg1 & Alfredo V. Peretti2
1
Laboratorio de Etología, Ecología y Evolución, Instituto de Investigaciones Biológicas
Clemente Estable, Montevideo, Uruguay, aisenber@iibce.edu.uy
2
CONICET - Laboratorio de Biología Reproductiva y Evolución, Cátedra de Diversidad
Animal I, F.C.E.F.N., Universidad Nacional de Córdoba, Córdoba, Argentina
Burrow digging on the sand has been reported as an energetically expensive

activity in spiders mainly due to three factors: silk production, digging activities
per se and predation risk while constructing the burrow. Allocosa brasiliensis is a
nocturnal wolf spider that constructs burrows along South American sandy coasts
of rivers, lakes and the Atlantic Ocean. Males construct burrows in the sand of
approximately 10 cm length, while females dig short refuges. The species shows a
reversal in typical sex roles and sexual dimorphism expected for spiders: females
are smaller than males, they locate their sexual partners and initiate courtship.
Copulations occur exclusively inside male burrows and females prefer those males
showing deeper burrows. Males provide females with their own burrows after
copulation that will serve as nests for the future progeny. According to this, we
could expect high selective pressures acting on male digging performance. The
objectives of the study were to describe in detail burrow digging in A. brasiliensis
and estimate the costs of male digging by quantifying immune response and fat
reserves in males that had constructed and males and females that had been
prevented to construct burrows.
We captured 10 adult males between January 2 of 2007 and March 7 of
2008, in Marindia (34º46'52.3"S, 55º49'29.6"W), Canelones, Uruguay. One week
after their capture at the field, each male was individually introduced in a glass
cage of 30 cm and 16 cm of base, and 20 cm height, with sand as substrate. We
video-recorded digging behaviours for one hour, since we detected sand extraction
(n=10). For testing immunity responses, we captured 19 adult females and 21
adult males of A. brasiliensis between March 2 and May 12, 2009, in the coastline
of Arroyo Copina (31°34.91'S, 64°39.46'W), Córdoba, Argentina. All the spiders
were fed ad libitum. We created three experimental groups: group 1) adult males
that had constructed burrows (n=10); group 2) adult males that had not constructed
burrows (n=11); group 3) females that had not constructed burrows (n=18). Males
of the first group were placed on plastic containers of 9 cm diameter and 14 cm
height with sand as substrate; individuals from the other groups were maintained
in Petri dishes with sand as substrate, where they could bury themselves to avoid
dehydration, but they could not dig a burrow. Twenty four hours later, they were
anesthetized with CO2 and taped laterally onto separate glass slides. A single sterile
1 mm long and 0.08 mm diameter implant was inserted on each spider abdominal
wall, between the epigastric furrow and the spiracles. Twelve hours later, each
53
implant was removed and photographed under a dissecting microscope. Immune

response was quantified by melanin percentage of encapsulation on each implant.
For fat measurements, we removed legs and carapace and used the spider abdomen
and for the extraction we used chloroform and weighed individuals before and after
the extraction (the difference was considered as fat content).
We distinguished eight behavioural units during digging: sand extraction,
exiting from the burrow, entering, silk deposition on the walls, silk deposition
around the burrow entrance, turning, blocking the burrow entrance and rest. We
appreciated a high relation among sand extraction, entering and exiting the burrow
and a high number of occurrences of these behavioural units. We detected high
duration of silk deposition on the walls and around the burrow entrance. We did not
find significant differences between males that had or had not constructed a burrow,
either in encapsulation rate (U=36.0, N1=10, N2=11, p=0.19), or fat reserves
(U=60.0, N1=10, N2=11, p=0.98). However, males showed higher encapsulation
rates and fat reserves compared to females (encapsulation rate: U=72.0, N1=15,
N2=21, p=0.01; fat reserves: U=44.0, N1=19, N2=21, p=0.0001).
Burrow digging by A. brasiliensis males was very stereotyped, showing complex
connections among the behavioural units. The high frequencies of some units as
sand extraction and the high duration of silk deposition suggest that digging
behaviour would be an energetically expensive activity. Silk production in spiders is
considered costly and spiders require multiple deposition of silk layers to maintain
stable burrows in the sand. The high frequencies with low duration of entering and
exiting the burrow could be related with avoiding predation risk during this activity.
We did not find significant differences in immune response by encapsulation on the
implant or fat reserves in males that had or had not constructed a burrow. This fact
could be due to the low sample size, period considered for construction, stage of the
reproductive period or homogenization of the sample due to laboratory foraging
conditions, hypotheses that require further studies. However, males showed higher
encapsulation rates and fat reserves compared to females. In invertebrates, fat bodies
are responsible for synthesizing anti-microbiotic peptides, so both encapsulation
rates and fat reserves agree with a higher immune response in males compared to
females. Males of A. brasiliensis are the sedentary sex that constructs the breeding
nest and stays for long periods buried without foraging, waiting for the mobile sex.
Divergences in life histories between the sexes would be determining contrasts in
energetic requirements and their management by each sex in A. brasiliensis. Future
studies will test variations in immune response between the sexes along the
reproductive period, relation between immune response and sexual activity and the
possible links between female and male choice in this species.
54
Spatial distribution along the sand dunes and temperature

buffering in two burrowing Allocosa species with sex role
reversal (Araneae: Lycosidae)
Anita Aisenberg1, Macarena González1, Álvaro Laborda2,

Rodrigo Postiglioni1,2 & Miguel Simó2
1
Clemente Estable, Montevideo, Uruguay, aisenber@iibce.edu.uy
2
Sección Entomología, Facultad de Ciencias, Montevideo, Uruguay
Allocosa brasiliensis and Allocosa alticeps are two sympatric and

synchronic wolf spiders that construct burrows along the sandy Uruguayan
coastline. Previous studies proposed a reversal in sex roles and sexual size
dimorphism for both Allocosa species. Females initiate courtship and copulation
takes place inside male burrows. Females prefer those males that show longer
burrows. After final dismount, the male donates his burrow to the female, and
she will stay inside, oviposit and exit for spiderling dispersal. The larger size in
males compared to females in A. brasiliensis could be associated with
competition for access to territories where males can construct more stable and
deeper burrows. The objectives of the study were to examine burrow density and
its distribution along the sand-dune profile and determine whether burrow depth
is related with temperature buffering in both Allocosa species.
To evaluate the spatial distribution of the burrows, we performed five
monthly samplings in the coastal sand dunes of Marindia (34º46'52.3"S,
55º49'51.5"W), Canelones, Uruguay. Samplings were performed in square plots
of 1 m2 drawn parallel to the line of the coast on the first line of dunes, two plots
on the sea-side and two on the land-side. Spiders were collected by sifting the
sand of each plot down to 20 cm depth and recording the location of all Allocosa
spp. spiders. The samplings were performed during day-light by four collectors,
using geological sieves with a mesh size of 4 mm. To evaluate temperature
buffering of the burrows, we recorded ninety eight burrows of A. brasiliensis
and A. alticeps located at the first line of dunes in Lagomar, Canelones, Uruguay
(34º50'59"S, 55º58'35"W). Measurements were performed by two researchers
during one hour, after sunset. Temperatures inside the burrows were recorded at
the bottom of each burrow, using a digital thermocouple. The air temperature
was registered approximately 5 cm away from the burrow entrance and 10 cm
away from the surface. We measured burrow depth in all the cases.
During the sifting, we found two females, four males and 66 juveniles of
A. alticeps; and four females, two males and 36 juveniles of A. brasiliensis.
When we compared the distribution of A. alticeps and A. brasiliensis (land-
side vs. sea-side) we did not find significant differences, considering the total
number of spiders (χ2=0.22, P=0.64). In A. alticeps, juveniles were more
frequent at the base of the dune compared to adults of the same species
(χ2=6.33, P=0.01). We did not find significant differences in the distribution
55
according to the stage in A. brasiliensis (χ2=4.92, df=2, P=0.09). A. alticeps

adults were more frequent on the slope and A. brasiliensis adults on the base
(Fisher, P=0.04). Temperatures outside the burrow averaged 17.5±2.4°C (range:
13.8-20.6), and inside the burrow 20.6±3.6°C (range: 13.2-27.0), and we found
significant differences between them (t97=16.83, P<0.001). Burrow depth
averaged 58.9±2.7 mm (range: 9.4-15.5). Temperature buffering (difference
between temperature outside and inside the burrow) increased with burrow depth
(R=0.76, F3,94=43.38, P<0.0001).
The results confirm that A. brasiliensis and A. alticeps burrows are
distributed differently on the sand dune according to the developmental stage
and species. Preferential areas for digging in adults seem to be the base of the
dune in A. brasiliensis and the slope in A. alticeps. Adults are expected to be
more selective when choosing the burrowing sites compared to juveniles
because females prefer to mate with males that have longer burrows. This fact
promotes that males will be selected to construct long and stable burrows. The
preference of A. brasiliensis adults for the base of the sand dune could be
associated with more protection from Southern winds that frequently blow in
these areas and possibly higher prey abundance. Individuals of both Allocosa
species are drastically constrained by humidity levels, both for constructing
more stable burrows and for their own survival. On the base of the dune, high
humidity levels are reached nearer to the surface, diminishing the energetic costs
of digging deeper burrows. On the other hand, the location of A. alticeps adults
in the slope observed in the present study could be interpreted as a mechanism to
avoid predation by A. brasiliensis adults. The larger size and consequently
higher predator capacity of A. brasiliensis adults seems to be driving the spatial
exclusion of A. alticeps females and males from the former species’ burrowing
areas. Female and male burrows were located on the same sites of the dune, on
each Allocosa species so present results could be in conformity with the
hypothesis of territorial males defending female harems, as has been cited for
other arthropods. The positive correlation between temperature and burrow
depth, in addition to the differences observed between the temperatures inside
and outside the burrows of Allocosa individuals, suggest that these refuges
provide thermal buffering against the highly variable environmental conditions.
Larger burrows would provide higher temperature buffering to help individuals,
and most importantly the females that will egg-lay, face the drastic
environmental thermo changes of coastal areas.
56
Fine structure of dimorphic sperm in mite harvestmen

(Opiliones: Cyphophthalmi)
Gerd Alberti1, Gonzalo Giribet2, Melanie Gutjahr3
& Elisabeth Lipke1,4
1
Ernst-Moritz Arndt University, Zoological Institute & Museum, Department of General
Zoology and Zoological Systematics, Greifswald, Germany,
alberti@uni-greifswald.de
2
Department of Organismic and Evolutionary Biology & Museum of Comparative
Zoology, Harvard University, Cambridge, MA, USA
3
Ernst Moritz Arndt University, Interfaculty Institute for Genetics and Functional
Genomics, Department of Functional Genomics, Greifswald, Germany
4
RWTH Aachen, Institute for Biology II, Department of Developmental Biology and
Morphology of Animals, Aachen, Germany
The spermatozoa of mite harvestmen have received special attention in

recent years with data currently available from representatives of three of the 6
families but only few species have been studied. In these studies it has been
shown that dimorphic spermatogenesis, resulting in aflagellate fertile eusperm
and infertile parasperm, seems to occur widely in the group. However, the
spermatozoa, in particular the eusperm, differ considerably in structure even in
representatives of the same family. Thus, it could be expected that sperm
morphology might present characters which could be informative for
phylogenetic and systematic interpretations at the family-level. Furthermore, the
peculiar characteristics, not known from other Opiliones, such as sperm
dimorphism, a transitory flagellum, formation of peculiar sperm balls together
with the plesiotypic mode of sperm transfer via a spermatophore urgently
required further studies on a broader taxon sampling to provide a profound basis
for comparisons with other Opiliones or even other arachnids. In the present
study we summarize our results on sperm ultrastructure based on observations
on more than 14 species of Cyphophthalmi totaling 5 of the 6 families.
57
Male meiosis: sex chromosomes and heterochromatin

behaviour in three nuptial gift-giving spider species
(Trechaleidae)
Maria José Albo1 & Alicia Postiglioni2
1
Clemente Estable, Uruguay, mjalbo@iibce.edu.uy
2
Departamento de Genética y Mejora Animal. Facultad de Veterinaria, Universidad de
la República, Uruguay, alicia.postiglioni@gmail.com
Nuptial gifts are heritable characters associated with reproduction, and

probably highly affected by environmental factors. These factors play an
important role in gene expression and are specially associated to tissue-specific
genes existing in G-banding chromosomes. In spiders, the families Pisauridae
and Trechaleidae have species in which males offer a prey wrapped in silk to
females during courtship and mating. In Uruguay, nuptial gifts have been
reported for Paratrechalea ornata and Trechalea bucculenta but not for
Trechaleoides biocellata (Trechaleidae).
The aim of this study was to investigate male meiosis in these three
species, determinate their sex chromosome system, and their -heterochromatin
distribution as an effect of G-banding, and compare and elucidate genetic
differences that traduce into their phenotypes. We dissected fourteen adult
gonads of Paratrechalea ornata, six of Trechaleoides biocellata and three of
Trechalea bucculenta. Some slides of each species were stained with Giemsa
while others were exposed to G-banding treatment.
We observed a diploid number of 2n=22 in Paratrechalea ornata, and
2n=26 in both Trechalea bucculenta and Trechaleoides biocellata. The three
species presented X1X20 as sex chromosome system. In the two formers, both
sex chromosomes placed together and aligned in parallel until the segregation in
anaphase, but stayed distant between each other in T. biocellata, and from
diakinesis these chromosomes were observed together in a parallel disposition.
Interstitial G-bands were similar in P. ornata and T. bucculenta, and both
differed from those in T. biocellata.
We found a relatively stable complement number among species of the
same family, indicating karyotypes are conserved. As X1X20 formula had been
suggested as ancestral in Araneae, the independence of both sex chromosomes in
T. biocellata added to the differences in G-banding suggests the existence of
innovative modifications in this species.
58
Nuptial gift implies a high cost for males of the nursery

web spider Pisaura mirabilis (Araneae: Pisauridae)
Maria José Albo1, Søren Toft2 & Trine Bilde2
1
2
Department of Biological Sciences, Ecology and Genetics, Aarhus University,
Denmark, soeren.toft@biology.au.dk, trine.bilde@biology.au.dk
In both vertebrates and invertebrates poor feeding or an unbalanced

nutritional state can affect reproductive success negatively. This can happen
through decreased courtship ability, or selection against males in poor feeding
condition by choosy females. Nuptial gifts, for instance, increase male
attractiveness during courtship and mating and gift characteristics could reflect
male nutritional condition. Males from the spider Pisaura mirabilis (Pisauridae)
offer a nuptial gift in the form of a prey wrapped in silk that functions as a
mating effort increasing male reproductive success. The female silk elicits gift
construction, however, males can offer the prey gift without silk covering.
Because silk mainly consists of proteins, gift construction could involve high
costs for males, and could determine decisions of males whether to wrap or not
to wrap. Under this hypothesis we predict that males in poor feeding condition
are less likely to wrap the prey when they perceive sexual stimuli from female
silk or in the presence of a female. Moreover, according to the hypothesis of
adaptive foraging we could expect that males in poor feeding conditions will
consume the prey instead of wrapping it.
The aim of this study was to investigate gift construction under different
sexual stimuli and different male feeding conditions. We analyzed gift
construction behaviour of males exposed sequentially to three treatments: 1)
female silk (S), 2) female silk plus the female herself (SF), and 3) no sexual
stimuli (N); first under good feeding conditions (N=17) and then under poor
feeding conditions (N=11). Because poorly fed males were also older, a control
group of same age (N=10) using well fed males was also tested.
Our results show that 88% in S, the 94% in SF and 13% in N of well fed
males wrapped the prey in silk, and 100% in S, 80% in SF and 13% in N
wrapped the prey in the control male age group, while 54% in S, 54% in SF and
11% in N wrapped the prey in the poorly fed group. No statistical differences
were found in the occurrence of gift construction between the treatments S and
SF in each group, but we observed males in N constructed significant fewer gifts
than males in S and SF. Because the absence of statistical differences we
grouped data from S and SF in each group to performed comparisons within
groups. Males from well fed group and the control group showed similar
frequencies of gift construction (91% and 90% respectively) and both
constructed significant more gifts than males from the poorly fed group (54%).
The poorly fed males also spent less time on gift construction (5.4±2.4 min)
59
compared with males from the well fed group (13.2±8.2 min) and the control
group (11.3±6.6 min). Poorly fed males that wrapped they prey in silk had
difficulties in prey manipulation and silk deposition, whereas poorly fed males
that not wrapped the prey, remained eating it.
Our findings show that the female silk is as strong a trigger as the presence
of a female on male gift construction behaviour. Response to female silk was
also found in Paratrechalea ornata, another gift giving spider species, and
supports the idea that early gift construction may be an adaptive strategy which
allows males immediate copulation once a female is located. Our data also show
that gift construction is costly and hence provides information on male foraging
state as an indicator of male quality. Hungry males invest in nuptial gift
construction rather than consuming the prey, hence the nuptial gift giving trait is
seems to be under strong sexual selection.
60
Worthless donations as a male reproductive tactic

in the nuptial gift giving spider Pisaura mirabilis
(Araneae: Pisauridae)
Maria José Albo1, Gudrun Winther2, Marie Rosenstand2,

Cristina Tuni2, Søren Toft2 & Trine Bilde2
1
2
Department of Biological Sciences, Ecology and Genetics, Aarhus University,
Denmark, trine.bilde@biology.au.dk
Nuptial gifts such as captured prey, regurgitations, glandular fluids, part of

or even males own body, have been well documented in insects. In some
species, however it has been reported that males may offer non-nutritive gifts.
For instance in some dance flies, males could use inedible token gifts to obtain
mates, and these are as successful as small genuine gifts.
In the spider Pisaura mirabilis males offer an insect prey wrapped in silk
as a mating effort. Field observations show that males sometimes wrap an
invaluable gift such as a flower head or an empty arthropod exoskeleton.
We investigated the possible use of worthless gifts by males and its effect
on male reproductive success. We staged experimental mating trials where males
could present a fly gift (FG treatment), no gift (NG), or a symbolic gift (SG;
cotton ball, flower head or sucked out prey) to the female, recording male
mating success and copulation duration.
Males with FG courted and copulated in 100% of the cases and gained the
longest copulations (76.3±31.5 min). In NG all males courted but only 45% of
males copulated and obtained the shortest copulations (14.5±13.6 min). In SG
70% of males wrapped an item and courted the female, while 30% courted
without a gift. Males offering symbolic items copulated in 86% of the trials and
obtained copulations of intermediate duration (54.8±38.5 min).
Our findings suggest that the gift giving trait is under strong sexual selection,
and that males may exploit female preference for the nuptial gift to gain
reproductive advantages with symbolic worthless gifts when no prey is
available, as in some insects. Because this kind of donations allows males to
increase the time transferring sperm and/or associated substances, without
offering actually a direct benefit to female, sexual conflict could be operating.
61
Cladistic analysis of the spider family Titanoecidae

(Arachnida: Araneae)
Lina M. Almeida-Silva1 & Antonio D. Brescovit2
¹ Universidade de São Paulo, Brazil, linamas@gmail.com

² Laboratório de Artropodes, Instituto Butantan, Brazil, adbresc@terra.com.br
Family Titanoecidae was proposed by Lehtinen to include five genera:

Titanoeca Thorell (31 spp.), Nurscia Simon (4 spp.), Goeldia Keyserling (9
spp.), Pandava Lehtinen (2 spp.) and Anuvinda Lehtinen (1 sp.). The family is
diagnosed by the presence of a complex dorso-apical fold on the male palpal
tibia, a triangular process at the base of the embolus and a tegular groove that
acts like a conductor. Recent phylogenetic hypothesis have not reached a
consensus concerning the relationships of Titanoecidae and the remaining
families of the RTA clade or the relationships between the genera included in the
family. In this study, a cladistic analysis based on parsimony was carried out to
test the monophyly of Titanoecidae and propose a relationship hypothesis for its
genera and species, emphasizing the Neotropical genus Goeldia. The matrix was
composed of 32 terminal taxa and 92 morphological characters. The in-group
included the type species of all Titanoecidae genera and additional 12 Goeldia
species, five Titanoeca and three Nurscia. The out-group was composed of three
species of Phyxelididae Lehtinen, one Amaurobiidae Thorell, one Tengellidae
Dahl, one Zoropsidae Bertkau and two Nicodamidae Simon. Novodamus
nodatus was used to root the trees. The analysis was carried out on the computer
program TNT and resulted in a single most parsimonious tree (L=302; CI=65;
RI=82). Titanoecidae arises as a clade supported by 12 synapomorphies: chilum
entire and surpassing the external borders of the AME; stridulatory apparatus of
the ectal surface of the chelicerae bearing spines; two retromarginal teeth on the
chelicerae; retrolateral lobe of the dorsal tibial apophysis ear-shaped; median
lobe of the tibial apophysis with distinct base and apices; absence of a
conductor; tegular grove functioning as a conductor; sperm ducts forming loops
in the tegulum; tegular loop of the sperm duct Pandava-type; median apophysis
bifid; and presence of a rim at the copulatory openings of the female epigynum.
The family is divided in two large clades: Goeldinae subfam. nov., including
Goeldia+Anuvinda, sister to Pandava, and Titanoecinae Lehtinen including
Nurscia + Titanoeca. The species of Goeldia are divided in two clades: ((G. diva
sp. nov. + G. utcuyacu sp. nov.) (G. camachoi sp. nov. + (G. mexicana (G.
luteipes + G. santosi sp. nov.)))) and (T. guayaquilensis (G. zyngierae (G.
yamamotoi sp. nov. (G. arnozoi (G. mirim sp. nov. (G. patellaris + G.
chinipensis)))))). Based on the results of the cladistic analysis Titanoeca
guayaquilensis Schmidt, from Ecuador, is transferred to Goeldia: Goeldia
guayaquilensis (Schmidt) comb. nov. In addition to the cladistic analysis we
present the taxonomic revisions of the genera Anuvinda, with Oriental
distribution, and Goeldia, with Nearctic and Neotropical distribution. The genus
62
Anuvinda remains monotypic. The female of Anuvinda escheri Reimoser is

redescribed based on specimens recently collected from Laos and Thailand and
the male is newly described. Six new species belonging to the genus Goeldia are
described, increasing the species composition from nine to fifteen species.
Goeldia santosi sp. nov., is described from Minas Gerais, G. diva sp. nov., from
Bahia and G. mirim sp. nov., from Piauí and Mato Grosso, all in Brazil, G.
camachoi sp. nov., from Huila, G. yamamotoi sp. nov., from San Sebastian de
Rabago, both in Colombia, and G. utcuyacu sp. nov., from Utcuyacu, Peru. Also,
the male of G. nigra (Mello-Leitão) and the female of G. guayaquilensis are
described for the first time.
63
Interfamilial phylogenetic relationships of goblin spiders,

based on morphological data (Araneae: Oonopidae)
Fernando Álvarez-Padilla1, Darrell Ubick2 & Charles Griswold3
1
Universidad Nacional Autónoma de México, Facultad de Ciencias, Dept. Biología
Comparada, Lab. Acarología, Mexico, fap@ciencias.unam.mx
2
Arachnology Lab, California Academy of Sciences, Department of Entomology, San
Francisco, CA, USA, dubick@calacademy.org
3
Francisco, CA, USA, cgriswold@calacademy.org
Cybertaxonomy has made possible simultaneous collaboration among

researchers in species description and construction of phylogenetic data sets.
These taxonomic descriptive data bases, in addition to producing species and
generic descriptions, can provide data matrices that are phylogenetically
informative. We present the first phylogenetic analysis of goblin spiders with a
matrix of ca. 430 species of oonopids, plus two species of orsolobids as
outgroups; coded for ca. 450 morphological characters. The character scoring for
this data set represents a collaborative effort of the Oonopid PBI (Planetary
Biodiversity Inventory) team of researchers, including postdoctoral fellows and
graduate students. A phylogenetic analysis using equal weights parsimony was
performed and Jackknife character support measures evaluated with these data.
Our analyses recovered as monophyletic and well supported some new and old
genera that have been recently revised or described; however, other sections of
the cladogram rendered several genera polyphyletic. Finally, we will be
discussing whether or not these polyphyletic genera, are indeed real or just
artifacts of a taxonomic descriptive data base that is analyzed with phylogenetic
algorithms.
64
Three new genera of goblin spiders from Madagascar

(Araneae: Oonopidae)
Fernando Álvarez-Padilla1, Darrell Ubick2 & Charles Griswold3

1
Universidad Nacional Autónoma de México, Facultad de Ciencias, Dept. Biología
Comparada, Lab. Acarología, Mexico, fap@ciencias.unam.mx
2
Francisco, CA, USA, dubick@calacademy.org
3
Francisco, CA, USA, cgriswold@calacademy.org
Currently goblin spiders, the spider family Oonopidae, are subject to

considerable taxonomic research by a large group of colleagues in many countries as
part of the Planetary Biodiversity Inventory (PBI). The work presented here
contributes to this effort to document and describe oonopid diversity world wide.
We have focused on the oonopid fauna of Madagascar, probably totaling 100
species, from which no species have previously been described. In this presentation
a total of 12 species from Madagascar and the Seychelles are described and
documented with ca. 534 compound digital images and 970 scanning electron
microscope pictures. These species are organized in three new genera, the largest
with eight species and the other two with a pair species each. All species are new
except Silhouettella assumptia, which was previously known from the Seychelles.
The core of the taxonomic descriptions for these twelve species and three genera,
were done automatically by coding character observations to the PBI website using
only a regular Internet browser and based mainly the images mentioned before. In
addition, we used a novel feature of this taxonomic descriptive data base to export a
phylogenetic data matrix of ca. 430 species of oonopids, plus orsolobids as
outgroups; coded for ca. 450 morphological characters. A phylogenetic analysis
recovered the twelve Malagasy species mentioned above as three different clades,
corresponding to our new genera and corroborating their diagnostic characters as
synapomorphies, but their interrelationships remain unstable. We emphasize that the
character scoring for this 400+species data set represents a collaborative effort of the
PBI team of researchers, postdoctoral fellows and graduate students. Finally, the
geographic distribution for all specimens (2158) will be added to the PBI Database
for Entering Collection Data (DEC) and the respective species web pages created.
All this information will be available online for all people and free of charge.
65
Shooting stars in an Asian tropical forest:

structural colours of jumping spiders
Diego P. Araujo1,2 & Daiqin Li1,3
1
Department of Biological Sciences, National University of Singapore, Singapore,
2
dbspdad@nus.edu.sg
3
dbslidq@nus.edu.sg
Animal colours are achieved by two processes: chemical (in case of

pigments) and physical or structural (the structure by itself affects the properties
of the light). Iridescent and structural colour, including ultraviolet, blue or green
result from the interaction of light with the nanostructure in the cuticular scales.
Although chemical colours are by far the most common, structural colours have
been documented in a wide range of animals. Many jumping spiders (Araneae:
Salticidae) have strikingly diverse iridescent and metallic colours, making them
attractive to their mates or difficult to detect by their predators. However, the
structural colouration and the mechanisms of structural colouration production
are poorly understood in salticids. In this study, we characterized and compared
the structural colours of several species of jumping spiders from Southeast Asia
using spectrophotometry. We also examined the cuticular scales that produce
structural colours in these species by using Scanning Electron microscopy. Some
salticid species (Portia labiata, Bathippus sp., Pancorius sp., Menemerus
bivittatus, Hasarius adansoni; Ptocasius sp., Thorelliola ensifera and Telamonia
sp.) have pigment colours that are usually dull dark browns, black and grey, with
some yellow and red markings. Their spectral pattern is typical of pigment
colours, with wavelengths reflected at low intensities and less pure colours (a
low chromaticity is achieved). However, Heliophanus sp, three species of
Phintella and Thiania bahomensis have structural colours including ultra-violet,
violet, blue, green, golden and white. The SEM revealed a series of scales for
each species, many of which are different from the previous described scales for
jumping spiders. Structural colouration and scales morphology are discussed in
the context of salticidae evolution.
66
Does karyotype change drive speciation in the woodlouse

hunter spider Dysdera? Multilocus phylogeographic
analyses of the Dysdera erythrina species complex
Miquel A. Arnedo1, Věra Opatová1, Milan Řezáč2 & Jiří Král3
1
Biodiversity Research Institute and Department of Animal Biology, Universitat de
Barcelona. marnedo@ub.edu, Vera.Opatova@seznam.cz
2
Crop Research Institute, Prague, Czech Republic, rezac@vurv.cz
3
Laboratory of Arachnid Cytogenetics, Department of Genetics and Microbiology,
Faculty of Science, Charles University in Prague, Prague, Czech Republic,
spider@natur.cuni.cz
With more than 250 described species, the woodlouse-hunter spider genus
Dysdera is one of the most species-rich genus in the western Paleartic. Several
factors may account for such a high level of species diversity. Dysdera species
are specialised predators of terrestrial isopods. The large variation observed in
chelicerae size and shape of Dysdera species, unparalleled among spiders,
reflects unique predatory tactics. Phylogenetic analyses conducted on Canarian
Dysdera endemics have demonstrated that coexisting species with divergent
cheliceral morphology are frequently closest relatives. Therefore, prey
segregation might have promoted diversification by preventing competition
among sympatric species. Dysdera also exhibits an unusual interspecific
diversity in diploid chromosome number, which ranges from 9 to 40, and hence
chromosome rearrangements may have driven speciation in Dysdera. In
addition, the restricted distributions of most Dysdera species suggest low
dispersal capabilities, which opens the door to the involvement of geological and
climatic factors in Dysdera diversification.
The Dysdera erythrina species complex provides an ideal model to
investigate the role of different biological and environmental factors in the
unusual diversification of the genus. This complex includes closely related
species that differ not only in genitalia but also in cheliceral morphology.
Dysdera erythrina has a large distribution range that spans from NE Iberian
peninsula to central Europe. Some of the species in the complex, however, have
much narrower distributions, circumscribed to well-known Mediterranean areas
of endemism. Finally, there is evidence that at least some of the species included
in the complex have different chromosome numbers.
We investigate the evolution and historical association between
chromosome numbers, phenotypic differentiation and geological factors by
inferring a thoroughly sampled phylogeny and conducting a multilocus
phylogeographic analysis of the species complex. Our results reveal an
unexpected diversity of distinct genetic lineages, with overlapping distribution.
Distinct chromosome numbers characterize some of the genetic lineages, while
67
cheliceral differentiation evolved mostly at the base of the tree. Relaxed clock
dating reveals that genetic lineages originated mostly during the onset of the
glacial cycles. All together, our data suggest that both climatic and chromosome
changes may have contributed to the establishment of reproductive barriers
between previously interbreeding populations, while differentiation in size
evolved subsequently, probably driven by interspecific competition.
68
Intrinsic and extrinsic factors in social evolution and

the geographical distribution of spider sociality
Leticia Avilés
Department of Zoology, University of British Columbia, Vancouver, Canada,

laviles.ubczool@gmail.com
The geographical distribution of species with different social systems

should provide clues as to the factors responsible for social evolution. Social
spiders are notable for having a distinctly tropical distribution. In the genus
Anelosimus, which contains the largest number of social species of any spider
genera, this latitudinal pattern is replicated altitudinally: social species are
restricted to wet low to mid-elevation tropical areas, while subsocial species
predominate at higher elevations and latitudes. We postulate that this pattern
results from an interaction between the dense three-dimensional webs
characteristic of social spiders with two separate environmental gradients. That
social species are restricted to the lower elevation wet tropics may reflect a
gradient in insect size, which we show decreases with elevation and latitude.
Large insects compensate for a decline with increasing colony size in the
number of prey caught per capita that results from a declining surface area to
volume ratio of the prey capture snares. Where large insects are abundant the
spiders make up for this decline by cooperatively capturing larger insects in
larger colonies. The result is a biomass per capita that is maximal at intermediate
colony sizes. Absence of subsocial species in the lowland rainforest, on the other
hand, may reflect gradients on the intensity of precipitation and abundance of
potential ant predators, which we show increase with proximity to the rainforest.
Through transplant and rain exclusion experiments we show that dense 3D webs
may be unsustainable for solitary living spiders in environments where intense
rains cause their frequent destruction. Dense 3D webs may also provide better
protection against predators, especially when large. Overall, these findings
illustrate how broad scale patterns of sociality, and social evolution itself, are the
result of an interaction between intrinsic features of organisms and the
environments in which they live.
69
Aelurillinae (Araneae: Salticidae) of the world

Galina Azarkina
Siberian Zoological Museum, Novosibirsk, Russia, urmakuz@gmail.com
Aelurillinae is one of 20 subfamilies of Salticidae with ca 5000 described

species (Platnick 2010) belonging to eight genera: Aelurillus Simon, 1884,
Asianellus Logunov et Hęciak, 1996, Langona Simon, 1901, Langelurillus
Próchniewicz, 1994, Microheros Wesołowska et Cumming, 1999, Phlegra
Simon, 1876, Proszynskiana Logunov, 1996, Rafalus Prószyński, 1999 and
Stenaelurillus Simon, 1885 (Prószyński 2007). The subfamily is known
exclusively from the Old World, Eurasia and Africa, except one species, Phlegra
hentzi (Marx, 1890).
The subfamily can be recognized by presence of cymbial pocket and
epigynal wings, the characters unknown among members of other subfamilies
(Logunov 1996). Beside these diagnostic characters I found two new
synapomorphies: a socket on outer side on endite and presence of joint
(common) teeth base on the promarginal side of cheliceral furrow. On the basis
of abovementioned affinities I include the tenth’s genus Mashonarus
Wesołowska et Cumming, 2002 to this subfamily.
Taxonomically Aelurillinae is one of the most difficult subfamily of
Salticidae in respect of species recognition because of very uniform shape of
palp, embolus hidden by tegulum and cymbium, variability of epigyne.
Study of the numerous material of Salticidae from Africa and Central Asia
reveals at least five more new genera of Aelurillinae. Well known genera such as
Aelurillus, Langona, Phlegra and Stenaelurillus require revisions. Numerous
new species, especially from Africa are awaiting descriptions.
References
Logunov D.V. 1996. Salticidae of Middle Asia. 3. A new genus, Proszynskiana
gen. n., in the subfamily Aelurillinae (Araneae, Salticidae). Bulletin of the
British Arachnological Society, 10: 171-177.
Logunov D.V. & Hęciak S. 1996. Asianellus, a new genus of the subfamily
Aelurillinae (Araneae: Salticidae). Entomologica Scandinavica, 26: 103-117.
Platnick N. 2010. The world spider catalog, version 10.5. AMNH, at www:
history. http://research.amnh.org/entomology/ spiders/catalog.
Prószyński J. 2007. Salticidae (Araneae) of the world.
http://salticidae.org/salticid/main.htm.
70
Host-associated differentiation in the relative leg length

of the kleptoparasitic spider Argyrodes kumadai
(Araneae: Theridiidae)
Yuki Baba1 & Tadashi Miyashita2
1
National Institute for Agro-Environmental Sciences, Tsukuba, Japan,
ybaba@affrc.go.jp
2
Laboratory of Biodiversity Science, School of Agriculture and Life Sciences,
University of Tokyo, Japan, tmiya@es.a.u-tokyo.ac.jp
Introduction
The kleptoparasitic spiders Argyrodes exhibit remarkable inter-specific
variations in foraging behaviour and morphology. Because Argyrodes
exclusively depends on host webs in its food acquisition and physical habitat,
host web traits seem to play important roles for trait diversification of
Argyrodes. However, this possibility has never been explored. A species of
the kleptoparasitic spider A. kumadai that mainly distributes in Japan utilizes
phylogenetically unrelated host spiders that predominates in their respective
regions; Cyrtophora moluccensis (Araneidae) is restricted to the South-west
Islands of Japan and Agelena silvatica (Agelenidae) is found on Japanese
mainland (Baba & Miyashita 2005). Due to the differences in web structure
and foraging behaviour of host spiders, opportunities for A. kumadai to
acquire prey is significantly limited in Agelena web (Baba et al. 2007), i.e.,
A. kumadai could not steal prey once captured by host spiders on Agelena
web. This may impose strong natural selection on walking performance of A.
kumadai to access prey quickly before host notice it. To test this possibility,
we focused on the relative leg-length (corrected for the effects of body size)
as a target of natural selection that seems to be associated with foraging
including movement across the web. Because Agelena constructs more
complex web with a higher thread density than Cyrtophora, the relatively
short leg-length seems to have an advantage in walking in narrow space on
Agelena web.
Material and methods

We conducted inter-population comparisons to test whether host trait
differences promoted differentiation in the relative leg-length of A. kumadai.
Firstly, we compared the leg-length among 13 populations (3 Cyrtophora and
10 Agelena populations) with ANCOVA using body size as a covariate. To
confirm the genetic background of the population difference, we compared
relative leg-length of A. kumadai reared in the constant room condition
between two populations where the host use differed (common garden
experiment). Finally, we conducted web transplant experiment to compare
71
the walking speed of A. kumadai between the population on experimental

webs with different thread densities representing the webs of the two host
spiders (simple and complex web).
Results and discussion

Relative leg-length was shorter in the population using Agelena than
that using Cyrtophora. Common garden experiment supported the genetic
background of above pattern, suggesting that different host use promoted
genetic differentiation of relative leg-length. The web transplant experiment
showed that walking speed did not differ between populations on the simple
web, but individuals with relatively shorter leg-length walked significantly
faster than those with relatively longer leg-length on the complex web. These
results supported the hypothesis that shorter leg-length was favoured in
walking performance on the complex web of Agelena. By contrast, longer
leg-length of the population using Cyrtophora seemed to have no advantage
in walking performance on Cyrtophora web, reflecting weak natural
selection on walking speed in Cyrtophora web.
References
Baba Y.G.. & Miyashita T. 2005. Geographical host change in the
kleptoparasitic spider Argyrodes kumadai associated with distribution
of two host species. Acta Arachnologica, 54: 75-76.
Baba Y.G., Walters R.J. & Miyashita T. 2007. Host-dependent differences in
prey acquisition between populations of a kleptoparasitic spider
Argyrodes kumadai (Araneae: Theridiidae). Ecological Entomology,
32: 38-44.
72
Who are these Goblin Spiders?

A new Australasian genus? (Araneae: Oonopidae)
Barbara C. Baehr1 & Mark Harvey2

1
Queensland Museum, South Brisbane, Qld, Australia, BarbaraB@qm.qld.gov.au.
2
Western Australian Museum, Welshpool DC, WA, Australia,
mark.harvey@museum.wa.gov.au
In 2002 Burger, Nentwig and Kropf described a quite unusual Opopaea

species from Sumatra as O. fosuma. It shares the swollen pedipalpal patella with
Opopaea but the bulb and the cymbium are clearly separated whereas in the
genus Opopaea cymbium and bulb are completely fused. The swollen pedipalpal
patella of O. fosuma is connected to the femur subbasally but the connection in
all other Opopaea species is more medially. The female genitalia of O. fosuma
lack the paddle-like sclerite (PSc) and have no nail-like process (Na) fitting into
posterior situated globular appendix (GAp) which are present in all Opopaea
females. Opopaea fosuma, instead, has a straight copulatory duct. On the other
hand males of O. fosuma resemble Camptoscaphiella infernalis, a blind species
described by Harvey & Edward in 2007 from Western Australia, in most
pedipalpal features such as a swollen pedipalpal patella and a well separated
cymbium and bulb, and the subbasal connection with the femur.
Camptoscaphiella infernalis shares with the type species C. fulva Caporiacco,
1934 from Pakistan the separated cymbium and bulb and the swollen palpal
patella, but both sexes of C. fulva have pairs of long spines on the first two pairs
of legs which are lacking in C. infernalis. Camptoscaphiella fulva has a huge
patella in males which is at least 7 times longer than the femur whereas the
patella of C. infernalis is only about twice as long.
As both species did not fit well in their original genera but share the main
characters we decided to establish a new Australasian genus for O. fosuma
(Burger et al. 2002), C. infernalis, and 26 new species from the Himalayan
Mountains in Nepal, India, as well as Malaysia, Indonesia, Papua New Guinea
and Australia. Most species are recorded from single locations and only two
species are more widely distributed.
73
Spider senses: how it all fits together

Friedrich G. Barth
Life Sciences, Department of Neurobiology, University of Vienna, Austria,

friedrich.g.barth@univie.ac.at
Spiders have magnificent sensory systems for the guidance of their

complex behaviour. Some of these senses show outstanding performance when
studied from an engineering point of view. However, a full appreciation of their
perfection and adaptedness is only made possible by a multifaceted approach
which brings together knowledge about (i) the ecologically relevant stimulus
patterns, (2) the detailed physics of stimulus uptake and transformation, and (iii)
behaviour under natural conditions.
The examples chosen to demonstrate this are vibration detection in the
context of courtship behaviour on the one hand and air flow sensing in the
context of prey capture on the other. The application of computational
mechanics and advanced technologies like atomic force microscopy and particle
image velocimetry provide ample evidence of a perfect match between the
biologically relevant stimulus patterns and the prominent selectivity and the
structural and functional details of the sense organs responsible for it.
74
The phenology of sympatric populations of Yllenus arenarius

(Araneae: Salticidae) isolated by the year of maturation
Maciej Bartos
University of Łódź, Department of Teacher Training and Biodiversity Studies, Łódź,

Poland, bartos@biol.uni.lodz.pl
The previously carried out life history research of Yllenus arenarius

suggested that in inland dunes in Poland there are two sympatric populations
isolated by the year of maturation. One of the populations produces young in
odd years, while the other population produces young in even years. For a short
period of time, in June, three age groups cohabit (each group older by one year
from the other). The time gap separating sexually mature individuals from both
groups makes gene flow between them highly limited. The spider’s phenology
was analyzed to check to which extent such isolation is possible. The research
also aimed at determining key periods in the life cycle of the spider and possible
deviations from the cycle pattern. Current study presents data collected over the
last thirteen years.
Limited food-niche overlapping among jumping spiders

from three coexisting age groups
Maciej Bartos

Intraspecific competition is a phenomenon thoroughly studied in animals.

Research on spider communities is also well documented. Such an interest is a
result of spider influence on prey populations and a potential application of
spiders as agents of pest control. Little information is available, however, on
such competition among cursorial, non-web-building spiders, especially those
studied in natural conditions. A suitable model provides Yllenus arenarius – a
dominant day-active spider predator in sandy habitats. This stenotopic species
dwells in dunes of Central and Eastern Europe. The spider’s populations reach
relatively high densities due to cohabitation of two spider populations hatched in
successive years. In June, there are three populations coexisting simultaneously
in the same area. High concentration of conspecifics in different age, size and
hunting experience allows studying how such high spider densities can be
maintained in a highly food-limited habitat. The results show that spiders from
three age groups select different insects as prey. The spiders in the first month of
75
life have relatively narrow food niche and prey mainly upon insects rarely found
in the diet of older individuals. Spiders older by one and two years have much
more diverse diet, but in the oldest specimens the index of prey diversity is the
highest. Individuals from all age groups maintain a fairly constant prey size to
predator size ratio.
The role of head identification in prey capture

by jumping spiders (Salticidae)
Maciej Bartos

For a cursorial predator, actively hunting its prey, there are several
questions it has to answer when it perceives any new object. It has to be decided
whether the observed object is a potential prey, a sexual partner, an enemy or
neither of those. If the object is identified as a prey, however, the question of
“how it should be approached?” arises. Numerous predators (including jumping
spiders) possess hunting tactics that are tailored to a prey. Their tactics are
characterized by e.g.: prey-specific direction and speed of approach, distance of
attack and other prey-specific behaviours. Some hunting patterns are inadequate
for certain prey and may significantly decrease the chances of success. Therefore
precise prey identification is crucial for the hunting success. The predator also
has to choose the area on its prey’s body which it will strike. The choice of
suitable point of strike allows firm prey grasping and precise venom injection,
which may considerably increase chances of subduing dangerous, large or motile
prey. It may also decrease the risk of injury caused by the prey and considerably
shorten the time, when the prey struggles to release. For these and other reasons
most animal predators (not only spiders) overpowering their prey try to grasp
their victims by the head or thorax, where ganglia responsible for movement
coordination are present. Predators possessing venom typically inject it in this
area, which allows immobilizing the prey relatively quickly. In current research I
analyze the characteristics used by jumping spiders in identification of their
prey’s head. Experiments with natural and computer-generated models are
compared.
76
Different levels of diversity within the spider family

Psechridae (Araneae)
Steffen Bayer
Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt am Main, Germany,

Steffen.Bayer@senckenberg.de
The cribellate spider family Psechridae comprises two genera: Psechrus

Thorell, 1878 and Fecenia Simon, 1887 distributed in Southeast Asia. They
differ in the arrangement of the eyes, the relative length of legs IV, the
colouration of the ventral opisthosoma and the clypeus height. But even more
remarkable are the differences in the shapes of webs used for prey capture.
Psechrus species build a horizontal, somewhat dome-shaped sheet web and hang
upside down underneath, whereas adult Fecenia produce vertical pseudo orb
web with an enrolled leaf as retreat in the hub. On the other hand juvenile
Fecenia create a 3-dimensional cone-shaped web without an enrolled leave, but
with a narrow cone-shaped retreat consisting of prey remains and particles.
Fecenia webs can be found in vegetation, mostly shrubs, whereas Psechrus
specimens build their retreats in crevices in soil, rocks, trees, dead wood etc.,
mostly near the ground. Up to now there are 30 valid species described, 25 of
which are belonging to the genus Psechrus.
Considering the amount of new descriptions of Psechrus species in the last
10 years, it is supposed that the real diversity of that group is much higher than
known so far (Platnick 2010). In contrast, the genus Fecenia, similarly
widespread as Psechrus, comprises only five species. The explanation for such a
distinct difference in species richness of these two genera can not be explained
sufficiently at the moment.
Since 2009 the author has focused on the Psechridae with their fascinating
and highly interesting webs and prey capturing behaviour. He started a detailed
revision of this family which includes not only morphological taxonomy but also
molecular aspects. The relationships of species within the genera, especially
within Psechrus, and furthermore the relationship of the Psechridae and other
members of the Lycosoidea are aim of the investigations. By now, almost all
type material of Psechrus and Fecenia species has been examined. Also a lot of
additional material from all SE Asian countries has been checked. In this context
some interesting findings could be made: First descriptions of three new species
from Laos were provided (Bayer & Jäger 2010) and the ones of seven further
species (two from Laos, two from Thailand, and each one from Malaysia,
Indonesia [Kalimantan] and the Philippines) are in progress; the male of Fecenia
travancoria Pocock could be discovered and will be described for the first time;
one species was synonymised erroneously (Psechrus annulatus Kulczyński), one
was synonymised with the wrong species (Psechrus libeltii Kulczyński, which is
actually a synonym of P. argentatus Doleschall and not of P. singaporensis
Thorell). It is striking that a lot of misunderstandings and misidentifications in
77
female Psechrus specimens were caused by the misinterpretation of the

epigynes, since in Psechrus their shape often varies strongly within one species.
In contrast, within all known species the vulvae look rather uniform. This
finding led to a new taxonomical evaluation of the female vulva in Psechrus,
which constitutes the morphological species discrimination character with the
highest priority. However, in Fecenia the shape of the vulvae rather varies,
whereas the epigynes mostly are specific and well assignable to each species.
In the future it is intended to focus on the molecular work to reconstruct
phylogeny within the family Psechridae and within the Lycosoidea.
References
Bayer S. & Jäger P. 2010. Expected species richness in the genus Psechrus in
Laos (Araneae: Psechridae). Revue suisse de Zoologie, accepted, 16 pp.
Platnick N.I. 2010. The world spider catalog, version 11.0. AMNH, at www:
http://research.amnh.org/entomology/spiders/catalog/.
78
Advances on the phylogeny of pirate spiders

(Araneae: Mimetidae), a molecular approach
Ligia Rosario Benavides & Gustavo Hormiga
Department of Biological Sciences, The George Washington University, Washington

DC, USA, ligia@gwmail.gwu.edu, hormiga@gwu.edu
Pirate spiders of the family Mimetidae are well known for their
araneophagic behaviour. The family has a worldwide distribution and currently
contains 156 species grouped in 13 genera. One of the fundamental problems
concerning the higher level systematics of mimetid spiders is the placement of
the family within Araneae. The placement of the family within the order has
been studied previously using morphological data. In this talk we present the
first molecular phylogeny of Mimetidae. We also address some of the
intrafamiliar phylogenetic relationships at the genus level. Our analyses include
sequence data from five loci (18S, 28S, 16S, COI and H3). We analyzed the data
under direct optimization and investigated clade sensitivity to different
combinations of analytical parameters (indel-to-change cost and the
transversion-to-transition ratio).
79
Phylogeography of a montane spider genus in

the Iberian Sistema Central mountain range,
insights from mitochondrial and nuclear markers
Leticia Bidegaray-Batista1, Rosemary G. Gillespie2

& Miquel A. Arnedo1
1
Biodiversity Research Institute & Department of Animal Biology, Av. Diagonal 645,
08028 Barcelona, Spain. letigaray@yahoo.com
2
Department of Environmental Science Policy and Management, University of
California, Berkeley, CA, USA
The role of Pleistocene climatic oscillations in driving species

diversification is a matter of debate. To date, the effect of climatic shifts on the
biodiversity of European mountain ranges has been studied mostly on species
with alpine and arctic-alpine distributions. The spider genus Harpactocrates
(Dysderidae) provides an excellent model to investigate the influence of climatic
change on the diversification and distribution of montane species in the southern
European refugia. The genus includes thirteen species distributed throughout the
mountain ranges of the Iberian Peninsula (10 species), the Alps and the
northernmost Apennines (3 species). They are most often found at high elevation
(above 1000 m), in temperate and moist forests, which indicates a preference for
cool and humid environments. Previous phylogenetic and phylogeographic
studies have shown that most speciation events in the genus trace back to the
Tertiary period, and that Plio-Pleistocene climatic oscillations served to shape
intraspecific variation. The present study focuses on 3 closely related
Harpactocrates species with non-overlapping distributions restricted mostly to
the Sistema Central mountain range: H. gredensis, H. globifer and H. gurdus.
We hypothesize that these species have undergone population range expansion
during cooler periods, whereas at interglacial periods species retreated to high
elevation refuges, which then led to population fragmentation. To test this
hypothesis, 126 individuals from the three species were sampled from 28
localities distributed throughout their range. Population and phylogeographic
analyses were conducted on DNA sequences from 3 mitochondrial markers
(16S, L1 and nad1) and 1 nuclear intron (srp54). The mitochondrial markers
alone reveal deep population structure in the western species H. gredensis and
the central species H. globifer, dating to around the Plio-Pleistocene epoch. The
eastern species H. gurdus shows low levels of genetic diversity and population
structure, and shallower population divergence times than its two relatives. The
preliminary inference based on the mitochondrial phylogeography is in
agreement with patterns found in other organisms with similar distributions and
ecological requirements, which suggests that the existence of multiple sky-
islands in the Iberian Sistema Central that acted as refugia during interglacial
periods. However, historical inferences based on single or linked markers are
hampered by natural selection, migration and stochastic coalescent processes,
80
which may results in large difference between the gene tree and the population
tree. In particular, phylogeographic patterns based on mitochondrial data alone
are notoriously biased in their ability to reflect the entire history of populations.
Therefore it is desirable to use multilocus approaches based on unlinked nuclear
markers, to recover a more accurate view of the evolutionary process underlying
population history. Unfortunately, spider phylogeography has been seriously
limited by the shortage of nuclear markers available for population level
analyses. The isolation and design of new nuclear markers in non-model
organism with limited genomic information is a not a trivial task. We have
circumvented this problem by isolation new anonymous markers from a
genomic library of H. globifer. We will present and discuss preliminary results
of the insights gained on the phylogeographic patterns of the Iberian Sistema
Central species by the analysis of these anonymous nuclear markers.
81
The geography of venom diversity: biogeography

and venom diversity in New World Loxosceles
(Araneae: Sicariidae)
Greta J. Binford, Andrew V. Merrell & Pamela A. Zobel-Thropp
Lewis & Clark College, Portland, OR, USA
The complexity of venom chemistry within individual spiders makes

understanding patterns of variation in venoms among species a challenge. This is
especially complicated by groups of gene families that have multiple members
expressed in a single venom. The puzzle of venom diversity is particularly interesting
in parallel with the puzzle of species-level biogeography. Sicariid spiders, including
brown spiders (Loxosceles) and 6-eyed sand spiders (Sicarius), are famous for bites
that cause dermonecrotic lesions in mammals. The toxin sphingomyelinase D (SMase
D) in venoms is a sufficient causative agent for lesion formation in animal models and
is also highly toxic to insects. This toxin is a member of a gene family (SicTox) and
multiple different SicTox genes are expressed in venoms of a single species. We have
used molecular evolution and molecular systematics techniques to present combined
biogeographic and venom diversity patterns in North American Loxosceles and South
American members of the laeta species group. Species relationships of Loxosceles
indicate that this genus colonized southern North America from South America via the
proto-Caribbean and has likely been present on North America for at least 33 million
years. Two lineages (Eastern and Western) migrated northward leading to at least two
lineages in the United States that are more closely related to species in Mexico than
they are to other US species. The laeta species group is more closely related to the
reclusa group than it is to the four other species groups in South America. The most
recent common ancestor of these two species groups was likely in NW South America
and diversification of the laeta group was southward into the region of the modern
Andes and beyond. While some SMase D genes of closely related species are more
similar to one another than they are to distantly related species, frequent duplications
add complexity to patterns of similarity of expressed SicTox genes in venoms of close
relatives. From L. arizonica alone we have isolated genes that are from 5 independent
SicTox lineages. The South American laeta species group is interesting in that their
SMase D genes are more different from those of other New World species than would
be expected based on species relationships. While our sampling is not yet thorough
enough to infer the timing of the evolutionary change that led to divergence of key
proteins in these venoms, we have found that the divergent laeta genes are present in a
broad range of species in the laeta species group.
82
Darwin’s dilemma: natural selection on orb web

performance opposes fecundity selection for extreme
female gigantism in spiders
Todd A. Blackledge1, Andrew Sensenig2, Nikolaj Scharff3,
Jonathan Coddington4 & Ingi Agnarsson5
1
University of Akron, Akron, OH, USA, blackledge@uakron.edu
2
University of Akron, Akron, OH, USA
3
Zoological Museum, University of Copenhagen, Copenhagen, Denmark
4
National Museum of Natural History, Washington DC, USA
5
Department of Biology, University of Puerto Rico, San Juan, PR, USA
Body size evolution is fundamental to the diversification of life. Sexual

size dimorphism (SSD) is pervasive throughout the animal kingdom and results
from differences in how sexual and fecundity selection operate on males versus
females. Female-biased SSD commonly involves fecundity selection while
male-biased SSD is driven in large part by competition for mates. Both
mechanisms imply strong directional selection driving the evolution of body
size, and are well-supported. However, the mechanisms opposing continued
change in body size are largely unknown, even though they play an essential role
in shaping SSD. Orb spiders present some of the most extreme examples of
female-biased SSD in the animal kingdom. In some species, female spiders are
up to nine times longer than males. Female gigantism results largely from
increases in fecundity, which in turn demands extremely high prey capture.
Here, we demonstrate that the evolution of giant female body size decreases orb
web function, such that the increased need for food by these giant spiders is
accompanied by decreased ability to gain that food. We use both empirical data
and a model to show an allometric relationship in how spider body size scales
with the stopping power of orb webs and the kinetic energy of preferred prey
such that relative web performance declines as female size increases. This
decline in web performance occurs despite overall improvements in silk
performance and total investment of silk by spiders. Thus, natural selection
ultimately places an upper limit on female gigantism and the evolution of SSD in
spiders.
83
Structure and spider diversity in the Churchill Area,

Manitoba, Canada
Gergin A. Blagoev & Sarah J. Adamowicz
Biodiversity Institute of Ontario, University of Guelph, Guelph, Ontario, Canada,

gblagoev@uoguelph.ca, sadamowi@uoguelph.ca
The Churchill area, lying at the transitional zone between the northern
boreal forest and arctic tundra, represents an interesting model for studying the
patterns of faunal transition between adjacent ecoregions. Arctic ecosystems and
especially transitional zones are sensitive areas where the impacts of climate
change are expected to be manifested first. During 2006 and 2009, spiders from
different habitats located within “islands” of the forest, tundra, and grass
meadows near Churchill, Manitoba were studied. A total of 160 species from 14
families was established and identified, based on morphological characters and
DNA barcoding employing the mitochondrial cytochrome c oxidase subunit I
(COI) gene. Intra- and interspecific genetic divergences of all described species
as well as a potential new species in the spider genus Alopecosa (Araneae,
Lycosidae) are presented and related to the species designations. In general,
close correspondence was detected between species and genetic clusters, with
most species displaying negligible intraspecific morphological variation, but
some of them show considerable variation. We also discuss species composition
and overlap among the different biomes present in the Churchill region.
84
Orb spider webs as models of prey-induced

phenotypic plasticity
Sean J. Blamires & I-Min Tso
Department of Life Science, Center for Tropical Ecology and Biodiversity, Tunghai
University, Taichung 407, Taiwan, sblamires@thu.edu.tw, spider@thu.edu.tw
Phenotypic plasticity is the ability for phenotypic change by a given

genotype. Phenotypic plasticity has been identified as a driver of evolution. The
ability for traits to be plastic is itself subject to selection. The organism’s
lifestyle influences its tendency for plasticity responses; the traits of long-lived
organisms tend to be more canalized, whereas those of short-lived organisms
more plastic. Prey-induced phenotypic plasticity is an important, yet poorly
understood, driver of predator-prey evolution. The spider orb web is an extended
phenotype. As orb web spiders tend to be short-lived there is selection pressure
toward plasticity of this phenotype with many studies suggesting web geometry,
silk biochemistry and silk mechanics are plastic in response to prey variations.
We studied the plasticity responses of orb web spiders by constructing reaction
norms for web geometry and silk mechanics in a range of Orbicularidae spiders
exposed to multiple prey-induced parameters. We manipulated the nutrients in
Nephila spp., Argiope spp. and Cyrtophora spp. food to determine the role of
nutrient availability in prey on web design evolution. We also performed
experiments to uncouple the relative influences of nutrients and vibratory stimuli
in web and silk plasticity in Nephila pilipes and Argiope keyserlingi. While all
orb web spiders have steep reaction norms, hence are highly plastic, identifying
the proximal cues used to respond to prey variations is complex because prey
traits are auto-correlated. Nutrients only partially explain plasticity patterns in
individual species. Nephila pilipes, for example, alters web geometry in response
to a combined influence of prey nutrients and web vibratory stimuli. Argiope
keyserlingi, likewise responds to both specific nutrients (e.g. protein), energy, as
well as prey size, feeding frequency and vibratory stimuli. We propose using
path models to statistically uncouple the multiply acting mechanisms and
identify the prey-mediated causal factors over orb web plasticity and describe
phenotype-environment feedback mechanisms, whereby web plasticity alters
spider niches, thereby impacting spider physiology, impacting back on web
plasticity.
85
Ground-dwelling spider fauna in Natural Forest Reserves

in Hesse, Germany
Theo Blick
Projekt Hessische Naturwaldreservate, Senckenberg Forschungsinstitut und

Naturmuseum, Frankfurt am Main, Germany, theo.blick@senckenberg.de
Strict Forest Reserves are forests where no forest operations are carried
out. In Hesse (Germany) 31 such reserves have been established, 22 of them
with a reference area where forest management is continued. Seven groups of
animals (Lumbricidae, Araneae, Heteroptera, Coleoptera, Hymenoptera
Aculeata, Macrolepidoptera, Aves) have so far been investigated in eight
reserves using a wide range of methods over two whole years (incl. winter).
Here, the ground-dwelling spiders of five Strict Forest Reserves in Hesse are
determined and analysed.
Data on spiders from pitfall traps are the topic of this talk. About 2/3 of
the spider species (>200) were trapped with the pitfalls; a slightly smaller
number than with trunk eclectors (in total > 300 species). In a single beech forest
(4 of the 5 areas are beech forests) 26% of the spider species known from Hesse
and 18% of the species known from Germany can be found. On average 110
spider species were recorded on the ground in every area.
The spider species are categorised according to their degree of affiliation
to forests: strictly in forests; mainly in forest habitats; also in forests but mostly
living in special habitats like swamps or warm slopes; eurytopic in open land
habitats. 76% of the species and 94% of the individuals belong to the first two
groups, i.e. the spiders show a high affiliation to forest habitats.
Central European beech forests are inhabited by a much larger number of
species than previously estimated. The seven investigated animal groups
comprise 1480±170 species, representing 25% of all animal species which
occur. Extrapolation yields a total of 5800 species for a beech forest with a size
of about 70 ha (reserve & reference area). This is 3-4 times more than the 1500-
2000 species hitherto assumed to occur in such an area.
86
Spider diversity in Lonar crater sanctuary and a new

species of Ariamnes (Araneae: Theridiidae) from India
Atul Bodkhe
J. D. Patil Sanludkar Mahavidyalaya, Daryapur, Distt. Amravati, Maharashtra, India,

atulecologia@yahoo.co.in
Introduction
Lonar is World No. 3 crater, formed 50 million years ago due to meteorite
impact. Government of India has declared Lonar as Wild life Sanctuary on dt.8th
June 2000.
Area details: total = 383.22 ha, reserved forest = 266.08 ha, water body =
77.39 ha, cultivation = 39.75 ha, It is the smallest sanctuary in India but the
habitat is very typical and it is very close to human settlement.
I have reported 100 spider species from Lonar Sanctuary from 20 Families
and 36 genera (June 8, 2009).
Family genera species

Araneidae 4 15
Clubionidae 1 7
Corinnidae 1 1
Eresidae 1 1
Gnaphosidae 1 4
Hersillidae 1 5
Lycosidae 1 4
Miturgidae 1 2
Oonopidae 2 3
Oxyopidae 2 10
Philodromidae 2 6
Pholcidae 1 2
Pisauridae 1 2
Salticidae 5 19
Scytodidae 1 1
Sparassidae 1 2
Tetragnathidae 1 1
Theridiidae 2 3
Thomisidae 6 11
Uloboridae 1 2
Total: 20 families 35 genera 100 species
87
Ariamnes sp. nov.

Diagnosis. The species differs from Ariamnes huinakolu and Ariamnes
waikula in eye arrangement. In the present species posterior medians are located
quite behind the posterior lateral eyes forming procurved arrangement while in
Ariamnes huinakolu and Ariamnes waikula the posterior medians are in straight
line with posterior laterals. Legs are too long with length of Femur longest, first
pair of leg is very long, No spines nor hairs on the legs. Abdomen much bulky
and tapers posterior forming a tubular structure ending in to a narrow tip. In
Ariamnes huinakolu and Ariamnes waikula abdomen is not ending in a tip.
Description. Cephalothorax and legs light yellowish-brown, legs light
yellow. Total length 6.00 mm, Carapace 2 mm. long, 1.5 mm. wide, abdomen 4
mm. long and 3 mm wide. Cephalothorax narrowing anteriorly and broadest at
coxa IV, without fovea, posterior end with two triangular black patches, laterally
provided with very small blunt spines. Eyes in two rows, anterior row of eyes
recurved (as seen from in front), anterior medians black, anterior laterals white,
oval or elliptical in shape, medians closer to laterals than to each other. In front
of anterior median eyes two transverse brown patches up to end of carapace are
present. Posterior row of eyes procurved, medians larger than laterals, pearly
white, medians closer to laterals than to each other. Median ocular quad as long
as wide. Sternum elliptical in shape, narrowing behind, provided with two
longitudinal brownish bands. Labium broader than long, nearly hexagonal in
shape, chelicerae longer than wide provided with small spines, anterior end is
provided with scopulae. Legs relatively long and slender provided with hairs and
some spine like hairs. Metatarsi and tarsi of all legs provided with small spines.
Leg formula is 1,4,2,3. Abdomen longer than wide, broadest in the middle,
tapering, posteriorly provided with mid-dorsal longitudinal band starting from
anterior to posterior end terminating in a pointed tip and mid-dorsally brownish
patches. Ventral side slightly longer than dorsal. Mid-ventrally provided with
two longitudinal bands starting from spinnerets. Spinnerets, conical in shape,
located at 1/3rd distance from anterior tip of abdomen.
Type specimens: holotype female, paratype female in spirit.

Type locality: Lonar, Dist- Buldhana, Maharashtra, India.
Acknowledgements
PCCF, Department of Forest, Nagpur, gave permission to carry out
research on spiders from Lonar crater.
88
Taxonomy and phylogeny of the European

Tegenaria/Malthonica-complex
Angelo Bolzern
Naturhistorisches Museum Basel, Basel, Switzerland, angelo.bolzern@arachnodet.com
Malthonica Simon and Tegenaria Latreille represent two species-rich

genera of the family Agelenidae. They are predominantly Palaearctic in
distribution. Currently 43 species and one subspecies are described in
Malthonica and 101 species in Tegenaria, respectively. In Europe, 33
Malthonica species and 55 Tegenaria species and one subspecies are described.
The group is notorious for its taxonomic problems: lack of diagnoses of the two
genera, arbitrary generic assignment of species, availability of information for
only one sex in many species and unknown internal phylogenetic relationships.
During the last 5 years much work has been done to improve the
knowledge of these taxa in focus. On species level, many recent publications
provide more information about Malthonica or Tegenaria species (Bolzern et al.
2008, Croucher et al. 2007, Gasparo 2007, Král 2007, Seyyar et al. 2008) or
include descriptions of new species (Bolzern et al. 2009, Bolzern et al. accepted,
Bolzern & Hervé 2010, Deltshev 2008, Guseinov et al. 2005, Kovblyuk 2006,
Kovblyuk & Ponomarev 2008). On the genus level, the problem to reconstruct
monophyletic groups is very demanding. A first attempt is the rearrangement of
the species of both genera based on the embolus length (Guseinov et al. 2005), a
character which is on its own not sufficient for phylogenetic reconstructions. A
further study provides a more detailed definition of the genus Malthonica, which
results in the exclusion of all but 2 species previously affiliated with this genus
(Barrientos & Cardoso 2007). The here presented study is a further step toward a
solution of these problems.
The exhaustive examination, including morphological and molecular
methods, of specimens of many European Malthonica and Tegenaria species
showed that numerous taxonomical changes are required in order to obtain
monophyletic groups. As a first step, the revision of the involved species
resulted in 14 proposed synonyms, 11 new species, 1 taxon newly treated as
nomen dubium and 4 species which can not be accurately placed. The
relationship between all these species was investigated performing different
phylogenetic analyses based on morphological characters or 3 different gene
sections. The resulting trees allow to formulate a new hypothesis, regrouping the
European species of Malthonica and Tegenaria into 4 genera: 1) Malthonica (2
spp.), 2) Tegenaria (50 spp.), 3) Aterigena n. gen. (5 spp., 4 European and 1 east
Asiatic species), 4) Eratigena n. gen. (19 spp.).
The following aspects are presented: 1) results from the morphological
and molecular analyses 2) presentation of the three genera Tegenaria, Eratigena
n. gen. and Aterigena n. gen. and 3) remaining questions and notes on species
outside Europe.
89
References
Barrientos J.A. & Cardoso P. 2007. The genus Malthonica Simon, 1898 in the
Iberian Peninsula (Araneae: Agelenidae). Zootaxa, 1460: 59-68.
Bolzern A., Crespo L.C. & Cardoso P. 2009. Two new Tegenaria species
(Araneae: Agelenidae) from Portugal. Zootaxa, 2068: 47-58.
Bolzern A., Hänggi A. & Burckhardt D. 2008. Funnel web spiders from
Sardinia: taxonomical notes on some Tegenaria and Malthonica spp.
(Araneae: Agelenidae). Revue Suisse de Zoologie, 115: 759-778.
Bolzern A. Hänggi A. & Burckhardt D. (accepted) Aterigena, a new genus of
funnel-web spider, shedding some light on the Tegenaria-Malthonica
problem (Araneae, Agelenidae). Journal of Arachnology.
Bolzern A. & Hervé C. 2010. A new funnel-web spider species (Araneae:
Agelenidae, Tegenaria) from Mercantour National Park, France. Bulletin
of the British Arachnological Society, 15: 21-26.
Croucher P.J.P., Jones R.M., Searle J.B. & Oxford G.S. 2007. Contrasting
patterns of hybridization in large house spiders (Tegenaria atrica group,
Agelenidae). Evolution, 61: 1622-1640.
Deltshev C. 2008. Two new spider species, Malthonica bozhkovi sp. nov. and
Tegenaria paragamiani sp. nov. from Rhodopy Mountains (Bulgaria and
Greece) (Araneae: Agelenidae). Zootaxa, 1872: 37-44.
Gasparo F. 2007. Note su Tegenaria percuriosa Brignoli, 1972, con descrizione
del maschio (Araneae, Agelenidae). Atti e Memorie della Commissione
Grotte “E. Boegan”, 41: 95-103.
Guseinov E.F., Marusik Y.M. & Koponen S. 2005. Spiders (Arachnida: Aranei)
of Azerbaijan. 5. Faunistic review of the funnel-web spiders (Agelenidae)
with description of new genus and species. Arthropoda Selecta, 14: 153-
177.
Kovblyuk M.M. 2006. Malthonica podoprygorai sp.n. from the Crimea (Aranei:
Agelenidae). Arthropoda Selecta, 15: 23-37.
Kovblyuk M.M. & Ponomarev A.V. 2008. New and interesting spiders (Aranei:
Agelenidae, Corinnidae, Gnaphosidae, Nemesiidae, Thomisidae) from the
West Caucasus. Caucasian Entomological Bulletin, 4: 143-154.
Král J. 2007. Evolution of multiple sex chromosomes in the spider genus
Malthonica (Araneae: Agelenidae) indicates unique structure of the spider
sex chromosome systems. Chromosome Research, 15: 863-879.
Seyyar O., Demir H. & Topçu A. 2008. A futher faunistic study on two species
of the genus Malthonica Simon, 1898 (Araneae: Agelenidae) from
Turkey. Turkish Journal of Arachnology, 1: 120-127.
90
Life history of three sympatric wolf spiders

from the Yukon Territory, Canada
Joseph J. Bowden & Christopher M. Buddle
McGill University-Macdonald College, Canada, joseph.bowden@mail.mcgill.ca
Arctic arthropod assemblages are predicted to undergo substantial changes in

response to global climate change, yet we know little about the ecology of many
groups in the north. During the summer 2005 we sampled along a latitudinal transect
in the northern region of Yukon Territory, Canada, to determine which factors
(climate, vegetation, or space) best explain patterns in ground-dwelling spider
species assemblages (Bowden & Buddle, in press). We also used a regional scale
approach during summer 2006 to sample and determine patterns of spiders across
elevation and latitude (Bowden & Buddle, in review). The results of these studies
pointed to the importance of vegetation in determining spider assemblages across
spatial gradients in the north. The most dominant species collected over the two
years of sampling were wolf spiders; the dominant tundra species were Pardosa
lapponica Thorell, Pardosa sodalis Holm and Pardosa moesta Banks on the tundra,
while Pardosa uintana Gertsch was dominant in forested areas.
Because the reproductive traits of individuals form the basis for adaptation to
changing and novel environments and are explicitly linked to local population and
community dynamics (Tokeshi 1999, Reynolds 2003), we wanted to characterize
the reproductive relationships of these species. During the summer 2008 (late June
to early August) we collected female wolf spiders from three tundra sites
(Tombstone, Ogilvy and Richardson mountain ranges) located along the Dempster
Highway in the Yukon. Using live pitfall trapping and hand collecting techniques
we collected females of the following species: P. lapponica, P. sodalis and P.
moesta. All females and their respective egg sacs were weighed alive in the field and
then preserved in 70% ethanol. In the laboratory the width of the carapace was
measured using an ocular micrometer in a stereomicroscope. We calculated body
condition using the residual index (Jakobs et al. 1996). We sought to determine
whether body size or mass independent of size (body condition) was a better
predictor of two different measures of reproductive fitness: fecundity and relative
reproductive output (RRO). We were also interested in whether a trade-off existed
between the number of offspring produced (fecundity) and the investment (average
offspring mass) in each propagule. We used linear mixed models with selection for
the most likely models of female fecundity and RRO using log-likelihood ration
tests and simple linear regressions to test the significance of egg size-number
tradeoffs within species.
While dissecting egg sacs to determine fecundity for these species we noticed
several egg sacs had been parasitized by Gelis sp. (Hymenoptera: Ichneumonidae)
We wanted to know whether these parasitoids were selecting larger females and
whether they demonstrated a preference for a particular host species. We used a
91
linear model with a binary response to test whether Gelis sp. was selecting larger
individuals within each species
We collected a total of 574 P. lapponica 257 P. sodalis and 118 P. moesta
females to analyze reproductive characteristics. We detected significant differences
among sites for many of our explanatory and response variables so we divided our
data by site and by species. The Tombstone site yielded the lowest body size
(condition) for P. lapponica and P. sodalis. Interestingly, this site also yielded the
most significant trade-off effects in these species. We also found that while Gelis sp.
were not selecting larger individuals within a species, they were selecting larger
individuals across species and specifically P. sodalis. We also found that up to 50%
of individuals in a given population can be parasitized by Gelis sp. resulting in a
reproductive fitness of zero.
Our work represents the first to quantify determinants of spider assemblages
and life history patterns of many northern spiders at a regional scale. We found that
vegetation composition and the structural characteristics it represents, best explained
patterns of northern spider assemblages. The boreal forest-tundra transition zone
should become an important focus for research on diversity and distribution of
arthropod species given the susceptibility of arthropods to climate changes
(Callaghan et al. 2001) and future climate predictions (e.g., Hansell et al. 1998,
Danks 2004). Determining patterns of terrestrial arthropod diversity and life history
patterns across the boreal-tundra is the first step to determining the effects that
climate changes will have on the Arctic fauna.
References
Bowden J.J. & Buddle C.M. Determinants of ground-dwelling spider assemblages at
a regional scale in the Yukon Territory, Canada. Écoscience, in review.
Bowden J.J. & Buddle C.M. Spider assemblages across elevational and latitudinal
gradients in the Yukon Territory, Canada. Arctic, in press.
Callaghan T.V., Bjorn L.O., Chernov Y., Chapin T., Christensen T.R., Huntley B.,
Ims R.A., Johansson M., Jolly D., Jonasson S., Matveyeva N., Panikov N.,
Oechel W., Shaver G. & Henttonen H. 2004. Effects on the structure of arctic
ecosystems in the short- and long-term perspectives. Ambio, 33: 436-447.
Danks H.V. 2004. Seasonal adaptations in arctic insects. Integrative and
Comparative Biology, 44: 85-94.
Hansell R.I.C., Malcolm J.R., Welch H., Jefferies R.L. & Scott P.A. 1998.
Atmospheric change and biodiversity in the Arctic. Environmental
Monitoring and Assessment, 49: 303-325
Jakob E.M., Marshall S.D. & Uetz G.W. 1996. Estimating fitness: A comparison of
body condition indices. Oikos, 77: 61-67.
Reynolds J.D. 2003. Life histories and extinction risk. In: Macroecology, T.M.
Blackburn & K.J. Gaston (eds), Blackwell Publishing, Oxford.
Tokeshi M. 1999. Species coexistence: Ecological and evolutionary perspectives.
Blackwell Science, Oxford.
92
Cladistic analysis of subfamily Mitobatinae with emphasis

on the polymorphic character analysis
(Opiliones: Gonyleptidae)
Cibele Bragagnolo & Ricardo Pinto-da-Rocha
Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Brazil,

cibrag@usp.br, ricrocha@usp.br
A cladistic analysis of the subfamily Mitobatinae is presented. All 10 of

the currently valid species of the genus Promitobates Roewer, 1913 were taken
into account. Promitobates ornatus (Mello-Leitão, 1922), a polymorphic and
widespread species, was initially split into four sub-units and its taxonomy
addressed with the results of the cladistic analysis. The other two polymorphic
species, P. hatschbachi H. Soares, 1945 and P. viridigranulatus Soares &
Soares, 1946, were also split into two sub-units. In addition to these species, the
cladistic analysis was performed with 20 other Mitobatinae species, representing
the 11 genera of the subfamily, and 5 species of other Gonyleptidae genera. The
character matrix comprised 75 characters: 19 from male genitalia, 27 from the
general external morphology, 20 from male legs and 10 from colouration. Two
equally parsimonious trees were obtained (L=257, C.I=0.38; R.I=0.72).
Mitobatinae arose as a monophyletic group, divided into two major
groups: [1] one that possesses the body roughly rectangular and males and
females with coxa and trochanter IV without large apophysis, including the
genera: Ischnotherus Kury, 1991, Encheiridium Kury, 2003, Metamitobates
Roewer, 1913, Ruschia Mello-Leitão, 1940, Mitobatula Roewer, 1931 and
Mitobates Sundevall, 1833 and [2] one that possesses the body roughly piriform
and an evident sexual dimorphism of coxa and trochanter IV, with males with a
large apophysis on coxa and tubercles on trochanter IV, including the genera:
Discocyrtoides Mello-Leitão, 1923, Longiperna Roewer, 1929, Neoancistrotus
Mello-Leitão, 1927 and Promitobates.
93
A new species of the sun-spider genus Mummucia

(Arachnida: Solifugae: Mummucidae) from Piauí,
northeastern Brazil
Leonardo Sousa Carvalho1, David F. Candiani2,

Alexandre B. Bonaldo2, Lincoln Suesdek3,4 & Paulo Roberto R. Silva5
1
Universidade Federal do Piauí, Floriano, Piauí, Brazil, carvalho@ufpi.edu.br
2
Museu Paraense Emílio Goeldi, Coordenação de Zoologia, Laboratório de Aracnologia,
Belém, Pará, Brazil
3
Instituto Butantan, Laboratório de Parasitologia, São Paulo, São Paulo, Brazil
4
Programa de Pós-Graduação “Biologia da Relação Patógeno-Hospedeiro”, Instituto de
Ciências Biomédicas da Universidade de São Paulo, São Paulo, Brazil
5
Universidade Federal do Piauí, Campus da Socopo, Centro de Ciências Agrárias,
Departamento de Fitotecnia, Teresina, Piauí, Brazil
Introduction
Despite the fact that the order Solifugae is several times less diverse than
Acari, Araneae, Opiliones, Scorpiones and Pseudoscopiones, this group is still
poorly known in South America. It is possible that, the low diversity currently
recorded to the Neotropical Region could be merely reflecting the lack of
taxonomists currently working on the group, which resulted in a small number of
species described in the current decade. The most important studies on South
American species were done by Roewer and, specially, Maury; but a few recent
contributions have been published, extending the geographical distribution of
Brazilian species, describing new species from Neotropical Biomes such as
Caatinga and Cerrado, or redescribing species based on recent collected
specimens.
The known distribution of Solifugae species in Brazil is still fragmentary,
with large areas without records. Presently, there are only two records in the
Northeastern Brazil: a single juvenile Ammotrechidae collected at Balsas, State
of Maranhão, and the type series of Mummucia mauryi Rocha 2001, which type-
locality is Ibiraba, Bahia State.
Herein, we present the first records of Solifugae from the State of Piauí,
Northeastern Brazil, with the description of a new species of the genus
Mummucia (Mummucidae), from the Serra das Confusões National Park, in the
Caatinga Biome. We also performed geometric morphometrical analyzes to
compare the new Mummucia species with Metacleobis fulvipes Roewer 1934,
owing to their cheliceral dentition similarities.
Methods
Adult individuals (males) of Mummucia n. sp. (n=14) were compared to
M. fulvipes (n=10) regarding the geometry of propeltidium. Digital images of
propeltidium of both species (males) were captured by a Leica DFC320 digital
camera coupled to a Leica S6 stereoscope equipped with plain lenses which
94
avoid image distortion. For each propeltidium, coordinates of eight “type I”

landmarks were digitized and assembled into matrices.
Geometric morphometrical analyzes were performed as described in
literature and are summarized as follows. Propeltidium shape was assessed after
discarding isometric size variation and its relative warps (principal components)
were plotted in graph describing the morphological space of the comparison
between both species. Consensus configurations of propeltidium were built from
shape coordinates after translating, scaling and rotating each specimen and were
compared between the species.
To test the accuracy of the morphometric classification, each individual
was reclassified according to its propeltidium similarity to the average shape of
each species. Mahalanobis distances were used to estimate metric distance in
this discriminant analysis. Landmark digitizing, data analyzes and graphs were
done using software BAC and PAD and TPS software pack.
The specimens are deposited in the Museu Paraense Emílio Goeldi
(MPEG, curator: A.B. Bonaldo) and Universidade Estadual de Feira de Santana
(UEFS, curator: F. Bravo). The formulae of cheliceral dentition and leg
spination, as well as the chetotaxy terminology followed recent taxonomical
publications of Neotropical Solifugae. Scanning Electronic micrographs were
obtained with a Zeiss LEO (1450 VP) scanning electron microscope from the
Laboratório Institucional de Microscopia Eletrônica de Varredura from MPEG.

The new species, Mummucia n. sp. resembles M. coaraciandu Pinto-da-
Rocha & Rocha, 2004 by the pleurite colouration, and M. mauryi by the number
of parallel narrow grooves on the cheliceral stridulatory apparatus, pleurite
colouration and number of anterior tooth on the fixed finger. It differs from these
two species by the combination of the following characters: stridulatory
apparatus on chelicerae mesal face with seven parallel narrow grooves, male
cheliceral fixed finger with two anterior teeth and cheliceral movable finger with
one anterior, one intermediate, and one principal tooth, graded in size from distal
to proximal II, III and I. In addition, pleurital brown spots containing sockets of
bifid bristles are arranged in a pattern which is nearly similar among the
individuals. Such pattern is apparently specific, since it is distinct from the
others species of Mummuciidae.
Variation. A female used for SEM pictures had only four ectal fondal
teeth, while the remaining females have five ectal fondal teeth.
Geometric morphometrical analyses. Shape analyses showed the
individuals arranged in distinct plot groups at the graphical morphospace defined
by principal components 1 and 2. Each group corresponded to one species, with
no overlapping between them. The accuracy classification of females based on
the Mahalanobis distances was 100% for Mummucia n. sp. and 92% for M.
fulvipes.
Superimposition of consensus configurations showed that landmarks 2-8
are distinct between the species. Mean values of ratio propeltidium length/width
95
were: Mummucia n. sp. = 0.89; M. fulvipes = 0.81. Ratio propeltidium

length/distance LM1-LM7: Mummucia n. sp. = 6.31; M. fulvipes = 5.34.
Remarks
The new species Mummucia n. sp., is assigned to the type genus
Mummucia, solely because at the present time it is impossible to reliably
distinguish the genera of Mummuciidae, as already pointed out by Maur y. The
same decision was taken by other authors. The family Mummuciidae still
requires an entire taxonomic and phylogenetic revision to better understand the
genera limits and define the most useful characters for species definition.
However, one character appears to be constant in the family: the flagellum
longitudinal ectal opening, which was already reported only for Mummucia
coaraciandu, M. mauryi, M. taiete Rocha & Carvalho, 2006 and Metacleobis
fulvipes.
Mummucia n. sp. resembles M. coaraciandu in the colour pattern of the
pleurites, which was proposed to be species-specific in the family
Mummuciidae. This feature appears to be convergent on these two species, as
they differ in the chelicerae dentition, a currently character to support species
recognition. Besides, the geometry of propeltidium showed that males of
Mummucia n. sp. may be diagnosed with >91% accuracy when compared to M.
fulvipes. Landmarks 2-8 comprise most the shape variety between the species.
Natural History. The individuals were collected at Serra das Confusões
National Park (9º27'-9º31'S, 43º05'-43º56'W, Guaribas and Caracol
municipalities) in Piauí State, Northeastern Brazil. The climate is hot tropical
semi-arid, with temperature ranging from 18ºC and 38ºC (average 25ºC). The
Serra das Confusões National Park is a 500.000ha reserve, covered by Caatinga
phytophysionomies (arboreal Caatinga, shrubland Caatinga, and enclave forests)
inserted on the arenilitic plateaus (chapadas) and depressions of the Parnaíba
River Basin. The sampling at Serra das Confusões National Park occurred in
October 2006 and July 2007, with pit-fall traps with drift fences (30 blocks of
four 60L plastic buckets, arranged like a “Y”, with drift fences of 60 cm high; 24
h samples). We collected fourteen specimens (eleven males and three females),
all in July 2007, when the enclave area was not sampled. A total of seven
individuals were collected on arboreal Caatinga and six individuals on shrubland
Caatinga. The low number of individuals sampled and the punctual sampling
events prevent us to infer about habitat selection.
96
Abundance and microhabitat use by the whip spider

Heterophrynus longicornis (Arachnida: Amblypygi)
in forest fragments formed by Tucuruí dam’s lake,
Pará, Brazil
Leonardo S. Carvalho1, Jerriane O. Gomes2, Selvino Neckel-Oliveira3,

David F. Figueiredo2 & Nancy F. Lo-Man-Hung2
1
Universidade Federal do Piauí, Campus Amílcar Ferreira Sobral, Floriano, Piauí,
Brazil, carvalho@ufpi.edu.br
2
Museu Paraense Emílio Goeldi, Coordenação de Zoologia, Belém, Pará, Brazil
3
Instituto de Ciências Biológicas, Universidade Federal do Pará, Belém, Pará, Brazil
Introduction
Large-scale habitat fragmentation and ecosystem change are some of the
consequences of dam constructions, affecting both terrestrial and aquatic wildlife.
The land-bridge islands created by damming rivers, result in alterations of the
forest structure, physical conditions, and species diversity: often resulting in the
extinction of some species and changes in the dominance of others.
An increasing number of studies have assessed the effects of forest
fragmentation on the tropical biota, but most of them were conducted in
landscapes for which the matrix was secondary forests, selective logging forest, or
exotic plantations. Until recently, only three studies have investigated the
association between whip spiders and microhabitat use and only one was carried
out in Brazil. Dias & Machado investigated the microhabitat use by
Heterophrynus longicornis (Butler 1973) in an area of continuous forest in Central
Amazonia. In this study, the authors concluded that H. longicornis prefers large
trees bearing buttresses for mating and hunting; and trees with burrows at their
base where the individuals hide during daytime.
The aim of the present study was to test the differences between the
abundance of H. longicornis in edge and interior island plots in the Tucuruí dam’s
lake in Eastern Amazonia; and also provide some information on the abundance
and microhabitat use by age-sex class of this whip spider. Besides, we tested the
influence of the mean basal area on the whip spider abundances in the sampled
plots. As H. longicornis prefers large trees bearing buttresses, and larger trees have
a higher mortality rate near forest fragments periphery, we expect that this species
would be less likely to reside on the periphery of the forest.

This study was carried out in the right margin of Tucuruí dam’s lake
(3º43'-5º15'S, 49º12'-50º00'W), in the municipality of Tucuruí, state of Pará,
eastern Amazonia, Brazil. This artificial lake was formed in 1984 and 1985, and
flooded an area of upland forest of approximately 2400 km2, creating more than
1600 forest islands of different sizes and isolation levels. These islands represent
97
the higher portions of the mounds and hills, which were not flooded by the lake,
therefore they exhibit a highly accidental topography. The annual mean rainfall
in the area is 2500 mm, with a dry season of three to five months long, normally
from July to November. The mean temperature is 24ºC.
Eight land-bridge islands on the right periphery of the Tucuruí dam were
chosen, using a satellite images (ETM/Landsat 7 of 2005). The criteria for
choosing the islands were size, shape, degree of isolation, and conservation
status of the island. The sizes of the islands varied from 12.9 to 91.3 ha, the
elevations of the islands from the periphery to the interior ranged from 8 to 31
meters (19.5±6.2 m; determined using GPS Garmin Map 76CS) and the shape
index varied from 1.074 to 2.09 The areas with shape index lower than 2.5,
theoretically presented areas that do not experience edge effect (core areas), and
could thus be selected for our study. On each island, two plots of 5 x 100 m were
laid out, one 30 m from the island edge, and the other in the interior of the island
at 100-150 m from the island edge. The classification of plots into periphery and
interior was based on previous studies of forest dynamics indicating that most
edge effects penetrate up to 100m from the edge. We actively searched for the
whip spiders along the transects, on tree trunks, inside natural cavities on fallen
logs, on the ground and inside termite nests. Each plot was searched only once,
between 19:00 and 24:00 h, on six consecutive nights during July 2006. In the
same we measured the diameter at breast height (DBH) of all trees and vines
with a DBH≥5 cm. This measurement was used to calculate the mean basal area
(BA) of each plot, using the formula BA=(π*DBH2)/4.
All individuals were captured and recorded as males, females or juveniles,
and then were released. Body measurements were not collected and juveniles
were not classified in sizes or age class. For each collected individual, the
substrate was recorded as: tree, ground or termite nest. The diameter at breast
height (DBH) of all trees with whip spiders was measured; and the presence (or
absence) of termite nests at the base of the trees was categorized. As the species
H. longicornis is widely distributed over the Brazilian Amazon, is easily
recognized by morphological characteristics and there are many records from
Tucuruí dam’s area, we considered unnecessary to collect and deposit voucher
specimens on research collections.
To test the null hypothesis of no significant differences between the
abundance of whip spiders in edge and interiors plots, and also the effect of the
mean basal area inside each plots we conducted a covariance analysis. The mean
basal area and the whip spider abundances were Ln transformed for this analysis.
To compare the abundances of whip spiders between the microhabitats classes,
we used a Kruskal-Wallis test. The statistical analyses we performed using
SYSTAT 12.0, with a significance level of 5%. Abundance data are expressed as
means ±1 SD.

We found 110 H. longicornis, of which 20 males, 38 females and 52
juveniles. In the periphery plots we found 63 (7.875±6.424) and in the interior plots
we found 47 (5.875±4.581) individuals. There was no significant difference between
98
the abundance of whip spiders on edge and interior plots (F=1.385; p=0.264), and no
covariate effect between the mean basal area and the habitat type (periphery or
interior plots) was observed (F=1.043; p=0.329). On the other hand, the abundance
of whip spiders directly increased with the mean basal area of the plots (F=8.431;
p=0.014).
The abundance of whip spiders was higher on trees with termite nests at the
base (n=51), than on trees without termite nests (n=28), or on termite nests far from
trees (n=19). Only 10 individuals were seen wandering or hunting on the ground.
There was no significant difference between the abundance of whip spiders in the
substrate classes (H=5.743; p=0.125). On eleven occasions, more than one
individual (from two to six specimens) was seen on the same tree, but no
intraspecific interaction was observed in the field. Five mother-offspring groups
were recorded on trees, and a male-male pair or an offspring group was seen only
once. Three mother-father-offspring groups and one male-female pair were
recorded. The whip spiders were found more frequently (84.81%) in tress with 10
cm and 50 cm of DBH. On termite nests, the whip spiders were seen only alone,
never in pairs or groups. Four females bearing egg sacs were recorded, all on trees
with termite nests at the base, and with DBH larger than 18 cm. A female preying on
a Lepidoptera (Arthropoda, Insecta) was recorded.
Many studies have shown that different invertebrate groups respond
differently to forest fragmentation and edge effects and concluded that some species
can increase their abundance toward the periphery and others can decrease it. The
absence of significant differences between the abundance of whip spiders in
periphery and interior plots of the islands might be explained by two reasons: either
this fact reflects the disturbed situation of the entire fragment, which could not have
core areas (places that do not suffer the edge effects) due to its topography; or this is
an indication that Amblypygi are not affected by environmental conditions imposed
by the edge effects.
The presence of whip spiders on/near termite nests appears to be related to the
availability of shelters, foraging surface and places that can be used as arena for
courtship, acting like tree buttress. Thus, the presence of termite nests might be
another variable to be evaluated in studies on whip spider’s microhabitat selection.
This substrate can suppress the absence of trees with buttress for the behavioural
activities of whip spiders.
Most studies state that adult Amblypygi are generally solitary and intolerant
predators of congeners, at least during part of their lives; although a few authors
argued that H. longicornis from the Brazilian Amazon are especially tolerant of one
another and the male-female pairs can share the same hiding place. This tolerance
appears to be higher than expected, as 49.4% of the individuals observed on trees,
were sharing the same tree with at least one conspecific. This can be explained by
the low degree of aggressiveness towards conspecifics. Future studies are necessary
to understand the population dynamics and structure of Heterophrynus longicornis
in natural and altered habitats.
99
Ecology of grass dwelling spiders in the coastal Randa

meadows in the eastern part of Gulf of Riga, Latvia
Inese Cera
University of Daugavpils, Latvia, inese.cera@gmail.com
Introduction
Coastal meadows are unique because of specific soil conditions, flora and
fauna communities and rare habitats with high conservation value. The need of
extensive management of these habitats is important to minimize overgrowing
by reed and shrubs (Melecis et al. 1997). As management of meadows has
decreased more attention has been paid to study of influence of harvesting
methods on flora and fauna. The impact of changes in harvesting methods of
meadows on spider distribution is discussed in several publications (Cattin at al.
2003, Humbert at al. 2009 – in inland meadows), about spider biodiversity
potential in inland salt meadows (Zulka et al. 1997). Additionally, about web-
spinning spider diversity and vegetation structural diversity versus pray
availability – Greenstone (1984) and spider dependence on the whole habitat
structure (also vegetation height) (Duffey 1966).
Randa meadows are protected and included in the North Vidzeme
Biosphere Reserve and are Natura 2000 site (Anonymous 2010). Historically the
territory of meadows was maintained by cattle grazing and hay harvesting.
During the last decades the management intensity has sharply decreased and the
majority of territory has overgrown by shrubs and reed. Brackish marine waters
irregularly overflow these meadows by during heavy storms.
The grass-dwelling arthropod communities have been researched since
1994 (Melecis et al. 1998). Results showed that spiders had higher abundance in
the xerophytic meadows. Studies of grassland arthropods showed the decrease of
spider total abundance from inland to seashore since 1996 (Melecis et al. 1997).
The aim of the current research is to investigate a spider fauna in the
Randa meadows and to identify species communities in the particular coastal
meadow habitats and connection with the vegetation characteristics during long-
term observations.

Eighteen sampling plots (2 x 25 m) were selected irregularly in the whole
territory of Randa meadows in all types of meadows. Sampling plots included
xerophytic meadows (4 plots), mesohygrophytic meadows (10 plots) and
xeromesophytic meadows (4 plots) (Melecis et al. 1997, 1998). Grass dwelling
spiders were collected by use of entomological sweep net four times per season
in May, June, July and August from 1997 to 2008. One sample per plot included
50 sweeps along the sampling plot. Killed arthropods were sorted in laboratory.
Spiders were put in the vials with alcohol (70%). Species were identified
100
according to Locket & Millidge (1953), Nentwig et al. I2003) and Almquist
(2005, 2006). Nomenclature follows Platnick (2010).
The seasonal data of every year and data of the 12-year study in the every
meadow type were pooled thus giving 18 data sets used in the analysis. Only
data of adult spiders were used. Detrended Correspondence Analysis (DCA) and
Indicator species analysis (significance level p>0.05) was used to find
differences among sampling plots (habitats), Cluster analysis – for plot
grouping. Engelmann’s (1987) domination classification was used to
characterise the dominance structure.
Results
In total, 644 adult spiders belonging to 13 families, to 75 species and 22
species identified to genus or family level. Spiders of the families Araneidae
(20.34% of all collected spiders) and Linyphiidae (28.57%) dominated.
Theridiidae (12.11%), Tetragnathidae (6.67%), Lycosidae (5.28%),
Philodromidae (7.92%), Thomisidae (9.94%) and Salticidae (4.19%) were
subdominants, while Pisauridae, Dictynidae, Miturgidae, Clubionidae and
Gnaphosidae were recedents or subrecedents.
No spider species dominated. Theridion impressum (7.61% of all collected
spiders), Microlinyphia pusilla (4.5%), Araneus quadratus (5.59%), Tibellus
oblongus (4.19%) and Xysticus cristatus (5.12%) were subdominants.
Individuals of the genus Tetragnatha spp. (4.81%), Bathyphantes spp. (4.34%)
and Pardosa spp. (4.04%) were subdominants.
Discussion
Spiders in the coastal meadows in Latvia have been investigated for the
first time and no comparative studies are available.
Pétillon et al. (2007) investigated impact of cutting and sheep grazing on
spiders and beetles in intertidal salt marshes with use of pitfall traps and
associated it with some halophilic species. 21.7% of species were the same as in
our research, but none of them was halophilic. Finch et al. (2007) studied spider
and beetle zonation in the salt marshes at North Sea coast. Positive connection of
increase of mean high tide and total spider number was found. Because of
different method and time scale used (transect, pitfall traps, two years) only
17.7% of species were as in our study. Irmler et al. (2002) obtained similar
results in the research in two salt marshes at North and Baltic Sea. Abundance of
spiders increased at North Sea, but species richness – at Baltic Sea (12.1%) and
connected with the increase of mean high tide.
Among dominant and subdominant species Xysticus cristatus can be found
on grass and mosses, Tibellus oblongus – at seashore and in wet meadows,
Theridion impressum – in heatlands, gardens, on the bushes, Araneus quadratus
– wet meadows with high vegetation (Almquist 2005, 2006), Microlinyphia
pusilla – on low vegetation in moist and dry habitats (Benjamin et al. 2002). In
comparison with literature data only Tibellus oblongus and Theridion
impressum could be regarded characteristic for coastal meadows.
101
Year-to-year fluctuations of spider fauna can play significant role in

distribution of species in the sampling plots.
References
Almquist S. 2005. Swedish Araneae, part 1 – families Atypidae to Hahniidae. Insect
Systematics & Evolution, 62, Sweden, 284 pp.
Almquist S. 2006. Swedish Araneae, part 2 – families Dictynidae to Salticidae.
Insect Systematics & Evolution, 63, Sweden, 285-603 pp.
Anonymous 17.02.2010. [Randa meadows, restricted area of nature], online:
http://www.daba.gov.lv/index.php?objid=502.
Benjamin S.P., Düggelin M. & Zschokke S. 2002. Fine structure of sheet-webs of
Linyphia triangularis (Clerk) and Microlinyphia pusilla (Sundevall), with
remarks on the presence of viscid silk. Acta Zoologica, 83: 49-59.
Cattin M.-F., Blandenier G., Banašek-Richter C. & Bersier L.-F. 2003. The impact
of mowing as a management strategy for wet meadows on spider (Araneae)
communities. Biological Conservation, 113: 179-188.
Duffey E. 1966. Spider ecology and habitat structure. Senckenbergiana biologica,
47: 45-49.
Engelmann A.-D. 1978. Dominant klassifizierung von Bodenarthropoden.
Pedobiologia, 18: 378-380.
Finch O.D., Krummer H., Plaiser F. & Schultz W. 2007. Zonation of spiders
(Araneae) and carabid beetles (Coleoptera: Carabidae) in island salt marshes
at the North Sea coast. Wetlands Ecology and Management, 15(3): 207-228.
Irmler U., Heller K., Meyer H. & Reinke H.-D. 2002. Zonation of ground beetles
(Coleoptera: Carabidae) and spiders (Araneidae) in salt marshes at the North
and the Baltic Sea and the impact of the predicted sea level increase.
Biodiversity and Conservation, 11: 1129-1147.
Humbert J.-Y., Ghazoul J. & Walter T. 2009. Meadow harvesting techniques and
their impacts on field fauna. Agriculture, Ecosystems and Environment, 130:
1-8.
Locket G.H. & Millidge A.F. 1953. British Spiders. 2. Metchim & Son LTD.,
London: 449 pp.
Nentwig W., Hänggi A., Kropf C., & Blick T. 2003. Spinnen Mitteleuropas/Central
European Spiders. Version 8.12.2003. An Internet identification key,
http://www.araneae.unibe.ch.
Pétillon J., Georges A., Canard A. & Ysnel F. 2007. Impact of cutting and sheep
grazing on ground-active spiders and carabids in internal salt marshes
(Western France). Animal Biodiversity and Conservation, 30(2): 201-209.
Platnick N.I. 2010. The World Spider Catalog. Version 10.0. AMNH, at www:
http://research.amnh.org/entomology/spiders/catalog.
Relys V. & Spunģis V. 2002. Check list of spiders (Arachnida, Araneae) of Latvia,
http://leb.daba.lv/Aranea.htm.
Melecis V., Karpa A., Kabucis I. & Savičs F., Liepiņa L. 1997. Distribution of
grassland arthropods along a coenocline of seashore meadow vegetation,
Latvia. Proceedings of the Latvian Academy of Sciences, Section B,
51(5/6): 222-233.
102
Melecis V., Karpa A. & Spuņģis V. 1998. The grass-dwelling arthropod

communities of the coastal reserve „Randu pļavas”. Latvijas Entomologs,
36: 55-65.
Zulka K. P., Milasowszky N. & Lethmayer Ch. 1997. Spider biodiversity
potential of an ungrazed and grazed inland salt meadow in the national
Park “Neusiedler See-Seekwinkel” (Austria): implication for management
(Arachnida: Araneae). Biodiversity and Conservation, 6: 75-88.
103
Courtship behaviour in the genus Pardosa

(Araneae: Lycosidae): what do we know about it?
Alberto Chiarle* & Marco Isaia
Dipartimento di Biologia Animale e dell’Uomo (DBAU), Università di Torino, Italy

*
Corresponding author: alberto.chiarle@unito.it
Communication in spiders is multimodal, and generally implicates visual,

vibratory/acoustical, tactile and chemical signals. Despite the good visual system
of several spider families (like for example Salticids), many spiders are
nocturnal and the major roles in communication are played by vibrations,
pheromones and tactile signals (Huber 2005).
Mechanical cues could be propagated by air (acoustic signals, detected by
slit sensilla and lyriform organs) or by the substrate (vibratory signals, detected
by trichobothria). Vibratory signals like web-plucking, percussion, stridulation
and body vibration are quite common in spiders. This communication system is
generally s+pecie-specific and used frequently in mate recognition and females
choice (Uetz & Roberts 2002, Huber 2005). An example in this sense is
provided by the wolf spider Hygrolycosa rubrofasciata (Lycosidae) in which the
male generates percussive sounds tapping rapidly the abdomen against leaf litter.
This kind of signals can propagate trough air over long distances and can even
be audible by human ear (Kronestedt 1996).
Chemical signals are also well developed in spiders (Foelix 1996). Spiders
may use chemical cues for the detection of the prey or predator as well as for
species recognition. Pheromones are often deposited by females to attracts
potential mates and trigger male courtship (Uetz and Roberts 2002). In Metellina
segmentata (Tetragnathidae) for example, the assessment of female quality by
males seems to be based on pheromonal cues (Huber 2005).
In most species of Lycosids all these cues are combined in specie-specific
courtship behaviours. This aspect aroused the interest of arachnologists from
taxonomical, ethological and evolutionary points of view. Within Lycosids, one
of the most investigated genus is Schizocosa. In this genus males produce
courtship displays characterized by vibratory and visual signals, associated with
the presence of ornamentation on the first pair of legs. The complex courtship
behaviour of the brush legged spider S. ocreata has been studied extensively
(Uetz 2000, Uetz & Roberts 2002, Taylor et al. 2005, Gibson & Uetz 2008).
Similarly, the wolf spider genus Pardosa shows the same potential
interest. This genus is widespread all through the Holarctic region and has
undergone a great deal of diversification (Tongiorgi 1966a, Topfer-Hofmann et
al. 2000). Species belonging to this genus can be sorted in several groups
according to morphological features such as habitus or shape of genitalia. An
example is provided by P. monticola group, the largest species-group within the
genus with at least 20 species (Tongiorgi 1966b). Even though it is easy to
assign species to the different groups, identification at species level within the
104
group often turns out to be very hard. Despite the courtship displays of the many
Pardosa species has never been observed, it may represent a useful diagnostic
tool to separate species. This is particularly true when considering sibling
species occurring in syntopy, as demonstrated by Den Hollander and Dijkstra
(1974), who described the first ethospecies, P. vlijmi, distinguished from the
sibling P. proxima on the basis of the courtship display. Another example within
the same genus is provided by Topfer-Hofmann et al. (2000) with the description
of two new ethospecies within the P. lugubris group: P. saltans and P. pertinax.
Our research focus on the study of the courtship behaviour of different
species of the genus Pardosa, providing new methods for the description and the
analysis of courtship displays aiming to outline the undescribed ones and to
study their variability within species. We collected spiders in NW Italy and
Belgium. All the specimens were adult or sub-adult reared in tubes in standard
condition and fed with Drosophila sp. ad libitum.
The courtship behaviour of the males has been obtained by putting it in
contact with the female into a glass arena. All the behaviours were recorded for
one hour with two high definition cameras at recording speed of 50i (50 frame
per second interlaced). The movies were acquired and edited with specific
software.
The first step of the analysis was to identify the behavioural acts that
characterized the behaviour type. The different behavioural acts compose a
behavioural pattern that is specie specific. On the other hand, each act is
composed by the movement of the different anatomical parts involved in the
behaviour (Lehner 1998) namely the palps, the abdomen, the first pair of legs
and the general movement of the body. In this way, a list of acts that can be
analyzed with a software can be obtained for each specimen. The software is
able to find linkages among different behavioural acts within a defined
behavioural pattern. The second type of analysis is the Optical flow analysis.
This mathematical method can measure the proprieties of the video (this analysis
was presented by Elias et al. (2005) working on the courtship behaviour of some
Salticids). The result is a speed profile plot of the local speed estimation of the
pixel that compose the image. The plot is analyzed with a second program that is
able to measure the duration of each behavioural act selected by the user, the
time interval between behavioural acts, the number of peaks that compose the
speed profile plot and their characteristics such as the mean period and
coefficient of variation.
By means of these methods we studied the courtship display of about
twenty species of Pardosa, more than half of them never observed before. In this
work we present an overview of the results obtained in the first two years of the
research.
The first study dealt with Pardosa wagleri and P. saturatior, sibling
species ecologically separated, previously studied by Tongiorgi (1966a) and
Barthel & von Helversen (1990). We described the courtship displays of the two
species for the first time, highlighting differences and similarity of the two
behaviours (Chiarle et al. in press). Our study deepened the work of Tongiorgi
105
(1966a) and Barthel & von Helversen (1990), providing new insights
(description and confront of the two displays) on the biology of the two species.
We also studied syntopic occurrence and mixed populations of P. proxima
and P. vlijmi that we recorded in Italy for the first time. A detailed description of
the courtship display in comparison with the one observed by den Hollander and
Dijkstra (1974) was provided. Additionally we compared the two behaviours and
the variability within species, adding some remarks about the morphology of
male palps, that may be useful for a correct species identification (Chiarle &
Isaia in prep.). In the perspective of using courtship displays to identify species,
we analyzed the courtship displays of P. blanda, P. mixta and P. torrentum
(showing clear difference in their courtship despite morphological similarity).
We also described the courtship displays of P. nigra, a species that stands out for
its unclear taxonomical position (several authors placed it in the genus
Acantholycosa) but also for the peculiarity of its courtship display. A possible
new ethospecies from an Alpine meadow in south western Alps has also been
observed.
To deepen the work we also investigated the degree of reproductive
isolation of these species, using molecular markers. This approach has been
successfully used in the study of phylogenetic divergence in sibling species
complexes (De Busschere et al. in prep). In our case we compared the
ethological results to the genetic ones, highlighting some interesting aspects in
terms of allopatric speciation (P. wagleri vs. P. saturatior), syntopic close-
related species (P. monticola group) and genetic diversity (P. proxima vs. P.
vlijmi).
References
Barthel J. & von Helversen O. 1990. Pardosa wagleri (Hahn 1822) and Pardosa
saturatior Simon 1937, a pair of sibling species (Araneae, Lycosidae).
Bullettin de la Société européenne de Arachnologie, Hors Série, 1: 17-23.
Chiarle A., Isaia M. & Castellano S. (in press). New findings on the courtship
behaviour of Pardosa wagleri (Hanh, 1822) and P. saturatior Simon,
1937 (Araneae, Lycosidae), a pair of sibling species. Contribution to
Natural History (Bern).
den Hollander J. den & Dijkstra H. 1974. Pardosa vlijmi sp . nov., a new
ethospecies sibling Pardosa proxima (C. L . Koch, 1948), from France,
with description of courtship display (Araneae, Lycosidae). Beaufortia,
22: 57-65.
Elias D.O., Land B.R., Manson A.C. & Hoy R.R. 2005. Measuring and
quantifying dynamic signals in jumping spiders. Journal of Comparative
Physiology, A, 192: 785-797.
Foelix R.F. 1996. Biology of spiders. Oxford University Press, Berlin.
Gibson J.S. & Uetz G.W. 2008. Seismcommunication and mate choice in wolf
spiders: components of male seismic signals and mating success. Animal
Behaviour, 75: 1253-1262.
Huber B.A. 2005. Sexual selection research on spiders: progress and biases.
Biological Review, 80: 363-385.
106
Kronestedt T. 1996. Vibratory communication in the wolf spider Hygrolycosa

rubrofasciata (Araneae: Lycosidae). Revue Suisse Zoologie, vol. hors
serie: 341-354.
Lehner P.N. 1998. Handbook of ethological methods. 2nd ed. New York,
Cambridge University Press, 672 pp.
Taylor P.W., Roberts J.A. & Uetz G.W. 2005. Flexibility in the multimodal
courtship of a wolf spider, Schizocosa ocreata. Journal of Ethology, 23:
71-75.
Tongiorgi P. 1966a. Italian wolf spiders of the genus Pardosa (Araneae: Lycosidae).
Bullettin of the Museum of Comparative Zoology, 134: 275-334.
Tongiorgi P. 1966b. Wolf spiders of the Pardosa monticola group (Araneae:
Lycosidae). Bullettin of the Museum of Comparative Zoology, 134: 335-359.
Töpfer-Hofmann G., Cordes D. & von Helversen O. 2000. Cryptic species and
behavioural isolation in the Pardosa lugubris group (Araneae, Lycosidae),
with description of two new species. Bulletin of the British
Arachnological Society, 11(7): 257-274.
Uetz G.W. & Roberts J.A. 2002. Multisensory cues and multimodal
communication in spiders: insights from video/audio playback studies.
Brain Behaviour and Evolution, 59: 222-230.
Uetz G.W. 2000. Signals and multi-modal signaling in spider communication.
In: Espmark Y., Amundsen T. & Rosenqvist G. (eds), Animal Signals:
Signalling and Signal Design in Animal Communication. Trondheim,
Norway: Tapir Academic Press, pp. 387-405.
107
Pseudoscorpions in the nests of birds

Jana Christophoryová1 & Zuzana Krumpálová2
1
Department of Zoology, Faculty of Natural Sciences, Comenius University, Bratislava,
Slovakia, christophoryova@gmail.com
2
Institute of Zoology, Slovak Academy of Sciences, Bratislava, Slovakia,
zuzana.krumpalova@savba.sk
Introduction
The fauna of bird nests has been mainly studied from the parasitological
point of view. Pseudoscorpions as typical nest predators represent a less
examined group.
The differences between the opinions about the relationship of
pseudoscorpions to bird-nest as a type of environment have inspired us to carry
out this research. In particular, the specific objectives were: 1) to evaluate
pseudoscorpion presence in different types of nests, 2) to monitor the occurrence
of developmental phases during the nesting season and 3) to characterize and
categorize pseudoscorpion species according to their relationship to the hosts.

A total of 171 positive nests of 28 bird species were collected in Slovak
Republic, Austria and Czech Republic. Nests were removed after fledging of
chicks and immediately sealed in polyethylene bags in field; only 18 of them
were picked up 20 days after fledging. The nests were divided into 6 categories:
A – nests in hollows and boxes, B – open nests on the ground, C – open nests in
reeds, D – open nests on trees and shrubs, E – nests on or in buildings, F – nests
in burrows. The similarity of the pseudoscorpion species representation in the
bird-nests was evaluated according to the hierarchical cluster analysis (single
linkage) based on the Jaccard similarity. Pseudoscorpions were also analyzed by
using the principal component analysis (PCA) utilizing Past exe.
Results
The highest number of species belonged to the family of Chernetidae (318
specimens) – Chernes hahnii, Dendrochernes cyrneus, Allochernes wideri,
Lamprochernes chyzeri, L. nodosus and Pselaphochernes scorpioides. Two
species belonged to the family of Cheliferidae (98 specimens) – Chelifer
cancroides, Dactylochelifer latreillii and one species represented each family of
Neobisiidae (4 specimens) – Neobisium carcinoides, Larcidae (4 specimens) –
Larca lata and Cheiridiidae (56 specimens) – Cheiridium museorum. 5 species
were recorded in Austria, 3 species in the Czech Republic and 11 species were
recorded in Slovak Republic.
Based on analysis of 480 specimens from 171 nests of 28 bird species the
pseudoscorpion community was characterized by the frequent presence of
species Ch. museorum, D. latreillii, D. cyrneus, A. wideri and P. scorpioides.
Almost 72% of individuals were present in hollow nests and nest boxes, the
108
lowest number of individuals were present in open nests in reeds and burrows
(0.2%). Pseudoscorpions living under tree bark or in tree hollows searched for
hollow nests respectively open nests on trees, epigeic species for nests on the
ground and synanthropic species for nests in or on the buildings.
PCA confirmed the high proportion and influence of P. scorpioides in the
Hoopoe nests, D. cyrneus in the Eurasian Tree Sparrow nests and A. wideri in
the nests of Tawny Owl, European Scops Owl and European Roller. In contrast,
D. latreillii influenced the nest fauna of Blackbirds and Song Thrushes and Ch.
museorum the nests of White Wagtails.
The development stages of nine pseudoscorpion species were confirmed in
some types of nests. The first females with eggs appeared in May, they reached a
peak in June and then they gradually declined in number until August. The
protonymphs reached the maximum in July afterwards their frequency continued
to decrease until September. The deutonymphs occurred in a higher number in
July, their maximum was in August. The abundance of tritonymphs gradually
increased from April and reached a peak in August. The adults achieved the
maximum in August.
Based on the analysis of the pseudoscorpions, they were grouped into
two basic categories:
A. ixenous species – pseudoscorpions occurred accidentally in bird-nests,
the developmental stages are represented sporadically – N. carcinoides, L.
chyzeri, L. nodosus and probably L. lata.
B. ilous or nidicolous species – pseudoscorpions occurred regularly in
bird-nests, the developmental stages are represented numerously. These species
prefer nests in certain habitats with specific building and microclimate
conditions:
- synanthropic nests: Ch. museorum, Ch. cancroides;
- hollow nests: D. cyrneus, A. wideri;
- open and hollow nests on trees and shrubs: D. latreillii, Ch. hahnii;
- nests with decomposed material: P. scorpioides.
We confirmed the occurrence of 11 pseudoscorpion species in bird nests.
Seven of them were nidiphilous or nidicolous. Microclimatic conditions,
conditions for individual development and trophic offers affect the
pseudoscorpion fauna in nests in extenso.
Acknowledgements
We would like to express our gratitude to all our colleagues for collecting
the bird-nests and Ing. Martin Fris and Doc. M. Holecová for their technical
assistance. This study was financially supported by VEGA grants No. 1/0176/09,
02/0067/08 and KEGA grant No. 3/6235/08.
109
A tangled web: bias sex ratios, bacteria and

mating behaviours of a solitary sheet web spider,
Pityohyphantes phrygianus (Linyphiidae)
Melanie A. Cotterill
University of Nottingham, United Kingdom, Plxmac2@nottingham.ac.uk
Pityohyphantes phrygianus (Linyphiidae) is a solitary sheet web spider

commonly found in coniferous forests. Fisher’s theory predicts that natural
populations should maintain a stable sex ratio of 1:1. However, P. phrygianus
populations are frequently observed to have a female biased sex ratio averaging
around 1:3. Predation and over-wintering mortality rates do not seem to account
for this bias alone which is observed even at the embryo stage.
Studies by Gunnarsson et al have shown that females tend to produce a 1:3
primary sex ratio.
A more recent study provided evidence that P. phrygianus females have
the potential to influence the sex ratio of their offspring by altering their post
copulatory position. Experiments manipulating the post copulatory position of
females, have shown that brood sex ratios near 1:1.
Interestingly P. phrygianus also harbours Wolbachia a maternally
inherited bacterial endosymbiont known to distort host offspring sex ratios in
numerous solitary arthropods. At least one other endosymbiont is present
(Rickettsia).
By altering its host’s behaviour, could the presence of Wolbachia be the
driving force in the maintenance of an otherwise paradoxical sex ratio bias?
110
Areas of endemism analysis and distribution modelling of

harvestmen (Laniatores) to discover biogeographical
history of Neotropical Atlantic Forest
Marcio B. DaSilva
Departamento de Zoologia, Universidade de São Paulo, São Paulo, Brazil,

1940@uol.com.br
Neotropical harvestmen have recently and successfully been used to

biogeography studies. This is a good group for a historic biogeographical research
due to its high levels of endemism. In this work I used 180 species ranges of
harvestmen living in Atlantic Rain Forest to address main patterns of endemic
distribution congruence and delimit areas of endemism. I used species of Stygnidae
Pickeliana (3 spp.), Protimesius (4 spp.), Stygnus (1 sp., the only one living in
Atlantic Forest), and Gaibulus (1 sp.); Cosmetidae Metavononoides (15 spp.);
Gonyleptidae Bourguyiinae (7 spp.), Caelopyginae (23 spp.), Goniosomatinae (37
spp.), Hernandariinae (18 spp.), Heteropachylinae (10spp.), Gonyleptinae
Mischonyx (9 spp.) and Roeweria (2 spp.), Mitobatinae Promitobates (10 spp.),
Pachylinae Eusarcus (23 spp.), Progonyleptoidellinae (12 spp.), and Sodreaninae (5
spp.), all of them with at least two geographical records. It was used NDM/VNDM
2.5 (Programs for identification of areas of endemism) to search for spatial
concordance among species ranges, with 30’x30’ and 1ºx1º cell-sized grid.
From all possible results of the program, I chose those that consider areas of
endemism as mutually exclusive spatial entities, i.e., areas which do not overlap any
other. I found 12 exclusive areas of endemism for coastal Atlantic Forest, related to
mountain range slopes and coastal plain. Species living in interior forest did not
agree spatially, i.e., there are not areas of endemism for harvestmen in the drier
Atlantic Forest, except in slopes of Serra do Espinhaço mountain range, a
Semidecidous forest area.
Thus I classified all species in three levels: endemic, endemic but with partial
congruence with other endemic, and widespread. I chose two species of each type to
run a distribution modelling analysis to aim possible causes of present range
restriction of harvestmen species. I used annual mean and seasonality (four values in
year) of temperature and precipitation with BIOCLIM algorithm in DIVA-GIS
program. Modelling of six species showed that present ecological conditions would
allow much wider distributions of them, i.e., geographical variation in present
climate do not explain those distributions.
Therefore, historical causes can explain those patterns since there is high
congruence in endemism of species and the niche modelling did not show ecological
restriction. Possible causes are related to uplifting of mountain ranges in Tertiary,
and marine transgressions in major rivers in warmer periods and separation of forest
refuges, which probably were located in mountain slopes facing the sea, in colder an
drier periods during Quaternary fluctuations.
111
A faunistic and zoogeographical review of spiders

in Albania (Arachnida: Araneae)
Christo Deltshev1, Blerina Vrenosi2, Gergin Blagoev3
& Stoyan Lazarov1
1
Institute of Zoology, Bulgarian Academy of Sciences, cdeltshev@institutezoology.com,
st.lazarov68@gmail.com
2
Museum of Natural Sciences, Faculty of Natural Sciences, Tirana University, Albania,
blerinavrenozi@yahoo.com
3
Biodiversity Institute of Ontario, University of Guelph, Canada,
gblagoev@uoguelph.ca
The spider fauna of Albania is represented by 168 species from 34

families. In this number, 54 species are new announced for the country. This
number was established after a critical review of the existing literature data and
original collection made in last 15 years during the field survey covering mostly
the coastal parts, caves and some mountain areas. This number of species is not
final, because the territory of Albania is poorly explored. According to their
current distribution, the Albanian species can be classified in 17 zoogeographical
categories, grouped into 4 complexes: widely distributed, European, Balkan
endemics and Mediterranean. Widely distributed species are dominants (61
species), but the most characteristic are Balkan endemics (30 species). The
number of endemics is really high and reflects the local character of the fauna.
112
Four new species of spiders of the genus Stegodyphus

Simon, 1873 from Satpuda, Maharashtra, India,
(Arachnida: Araneae: Eresidae)
Ujjwala S. Deshmukh
Department of Zoology, Government Vidarbha Institute of Science & Humanities,

Amravati, India, ujjwaladeshmukh@rediffmail.com
The spiders of the family Eresidae are represented poorly in Indian fauna.
This study examines species in a relatively diverse lineage of Stegodyphus
spiders in the Satpuda highlands, where they appear to have undergone adaptive
radiation, with several species generally co-occurring at any one locality. Four
new species of spiders from family Eresidae are recorded from outskirt of
Amravati city (Maharashtra) during 2009.
Keywords: new species, taxonomy, Stegodyphus, India.
113
Rounding up the usual suspects: molecular phylogeny

of orb weaving spiders (Araneoidea)
Dimitar Dimitrov1,6*, Lara Lopardo2,6, Gonzalo Giribet3,

Miquel Arnedo4, Fernando Álvarez-Padilla5
& Gustavo Hormiga6
1
Zoological Museum, University of Copenhagen, Copenhagen, Denmark,
dimitard.gwu@gmail.com
2
Zoologisches Institut und Museum, Allgemeine und Systematische Zoologie,
Greifswald, Germany, laralopardo@gmail.com
3
Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA,
ggiribet@oeb.harvard.edu
4
Biodiversity Research Institute UB, Departament Biologia Animal, Universitat de
Barcelona, Barcelona, Spain, marnedo@ub.edu
5
Universidad Nacional Autónoma de México, Facultad de Ciencias, Departamento de
Biología Comparada, Mexico D.F, Mexico, fap@ciencias.unam.mx
6
DC, USA, hormiga@gwu.edu
* Presenting author
We present a revised hypothesis for the relationships of the families in the

superfamily Araneoidea. The present study is based on a combination of newly
collected sequence data and information already available in GenBank for six
loci for over two hundred taxa. These data have been analyzed using parsimony
(both under direct optimization and static homology) and model based
phylogenetic methods. We also explore the contribution of various data
partitions to the phylogenetic pattern. The addition of new taxa and the
availability of data for some poorly studied groups provide new insights on
araneoid phylogeny and evolution.
114
Spinning apparatus of two rare wolf spiders

(Araneae: Lycosidae) – preliminary results 1
Petr Dolejš1,2, Jaroslav Smrž1 & Jan Buchar1
1
Department of Zoology, Faculty of Science, Charles University in Prague, Czech
Republic, dolejs@natur.cuni.cz
2
Department of Zoology, National Museum, Prague, Czech Republic
The spinning apparatus (spinning glands and spinnerets) is one of the most
outstanding features in spiders. Its product, a silk thread, plays an important role
in all spiders, at least in several stages of their life. Till now, mostly the spinning
apparatus of araneid spiders has been studied. On the contrary, there are only a
few papers dealing with the lycosid spinning apparatus. Jaworowski (1896),
Richter (1970a) and Traciuc (1971) were among the firsts who studied the
spinning apparatus of wolf spiders. Richter (1970b) focused on histological and
morphological changes of spinning glands related to maturation of Pardosa
amentata whereas Wąsowska (1977) described spinnerets development in P.
lugubris. Bílek (1992) compared piriform spigots of four wolf spiders with those
of other araneomorph spiders, and finally Townley & Tillinghast (2003)
investigated the role of lycosid ampullate gland silks. However, a complex
study, combining histology of spinning glands and morphology of spinnerets in
all instars of wolf spiders is still lacking.
The aim of the present study is to describe the spinning apparatus of two
rare European wolf spiders (Lycosidae), to investigate its changes during the
spiders’ ontogeny and to compare a spinning apparatus of a burrowing spider
with that of a vagrant one. Tricca lutetiana (Simon, 1876) is a burrowing wolf
spider inhabiting entirely enclosed, not silk lined burrows (Dolejš et al. 2008)
whereas Arctosa alpigena lamperti Dahl, 1908 is a vagrant species. The spiders
were fixed in modified (Smrž 1989) Bouin-Dubosque-Brasil fluid, embedded in
Paraplast plus (Fluka) and sectioned by rotary microtome (Leica RM 2155) at 6
– 8 μm. Sections were stained in triple stains (Masson’s, Gomori’s, Pollak’s and
Domagk’s trichrome). The preparations were inspected under a light microscope
(Olympus Provis AX 70). Selected plates were photographed using a 3CCD
colour video camera (Sony DXC-950P). The external morphology of spinnerets
was examined using a scanning electron microscope (JEOL JSM-6380 LV).
Here, results of the first three and last two instars are presented. Number
of half of the spinning apparatus is given. The spinning apparatus of the first
instar (T. lutetiana/A. a. lamperti) consists of 5/6 piriform and 7/15 aciniform
glands only. During the second and third instar, their number increase (more
rapidly in A. a. lamperti), and a pair of ampullate glands appears. In both
species, subadult males, compared with adult males, possess similar number of
1
The research founded by the Grant Agency of the Charles University: GA UK140907
115
spinning glands. In females of both species, only a few tubuliform glands appear
after the final moult.
T. lutetiana and A. a. lamperti have four types of spinning glands as other
lycosid species studied so far. However, piriform and aciniform glands differ in
number and size, probably reflecting different life strategies of both species.
Number of spinning glands of A. a. lamperti resembles those of vagrant Pardosa
lugubris (Wąsowska 1977), whereas number of spinning glands of T. lutetiana is
lower as the species does not spin very often given its lifestyle (Dolejš et al.
2008). In further study, the spinning apparatus of the remaining instars of both
species will be investigated. For comparison, developmental changes of spinning
apparatus in two more species – Pardosa amentata (Clerck, 1757) and
Xerolycosa nemoralis (Westring, 1861) – will be studied.
References
Bílek P. 1992. Cuticular spinning structures reveal evolutionary relationships in
araneomorph spiders. Sbor. věd. Prací LF UK Hradec Králové, 35(4): 353-370.
Dolejš P., Kubcová L. & Buchar J. 2008. Subterrestrial life of Arctosa lutetiana
(Araneae, Lycosidae). Journal of Arachnology, 36(1): 202-203.
Jaworowski A. 1896. Die Entwicklung des Spinnenapparates bei Trochosa
singoriensis Laxm. mit Berücksichtung der Abdominalhänge und der
Flügel bei den Insekten. Zeitschrift fűr Naturwissenschaften, 30(23): 39-74.
Richter C.J.J. 1970a. Relation between Habitat Structure and Development of
the Glandulae ampullaceae in Eight Wolf Species (Pardosa, Araneae,
Lycosidae). Oecologia, 5: 185-199.
Richter C.J.J. 1970b. Morphology and Function of the Spinning Apparatus of
the Wolf Spider Pardosa amentata (Cl.) (Araneae, Lycosidae). Zeitschrift
fűr Morphologie der Tiere, 68: 37-68.
Smrž J. 1989. Internal Anatomy of Hypochthonius rufulus (Acari: Oribatida).
Journal of Morphology, 200: 215-230.
Townley M.A. & Tillinghast E.K. 2003. On the use of ampullate gland silks by
wolf spiders (Araneae, Lycosidae) for attaching the egg sac to the
spinnerets and a proposal for defining nubbins and tartipores. Journal of
Arachnology, 31(2): 209-245.
Traciuc E. 1971. L’appareil séricigèna chez Pardosa lugubris (Lycosidae,
Araneae). Revue Roumaine de Biologie, Zoologie, 16(3): 171-174.
Wąsowska S. 1977. Studies on the spinning apparatus in spiders. Postembryonic
morphogeny of the spinning apparatus. Zoologica Poloniae, 26(3-4): 55-407.
116
“Cteniza” bavincourti and arachnid trace fossil nomenclature

Jason A. Dunlop1 & Simon J. Braddy2
1
Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity
at the Humboldt University Berlin, Berlin, Germany, jason.dunlop@mfn-berlin.de
2
Department of Earth Sciences, Wills Memorial Building, University of Bristol, Queen’s
Road, Bristol, UK, S.J.Braddy@bristol.ac.uk
Sabella bavincourti Vaillant 1909 from the Eocene of northern France is a

little-known trace fossil subsequently attributed – as Cteniza bavincourti – to the
burrowing activities of a trapdoor spider. It is thus an ichnospecies name and not
a body fossil. Its interpretation as the activity of a spider is questionable and its
original assignment to a worm burrow seems intuitively more likely. Irrespective
of the affinities of the producer, the ICZN also covers ichnotaxa such that
classifying these structures under a modern spider (or worm) genus name creates
a homonym. Theridium columbianum (Scudder 1878) from the Eocene of the
USA is based on fossilized (theridiid?) egg sacs. As such, this species should be
regarded as an ovotaxon; a nomenclatural grey zone between body and trace
fossils. Similar problems underlie fossilized galls attributed (probably correctly)
to mites, but assigned to living eriophyid mite genera. Fossil galls are the
preserved pathological reactions of plant tissue and are also not ichnotaxa sensu
stricto. We propose that these mite names also lie outside the bounds of
zoological nomenclature. Within the broader context of arachnid-related trace
fossils, we briefly review fossil spider webs and arachnid trackways.
117
Arachnid and myriapod types in

the Museum für Naturkunde, Berlin
Jason A. Dunlop & Anja Friederichs
Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at
the Humboldt University Berlin, Germany, jason.dunlop@mfn-berlin.de,
anja.friederichs@mfn-berlin.de
The Museum für Naturkunde Berlin, boasts over 5,000 type species of
arachnids, myriapods and stem-group arthropods. A preliminary Excel file
listing our known, suspected (and missing) types is now freely available from
the authors as an aid to systematic research, and will be regularly updated and
emended in the future. The raw data has also been cross-referenced by author,
locality and collector. This allows collection strengths to be identified and
should also facilitate wider studies of, say, biogeography or cultural-historical
aspects. Many of our types have already been databased in further detail using
the SysTax platform <http://www.biologie.uni-ulm.de/systax/>, developed under
the auspices of the German branch of GBIF (Global Biodiversity Information
Facility) <http://www.gbif.de/>. We encourage users to access our primary data
– and those of other collections – via these portals too.
118
Patterns in the composition of ground-dwelling spider

communities in the Pilbara bioregion, Western Australia
Bradley J. Durrant1, Mark S. Harvey2,4, Volker W. Framenau2,4,

Ricardo Ott2,3 & Julianne M. Waldock2
1
Department of Environment and Conservation, Wanneroo, Western Australia, Australia
2
Department of Terrestrial Zoology, Western Australian Museum, Locked Bag,
Welshpool DC, Western Australia, Australia,
julianne.waldock@museum.wa.gov.au
3
Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul, Rua Dr.
Salvador França, Porto Alegre, Rio Grande do Sul, Brazil
4
School of Animal Biology, University of Western Australia, Crawley, Western
Australia, Australia
Some of the patterns of species composition of the ground-dwelling spider

fauna of the Pilbara bioregion, Western Australia, particularly in terms of
cartographic, climatic and geological attributes are explored. The survey covered
24 areas chosen to represent the geographical extent and diversity of terrestrial
environments in the Pilbara region, an area of approximately 179000km2. A total
of 375 species comprising 14 families were recorded.
119
Recent studies of tropical orb webs: effects of

miniaturization and manipulation by parasitoid wasps
William G. Eberhard
Smithsonian Tropical Research Institute and Escuela de Biologia Universidad de Costa

Rica, Costa Rica, william.eberhard@gmail.com
Orb construction involves many behavioural decisions that require rapid,

precise coordination and orientation, memory, and sustained concentration. In
this talk I will emphasize two different kinds of challenges to spiders as they
make these decisions – the reduction in mental capacities likely to be associated
with tiny body size, and the effects of parasitic larvae of ichneumonid wasps,
which induce their host spiders to build special structures that promote the
survival of the wasp´s cocoon and pupa. Because there is a minimum size below
which neurons cannot transmit action potentials reliably, animals of very tiny
size must either possess a proportionally larger CNS to maintain similar numbers
of neurons, or reduce the numbers of neurons. The disproportionately high costs
of building and maintaining nervous tissue may select against the oversized CNS
option, while the need to maintain behavioural capacities could select against the
reduced neuron numbers option. What do tiny spiders do in such a situation? I
will present several types of morphological and behavioural data which suggest
that species with tiny adults, and larger species with tiny nymphs show
unreduced behavioural capacities, and adopt the oversized CNS option. The
behavioural manipulations produced by wasp larvae apparently result from
injection of a psychotropic substance or substances into the spider. In two spider
species the behavioural effects are reversible if the larva is removed, and in one
of these they appear to depend on the concentration of the psychotropic
substance within the spider´s body. Different wasp species show exquisite
adjustments of their effects on the host spider that exploit details of the spider`s
natural history; it is not yet certain whether these adjustments are the result of
differences between the psychotropic substances that they use.
120
Myrmarachne (Araneae: Salticidae) of the Philippines

G.B. Edwards
Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, FL, USA,
edwardg@doacs.state.fl.us
Historically, the Philippines had 23 species of Myrmarachne described

from within its political boundaries, and two other species described from Java
and reported from the Philippines. Of the 23 species described from the
Philippines, 20 are described from males (3 have essentially undescribed females
in the type series), 3 from females, and only 1 species is adequately described
from both sexes. One of these males, M. bellicosa, is resurrected from synonymy
with the apparently widespread Javan species, M. maxillosa. Two of the females
appear to be synonymous with 2 of the males, and 2 different males are
synonymous with 2 other males, reducing the overall number to 21 species.
Other synonymies are suspected, but some types are difficult to obtain to make
comparisons. Females are described for 9 of the species previously known only
from males.
The geographic makeup of these 21 species is about as unbalanced as the
previous sex ratio of described species. Luzon Island has 14 species (1 each of
which also occur on Mindanao and Panay), Panay and Palawan have 2 other
species each, and 1 additional species apiece uniquely occurs on Mindanao and
Negros. One species is only known from “the Philippine Islands.” Previously
undescribed species and new records help to restore this balance, as 5 new
species are described from Mindanao, and an additional 5 species (one new) are
reported from Palawan (including the first record of M. gisti from the
Philippines). Present thought is that island Myrmarachne faunas seem to mostly
consist of species unique to that island or island group. This is consistent with
recent evidence that Myrmarachne speciation is driven by adaptive radiation.
Conversely, other evidence suggests that perhaps many species of Myrmarachne
are widespread, with slight local variations from place to place that are
misinterpreted as new species, and which ultimately will result in numerous
synonyms.
121
Sit-and-wait predators: a misnomer for orb-weaving spiders
Mark A. Elgar
Department of Zoology, University of Melbourne, Vic., Australia,

m.elgar@unimelb.edu.au
Web-building spiders are frequently described in biology textbooks as

‘classic’ examples of ‘sit-and-wait’ predators, a term used to describe those
species that do not actively pursue their prey, but rather remain in the same place
waiting for prey to come within striking distance. One might infer from this
description that orb-weaving spiders take a relatively passive role in acquiring
food. However, it is increasingly clear that these spiders, and orb-weavers in
particular, employ all manner of strategies to improve their foraging success.
These strategies include strategic placement of the web, modifying the structure
of the web, attracting prey to the vicinity of the web, and reducing the risk of
losing prey to natural enemies. Evidence for these strategies are reviewed,
highlighting controversies and where there are avenues for addressing interesting
questions.
122
The systematics of the New World scorpion

genus Centruroides Marx, 1890 (Buthidae)
Lauren A. Esposito1 & Lorenzo Prendini2

1
American Museum of Natural History, New York, NY, USA, esposito@amnh.org
2
American Museum of Natural History, New York, NY, USA, lorenzo@amnh.org
The New World buthid scorpion genus Centruroides Marx, 1890 is a

morphologically diverse and highly venomous clade, which includes the only
scorpions of medical importance in North America and the Caribbean. In some
states in central and western Mexico, nearly 5 out of every 100,000 inhabitants
die annually from Centruroides envenomation. Despite their notoriety, little
progress has been made in the systematics of Centruroides in over a century, for
several reasons. The morphological characters traditionally used (morphometrics
and colour) for Centruroides taxonomy often overlap between closely related
species and are vaguely defined. Many commonly used characters vary within
populations, hindering species delimitation and identification. Several species
have extremely large distributions, suggesting they may be complexes of cryptic
species. These include “species” with very discontinuous distributions, but for
which traditional characters have proven unreliable. Various researchers have
revised small groups of species as part of regional treatments, but no modern
taxonomic revision of the entire genus exists. The only attempt at phylogenetic
analysis used molecular data from a single gene (16S rDNA) to infer a
phylogeny of only thirteen of the seventy-two currently recognized species, and
concluded that the genus was not monophyletic. We present the first
comprehensive phylogeny of Centruroides, based on five nuclear and
mitochondrial loci, sequenced from 60 species and 10 outgroup taxa. Results of
this analysis confirm the monophyly of Centruroides and reveal novel insights
into the systematics of New World buthid scorpions.
123
Cytogenetics of four corinnid species (Araneae: Dionycha)

with the fourth record of a X1X2X3X4 sex chromosome
system in spiders
Viviane Fagundes de Mattos1, Marcos José Wolf1,

Leonardo Sousa Carvalho2, Antonio Domingos Brescovit3
& Douglas Araujo1
1
Universidade Estadual de Mato Grosso do Sul, Unidade de Mundo Novo, Mato Grosso
do Sul, Brazil, vivianefagundesmn@hotmail.com, marcos.jwolf@hotmail.com
2
Universidade Federal do Piauí, Campus Amílcar Ferreira Sobral, Piauí, Brazil,
carvalho@ufpi.edu.br
3
Laboratório de Artrópodes, Instituto Butantan, São Paulo, Brazil,
adbresc@terra.com.br, daraujo@uems.br
The family Corinnidae consist of 956 species and 81 taxonomically

described genera, including, for example, Castianeira, with 131 species
worldwide; Corinna, containing 69 species predominantly distributed in South
and Central Americas; Falconina that includes only four species; and Xeropigo,
with only nine described species, both occurring mainly in South America.
Cytogenetical studies in Corinnidae are scarce, involving only six species of four
genera, Castianeira, Falconina, Oedignatha e Trachelas, showing diploid
number between 2n♂=22 and 2n♂=28, acrocentric chromosomes and sex
chromosome system of the type X1X2. The genera Corinna and Xeropigo were
not karyotyped until now. Considering the scarcity of chromosomal data to the
family, the aim of the present work is to characterize Castianeira sp., Corinna
gr. nitens, Falconina sp., and Xeropigo n. sp. in relation to the diploid number,
chromosomal morphology and type of sex chromosome system, comparing the
data with cytogenetic information on other dionychans available in the literature.
Two males of Castianeira sp., and one male and one female of Corinna
gr. nitens were collected in São Francisco Island (24°00'32.40"S,
54°09'52.18"W), Parque Nacional de Ilha Grande, border between the states of
Mato Grosso do Sul and Paraná, Brazil. The male specimen of Falconina sp.
was obtained in the Research Campus of Museu Paraense Emílio Goeldi
(01º27'03.03''S48º26'40.2''W), city of Belém, state of Pará, Brazil. Four males of
Xeropigo n. sp. were collected in the district of Morada do Sol (05º03'56.16"S,
42º46'01.02"W), city of Teresina, state of Piauí, Brazil. After the gonads
extraction, the specimens were preserved in 70% alcohol and those collected at
the Parque Nacional de Ilha Grande were deposited in the collection of Instituto
Butantan, São Paulo, state of São Paulo, Brazil, and those collected in Belém
and Teresina were deposited in the collection of Museu Paraense Emílio Goeldi,
Belém, state of Pará, Brazil.
The gonads were dissected in physiologic solution (7.5 g of NaCl, 2.38 g of
Na2HPO4, 2.72 g of KH2PO4, diluted in 1 litre of distilled water) and submitted to
124
a 0.16% colchicine solution (prepared with physiologic solution) during 2 hours.

The hypotonic treatment consisted in immersion of the gonads in tap water during
20 minutes, and the fixation was performed with Metanol: Acetic Acid solution
(3:1). The microscopy slides containing the preparations were stained with 3%
Giemsa solution. The mitotic and meiotic cells were photographed using a digital
capture system coupled to a light microscope.
Analyses of male diplotene cells of Castianeira sp. showed 12 autosomal
bivalents, with one terminal or interstitial chiasma, and two sex univalents, which
correspond to the X1 and X2 chromosomes that appears always positive
heteropycnotic and close one to another. In some cases is even difficult to
establish the limits between the two sex chromosomes. Two daughter metaphase II
cells presented n=12 and n=14=12+X1X2. In the nucleus with n=14 it is possible
to identify the sex chromosomes due to their positive heteropycnosis. Thus, the
meiotic formulae of Castianeira sp. is 12II+X1X2, indicating a diploid number of
2n♂=26. Based on some incomplete mitotic cells and in the metaphase II it is
possible to define the chromosomal morphology as telocentric.
Spermatocytes I of Corinna gr. nitens revealed 13 autosomal bivalents, with
one terminal or interstitial chiasma, and two sex univalents (13II+X1X2). Earlier
segregation in two autosomal bivalents was recorded in one metaphase I cell. The
observation of one nucleus in transition from metaphase II to anaphase II showed
n=15=13+X1X2. In this cell it is possible to visualize the segregation of the sister
chromatids and recognize the positive heteropycnotic and relatively small sex
chromosomes. A spermatogonial metaphase of C. gr. nitens revealed 2n♂=28,
confirming the meiotic findings above. Oogonial metaphases of C. gr. nitens
showed 2n♀=30, composed by chromosomes that, at least in majority, seems to be
telocentric. The female diploid number confirms the sex chromosome system of
the type X1X2 for the males and X1X1X2X2 for females.
Pachytene and diplotene nuclei of Falconina sp. presented 13 autosomal
bivalents and two positive heteropycnotic sex univalents that always appears side-
by-side and corresponds to the X1 and X2 chromosomes (13II+X1X2). The X
chromosomes are clearly of different sizes. In the case of the autosomal bivalents
in diplotene, only one terminal or interstitial chiasma was registered.
Spermatogonial metaphases showed 2n♂=28 telocentric chromosomes,
confirming the meiotic findings.
Xeropigo n. sp. presented male diplotene cells with 13 autosomal bivalents
and four sex univalents that always appeared close to each other and corresponds
to the X1, X2, X3, and X4 chromosomes. The autosomal bivalents showed only
one terminal or interstitial chiasma and the sex univalents appeared positive
heteropycnotic. The X chromosomes clearly presented different sizes in some
diplotene cells. Thus, the meiotic formulae for Xeropigo n. sp. is
13II+X1X2X3X4. Male mitotic metaphases of this species showed a diploid
complement composed by 30 telocentric chromosomes, that is,
2n♂=30=26+X1X2X3X4.
Initially, it is important to emphasize that these are the first cytogenetical
findings to the genera Corinna and Xeropigo. The other chromosomal records on
corinnids are on three Castianeira species, including that analyzed here, all with
125
2n♂=26; two Falconina species, including that studied in the present research, all
with 2n♂=28, as well as Corinna; one species of Oedignatha, with 2n♂=26; and
three species of Trachelas, with 2n♂=22 or 2n♂=24. Looking at these data, one
can notice that at least until now, the chromosomal data on corinnids are useful
from the cytotaxonomical point of view. Besides, Xeropigo has the highest diploid
number found to the family, 2n♂=30. The genera Corinna and Falconina showed
2n♂=28, while the genera Castianeira and Oedignatha presented 2n♂=26, and the
genus Trachelas is characterized by smaller numbers (2n♂=22 and 2n♂=24). In
relation to the chromosome morphology, the family seems to be extremely
conserved, showing elements exclusively acro/telocentric, which is in accordance
with the observed to other dionychans. The same constancy can be observed
regarding the sex chromosome system that, excepting by the X1X2X3X4
described here for Xeropigo, it is constituted only by the X1 and X2 chromosomes
in all other corinnids, the most common type among all spiders.
It is interesting to notice that the species Xeropigo (2n=30), that at first look
seems to presents a great difference in relation to Corinna and Falconina (2n=28),
actually diverges only in relation to the sex chromosomes, possessing the same 26
autosomes.
The sex chromosome system of the type X1X2X3X4 is extremely rare
among spiders, occurring only in three species belonging to the families
Tetragnathidae (Araneoidea), and Sparassidae, a Dionycha as well as the
corinnids. According with the literature, in Tetragnathidae, the karyotype
2n♂=24=20+X1X2X3X4 could have been arisen from a karyotype with
2n=25♂=22+X1X2X3 that originates from a karyotype with 2n♂=24=22+X1X2.
In the case of Corinnidae, it is possible to suggest that a complement composed by
2n♂=28=26+X1X2, as that found in Corinna and Falconina, could give rise to a
complement constituted by 2n♂=29=26+X1X2X3. This type of karyotype was not
found in Corinnidae up to now; however, its existence is extremely plausible,
considering that this configuration it was actually found in Selenopidae and
Saticidae, two other families of Dionycha. Afterwards, the karyotype composed by
three X chromosomes could give rise to a complement that consists of
2n♂=30=26+X1X2X3X4, as described for Xeropigo. In this way, there is no
change in the number of autosomes, which remains 26 during the karyotype
evolution, and only the number of sex chromosomes increases. According to the
literature, both the differentiation of a X1X2 system into a X1X2X3 system, and
the transformation of a X1X2X3 system into a X1X2X3X4, imply the non-
disjunction or duplication of a X chromosome, with the subsequent homology lost.
126
Comparison of development of book gills

in the horseshoe crab and book lungs in scorpions
Roger D. Farley
Department of Biology, University of California, Riverside, CA, USA,

roger.farley@ucr.edu
In 1872 the first papers were published suggesting that development in

horseshoe crabs resembles arachnids rather than crustaceans, and the book gills
of horseshoe crabs may be homologous with book lungs in arachnids. At the end
of the nineteenth century and in the early twentieth, there was substantial interest
in comparing the development of book gills and book lungs. These authors
agreed that spider and scorpion book lungs originate from an invagination of
epithelium posterior to limb buds in the anterior opisthosoma, and a spiracle
opening is retained at the invagination site. There was, however, substantial
disagreement about how the book lung lamellae are formed from the epithelium.
These early papers consist of interpretive diagrams from histological
sections; the investigations vary in thoroughness and were done on animals from
different taxa. Folds in the invaginated tissue were commonly interpreted as the
beginnings of book lung lamellae (e.g., Purcell 1909). These folds posterior to
the limb buds were thought to resemble the outgrowth folds that Kingsley (1893)
described for book gill development from the posterior surface of branchial
appendages in the opisthosoma of horseshoe crabs.
From their investigation of book lung development in spiders,
Montgomery (1909) and Janeck (1909), concluded that the initial folds behind
the limb buds are transitory and disappear, and the lamellae are formed by
alignment of cells from a cluster derived from the invaginated epithelium. As
described below, lamellar formation by cell alignment was also observed in my
recent study (SEM) of book lung development in scorpion embryos (Farley
2008).
The embryology is still unclear, but differences in lamellar development
do not preclude the possibility of book gill/book lung homology. In a review of
embryology and larval development, Wilmer (1990, p. 128) concluded that
modifications commonly occur for the benefit of the embryo or larva, and such
changes often obscure ancestral conditions. Some similar cell actions occur in
book gill/book lung development (Farley 2008, 2010), and there is evidence that
some of the same genes are expressed at the sites where book gills and book
lungs form (e.g., Damen et al. 2002).
My own investigation (SEM) of book gill development in horseshoe crabs
(Limulus polyphemus L.) are in agreement with Kingsley’s (1893) histological
observation that the lamellae arise as an outgrowth from the posterior surface of
the preceding branchial appendage (Farley 2010). Trabeculae are formed from
cell processes that extend into the appendage and lamellar lumen from the
hypodermis. For the development of gill lamellae, I have not seen the type of
127
cell alignment described for lamellae in the book lungs of spider and scorpion
embryos (Montgomery 1909, Janeck 1909, Farley 2008) .
In scorpion embryos, the mesosomal limb buds disappear (except for the
pectinal ones) well before spiracles and tissue invagination become evident near
the posterior margin of opisthosomal segments 4-7 (Farley 2008). Since there is
no pre-existing structure that can serve as a guide, the lamellae are initially
formed by alignment of cells (single file) in a series of parallel rows. It is
difficult to discern from the diagrams provided by Montgomery (1909) and
Janeck (1909), but there may be differences in the details of cellular formation
of lamellae in spider and scorpion embryos (Farley 2008), although cell
alignment from undifferentiated precursors appears to be a common feature.
I have examined book lung development in representatives from several
scorpion families. Some steps appear to be bypassed in the more derived
families (e.g., Scorpionidae, Liochelidae) as compared with the basal ones (e.g.,
Buthidae, Vaejovidae), but the cellular mode of lamellar formation is similar in
all those studied so far. Cells derived from the invaginated spiracle site align in a
vertical plane and secrete cuticle around themselves (Farley 2008). The secretion
often appears to be holocrine, releasing granules that aggregate spontaneously to
form the lamellar walls. The cells undergo cytolysis while sandwiched within
the cuticular walls they secreted. This leaves the channel open for air except for
bridging trabeculae (cross bridges) that help hold the walls in place. The source
of these initial trabeculae in the second instar is still unknown, either the
degenerating cells in the air channel or the epithelial (tissue) layer that forms on
the cuticle walls in the first and second instar. The epithelial layer produces the
replacement cuticle in the moults of juveniles.
In the book lungs of adult scorpions, the anterior (proximal) edges of the
lamellae (air sacs) have bridging trabeculae that attach to both walls of the air
channel (Kamenz & Prendini 2008, Farley 2008). The posterior (distal) regions
of the lamellae have other types of (non-bridging) trabeculae that are firmly
attached to only one wall of the air channel. This pattern of trabecular
distribution is apparently a result of the growth and moulting process of the
lamellae.
In juvenile scorpions, the bridging trabeculae are formed as part of the
new growth for enlarging the lamellae prior to and during the moult, as bridging
trabeculae are also formed in the air channels when book lungs are initially
formed in embryos and first and second instars (Farley 2008). The non-bridging
trabeculae are produced as part of the replacement cuticle for older, more
posterior regions of the lamellae that are shed in the moult.
References
Damen W.G.M., Saridaki T. & Averof M. 2002. Diverse adaptations of an
ancestral gill: a common evolutionary origin for wings, breathing organs
and spinnerets. Current Biology, 12: 1711-1716.
Farley R.D. 2008. Development of respiratory structures in embryos and first
and second instars of the bark scorpion, Centruroides gracilis (Scorpiones:
Buthidae). Journal of Morphology, 269: 1134-1156.
128
Farley R.D. 2010. Book gill development in embryos and first and second instars
of the horseshoe crab Limulus polyphemus L. (Chelicerta, Xiphosura).
Arthropod Structure and Development (in press).
Janeck R. 1909. Die entwicklung der blättertracheen und der tracheen bei den
Spinnen. Jenaische Zeitschrift für Naturwissenschaft, 44: 587-646.
Kamenz C., Prendini L. 2008. An atlas of book lung fine structure in the order
Scorpiones (Arachnida). Bulletin of the American Museum of Natural
History, 316, pp. 4-45.
Kingsley J.S. 1893. The embryology of Limulus. Part II. Journal of Morphology,
8: 195-270.
Montgomery T.H. Jr. 1909. On the spinnerets, cribellum, colulus, tracheae and
lung books of Araneids. Proceedings of the Academy of Natural Sciences
of Philadelphia, 61: 299-320.
Purcell W.F. 1909. Development and origin of the respiratory organs in Araneae.
Quarterly Journal of Microscopical Science, 54: 1-110.
Willmer P. 1990. Invertebrate Relationships, Patterns in Animal Evolution.
Cambridge, Cambridge University Press, 400 pp.
129
Spiders in apartment buildings of Ukraine cities

Mariia M. Fedoriak1 & Evgeni M. Zhukovets2
1
Department of Ecology and Biomonitoring, Chernivtsi National University, Chernivtsi,
Ukraine, m.m.fedoriak@gmail.com
2
"Stanlyuks" Ltd., Minsk, Belarus, emzhukovets@mail.ru
There is no comprehensive survey of species composition and ecological

peculiarities of many groups of synanthropic organisms, including spiders. Yet
they flourish in parks and squares of all Ukraine cities and also occur in different
types of buildings. Information on the spiders inhabiting buildings was
summarized earlier by Fedoriak & Rudenko (2009).
We inventoried the fauna of synanthropic spiders occurring in various
buildings of Ukraine cities during 2006-2009. The material was collected from
different types of buildings, including industrial enterprises, viz., from attics,
basements, stairways, etc. In the present work, we discuss the samples collected
only from stairways of multi-storey apartment buildings: viz., from walls,
ceilings and window frames. Taking into account that in warm seasons the areas
of the main entrance and ground floor are characterized by higher spider
diversity and density compared to those of top floors (Fedoriak & Brushnivska
2009), the material was mainly collected from the mentioned areas.
The territory of Ukraine is subdivided in 25 administrative units, each with
own centre where we carried out our investigation. The inventoried
administrative centres differ in their populations (from 116.5 thousand people in
Uzhhorod to 2718.1 thousand in Kyiv) and in peculiarities of their industrial
development. Each of the investigated administrative centres represents an
urboecosystem conditioned by an intensity of the passenger/freight
transportation, population density and accessibility of information. According to
the physiographic subdivision given in the National Atlas of Ukraine (2007),
Ukraine’s administrative centers belong to the following zones: the mountain
region – the Ukrainian Carpathians (Uzhhorod, Chernivtsi, Ivano-Frankivsk);
the mixed forest zone (Zhytomyr, Kyiv, Chernihiv); the deciduous forest zone
(Lutsk, Rivne, Lviv, Ternopil, Khmelnytskyi); the forest-steppe zone (Sumy,
Cherkasy, Poltava, Kharkiv, Vinnytsya); the steppe zone (Odessa, Kherson,
Mykolayiv, Zaporizhzhya, Kirovohrad, Luhansk, Donetsk, Dnipropetrovsk); and
the Crimean mountain region (Simferopol). Almost the entire territory of
Ukraine lies in the temperate zone, except for the southern macro-Crimean
mountains, which contain sub-tropical landscapes of the sub-Mediterranean
type.
On the basis of extensive material (15 978 specimens in total) it has been
established that over a hundred of spider species of 26 families inhabit apartment
buildings of Ukraine administrative centers. Of them, 107 were identified to
species level: viz., Agelenidae (11), Amaurobiidae (2), Anyphaenidae (1),
Araneidae (11), Dictynidae (4), Coriniidae (1), Dysderidae (1), Gnaphosidae (2),
Linyphiidae (21), Liocranidae (1), Lycosidae (4), Mimetidae (2), Miturgidae (1),
130
Nestisidae (1), Oecobiidae (4), Philodromidae (2), Pholcidae (6), Pisauridae (1),
Salticidae (4), Segestriidae (2), Scytodidae (1), Tetragnathidae (4), Theridiidae
(18), Thomisidae (2); also only immatures of Clubionidae and Oxyopidae were
collected. The majority of these spiders are likely to be ‘incidental’ synanthropic
species. We have calculated a relative number of each species’ samples within
the assemblages of all the studied administrative centers according to dominance
structure after Stöcker & Bergman, with the dominance classes as follows: 31.7-
100% – eudominant; 10.1-31.6% – dominant; 3.2-10.0% – subdominant; 1.1-
3.1% – recedent; less than 1% – subrecedent (Stöcker & Bergmann, 1977).
Representatives of only 14 species of 8 genera and 5 families are numerous
enough to belong to higher dominance classes (eudominant, dominant and
subdominant) at least in one of the inventoried cities. Such species form
dominant nuclei of the spiders’ assemblages of Ukraine apartment buildings.
In average, the percentage of Pholcidae and Theridiidae was higher than
that of other families, 57% and 32% correspondingly. Ph. phalangioides
(37.7%) is the most abundant species of Ukraine apartment buildings. It is an
obligatory component of synanthropic spider complexes of every administrative
center, but its proportion varies greatly from about 1% in Rivne to 67% in
Simferopol’. In 16 of the 19 explored cities (64%), Ph. phalangioides was the
most abundant species. It is worth mentioning that Ph. alticeps replaced the
former eudominant species in Lutsk, Rivne, Ternopil (the deciduous forest zone)
and Poltava (the forest-steppe zone). Before our investigations, the latter species
had been recorded in Ukraine only from Kherson region and from the Crimea,
but without exact localities (Dunin 1998). Ph. alticeps was described by Spassky
(1932) on the basis of the material taken from the city of Novocherkassk. Later,
this species was recorded from the European part of Russia, the Caucasus and
Tadjikistan, as well as from several localities of Iran and Afghanistan
(Mikhailov 1997; Senglet 1974, 2008). To date, Ph. alticeps has been reported
from all the physiographic zones of Ukraine except for the Crimea (Fedoriak
2007, Fedoriak & Rudenko 2009), as well as from Poland and Belarus (Fedoriak
2008). We assume that the lack of earlier information about Ph. alticeps was due
to misidentifications.
Except for the aforementioned species, a cumulative percentage of
Ph. ponticus, S. castanea, S. triangulosa and P. tepidariorum was higher than
that of other species, but their distribution was uneven and these species were
very numerous only in spider complexes of a few of cities. Species contributing
about 50% of the totally collected spiders are shown in the Fig. 1. Some poorly
known species have been found. For example, ‘Crustulina’ albovittata
(Theridiidae) was found in Simferopol. Earlier, the species was only known
from two specimens collected in buildings in 1860 (Thorell, 1875). So, our
finding is the first one after almost 150 years. This species does not belong to
Crustulina, but its correct belonging remains unclear (Marusik pers. comm.).
Another interesting species is the North American Agelenopsis potteri, which
has been recorded in recent years from the Far East, Kyrgyzstan and the east of
Ukraine. Thus, Uzhgorod is currently the westernmost locality of the species
distribution in the Palearctic Region.
131
Table. A relative number of species (%) forming dominant nuclei of the spiders’
assemblages of Ukraine apartment buildings, in accordance with the physiographic
subdivision given in National Atlas of Ukraine (2007). UK – the mountain region – the
Ukrainian Carpathians; MFZ – the mixed forest zone; DFZ – the deciduous forest zone;
FSZ – the forest-steppe zone; SZ – the steppe zone; CMR – the Crimean Mountain
region.
Taxon UK MFZ DFZ FSZ SZ CMR Total

Agelenidae
Malthonica ferruginea 0.06 5.35 1.08 0.64
Tegenaria domestica 1.00 3.12 2.61 1.35 0.51 1.37
Tegenaria s .l. spp. 2.36 3.42 3.41 2.12 1.52 8.68 2.58
Linyphiidae
Megalepthyphantes
0.40 1.11 1.37 0.83 0.06 0.62
nebulosus
Oecobiidae
Oecobius sp. 0.06 0.86 0.19
Pholcidae
Pholcus alticeps 6.22 8.62 26.79 19.40 1.40 11.29
Ph. phalangioides 52.03 29.64 28.31 24.94 36.24 66.86 37.69
Ph. ponticus 0.46 7.36 6.47 9.67 16.33 7.31
Scytodidae
Scytodes thoracica 0.44 0.22 0.08 0.23 0.86 0.59 0.41
Theridiidae
Parasteatoda tabulata 0.81 1.04 0.44 0.34 0.77 0.64
P. tepidariorum 3.09 2.75 5.58 2.04 2.02 0.79 2.92
Parasteatoda spp. 3.59 2.60 3.05 3.16 1.43 0.79 2.79
Steatoda bipunctata 1.02 0.31
St. castanea 12.92 14.04 11.53 23.08 23.59 5.33 17.02
St. grossa 2.53 4.38 3.29 2.27 2.14 2.17 2.65
St. triangulosa 1.69 8.62 0.48 5.31 7.14 12.43 4.36
Theridion melanurum 0.51 0.11
Other species 11.38 7.73 5.51 5.20 4.62 2.36 7.1
132
Fig. 1. The most abundant species, forming a half of the collected samples in each
administrative centre.
References
Dunin P.M. 1998. Review of the spiders family Pholcidae C.L. Koch, 1851
(Arachnida, Aranei, Haplogynae) of Eastern Europe. In: The Proceedings of
the I Intern. Conf. "Problems of Entomol. of the European part of Russia
and adjoining territories". Samara, pp. 139-144 [in Russian].
Fedoriak M.M. & Rudenko S.S. 2009. On the state of study of spiders (Aranei)
in dwelling and business premises of Ukrainian settlements. Scientific
Papers. Kherson, pp. 383-388 [in Russian].
Fedoriak M.M. 2008. Preliminary data about the distribution of Pholcus alticeps
(Aranei: Pholcidae) in Ukraine and adjoining territories. In: The
Proceedings of the Intern. Conf. "Fundamental aspects of biology in
solving environmental problems". Astrahan, pp. 243-246 [in Russian].
Fedoriak M.M. & Brushnіvska L.V. 2009. Species richness and density of
spiders communities (Araneae) of different floors of buildings. In: The
Proceedings of the II Intern. Conf. "Recent developments in biology,
ecology and chemistry". Zaporіzhzhya, pp. 82-83 [in Ukrainian].
Mikhailov K.G. 1997. Catalogue of the spiders of the territories of the former
Soviet Union (Arachnida, Aranei). – Moscow: Zoological Museum of the
Moscow State University, 416 pp.
National Atlas of Ukraine. 2007. Kyiv, 440 pp. [in Ukrainian].
Senglet A. 1974. Pholcus nouveaux d’Iran (Araneae: Pholcidae). Revue suisse
de Zoologie, 81: 803-812.
133
Senglet A. 2008. New species of Pholcus and Spermophora (Pholcidae,

Araneae) from Iran and Afghanistan, with notes on mating mechanisms.
Revue suisse de Zoologie, 115: 355-376.
Spassky S. 1932. Aranearum species novae II. Bull. Museum National
d’Histoire naturelle, Paris, 4(2): 972-979.
Stöcker G. & Bergmann A. 1977. Ein Modell der Dominanzstruktur und seine
Anwendung. Archiv für Naturschutz und Landschaftforschung, 17(1): 1-26.
134
Can’t you find me? Female sexual response in

Grammostola schulzei (Schmidt 1994),
an Argentinean tarantula
Nelson Ferretti1, Sofia Copperi2, Gabriel Pompozzi2

& Fernando Pérez-Miles3
1
Centro de Estudios Parasitológicos y de Vectores CEPAVE (CCT- CONICET- La
Plata) (UNLP), La Plata, Argentina, nelsonferretti@hotmail.com
2
Departamento de Biología, Bioquímica y Farmacia, Universidad Nacional Del Sur,
Bahía Blanca, Argentina, sofia_copp@hotmail.com,
gabrielpompozzi@hotmail.com
3
Facultad de Ciencias, Sección Entomología, Montevideo, Uruguay,
myga@fcien.edu.uy
Introduction
Despite wide distributed in tropical and subtropical regions of the world,
the biology of Theraphosidae is poorly known. Studies of mating behaviour are
limited to few species.
Vibratory sexual signals (acoustic and seismic) are frequently used in
spider courtship, especially in tarantulas (Uetz & Stratton 1982, Prentice 1992,
Quirici & Costa 2005, 2007). Seismic signals generated by male spiders can
reach 1 m in ctenids (Barth et al. 1988) and 1.30 m in theraphosids (Quirici &
Costa 2007). The usual indicator of effective male signals is the behavioural
change of the receptive female. The most unambiguous receptive response
usually is leg waving, leg tapping or body vibrations (Ferretti & Ferrero 2008,
Prentice 1992, Quirici & Costa 2005, 2007). Grammostola schulzei (Schmidt
1994) is a tarantula from Argentina that lives in burrows under stones in rocky
hills (Ferretti & Ferrero 2008). G. schulzei is the unique species of the genus
where females showed sexual response to courting males, but the context and
possible function of this response was not elucidated (Ferretti & Ferrero 2008).
In captivity, female sexual response could be observed allowing females to build
and occupy a retreat as in natural conditions but not in open arena (Bertani et al.
2008). The main objective of this work is to elucidate under what context
females responded to male’s courtships and to determine the possible function of
these behaviours.

Eight males and eleven females were collected in Sierra de la Ventana
(38º07'63"S, 61º47'30"W), Buenos Aires, Argentina during October - January
2008-2009. For all experiments, females of known reproductive history were
used. They were housed in glass jars of 13 cm diameter and 15 cm height, with 5
cm of soil as substrate and water provision. They were fed ad libitum with
Zophobas sp. larvae. We carried out three series of 20 experiments each with
135
different situations for the encounters male-female. In series “A”, males were
placed free in terrarium but with no access to female due to the heavy stone
covering the burrow. In series “B”, males were placed far off from the female
burrow (30 cm) and confined into a glass cup. In series “C”, males were placed
over the heavy stone that closed the burrow of female and confined into a glass
cup. The courtship of males was elicited by abundant silk of females inside the
glass cup. For the encounters we used glass terraria measuring 30 x 35 and 30
cm high, containing a 10 cm thick layer of soil as substrate. In each terrarium,
a burrow was constructed against the glass wall, allowing visual observations
of female behaviour inside burrow. Each terrarium was placed in different
tables in order to minimize ground vibrations and to prevent seismic signals
from passing between terraria. Females were placed in terraria and trials were
carried out for >5 days, allowing each female to adapt to the burrow. The
entrance of each burrow was closed with a heavy stone (collected from their
natural habitat) so females couldn’t leave the burrow and had no access to
males. We considered active each male which showed intense courtship of >30
min. The encounters were videotaped with a Handycam Panasonic SDR-S7
and video records were analyzed with a PC program.
Results
We observed two types of responses of females to male courtships: leg
tapping and body movements. In series A, three different females made their
sexual display tapping vigorously with palps and first pair of legs or only with
palps. In video analyses, females made leg flexing, lifting and lowering against
the substrate. The mean number of leg movements during leg tapping was
16.415.26 SD and the mean number of bouts was 20.512 SD. The mean
duration of leg tapping was 1.63s0.90 SD and the mean duration between
bouts of female leg tapping was 48.20s37.49 SD. All responses were after the
palpal drumming of males and long courtships (approximately 1 hour).
Moreover, females that responded with leg tapping scraped vigorously the
stone with first pair of legs. One male obtained responses from two different
females. After female tapping, males frequently oriented to the female burrow
and stayed courting nears her. Females responded with body movement in
series A and C, where males were confined. These displays consisted in high
frequency downward and upward movement of the entire body. In video
analysis, we observed a contraction of the third pair of legs during movements.
These responses were observed after male palpal drumming and the number of
the movements was highly variable, ranging from 2 to 61 (mean=16.6  24.96
SD). One female made body movements with two different males in series A
and C. Two more females performed this behaviour in series C and even one
of them made body movements with two different males. During series C one
female that performed body movements made three attempts of abrupt
emergence inside the burrow and the male quickly retreated.
136
Discussion
Our results suggest the presence of seismic communication by substrate
during courtship in G. schulzei in agreement with findings of Quirici & Costa
(2005) in other theraphosids. Female leg tapping as response to some male
courtship seems to indicate her receptive condition and his attractiveness, as
was found for Acanthoscurria suina and Eupalaestrus weijenberghi (Pérez-
Miles et al. 2007, Quirici & Costa 2005). However, female leg tapping
response was only observed in series A in which males freely walked in the
terrarium and after extremely long courtships. We suggested that additionally
female response could orient male towards her location. This orientation could
serve at long distance because when males were very near the burrow, females
did not respond (series C). Considering that female response is obtained after
an insistent long male courtship, we interpret that such response could act in
extreme cases where males could not find her burrow or if is difficult for the
female to exit from the burrow. Also insistence of males could be a positive
honest signal of males with good condition to mate, and consequently selected
by females. These females responded with leg tapping to males which present
long courtships together with active searching (walking) for female’s location
(series A). This was not the case in series B and C with males confined.
Although these males confined courted, they did not receive female tapping
response. Summarizing females respond with leg tapping only to males which
court and walk. The female body movements are interpreted here as a rejection
considering male retreated after this female behaviour in series C. Body
movements seems to be different from the piston behaviour observed for E.
weijenberghi (Pérez-Miles et al. 2007), because in G. schulzei this behaviour
consisted in downward and upward body movements, have higher frequency,
and are probably originated by third pair of legs. Piston behaviour was
proposed as a ritualized rejection response, considering that after that male
immediately escaped (Pérez-Miles et al. 2007). Female body movements are a
signal that could help the male to save time and energy spent in courtship and
also to reduce the risk of attracting predators. For the female this rejection of
male contributes avoiding her distraction from other biological activities
(Pérez-Miles et al. 2007). We observed body movements of females
predominated in series C, where males were confined in the nearest possible
location to female burrow.
References
Barth F.G., Bleckmann H., Bohnenberger J. & Seyfarth E.A. 1988. Spiders of
the genus Cupiennius Simon 1891 (Araneae, Ctenidae). II. On the
vibratory environment of a wandering spider. Oecologia, 77:194-201.
Bertani R., Fukushima C.S., & Silva Júnior P.I. 2008. Mating behaviour of
Sickius longibulbi (Araneae, Theraphosidae, Ischnocolinae), a spider that
lacks spermathecae. Journal of Arachnology, 36: 331-335.
Ferretti N. & Ferrero A. 2008. Short Communication: Courtship and mating
behavior of Grammostola schulzei (Schmidt 1994) a burrowing tarantula
from Argentina. Journal of Arachnology, 36: 480-483.
137
Pérez-Miles F., Postiglioni R., Montes de Oca L., Baruffaldi L. & Costa F.G.
2007. Mating system in the tarantula spider Eupalaestrus weijenberghi
(Thorell, 1894): Evidences of monandry and polygyny. Zoology, 110:
253-260.
Prentice T.R. 1992. A new species of North American tarantula, Aphonopelma
paloma (Araneae, Mygalomorphae, Theraphosidae). Journal of
Arachnology, 20: 189-199.
Quirici V. & Costa F.G. 2005. Seismic communication during courtship in two
burrowing tarantula spiders: an experimental study on Eupalaestrus
weijenberghi and Acnathoscurria suina. Journal of Arachnology, 33: 159-166.
Quirici V. & Costa F.G. 2007. Seismic sexual signal design of two sympatric
burrowing tarantula spiders from meadows of Uruguay: Eupalaestrus
weijenberghi and Acanthoscurria suina (Araneae, Theraphosidae). Journal
of Arachnology, 35: 38-45.
Uetz G.W. & Stratton G.E. 1982. Acoustic communication and reproductive
isolation in spiders. In: Witt P.N. & Rovner J.S. (eds), Spider
Communication: Mechanisms and Ecological Significance. Princeton
University Press, Princeton, New Jersey, pp. 123-159.
138
First notes on reproductive biology, burrows and egg sacs

of the tiny mygalomorph spider Xenonemesia platensis
(Araneae: Microstigmatidae)
Nelson Ferretti1, Gabriel Pompozzi2, Sofia Copperi2,

Fernando Pérez-Miles3 & Alda González1
1
Centro de Estudios Parasitológicos y de Vectores CEPAVE (CCT- CONICET- La
Plata) (UNLP), La Plata, Argentina, nelsonferretti@hotmail.com,
asgonzalez@cepave.edu.ar
2
Departamento de Biología, Bioquímica y Farmacia, Universidad Nacional Del Sur,
Bahía Blanca, Argentina, gabrielpompozzi@hotmail.com,
sofia_copp@hotmail.com
3
Facultad de Ciencias, Sección Entomología, Montevideo, Uruguay,
myga@fcien.edu.uy
Introduction
The Microstigmatidae are characterized by the rounded book-lungs
openings, in conjunction with extremely shortened posterior lateral spinnerets
(Goloboff 1995). This family comprises 15 species, nine of them distributed in
the New World (Platnick 2010). Microstigmatidae are ground-dwelling and free-
living spiders that appear to make minimal use of silk and could readily attack
and fed upon small insects (Griswold 1985). Some studies were done on
reproductive biology of Mygalomorphae (Coyle 1985, Coyle & O´Shields 1990,
Costa & Pérez-Miles 2002, Ferretti & Ferrero 2008, Yañez et al. 1999) but
sexual behaviour of Microstigmatidae is up to now unknown. Few records of
natural history and ecology were reported for the Old World microstigmatids
(Griswold 1985) and for a Brazilian species (Indicatti et al. 2008). Xenonemesia
platensis is the only microstigmatid species present in Argentina and only some
comments on their habitat have been provided (Goloboff 1988). In the present
study we describe the mating behaviour of X. platensis based on three successful
matings under laboratory conditions and give some notes on their burrows and
egg sacs.

Two males and six females were collected during August 2009 on the
Martín García Island (34º11'25"S, 58º15'38"W). The individuals were
maintained in plastic Petri dishes (9 cm diameter), with soil as substratum and a
wet cotton wool. All individuals were fed weekly with cockroaches (Blattela
germanica). We carried on five sexual encounters. The female dish was placed
in the centre of a larger glass cylindrical container (19 cm diameter and 10 cm
high) with a layer of soil of approximately 6 cm. Then, the male was gently
placed far off from the female position. The room temperature during the
experiments was 26.7°±1.52°C. The encounters were videotaped and analyzed
with a PC program (Sony Vegas 9.0).
139
Results
Males initiated courtship when contacted with the female body. Females
remain passive. The courtship started with body vibrations originated by
contractions of the first and second pair of legs followed by fast upward and
downward movements of palps in a simultaneous phase. The frequency was
33.5±3.8 SD bouts/min and the mean duration was 0.54s±0.06 SD (n=25). Also
males made palpal boxing that consisted in alternating movements of the pedipalp
touching the genital zone of the female. The males performed 6.66±4.04 SD
bouts/min with a mean duration of 1.92s±0.95 SD (n=6). After contact, males
vigorously hit the first and second pair of legs of females with his second pair of legs
extended. This behaviour consisted in tapping at high frequency in an alternating or
synchronous phase; male tarsi contacted with the metatarsi of female. Males
performed 6±1.73 SD bouts/min with a mean duration of 0.96s±0.38 SD (n=7).
After tapping, the males clasped with first pair of legs between palps and chelicerae
of the female. The distal portion of each male tibia without tibial apophyses or
megaspines was against the prolateral surface of each female pedipalp base and then
begins palpal insertion attempts. The mean number of insertions was 8±6.2 SD and
their mean duration was 17.61s±18.11 SD (range =3-37.88). During insertions,
males continued courting the female by tapping with legs II and body vibrations.
Males then added a new courtship behaviour; they raised the second pair of legs
with an angle of 90° between femur and patella, and quickly moved legs upward and
downward. Male tibia, metatarsi and tarsi remain extended and the tarsi beat and
scrape the second and third coxae of female. This behaviour had a mean duration of
8.62s±5.85 SD and the velocity was 14 beats per second. During the palpal insertion
attempts the second pair of legs was pushing the first pair of legs of female and the
female’s pedicel was flexed upwards so that the cephalothorax-abdomen angle was
30-50°. The mean duration of copulation was 4.04 min ±1.13 SD. After copulation
females retreated into the burrows. In one case, the male followed courting her in
contact making body vibrations. We observed one female rejection that consisted in
vigorous lateral abdominal oscillations with body elevated. After that male escaped.
All individuals were found in a shore forest occupying short burrows with
little silk covering the walls and the entrance closed with silk and debris. In nature,
the silk tube entrance ranged from 0.8 to 9.5 mm diameter and its length 16.15 mm
±7.7 SD; the short burrow measured approximately 12 mm. The temperature inside
the burrows was 15°C and the environment temperature was 14°C. In laboratory
they constructed the same shelters as in nature but with more silk covering the walls.
Two egg-clutches were observed in the laboratory. One female made the egg sac on
23 November 2009 and the other on 16 December 2009. The eggs were
agglomerated in a sphere without a well developed silk cover. They measured 5 mm
width x 8.6 mm length containing 17 eggs and 6.7 mm width x 8.4 mm length. Both
females abandoned their egg sacs between 7 January 2010 and 25 January 2010.
Discussion
As far as we know, these are the first reports on sexual behaviour of a
Microstigmatidae. Some uncommon behaviours are remarkable. Differing from
140
most known mygalomorphs the male did not started courtship when contact
female silk but only after contact with female. Early studies proposed that
mygalomorph spiders lacked chemical cues in sexual communication (Baerg
1958, Platnick 1971); however, more recent studies reported the presence of
pheromone associated with silk threads of females (Costa & Pérez-Miles 2002,
Ferretti & Ferrero 2008). Our findings in X. platensis could reveal the absence of
pheromone associated to silk.
The body vibrations observed in the courtship of X. platensis could be
similar to those observed in some theraphosids (Costa & Pérez-Miles 2002,
Ferretti & Ferrero 2008) but in X. platensis the vibration is generated by first and
second pair of legs instead of pair III involved in theraphosids. Others
remarkable behaviours in X. platensis were the brusque movements of the palp
and the scraping with legs II during courtship that were not reported in any other
mygalomorph spider. The brusque movements of palps could be similar to the
“twitching” observed in a diplurid (Coyle & O´Shields 1990) which consisted in
separate sudden flexions or extensions of one or more legs or palps. The male
courtship in copula observed in X. platensis was not reported in mygalomorphs
(Costa & Pérez-Miles 1998, Costa & Pérez-Miles 2002, Ferretti & Ferrero 2008,
Jackson & Pollard 1990). It is possible that the female may be testing male’s
copulatory ability, monitoring his performance not only in genital stimulation, as
suggested by Eberhard (1985). This male behaviour could additionally inform
the female about his good quality and consequently could be interpretable as
other mechanism for seduction.
References
Baerg W.J. 1958. The Tarantula. University of Kansas Press, Lawerence,
Kansas, 88 pp.
Costa F.G. & Pérez-Miles F. 1998. Behavior, life cycle and webs of
Mecicobothrium thorelli (Araneae, Mygalomorphae, Mecicobothriidae).
Journal of Arachnology, 26: 317-329.
Costa F.G. & Pérez-Miles F. 2002. Reproductive biology of Uruguayan
theraphosids (Araneae, Theraphosidae). Journal of Arachnology, 30: 571-587.
Coyle F.A. 1985. Observations on the mating behaviour of the tiny
mygalomorph spider, Microhexura montivaga Crosby & Bishop (Araneae,
Dipluridae). Bulletin of the British Arachnological Society, 6(8): 328-330.
Coyle F.A. & O’Shields T.C. 1990. Courtship and mating behavior of
Telochoris karschi (Araneae, Dipluridae), an African funnel web spider.
Eberhard W.E . 1985. Sexual selection and animal genitalia. Harvard Univ.
Press, Harvard, Massachusetts.
Ferretti N. & Ferrero A. 2008. Short Communication: Courtship and mating
behavior of Grammostola schulzei (Schmidt 1994) a burrowing tarantula
from Argentina. Journal of Arachnology, 36: 480-483.
141
Goloboff P.A. 1988. Xenonemesia, un nuevo genero de Nemesiidae (Araneae,

Mygalomorphae). Journal of Arachnology, 16: 357-363.
Goloboff P.A. 1995. A revision of the South American spiders of the family
Nemesiidae (Araneae, Mygalomorphae). Part I: species from Peru, Chile,
Argentina, and Uruguay. Bulletin of the American Museum of Natural
History, 224: 1-189.
Griswold C.E. 1985. A revision of the African spiders of the family
Microstigmatidae (Araneae: Mygalomorphae). Annals of the Natal
Museum, 27(1): 1-37.
Indicatti R.P., Lucas S.M., Ott R. & Brescovit A.D. 2008. Litter dwelling
mygalomorph spiders (Araneae: Microstigmatidae, Nemesiidae) from
Araucaria Forests in Southern Brazil, with the description of five new
species. Revista Brasileira de Zoologia, 25(3): 529-546.
Platnick N. 1971. The evolution of courtship behavior in spiders. Bulletin of the
http://research.amnh.org/entomology/spiders/catalog81-87/index.html.
Yañez M., Locht A. & Macías-Ordóñez R. 1999. Courtship and mating behavior
of Brachypelma klassi (Araneae, Theraphosidae). Journal of Arachnology,
27: 165-170.
142
DNA barcoding of the Balkan scorpions: preliminary results
Victor Fet1, Gergin A. Blagoev2, Natalia Ivanova3

& Marjan Komnenov4
1
Department of Biological Sciences, Marshall University, WV, USA, fet@marshall.edu
2
gblagoev@uoguelph.ca
3
nivanova@uoguelph.ca
4
Macedonian Ecological Society, Institute of Biology, Faculty of Natural Sciences &
Mathematics, Skopje, Republic of Macedonia, mkomnenov@yahoo.com
Our understanding of traditional scorpion taxa at all levels is changing

rapidly and considerably due to introduction of modern methods of analysis
(both morphological and molecular), discovery of new important characters,
intensive new collection, and study of old museum material. The genus
Euscorpius Thorell, 1876 (Euscorpiidae) has been problematic for decades
(Hadži 1930, Caporiacco 1950) but DNA markers show that this genus indeed
contained a not-so-hidden diversity. “DNA barcoding” (using mitochondrial
cytochrome oxidase I gene; Hebert et al. 2003) emerges as a fast and universal
DNA marker technique at species level; this method is now gaining momentum
in applied systematics and species identification of Arachnida. Our pilot survey
with optimized PCR and sequencing primers and automated barcoding protocol
included ca. 50 populations of Euscorpius sequenced, mainly from Bulgaria and
Macedonia, and also from Serbia, Montenegro, Croatia, Slovenia, and Romania.
The preliminary show the correct species-level recovery of several well-defined
species such as E. italicus and E. tergestinus; other, less understood groups such
as E. hadzii (Fet & Soleglad, 2002) likely include several species. At least three
Euscorpius species live in Bulgaria. Further studies are being conducted on
Greek fauna. The Western Balkans have relict and diverse populations of
subgenus Alpiscorpius (= “mingrelicus” complex), absent from most of
mainland Greece and Aegean islands. Introduction by humans likely plays role
for many Euscorpius (Fet et al. 2006); DNA markers show that E. tergestinus in
Austria was introduced from Slovenia. A picture that emerges of scorpions in the
Balkans is that of a diverse and rich group, with various trends of speciation and
endemism. We estimate that at least 15-20 species of Euscorpius inhabit the
Balkan Peninsula, often sympatrically. The collaborative international work is
under way to obtain a detailed molecular and morphological phylogeny of these
species and their populations.
143
Testing the revised gravity hypothesis for body size

evolution in a highly dimorphic orb-web spider:
larger dwarfs climb faster
Matthias W. Foellmer1, Michael Marson2 & Jordi Moya-Laraño3

1
Department of Biology, Adelphi University, New York, NY, USA, Foellmer@adelphi.edu
2
Department of Biology, Trent University, Peterborough, Ontario, Canada
3
Cantabrian Institute of Biodiversity, Departamento de Biología de Organismos y
Sistemas, Universidad de Oviedo, Oviedo, Asturias, Spain
The adaptive significance of sexual size dimorphism (SSD) in spiders is of

great interest for evolutionary biologists, because spiders are the terrestrial
animal taxon that exhibits the greatest range and the most extreme cases of SSD.
The gravity hypothesis for the evolution of extreme SSD has been particularly
controversial, and has recently been revised. The revised gravity hypothesis
(rGH) predicts a curvilinear pattern for the relationship between body size and
achievable climbing speed with an optimal climbing speed for spiders with a
body mass of about 43 mg. Below 43 mg spiders are predicted to show a
positive relationship between body size and climbing speed, above 43 mg this
relationship is predicted to become negative. Here we test the rGH using the
highly dimorphic orb-weaver Argiope aurantia by inducing males to run on
dowels suspended at five different angles (0°, 22.5°, 45°, 67.5°, 90°). In A.
aurantia, males fall well below the 43 mg threshold. As predicted, larger males
were faster climbers, and ran faster at all other angles as well. Further, the size
effect was largely due to bigger males having longer legs and thus being capable
of wider strides. This confirms a previous field study estimating selection in this
species and means that the period of mate search likely selects for large size in
this species with relatively small males, due to the superior locomotory
performance of larger males. These results are not in agreement with recent
studies on other orb-weavers. We discuss possible reasons for the discrepancies
as well as the broader evolutionary implications.
144
Towards a standardized and optimized protocol for rapid

biodiversity assessments: spider species richness and
assemblage composition in two savanna vegetation types
Stefan H. Foord1, Mulalo I. Muelelwa1,

Ansie S. Dippenaar-Schoeman2 & Edward M. Stam3
1
Department of Zoology, University of Venda, Thohoyandou, South Africa
2
ARC-Plant Protection Research Institute, Queenswood and Department of Entomology
and Zoology, University of Pretoria, South Africa
3
National Zoological Gardens of South Africa, Pretoria and the Mammal Research
Institute, University of Pretoria, South Africa
A semi-quantitative inventory of spider diversity was done in the Blouberg

Nature Reserve (BNR) and Western Soutpansberg Conservancy (WSC) situated
in the Savanna Biome of the Limpopo Province, South Africa. Two hundred
and ninety six samples of one person-hour work each, comprised of five
methods (vegetation beating, sweep netting, aerial hand collecting, ground hand
collecting and leaf litter sifting) were divided between four relatively
homogenous sites (plant communities) within a vegetation type of the BNR and
WSC, respectively, an average of 37 per site. In addition, 25 pitfall traps were
left open for a total of 20 days in each of the eight plant communities, 200 in
total. We collected 1328 adult spiders representing 186 species of which 31%
were singletons in the BNR vegetation type and 909 spiders in 222 species of
which 41% were singletons in the WSC vegetation type. The number of species
present was estimated using six estimators. The estimates varied between 233 –
307 for the BNR and 302 – 386 for the WSC. Inventory completeness was more
than 70%. The fit to a lognormal distribution suggests that there are 370 species
(750 000 individuals) and 445 (850 000 individuals) species in the universes (16
ha) sampled within the two vegetation types. Collector experience had no effect
on the results of the inventory, whereas time of day had a very small yet
significant effect. Seasonality only affected abundance and richness, but not
assemblage composition. Sampling methods used had the biggest effect on our
results. These results are used to design an optimized sampling protocol for
standardized inventories in the Savanna Biome.
145
The cytogenetic approach reveals speciation events

in social spiders Stegodyphus (Araneae: Eresidae) 2
Martin Forman1, Jiří Král1 & Charles R. Haddad2
1
Faculty of Science, Charles University, Prague, Czech Republic
2
Department of Zoology and Entomology, University of the Free State, Bloemfontein,
South Africa, formivelkejpan@seznam.cz
Sociality in spiders is a rare phenomenon reported in about 20 species

from seven families. Amongst these, the genus Stegodyphus is remarkable by
three independent origins of permanent sociality from subsocial ancestors (Kraus
and Kraus 1988, Johanessen et al. 2007). Hence, we compare our results with
published data on Stegodyphus karyotypes. In general, karyotypes of most
entelegyne lineages are conservative, showing similar diploid counts and
acrocentric morphology of all chromosome pairs. A common trend in the
evolution in entelegynes is the reduction of diploid numbers. Downsizing of 2n
is most likely achieved through tandem fusions. The evolution of social
behaviour in Stegodyphus is accompanied by an acceleration of chromosome
changes, which is reflected in the unusually high range of diploid counts. In all
eight species so far analyzed, in our study or published studies, the
chromosomes show acrocentric morphology. The ancestral male karyotype of
Stegodyphus consists of 30 chromosomes. Consistent with most entelegyne
spiders studied so far, Stegodyphus displays an X1X20 sex chromosome system
(resulting in X1X2 in males and X1X1X2X2 in females). This chromosome
constitution was found in most subsocial Stegodyphus species, namely S.
africanus, S. dufouri (Forman et al. 2007) and S. pacificus (Sharma and Singh
1957). Interestingly, two very different karyotypes have been found in two
species of social eresids. Populations of S. sarasinorum from central and south
India possess a 2n♂=30, X1X20 (Bole-Gowda 1958, Sharma & Parida 1987,
current study). In contrast to this, Mittal (1970) reported specimens with a
reduced chromosome number (2n♂=24, X1X20) from Punjab. We found an
analogical situation in African S. mimosarum. While the population from South
Africa (KwaZulu-Natal) exhibits the typical ancestral Stegodyphus karyotype of
2n♂=30, X1X20, males from Tanzania (Dar es Salaam) have only 24
chromosomes, including sex chromosomes X1 and X2. Hybridisation of such
different forms will not produce the fertile offspring. Therefore, cytogenetic data
suggest that the social species S. mimosarum and S. sarasinorum represent
complexes of at least two species. The presence of speciation events in social
eresids contradicts the traditional view that considers sociality in spiders to be an
2
The research was supported by Grant Agency of the Czech Academy of Sciences
(project no. IAA601110808).
146
evolutionary dead end. On the other hand, these findings are in agreement with
recent findings about the evolutionary stability of sociality and the high genetic
divergence in social Stegodyphus species (Johanessen et al. 2007, Johannessen et
al. 2009). In contrast to other social species of the genus, no karyotype
differences have been found in populations of S. dumicola from Namibia and
South Africa. Males of all populations display 26 chromosomes including an
X1X20 system (Avilés et al. 1999; current study). Interestingly, we found the
same diploid count in a related subsocial species, S. tentoriicola. Finally,
subsocial S. lineatus differs remarkably in its karyotype (2n♂=43, X1X2X30)
from other Stegodyphus species particularly the additional male sex chromosome
and diploid number. The chromosome constitution of S. lineatus is similar to
that which we found in three burrowing South African eresids (Gandanameno
sp. 2n♂=38, X1X20, Paradonea sp. 2n♀=46, and Dresserus kannemeyeri
2n♂=40, X1X20). Thus, its placement in Stegodyphus should be revised. Based
on our data, we speculate about the basal position of S. lineatus in the family
Eresidae.
References
Avilés L., Varas C., Dyreson E. 1999. Does the African social spider
Stegodyphus dumicola control the sex of individual offspring?
Behavioural Ecology and Sociobiology, 46: 237-243.
Bole-Gowda B.N. 1958. A study of the chromosomes during meiosis in twenty-
two species of Indian spiders. Proceedings of the Zoological Society of
Bengal, 11: 69-108.
Forman M., Král J., Musilová J., Lubin Y. 2007. Karyotype study of social
species of the spider genus Stegodyphus Simon, 1873 (Araneae: Eresidae).
In: Rocha R.P., (ed), 17th International Congress of Arachnology; São
Pedro, São Paulo, Brasil, p. 244.
Johannesen J., Lubin Y., Smith D.R., Bilde T., Schneider J.M. 2007. The age
and evolution of sociality in Stegodyphus spiders: a molecular
phylogenetic perspective. Proceedings of the Royal Society, Biological
Sciences, 274: 231-237.
Johannesen J., Wickler W., Seibt U., Moritz R. 2009. Population history in
social spiders repeated: colony structure and lineage evolution in
Stegodyphus mimosarum (Eresidae) Molecular Ecology, 18: 2812-2818.
Kraus O., Kraus M. 1988. The genus Stegodyphus (Arachnida: Araneae). Sibling
species, species groups and parallel origin of social living. Verhandlungen
des naturwissenschaftlichen Vereins Hamburg, 30: 151-254.
Mittal O.P. 1970. Karyological studies on the Indian spiders. IX. Chromosome
constitution in two cribellate species. Genetica, 41: 575-580.
Sharma G. P., Singh S. 1957. Cytological studies on the Indian spiders I.
Chromosome complement and male meiosis in Stegodyphus specificus.
Research Bulletin of Punjab University, 8: 389-393.
Sharma N., Parida B.B. 1987. Study of chromosomes in spiders from Orissa.
Pranikee, 8: 71-76.
147
Antimicrobial activity of scent gland exudates in

Cyphophthalmus duricorius
(Opiliones: Cyphophthalmi: Sironidae)
Petra Föttinger1,2, Günter Fauler3, Helmut Bergler4,
Günther Koraimann4, Hans-Jörg Leis2 & Günther Raspotnig1,2
1
Institute of Zoology, Karl-Franzens-University Graz, Austria, petra.foettinger@uni-
graz.at
2
Research Unit of Osteology and Analytical Mass Spectrometry, Medical University
Graz, Austria, hans.leis@meduni-graz.at, guenther.raspotnig@uni-graz.at
3
Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University
Graz, Austria, guenter.fauler@meduni-graz.at
4
Institute of Molecular Biosciences, Karl-Franzens-University Graz, Austria,
helmut.bergler@uni-graz.at, guenther.koraimann@uni-graz.at,
Introduction
Like all other harvestmen, Cyphophthalmus duricorius possesses a
prosomal exocrine gland system, the so-called defence or scent glands which
emit repellent exudates against predators. In case of being mechanically irritated,
the harvestman discharges large, brownish droplets of secretion from the gland
openings. Subsequently, the opilionid dips its legs II into the secretion and
transfers it to the offender, a behaviour referred to as ‘leg dabbing’ (Juberthie
1961).
Chemical analysis of scent gland secretion of C. duricorius exhibited a
bouquet of at least 24 components belonging to the two chemical classes of
methylketones and naphthoquinones (Raspotnig et al. 2005). Some of the
constituents, especially chloronaphthoquinones, proved to be unique among
exocrine exudates of arthropods. Naphthoquinones and their numerous
derivatives have already been known as biologically active agents since the late
1940ies (Hoffmann-Ostenhof et al. 1947, Silver & Holmes 1967, Ambrogi et al.
1970). The first opilionid scent gland secretion experimentally applied to
different strains of bacteria and to protozoa in order to elucidate its antimicrobial
potential was that of the South American laniatorean Acanthopachylus
aculeatus. Estable et al. (1955) found that the secretion’s alkylated
benzoquinones were highly effective against gram+and gram- bacteria as well as
against different murine intestinal parasites.
In the study at hand, we investigated 1) the antimicrobial potency of the
Cyphophthalmus-secretion against various microorganisms, and 2) the capability
of C. duricorius to spontaneously release secretion as preventive measure
against microorganisms.

The antimicrobial activity of both whole secretion and single synthetic
secretion constituents was studied qualitatively by Kirby-Bauer tests (zone of
148
inhibition tests) and quantitatively by the fold broth dilution method determining
the minimum inhibitory concentration (MIC).
We tested the main compounds 1,4-naphthoquinone, 2-undecanone, and 2-
tridecanone which were purchasable at Sigma Aldrich. 4-chloro-1,2-
naphthoquinone was synthesised according to the protocol of Perumal & Bhatt
1980 and purified to a degree of 95%. Absolute ethanol served as a blank in the
antibiotic assays.
The test strains Escherichia coli J5 (Prof. Leisinger, Zurich),
Staphylococcus aureus ATTC 29067, Klebsiella pneumoniae DSM 681
Salmonella typhimurium TR 5655, and the yeast Saccharomyces cerevisiae BY
4741 EUROCAT were provided by the Institute of Molecular Biosciences Graz,
Austria. Furthermore, five bacterial strains were isolated from a soil sample,
taken from one of the habitats of C. duricorius. These strains were identified by
16S rRNA-sequencing as Acinetobacter sp., Bacillus cereus,
Enterobacter/Citrobacter sp., Microbacterium arborescens, and Staphylococcus
carnosus.
In order to study a possible stimulation of secretion release triggered by
the respective microorganisms, individuals of C. duricorius were placed on
inoculated agar plates (and pure agar plates as a reference) and were then
monitored hourly.
Results
Kirby-Bauer tests using substance concentrations far above the
physiological level showed that all of the gram-negative bacterial strains were
unsusceptible to the whole secretion and the two methylketones tested. In
contrast, both 1,4-naphthoquinone and 4-chloro-1,2-naphthoquinone lead to
distinct zones of inhibition. Quantitative tests were performed with gram-
positive bacteria and yeast: MIC of whole secretion constantly amounted for 2
mg/ml for bacteria, and 1 mg/ml for yeast. These concentrations correspond to
the total secretion volume of ten individuals of C. duricorius. MIC of
undecanone was 16 mg/ml for all bacteria and 8 mg/ml for yeast. MICs of the
two naphthoquinones ranged from 0,3 (yeast) to 1,2 mg/ml (St. aureus/St.
carnosus). However, none of the gram-positive bacterial strains or yeast reacted
to tridecanone. Ethanol blanks did not lead to growth inhibition.
Preliminary studies revealed that - following mechanical irritation - ‘leg
dabbing’ and distribution of secretion over the body surface of C. duricorius
were simultaneous events. Also exposure to bacteria and yeast resulted in
permanent coverage with a liquid film, indicating constant emission of secretion.
Discussion
Naphthoquinones are well known for their antimicrobial activity,
especially if nucleus and/or side chains are halogenated (Ambrogi et al. 1970).
They act as bacterial growth inhibitors by functioning competitively in electron
transport with endogenous ubiquinone (Silver & Holmes 1968). Thus, it was not
surprising that naphthoquinones play a major role in the antibiotic activity of the
gland secretion of C. duricorius. Contrary to our expectations, also
149
methylketones inhibit growth of gram-positive bacteria and yeast which has

never been reported so far. It would be of high interest, whether also other
ketone constituents, particularly unsaturated ones, exhibit biological activity.
Usually, development of a secretion film over the harvestman’s body
corresponds to irritation by an offender, probably in order to maintain the
deterrent effect of the volatile exudates. First results of the bioassays suggest that
the opilionid also constantly releases secretion, if confronted with
microorganisms. Since C. duricorius typically occurs in the litter and soil of
mixed forests and therefore inhibits a microorgansim-rich environment, a
secretion film might serve a primary protective barrier against bacteria and
fungi.
References
Ambrogi V., Artini D., de Carneri I., Castellino S., Dradi E., Logemann W.,
Meinardi G., Di Somma M. & Tosolini G. 1970. Studies on the
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150
Salt lake cities underground – systematics and ecology

of the Australian wolf spider genus Tetralycosa
(Araneae: Lycosidae)
Volker W. Framenau1 & Peter Hudson2

1
Western Australian Museum, volker.framenau@museum.wa.gov.au
2
South Australian Museum, Peter.Hudson@samuseum.sa.gov.au
Australia is arguably the driest continent on earth and its arid interior is
characterised by ephemeral salt lakes with a very specialized fauna, including
wolf spiders of the genus Tetralycosa (subfamily Artoriinae). The genus
contains a total of 13 species and a phylogenetic analysis identified a
monophyletic clade of eight species that live permanently on the surface of salt
lakes suggesting that this habitat of extreme environmental conditions was
invaded only once during the evolutionary history of the genus. Of the five
basal, non-salt lake inhabiting representatives, T. oraria exclusively inhabits
coastal beaches and sand dunes, whereas the other four species are found at
inland water bodies with generally high salt content, including mound springs of
the Great Artesian Basin in South Australia and samphire flats.
The salt lake inhabiting species survive their extreme environment
possible by constructing characteristic off-set burrows in the muddy bed of dry
lakes that usually extend down to the saline water table. A plug of mud often
seals the burrows. In summer, the spiders usually backfill the entire upper
burrow with mud acquired during the process of excavating a new upper
entrance in the opposite direction to that of the old. Humidity is maintained by
the presence of water at the bottom of the burrow. During particularly hot
weather burrow entrances are plugged which also offers some protection from
predators and presumably flooding.
151
Effects of experimental fire regimes on the diversity

of cursorial spiders in Brazilian savannah
(cerrado vegetation)
Geraldo B. Freire Jr. & Paulo C. Motta
Universidade de Brasília, Brazil, geraldo_freire@yahoo.com.br, mottapc@unb.br
Introduction
The Brazilian savannah, called “cerrado”, covers some 2 million km2 of
Central Brazil and includes various types of vegetation, a mix of woody and
herbaceous plants. After the Amazon Forest, it is the second biggest Brazilian
biome and has well-defined climate with strong dry winter (approximately May
to September). Fire is the most prevalent form of disturbance in the cerrado and
it is extremely important for this environment that has a long and varied history
of burning. This kind of abiotic disturbance would affect patterns of diversity.
The basic idea of the “intermediate disturbance hypothesis” of Connell is that
the highest diversity is maintained at intermediate levels of disturbance by
preventing competitively dominant species from excluding others.
The aim of this study was to determine the effects of frequency and season
of the prescribed burns on cursorial spiders. We used abundance, richness,
equitability, community composition and diversity as parameters to measure
these effects.

We used pitfall traps in five experimental areas (200 m x 500 m) of
Cerrado sensu stricto (Reserva Ecológica do Roncador, Brasília/DF, Brazil)
submitted to different burning regimes (season and frequency): control area (CT)
preserved from fire for at least 36 years; early biennial (EB): burning every two
years in early dry season, late June; modal biennial (MB): burning in the middle
of the dry season, early August; late biennial (LB): burning at the end of
September; modal quadrennial (MQ): burning every four years in the middle of
the dry season (early August). The last burning of quadrennial plot occurred in
August 2007 and in the biennial plots in July, August and September 2008.
In each plot were installed ten sets of traps, each of four containers (35
litres) buried in the ground with its rim at surface level. The traps were placed in
a "Y" with drift fences (6m long) and were open for five days every month. Each
month is considered a sample (n=19, April 2007 to October 2008).
Statistical analysis
Sampling effort: rarefactions with the values of Mao Tau and the Jacknife 1,
with 1000 randomizations without replacement (Estimates).
Abundance: ANOVA (p<0.05) and the Tukey test to indicate which pairs are
different. The abundance values were transformed in logarithmic (log
(x+1)).
152
Richness: Kruskal-Wallis for testing hypotheses for Mao Tau and Jacknife 1,
and Behrens Fisher as pos hoc.
Equitability: Pielou index with ANOVA (R-2.10.0, R Development Core Team).
Diversity: Profile of Rényi (Ha), MANOVA and discriminant analysis.
Similarity: Hierarchical Cluster Analysis (Bray-Curtis distance).
Results
The total sampling effort, considering the 95 samples, was adequate to
capture most species of the community since the asymptote was obtained by the
50th sample as indicated by the rarefaction curve of Mao Tau. We collected
4.315 individuals (2.413 males, 637 females and 1.265 juveniles) were
Lycosidae (2.048 specimens) represents 47.5% of total abundance. We collected
50 species of 26 families and the most abundant species were Alopecosa sp. 2
(431 individuals), Leprolochus cf. birabeni (236), Trochosa sp. (183), Ctenidae
sp. 1 (147), Lycosidae sp. 11 (103), Actinopus sp. 1 (85) and Hogma gumia (67).
The control area, with 1457 individuals, was the most abundant, followed
by EB (879), MB (812), MQ (634) and LB (573). The treatments were
significantly different in relation to abundance (F4,90: 2.6; p < 0.05) and the
control plot has higher abundance compared to LB.
The plots have similar richness (EB, 44; MQ, 43; MB, 39; LB, 36; CT, 36;
F4,90: 0.19, p=0.94). The treatments were significantly different based on the
Jacknife 1 (Kruskal Wallis - H: 18.73; fd: 4; p<0.01) indicating that the plots EB
and MQ have more estimated species number than CT and LB (p<0.05).
The control plot presents the smallest equitability differing from all
biennial fire regimes (EB, MB e LB) (Pielou, F4,90: 5.4; p<0.001). The Rényi’s
equitability indicated LB and MB with more homogeneous distribution,
followed by EB and MQ with identical values and, like Pielou index, CT has
more dominance, that is, smaller equitability.
The diversity index (Rényi) among the treatments are significantly
different (α=2; F4,82: 2.94 ; p=0.02). The treatments LB, MB e MQ show more
diversity than the control area (p<0.05). There is a tendency to more diversity in
EB in comparison to control (p=0.06).
There are differences among the treatments in relation to species
composition (MANOVA – Pillai’s trace 2.55; p<0.01) and the first two principal
axes explain 80% of total variation. The control area is characterized by more
abundance of the three most representative species in first canonical axis
(Alopecosa sp. 2, Actinopus sp. 1 and Lycosa sp. 3). The LB area, on the other
hand, has the Hogna sp. 1, Cybaeodamus sp. 1 and Xeropigo sp. as the most
important species in explaining the second axis.
The hierarchical cluster analysis indicated the composition of the soil
spiders in two distinct groups (Mantel r=0.97, p<0.01), one containing the area
preserved from fire, and the other areas subjected to different regimes of
burning. Regarding the second group, there was the separation of LB area and
high similarity between EB with MB and MQ, the last two being very similar in
species composition.
153
Conclusions
There was no significant difference between the abundances of different
burning regimes (EB, MB, LB and MQ). However, there was significant
reduction of this parameter in the LB compared to CT. This suggests that the
time of burning may be more influential than the frequency on the reduction of
abundance.
Burning supports increasing the richness and evenness, with the largest
number of species of cursorial spiders in areas with burning at the beginning and
the middle of the dry season (EB and MQ) and without a clear differentiation
between the regimes of fire for evenness, although there is a trend to more
homogeneous distributions in areas with burning in the middle or the end of the
dry season.
There is no support to the Intermediate Disturbance Hypothesis for firing
frequency, since the intermediate area of burning (every four years), despite
showing a higher value of diversity in relation to the control area, does not differ
from the modal biennial area (MB).
Similarly, the frequency does not corroborate the Intermediate
Disturbance Hypothesis based on season of burning, since the area with burning
in the middle of the dry season (MB) did not have more diversity compared to
other treatments (EB and LB).
We found a greater similarity between the areas with different burning
regimes than between them and the control area. The high similarity between the
MB and MQ areas is indicative that compared to the frequency, timing of
burning has more influence on the cursorial spiders. Knowing that moisture loss
from the fuel available for combustion occurs throughout the dry season, it is
expected that burns in the same season promote similar effects on the vegetation
which indirectly influences the spiders similarity, as observed in this study.
154
On dwarfs and trees - the Savignia genus group

(Araneae: Linyphiidae: Erigoninae)
Holger Frick1, Wolfgang Nentwig2 & Christian Kropf1
1
Natural History Museum Bern, Department of Invertebrates, Bern, Switzerland,
holger.frick@students.unibe.ch, Christian.kroph@iee.unibe.ch
2
University of Bern, Institute of Ecology and Evolution, Bern, Switzerland,
wolfganf.nentwig@zos.unibe.ch
We show that morphology-based phylogenies can resolve long lasting

systematic problems of highly diverse and morphologically complex taxa. The
Savignia-group sensu stricto is a morphologically homogenous group of seven
genera (136 species in total) with obscure systematics and unknown
phylogenetic relations on species-level.
Former analyses on the genus level phylogeny of erigonine spiders
showed that the genera are closely related. Expansion of these analyses by
refining the taxon sampling within genera resulted in hundreds of most
parsimonious trees with ambiguous relations among the newly added taxa. The
limit of adding new taxa without adding new characters to these matrices has
been exceeded.
However, the copulatory organs of erigonine spiders and of members of
the Savignia-group especially, are very complex. We restrict the taxon sample to
the Savignia-group genera plus outgroup taxa and scored 216 new characters to
account for similarities among these species. Based on the principle of total
evidence, we included also 80 informative characters from former analyses. This
resulted in 36 most parsimonious trees resolving the phylogeny of 72 taxa with
ambiguous relations in only four polytomous nodes. We considered 60 ingroup
taxa out of 158 species belonging to these genera. The average Bremer-support
was 4.5 (median 3). Based on the synapomorphies of highly supported nodes we
redescribe the genera Savignia, Diplocephalus, Araeoncus, Erigonella,
Dicymbium and Glyphesis.
While the genera are well supported, their interrelationships are not.
Genera with simpler genital morphology emerged at the base of the cladogram.
Simplified, the ingroup generic relationships of the Savignia-group sensu stricto
are Glyphesis (Dicymbium (Araeoncus (Diastanillus (Diplocephalus (Erigonella
(Hemistajus, Savignia)))))).
155
Harvestmen (Opiliones) as hosts of terrestrial

Parasitengona (Acari: Actinotrichida) larvae
Grzegorz Gabryś1, Magdalena Felska2 & Joanna Mąkol2,3

1
Department of Zoology, University of Zielona Góra, Zielona Góra, Poland,
g.gabrys@wnb.uz.zgora.pl
2
Institute of Biology, Department of Invertebrate Systematics and Ecology, Wrocław
University of Environmental and Life Sciences, Poland,
joanna.makol@up.wroc.pl
3
Institute of Natural Sciences, Wrocław University of Environmental and Life Sciences,
Poland
Parasitengona constitute one of the most numerous and the most diverse
group of mites. At a rough estimate there are at least 9000 described (nominal)
species that represent more than 800 genera, 60 families, and 15 superfamilies.
More than half of them, i.e. circa 5000 species, belong to Parasitengona aquatica
(= Hydracarina, Hydrachnellae, or Hydrachnidia), the "water mites". The
remaining taxa belong to Parasitengona terrestria (= Trombidia), the "velvet
mites". Unquestionably, the chigger mites - Trombiculidae (comprising
Trombiculinae and Leeuwenhoekiinae) are the most abundant within
Parasitengona terrestria. Trombiculinae consist of more than 3000 species of 221
genera, whereas Leeuwenhoekiinae comprise 212 species of 35 genera. The
remaining several hundred species are the so called "non-chiggers terrestrial
Parasitengona". The life cycle of Trombidia consists of seven stages, including
three active ones: adult, deutonymph, and larva. The egg, prelarva, protonymph,
and tritonymph are inactive calyptostadia. The active postlarval instars of
Parasitengona terrestria are predatory and they feed on insect eggs, larvae, and
pupae as well as on other small arthropods. The majority of heteromorphic
larvae (excl. Calyptostomatidae) are parasites on invertebrates. The identified
host species belong to Coleoptera, Diptera, Orthoptera, Lepidoptera,
Trichoptera, Hemiptera, Homoptera, Hymenoptera, Thysanoptera, Dermaptera,
Isoptera, Psocoptera, Mecoptera, Odonata, Blattodea, Collembola,
Microcoryphia, Thysanura, Chilopoda, Diplopoda, Solifugae, Pseudoscorpiones,
Araneae, Acari, and Opiliones.
In the present work, an attempt to collect and systematize all data on
Parasitengona terrestria parasitizing Opiliones has been made. The literature data
was supplemented with new records of parasitic relationships, parasitic taxa, and
host species from various zoogeographical regions.
156
Influence of the historical structures of agricultural

landscape on diversity of the epigeic spider communities 3
Peter Gajdoš1 & Lenka Dankaninová2
1
Institute of Landscape Ecology, Nitra Branch, Slovak Academy of Sciences, Nitra,
Slovakia, nrukgajd@savba.sk
2
Faculty of Natural Sciences, Constantine the Philosopher University, Nitra, Slovakia
Authors summarise the research results on epigeic spider fauna in 3 model

areas representing 3 types of the traditionally used agricultural landscapes:
vineyards in Svätý Jur, mountain agricultural landscape with dispersed
settlement in Hriňová and the mountain meadows and pastures in Liptovská
Teplička. The spider communities have been investigated on 55 study plots by
pitfall traps in 2009. Selected investigated plots are typical historical structures
which were chosen as characteristic for 3 researched types of the Slovakian
landscape. From a structural perspective they represent a mosaic of small-scale
arable fields, vineyards, meadows, pastures and permanent agricultural
cultivations and their size, shape, orientation, distribution, utilization,
agricultural forms of relief (balks), margins of cross field tracks and other
characteristics as well as regional or local differentiations which are the results
of interactions between natural conditions, geographical position, cultural-
historical and economical development. During one year research more than 10
000 spider specimens belonging to more than 200 species were captured.
Ecological relationships of spider species to individual historical structures were
evaluated. These structures create very good living conditions for high
biodiversity of the arachnofauna and occurrence of many rare and threatened
spider species. Of the identified species, more than 20 species are listed in the
Red List of Spiders of Slovakia in different categories of threat (e.g. Dysdera
crocata, Ero tuberculata, Dipoena erythropus, Acartauchenius scurrilis,
Erigonella hiemalis, Gonatium rubens, Megalepthyphantes collinus, Meioneta
saxatilis, Meioneta simplicitarsis, Peponocranium orbiculatum, Pocadicnemis
juncea, Theonina cornix, Walckenaeria acuminata, Walckenaeria alticeps,
Altella lucida, Echemus angustifrons, Gnaphosa modestior, Haplodrassus
dalmatensis, Zelotes gracilis, Xysticus kempeleni, etc.). High richness of the
spider fauna and occurrence of the rare and threatened species for Slovakia
allocate on high biotic value of the model areas of agricultural landscape.
Presented results together with the results of further research concerning the
historical structures of agricultural landscape will serve as a background for
preparation of documentation or legislation aiming in protection of these
3
The research was supported by the financial mechanism of EEA, project No. 2008-03-
09 “Development scenarios of representative landscape ecosystems in the Slovak
Republic considering global changes“.
157
biotopes. The reason for it is the real risk that in near future, due to abandonment
of these structures, following succession of forest and urban pressure, decline or
even irreversible loss of biodiversity linked to such specific biotopes will be the
result.
158
Epigynal morphology of Entelegynae and

its evolutionary implications
Jie Gao & Lihong Tu
gaojiexyyt@163.com, tulh@ioz.ac.cn
Two types of spider epigynum have been recognized: haplogyne and

entelegyne. It is widely accepted that the haplogyne condition is plesiomorphic
for all Araneae. Several haplogyne groups nested with entelegyne clade made
the monophyly of entelegynae more ambiguous. Recently published
morphological data on “micronetine” female genitalia indicate that our
knowledge of spider epigynal morphology based on transmitted light
microscopy data is largely incomplete and somewhat misleading. The
entelegyne epigyna, at least in some groups, are formed by a pair of grooves,
rather than ducts. In present study, we sampled more than 30 representative
species from over 20 spider families including both duct-model and groove-
model epigyna. Using a comparative approach, we inferred the basic formation
of the entelegyne epigynum and variation patterns in different groups. Our
observations confirmed the widespread occurrence of epigynal grooves in
Entelegynae. The results will help to formulate new homology hypotheses for
epigynal structural elements across taxa and to provide new morphologic data
for phylogenic reconstruction.
159
Prey records of Oecobius navus Blackwall, 1859

(Araneae: Oecobiidae) associated to urban area
in Montevideo, Uruguay
Luis Fernando García1,2,3, Mariángeles Lacava1,2
& Carmen Viera1,2,3
1
Departamento de Entomología, Facultad de Ciencias, Universidad de la República,
Montevideo, Uruguay
2
Laboratorio Ecología del Comportamiento IIBCE, Montevideo, Uruguay
3
anelosimus@gmail.com
The spider Oecobius navus Blackwall, 1859 is a cosmopolitan species

frequently associated to urban areas (Voss et al. 2007). Although the taxonomy
of the family in South America is relatively well known (Santos & Olivera-
Gonzaga 2003), the biology of most of species remains poorly understood. For
the genus Oecobius only few studies on general aspects of the biology of O.
annulipes (Glatz 1967) and O. templi Cambridge 1876 (Debski 1923), life cycle
(Miyashita 1992) and habitat preferences on urban areas of O. navus are known
(Voss et al. 2007).
Until now there are no studies related to the prey composition on species
of this family. Here, we describe the diet for the population of O. navus living in
urban area of Montevideo, Uruguay.
Prey of O. navus was collected from 141 webs in four different localities
of the Instituto de Investigaciones Biológicas Clemente Estable. The spiders
were consuming the prey on the moment they were observed or the prey
carcasses of recently consumed items were attached to the web. Only one prey
per web and resident adult female spiders were collected, the latter being the
most abundant at the time of study. The prey was identified and classified into
walking and flying following Longhorn and Triplehorn (2004).
We recorded 141 preys, majority being represented by walking organisms.
This could result from the fact that spiders build web at the substrate level as
walkers. The principal prey items were ants (70%) and were followed by
lygaeids and aphids (6% each). The high consumption of ants agrees with
previous observations of Glatz (1967) and Voss et al. (2007). The high
percentage of this kind of prey could suggest behavioural and other adaptations
similar to those observed in myrmecophagic zodariid spiders, where ants even
several times bigger than spiders are consumed (Pekár 2004, Pekár & Lubin
2009, Pekár et al. 2005, 2008).
Although in our observations the spiders were closely distributed (up to 6
spiders observed on a distance of 4 cm), no cases of cannibalism were reported.
Only one specimen of the order Araneae was consumed, and although it could
not be identified due to poor preservation, but it was not an oecobiid spider. The
lack of cannibalism in spite of the spatial proximity could suggest conspecific
tolerance or anti-predatory mechanisms to avoid this behaviour. Aspects related
160
to the possible myrmecophagy and tolerance among individuals on this species

should be the object of further research.
Acknowledgments
The authors wish to thank Dr. Stano Pekár for his valuable comments on
the manuscript and help with literature.
References
Debski B. 1923. Quelques observations sur les moeurs de l’Oecobius templi
Cambridge 1876, retrouvé á Helouan (Arachnida). Bulletin de la Société
Entomologique D'égypte, 1922: 121-126.
Glatz L. 1967. Zur biologie und morphologie von Oecobius annulipes (Araneae:
Oecobiidae). Zoomorphology, 64: 185-214.
Johnson N.F. & Triplehorn C.A. 2004. Borror and DeLong's Introduction to the
Study of Insects. Brooks Cole Ed., 864 pp.
Miyashita K. 1992. Life cycle of Oecobius annulipes (Araneae: Oecobiidae)
under indoor conditions and the effect of photoperiod on nymphal
development. Acta Arachnologica, 41: 5-10.
Pekár S. 2004. Predatory behavior of two European ant-eating spiders (Araneae,
Zodariidae). Journal of Arachnology, 32: 31-41.
Pekár S. & Lubin Y.D. 2009. Prey and predatory behavior of two zodariid
species (Araneae, Zodariidae). Journal of Arachnology, 37: 118-121.
Pekár S., Král J & Lubin Y.D. 2005. Natural history and karyotype of some ant-
eating zodariid spiders (Araneae, Zodariidae) from Israel. Journal of
Pekár S., Toft S., Hrusková M., & Mayntz D. 2008. Dietary and prey-capture
adaptations by which Zodarion germanicum, an anteating spider (Araneae:
Zodariidae), specialises on the Formicinae. Naturwissenschaften, 95: 233-
239.
Santos A. & Gonzaga M.O. 2003. On the spider genus Oecobius Lucas, 1846 in
South America (Araneae, Oecobiidae). Journal of Natural History, 37:
239-252.
Voss S.C., Main B.Y. & Dadour I.R. 2007. Habitat preferences of the urban wall
spider Oecobius navus. Australian Journal of Entomology, 46: 261-268.
161
Description of a new extravagant species

of the genus Thorelliola (Araneae: Salticidae),
with remarks on generic limits
Joanna Gardzińska
Katedra Zoologii, Akademia Podlaska, Siedlce, Poland, gard@ap.siedlce.pl
The genus Thorelliola is known from Malaysia, New Guinea and some
Pacific Islands. It was hitherto considered remarkably homogenous and
distinctive due to the body shape, structure of genitalia and male clypeal
mechanoreceptive setae modified in different ways.
In a new species from New Guinea the clypeal configuration doesn’t
correspond with recent definition of Thorelliola. The males are distinctive by the
possession of clypeal fringe of lanceolate pallid hairs below AMEs and
extravagant anterolateral rows of cheliceral spines, unknown in the genus so far.
The females may be recognized by clypeal hairs and details of epigyne,
including position of copulatory openings and course of insemination ducts.
New criteria for the genus are proposed and the new species is diagnosed,
described and illustrated.
162
Mechanism and function of colour variation in

the crab spider Thomisus spectabilis (Thomisidae)
Felipe M. Gawryszewski1, Ana L. Llandres2, Debra Birch1

& Marie E. Herberstein1
1
Department of Biological Sciences, Macquarie University, North Ryde, NSW,
Australia, f.gawry@gmail.com
2
Department of Functional and Evolutionary Ecology, Estación Experimental de Zonas
Áridas (CSIC), Almería, Spain
Sit-and-wait predators do not actively hunt for their prey. For that reason
strategies that increase their probability of encountering prey are expected to
evolve. Such strategies include the building of traps, selection of profitable
patches and displaying cryptic colouration to avoid detection. In addition, many
sit-and-wait predators employ tactics that actively attract their prey. For
instance, several crab spiders (Thomisidae) that ambush prey on flowers are UV-
reflective, creating a colour contrast against the flowers that is attractive for
pollinators. These spiders are also known by their ability of changing colour,
including the UV component, over a few days. Therefore in this system spiders
that invest in a strategy of high UV reflectance are expected to capture more
prey and consequently increase their weight. Here we investigated the variation
of colouration in the spider Thomisus spectabilis (Thomisidae) during two
seasons and analysed the correlation between spider colouration and spider body
condition. Secondly, we examined the mechanism of colour variation in this
species, with emphasis in the UV component, using a series of histological and
spectrophotometric analyses. Our results showed strong variation in the UV
component, but not in other wavelengths. Also, the results suggest that high UV
reflectance has a benefit for spiders. In the year that spiders were on average
highly UV reflective there was a positive correlation between spider UV and
body condition, but not in the year when spiders were low UV. The histological
analyses suggest that high UV colouration is achieved by exposing UV-
reflective guanine crystals, present in storage cells bellow the epithelium,
through an UV-transmitting epidermis and cuticle. White non-UV and yellow
colouration are achieved by the presence of ommochrome pigments in different
stages within the epithelial cells, covering the UV-reflective guanine crystals. It
is not clear why spiders do not always invest in a strategy of high UV
reflectance, but we suggest that spiders are adjusting their colouration in
response to the presence of predators and/or quantity of different types of prey.
163
Systematics of the spider genus Stemonyphantes

(Araneae: Linyphiidae)
Efrat Gavish-Regev1, Gustavo Hormiga2 & Nikolaj Scharff1
1
Department of Entomology, Natural History Museum of Denmark, Zoological
Museum, University of Copenhagen, Copenhagen, Denmark,
efrat.gavish@gmail.com
2
DC, USA
Stemonyphantes Menge, 1866, includes 15 described species found in

temperate regions of the northern hemisphere, in at least two distinct species-
groups ("lineatus" group and "abantensis" group). Stemonyphantes has been
hypothesized to be the sister lineage of the remaining Linyphiidae, and treated as
a monotypic subfamily (Stemonyphantinae). In a recent phylogenetic analysis of
combined morphological and molecular data, Arnedo et al. (2009) found three
alternative phylogenetic placement of Stemonyphantes within ”linyphioids”: as
the sister group of Pimoidae, in an unresolved trichotomy with Pimoidae and the
remaining Linyphiidae, or as the sister group to all other linyphiids.
We study the genitalic and somatic morphology of Stemonyphantes and
discuss character homologies in relation to its basal phylogenetic placement.
164
Opiliones as models for evolutionary biogeographic studies

- vicariance, dispersal, translocations
Gonzalo Giribet
Museum of Comparative Zoology, Department of Organismic and Evolutionary

Biology, Harvard University, Cambridge, MA, USA, gonzalo.giribet@gmail.com
Due to the growing knowledge in the faunistic knowledge of several

regions of the World, Opiliones have been recently used for contrasting
biogeographic and evolutionary patterns that can be generalized to other
organisms and for other landmasses. Research being conducted in my laboratory
focuses on different groups of Opiliones with a variety of dispersal capabilities.
Cyphophthalmi have been used due to their old age and low vagility for
elucidating ancient vicariant patterns, but a few instances of possible dispersals
have been reported. In other cases, phylogenetic and biogeographic explanations
are at odds, and may only be justified through long-range dispersal events
(translocations), or wrong geological reconstructions. On the contrary, other
groups of Opiliones have shown immense dispersal capabilities, colonizing,
from the Neotropics, a large number of islands in the South Pacific and
Southeast Asia. These and other examples will be presented and discussed.
165
A new species of Mecynargus Kulczyński, 1894

(Araneae: Linyphiidae) from the steppe zone of Ukraine
Valery A. Gnelitsa
Sumy State Teacher’s Training University, Ukraine, Gnelitsa@mail.ru
Mecynargus foveatus (Dahl, 1912) is the only one of 14 spider species of

the Hypoarctic genus Mecynargus Kulczyński, 1894 (Platnick 2009) recorded
from Ukraine. It has been found in the meadow in the NW Ukrainian forest
zone.
Recently in the steppe places of both the Kherson district (South Ukraine)
and the Crimea tiny pale Mecynargus spiders have been collected by Barber
traps and by hand-held suction sampler. These spiders were caught sometimes in
relatively great amount during October - April in the open spaces with meadow
vegetation.
The pale colour, small general length and extremely tiny male palps
together with M. foveatus - like epigyne impede the correct determination. These
spiders can be mistaken with M. foveatus despite the dark colouration of the
latter. They are of the same size and prefer the same habitat conditions.
We compared the new Mecynargus with M. foveatus and found more
distinctive features besides the colouration. Male differs from M. foveatus by the
form of the carapace, tibia spination, metatarsus trichobothria position, and the
details of the palp such as the embolic division configuration, the absence of the
distal tooth on the tegulum and the shape of the palpal tibia as well. The female
differs by the epigyne (length/width ratio) and the shape of the receptacles.
Moreover the “steppe” Mecynargus is a winter mature species whereas M.
foveatus is a summer mature one. Thus, I consider the second Ukrainian
Mecynargus an undescribed species, closely related to M. foveatus.
166
Untangling the spiny clade of vaejovid scorpions: Evolution

of the largest radiation of North American desert scorpions
Edmundo Gonzalez1,2 & Lorenzo Prendini1
1
Scorpion Systematics Research Group, Division of Invertebrate Zoology, American
Museum of Natural History, New York, NY, USA, egs@amnh.org,
lorenzo@amnh.org
2
Ecology and Evolutionary Biology, City University of New York, New York, NY, USA
The Spiny Retro-Barbate Clade (SRBC) is the largest monophyletic group

in the endemic North American scorpion family Vaejovidae Thorell, 1876.
SRBC vaejovids are widely distributed in North America and range from the
southwestern USA to southern Mexico. The evolutionary relationships of the
SRBC within the family Vaejovidae recently became clearer in the course of an
NSF-funded RevSys grant: Revisionary Systematics of the North American
Scorpion Family Vaejovidae (http://www.vaejovidae.com). Although Soleglad
and Fet (2008) erected three new genera to accommodate species formerly
included in three groups of the genus Vaejovis (which comprises part of the
SRBC), phylogenetic analyses conducted as part of the RevSys grant, questioned
the monophyly of these genera. The work presented here aimed to (1) survey the
regions of greatest SRBC vaejovid diversity to discover new species, document
distributions, and gather fresh material for morphological and genetic studies;
and (2) revise the classification of the SRBC on the basis of monophyly. New
material was collected across the distributional range of the SRBC and
additional material studied in several museum collections. Tissue samples for
DNA isolation were obtained and sequenced for two nuclear and three
mitochondrial markers: Cytochrome Oxidase I, 12S rDNA, 16S rDNA, 18S
rDNA and 28S rDNA. 350 morphological characters were scored across the
same 78 species for which DNA sequences were obtained. Combined and
partitioned analysis of static alignments and morphology were analyzed with
parsimony and posterior probability as optimality criteria. Direct optimization of
the combined and partitioned data was also conducted. All analyses retrieved a
monophyletic SRBC and eight clades that contradict traditional and recently
proposed classifications of Vaejovidae.
References
Soleglad M.E. & Fet V. 2008. Contributions to Scorpion Systematics. III.
Subfamilies Smeringurinae and Syntropinae (Scorpions: Vaejovidae).
Euscorpius, 71: 1-115.
167
A new record of redback spider, Latrodectus hasselti

(Thorell, 1870) (Araneae: Theridiidae)
from south-western Iran
Hamid R. Goudarzi1 & Mohammad Abdigoudarzi2

1
Arthropods Lab. Venomous Animals and Antivenom Department, Razi Vaccine and
Serum Research Institute (RVSRI), Iran
2
Acarolology laboratory, Parasitology Department, Razi vaccine and serum research institute
Karaj Iran
So far, four species of the genus Latrodectus: L. tredecimguttatus, L.

dahli, L. pallidus, and L. geometricus have been reported by the author and other
investigators from Iran.
The redback spider is native to Australia, but in recent decades there have
been some reports from other parts of the world. During ecological and faunistic
investigations of medically important spiders of Khoozestan Province (SW Iran)
held from 1st to 4th March 2010, we have found a female of Latrodectus hasselti
and her three egg sacs, all collected beside the man-made debris, 40 km from
Ahvaz to Haftgel road.
This is the first record from Iran and it seems most likely that the shipment
of goods to the southern costal regions could be the source of the introduction.
168
Phylogeny and behaviour of enigmatic orbicularians:

Zygiellidae and the giant orbweaver Caerostris
Matjaž Gregorič1*, Ingi Agnarsson2, Todd Blackledge3

& Matjaž Kuntner1
1
Institute of Biology, Scientific Research Centre, Slovenian Academy of Sciences and
Arts, Ljubljana, Slovenia
2
3
Department of Biology, University of Akron, Akron, OH, USA
* Presenting author: matjaz.gregoric@gmail.com
The classical Araneidae contains a large diversity of orbweavers that do

not necessarily have much in common except primitively building orb webs. For
example, the Holarctic free sector spider genus Zygiella s. l. is taxonomically
controversial, was transferred between araneids and tetragnathids in the past, and
was recently even split into four genera proposed to belong to their own family,
Zygiellidae. Another enigmatic “araneid” is the bark spider, genus Caerostris.
Caerostris are large orbweavers that are widespread in the old world tropics, but
extremely understudied, and their phylogenetic affinities are controversial. Here,
we reconstruct the phylogeny of major orbweaving lineages based on nuclear
and mitochondrial data and show that the classical Araneidae is polyphyletic.
Zygiellidae, including several genera currently in Araneidae (Zygiella s. l.,
Phonognatha and Deliochus) receives strong support, and is not closely related
to Araneidae. Some web features including double radii characterize zygiellids
and distinguish them from araneids. Our phylogenetic results also support
Caerostris as a clade outside of Araneidae but do not place it conclusively,
grouping it sister to Araneidae, Nephilidae, or Zygiellidae. Our preliminary work
revealed high evolutionary diversity for Caerostris in Madagascar, where some
species even utilize a novel ecological niche by bridging rivers and lakes
producing webs suspended upon 25m long bridgelines. We broaden the
knowledge of Caerostris behaviour by explaining how the heavy bodied
Caerostris darwini build webs across rivers, what prey they catch, and how they
avoid destruction of webs by large flying animals.
169
The lace web spiders (Araneae: Phyxelididae) of

Madagascar: phylogeny, biogeography and taxonomy
Charles E. Griswold1,2,3,4, Hannah Marie Wood1,2

& Anthea Carmichael1
1
Arachnology Lab, Entomology Department, California Academy of Sciences, San
Francisco, CA, USA
2
University of California, Berkeley, Environmental Science, Policy and Management,
Berkeley, CA, USA
3
Department of Biology, San Francisco State University, San Francisco, CA, USA.
4
Corresponding author: cgriswold@calacademy.org.
The lace web spiders (Araneae, Phyxelididae) of Madagascar are revised.

Molecular phylogenetic analyses for 32 Malagasy phyxelidid exemplars, 9
confamilial outgroup taxa, and 7 other more distant outgroups are performed on
for 4 genes (28S, 18S, H3 and COI) utilizing Bayesian, maximum likelihood and
parsimony analyses. These analyses suggest that there are 15 species of
Phyxelididae that may be recognized from Madagascar, that these may be
divided into three genera, and that the Malagasy phyxelidids form a
monophyletic group, probably resulting from a single invasion of the island by
an ancestor from Africa. Implications of these findings for biogeography within
Madagascar and of biotic relations of Madagascar to other landmasses are
discussed. Observations on the courtship and mating behaviour of phyxelidids
are presented.
170
Cladistic analysis and biogeography of the genus

Oligoxystre Vellard, 1924 (Araneae: Mygalomorphae:
Theraphosidae)
José Paulo Leite Guadanucci
Universidade Federal dos Vales do Jequitinhonha e Mucuri, Diamantina, Minas Gerais,

Brazil, joseguadanucci@gmail.com
The genus Oligoxystre was originally established in 1924 as monotypic for

O. auratum from southern part of Goiás state, Central Brazil. In 1985,
Oligoxystre was considered senior synonymy of the genus Cenobiopelma and
subsequently comprised three species, including O. mimeticum and O.
argentinense. According to recent taxonomic revision, the two latter species
have been removed from Oligoxystre and six species in the genus have been
recognized: O. auratum (the type species), O. caatinga, O. bolivianum, O.
tucuruiense, O. rufoniger and O. dominguense. More recently, another new
species, O. diamantinesis, was described from Serra do Espinhaço, city of
Diamantina. The examination of collection material and field trips to different
localities of Serra do Espinhaço revealed two more new species from the
Atlantic forest domain.
The geographic distribution of the species are as follows: O. tucuruiense:
single record from Eastern Amazonia; O. caatinga: NE Brazil (Caatinga biome)
- partly sympatric with O. rufoniger; O. bolivianum: from Brazilian Central
Cerrado to S Bolivia; O. dominguense: Cerrado at northern state of Goias; O.
diamantinensis: Serra do Espinhaço; O. rufoniger: partly sympatric with O.
caatinga, extending to Meridional Serra do Espinhaço; O. sp. 1: E side of Serra
do Espinhaço; O. sp. 2: southern state of Bahia.
A cladistic analysis has been performed and the following topology
obtained: (O. caatinga (O. sp. 2 O. sp.1 (O. tucuruiense + O. rufoniger) (O.
diamantinensis (O. bolivianum + O. dominguense))))). An area cladogram was
obtained replacing the terminal taxa by its geographical area distribution.
According to the area cladogram, it is possible to draw the following
conclusions: the origin of Cerrado fauna is monophyletic; the occurrence of two
sympatric species in Caatinga is due to different events, a basal differentiation of
Caatinga from the rest of the genus, which originated O. caatinga, and the
separation between Eastern Amazonia and Caatinga, which originated O.
tucuruiense and O. rufoniger; Serra do Espinhaço, which does not represent a
vicariant event due to its pre-Cambrian origin, is the limit among three major
Brazilian biomes (Caatinga, Cerrado and Atlantic forest) and the sympatry
among three species (O. rufoniger, O. diamantinensis and O. sp. 1) mainly at the
meridional portion, is probably the result of the contact of the typical fauna of
each biome.
171
A troglophyle population of Diplura sp. (Araneae:

Mygalomorphae: Dipluridae) in a quartzitic cave
in Diamantina, Minas Gerais, Brazil
José Paulo Leite Guadanucci, Pilar L. Maia Braga,
Fernanda de Souza Sá & Rafael da Fonseca Ferreira
Universidade Federal dos Vales do Jequitinhonha e Mucuri, Diamantina, Minas Gerais,

Brazil, joseguadanucci@gmail.com
The cave Monte Cristo, located at18o17.822'S, 43o33.511'W, is an

approximately 200 meters long quartizic formation. Hypogean environments house
animals classified in three ecological-evolutionary categories according to their
dependency on the cave: trogloxene, troglophyle and troglobite. During an inventory
survey on cavernicolous arachnids in caves in Diamantina, state of Minas Gerais,
several representatives of Diplura sp were found. Considering the rarity of dense
populations of Mygalomorphae spiders in caves, this finding led us to conduct a
survey on the population dynamics of such species. Representatives of the genus
Diplura are easily recognized by its mid-sized body, long posterior lateral spinnerets
and presence of a maxillary lyra composed of few clavate setae. They build silky
webs with tunnels with sheet web at the entrance, what makes them easily found.
This work aims at studying this population and present data on the abundance of
individuals, spatial distribution within the cave, territoriality, phenology and
circadian rhythm. We have done three excursions from January to March of 2010,
when all webs with spiders were marked and numbered, and all spiders found were
marked with coloured ink on the carapace. A total of 38 individuals were marked,
what makes it the largest mygalomorph population recorded inside a cave, 13 in the
first trip, nine in the second and 16 in the third. Several spiderlings were found in the
third excursion, indicating the end of the reproductive season, when all juveniles
have already hatched and started to disperse and establish shelters in the habitat.
Twenty four individuals were found close to the entrance, and the rest were at the
aphotic region, where temperatures are lower and more constant and humidity is
higher. We found no significant difference in the abundance of potential preys in the
different regions of the cave, having no relation to spider distribution. Only one
spider changed its web location during the observations and built a new shelter less
than 1 meter away. Moreover, the number of old exuviae deep within the webs of
many spiders found indicates that once the spider has established its web, it remains
there for the rest of the life. Careful searches have been done in the surroundings of
the cave and no representatives or webs of Diplura sp were found, showing the clear
preference for the cave environment. Several other animals are known to inhabit
caves as troglophyles (e.g. bats, harvestman, pseudoscorpions, several insects), what
shows the importance for the preservation of such environments. Monthly
observations will be done during a year to evaluate aspects on the phenology and
circadian rhythms of this population.
172
Sociality and resource use: insights from a community

of social spiders in Brazil
Jennifer Guevara1, Marcelo O. Gonzaga2, João Vasconcellos-Neto3

& Leticia Avilés1
1
University of British Columbia, Canada, guevara@zoology.ubc.ca,
laviles.ubczool@gmail.com
2
Universidade Federal de Uberlândia, Instituto de Ciências Biomédicas, Instituto de
Biologia, Uberlandia, MG, Brazil
3
Sao Paulo, Brazil
Behavioural differences among closely related species, in addition to

morphological ones, can play a significant role in species coexistence and thus
the assemblage of natural communities. Even though closely related species are
expected to share many niche dimensions, they may differ in ways that allow
them to utilize different resources and thus alleviate competitive interactions.
Body size differences, for instance, often allow exploitation of food of different sizes.
Analogous to body size, for species living in groups, group size differences and level
of sociality may also contribute to resource partitioning, a possibility that has hardly
been considered. We tested this idea in a subtropical forest site in Brazil where five
spider species (Anelosimus) with levels of sociality ranging from almost solitary
to highly social coexist. We found that the range of insect sizes captured by the
species reflected their nest and colony size so that species with larger colonies
and webs captured larger insects than less social species. Yet, among those
species whose webs did not differ significantly in size—the two with the largest
and the two with the smallest webs—one captured significantly larger insects
than the other. This difference in prey size was apparently due to differences in
the extent to which nest mates cooperated in the capture of prey, as in only one
of the species in each pair did the size of the insects captured increased with
colony size. The four species were thus packed along the spectrum of available
insect sizes from least to most social. This pattern of resource use was more
over-dispersed than expected by chance, suggesting limited overlap between
contiguous species as would be expected if the species had been assembled to
avoid extensive dietary overlap. We suggest that social evolution can potentially
create large differences in resource use, thus contributing to the coexistence of a
greater number of similar species in ecological communities.
173
Revisions of the cryptic Castianeirinae (Araneae:

Corinnidae) of the Afrotropical Region: exceptional
species diversity, unusual biogeographical patterns and
evolutionary novelties 4
Charles R. Haddad
Department of Zoology & Entomology, University of the Free State, Bloemfontein,

South Africa, haddadcr@ufs.ac.za
Revisions of the Afrotropical Castianeirinae are underway, focusing

initially on four genera of cryptic castianeirines, i.e. not mimicking ants as in
most other genera in the subfamily.
Medmassa Simon, 1887 served as a dumping ground for unspecialised
corinnids, as resolution of the genus in the region resulted in retention of only
one of the eight species previously described, i.e. M. semiaurantiaca Simon,
1910, a primarily arboreal species widespread in the Afrotropical Region
(Haddad & Bosselaers 2010).
The revision of Copa Simon, 1885 indicates that one species, C.
longespina Simon, 1910, is misplaced and should be transferred to Echinax
Deeleman-Reinhold, 2001, a genus previously known only from South-East Asia
(Deeleman-Reinhold 2001, Marusik et al. 2009). Three new species have been
discovered in tropical and subtropical Africa. All of the species appear to be
primarily arboreal. This presents a very peculiar distribution, suggesting that the
genus should also occur in the Indian subcontinent.
Copa sensu stricto is itself represented in the region by only two
continental species, while the genus has speciated extensively on Madagascar,
with more than forty new species in addition to two described species. The
genus has also been recorded from Sri Lanka. The radiation on Madagascar is
coupled with several genitalic modifications: 1) considerable variation in male
embolus structure, 2) the evolution of single, double or triple cymbial
apophyses, and on occasion, tibial apophyses, and 3) reduction in many species
in the length of the copulatory ducts and spermathecae. Despite these genitalic
modifications to effect sexual isolation, somatic morphology has remained very
consistent with the continental type species. The cymbial and/or tibial apophyses
are evolutionary novelties found in most (if not all) Madagascan castianeirines
and not occurring in any continental species, suggesting that they must have
been adaptive in enforcing sexual isolation.
A revision of Messapus Simon, 1898 indicates that the type species, M.
martini Simon, 1898, represents two distinctly different species, one tentatively
placed in Corinninae (holotype female) and the other belonging to Castianeirinae
4
Research supported by the National Research Foundation of South Africa through its
Thuthuka programme (grant number TTK2008050500003).
174
(syntype male) (contra Bosselaers & Jocqué 2000). A new genus is created to
accommodate the misplaced male, its matching female is described, and the
correct matching male of M. martini is described. Copa lacustris Strand, 1916
and Castianeira kibonotensis Lessert, 1921 are transferred to the new genus, and
more than ten new species are described from southern, central and eastern
Africa. In contrast to Copa, the new genus has apparently not colonised
Madagascar (but does occur in the Comoros Islands, possibly introduced) and
radiation seems to be related to two main geological events: the formation of the
Great Rift Valley in east Africa (particularly leading to speciation in the Eastern
Arc Mountains of Tanzania), and the formation of the Maputaland coastal plain
in eastern South Africa and Mozambique.
The diagnostic characteristics of cryptic Afrotropical castianeirines are
discussed, as well as future directions for taxonomic research on the group.
References
Bosselaers J. & Jocqué R. 2000. Studies in Corinnidae: transfer of four genera
and description of the female of Lessertina mutica Lawrence 1942.
Tropical Zoology, 13: 305-325.
Deeleman-Reinhold C.L. 2001. Forest spiders of South East Asia: with a
revision of the sac and ground spiders (Araneae: Clubionidae, Corinnidae,
Liocranidae, Gnaphosidae, Prodidomidae and Trochanterriidae [sic]).
Brill, Leiden, 591 pp.
Haddad C.R. & Bosselaers J. 2010. A revision of the genus Medmassa Simon, 1887
(Araneae: Corinnidae) in the Afrotropical Region. Zootaxa, 2361: 1-12.
Marusik Y.M., Zheng G. & Li S. 2008. First description of the female of
Echinax panache Deeleman-Reinhold, 2001 (Aranei: Corinnidae:
Castianeirinae). Arthropoda Selecta, 17: 65-68.
175
Does any gradient of spider species diversity exist in a

river valley of the Eastern Europe?
Izabela Hajdamowicz1, Marzena Stańska1 & Maria Oleszczuk2

1
Department of Zoology, University of Podlasie, Siedlce, Poland,
hajdamo@ap.siedlce.pl, stanska@ap.siedlce.pl
2
Research Centre for Agricultural and Forest Environment in Poznan, Polish Academy
of Sciences, Poznań, Poland, oleszczukm@vp.pl
The gradients of species diversity and hypotheses explaining the

phenomenon are still important topic in the conservation biology. To describe a
model of species diversity distribution in a river valley, different hypotheses like
geographical, accessibility, naturalness - disturbance, environmental - humidity
and sun exposure were tested. Our studies have been conducted in the Bug River
Valley, located in the Eastern Europe. In the strongly changed European
landscape, the bed of the river has got a natural meandering character, creating
diversity of habitats, both in the water and on the land. Along the river, narrow
belts of riparian natural forest and rushes at river oxbows occurred. The edges of
the valley are covered with semi-natural xerothermic and mesoxerothermic
grasslands and thickets. In the biggest part of the valley, on flooded terraces,
mown meadows and sand grasslands, both used as pastures, were situated. In
spite of the human impact, the Bug River Valley belongs to the one of the most
important ecological corridors in Europe. Twenty six study plots were located in
four sites, representing different habitats of the valley. Between March and
November in 2007, spiders were collected by 10 pitfall traps in each study plot
and the traps were emptied twice a month. Approximately 50 000 individuals of
spiders belonging to 244 species were collected. To measure the species
diversity, the species richness and the Shannon-Wiener index were applied. The
geographical species diversity gradient was revealed. The spider species
diversity was decreasing from south-east to north-west down the river.
Moreover, shortly accessible flooded habitats like riparian forests, rushes and
wet meadows were characterized by lower spider species diversity than fresh
meadows, grasslands and thickets on the edge of the valley. In general, in the
environmental gradient of humidity, in dry habitats the species diversity was
higher than in wet habitats. Furthermore, the low-disturbed habitats, like
grasslands were characterized by higher spider species diversity in contrast to
natural and stronger disturbed habitats.
176
Pirates of the dark: systematics and first phylogeny

of Australasian pirate spiders (Araneae: Mimetidae)
Danilo Harms1,2 & Mark S. Harvey2
1
School of Animal Biology M092, The University of Western Australia, Crawley,
Western Australia, danilo.harms@museum.wa.gov.au
2
Department of Terrestrial Zoology, Western Australian Museum, Welshpool, Western
Australia, Mark.Harvey@museum.wa.gov.au
Pirate spiders (Arachnida: Araneae: Mimetidae) are vagrant predators of

other spiders and do not build webs on their own, but instead invade alien webs
in order to prey upon the host. They perform behavioural patterns of aggressive
mimicry and pull on strands of silk like a potential prey item, to entice and then
attack the host within its own web. In accordance with their highly specialized
ecology, mimetids show remarkable adaptations in somatic morphology, such as
conspicuous rows of raptorial spines on the forelegs, which are arranged to form
a basket around the victim during prey capture.
Pirate spiders are diverse in Australia and New Zealand and occur there
with some 30 species in three genera. Australomimetus traditionally included
most Australian mainland species and was apparently confined to the Australian
east-coast. In contrast, all Tasmanian and New Zealand species were dumped
into the large, cosmopolitan genera Ero and Mimetus. This classification made
little sense, both biogeographically and phylogenetically, and has never been
subject to critical testing. We have therefore performed the first phylogenetic
analysis of Australasian pirate spiders. Our analysis, using 29 species and some
90 morphological characters, supports an expanded concept of Australomimetus
which remains as the only native pirate spider genus in Australia and is found to
contain the entire Australian and New Zealand fauna.
Our analysis also provides evidence that Australomimetus is not an
Australasian endemic. Instead, the genus crosses Wallace’s Line in the north,
with additional species occurring in Japan, China and Indonesia.
177
Vicariance and the evolution of ancient pseudoscorpions:

subfamily Pseudotyrannochthoniinae (Arachnida:
Pseudoscorpiones: Chthoniidae)
Danilo Harms1 & Mark S. Harvey2

1
School of Animal Biology M092, University of Western Australia, Crawley, WA 6009;
Department of Terrestrial Zoology, Western Australian Museum, Welshpool DC,
WA, Australia, harmsd01@student.uwa.edu.au
2
Department of Terrestrial Zoology, Western Australian Museum, Welshpool DC, WA,
Australia, mark.harvey@museum.wa.gov.au
We are investigating the phylogeny and phylogeography of an ancient and

globally distributed lineage of pseudoscorpions, the Pseudotyrannochthoniinae.
These are small, predatory arthropods that belong to one of the oldest terrestrial
animal groups. Pseudotyrannochthoniines are found in relictual habitats on five
continents and have low vagility and naturally small distributions. By using
Pseudotyrannochthoniinae, we will be able to test biogeographic hypotheses on a
spatial and temporal scale rarely attempted before. The specific aims of our
study are:
1) To undertake a comprehensive phylogenetic analysis of the
Pseudotyrannochthoniinae, using representatives of all currently recognised
genera from at least five continents plus a variety of other pseudoscorpions of
the family Chthoniidae as outgroup taxa. Morphological characters and DNA
sequences will be analysed using cladistic methods, resulting in a systematic
revision of all currently recognised genera.
2) To test a vicariant model of Pangaean biogeography, under the testable
assumption that the current distribution of the Pseudotyrannochthoniinae is the
result of continental drift. We will conduct a phylogeographic analysis of the
molecular data and analyse spatial distributions and ages of these
pseudoscorpions across continents. This will lead to novel insights into their
geographic distribution, both past and present, and their evolution over space
and time.
3) To undertake a systematic and taxonomic revision of
Pseudotyrannochthoniinae in Australia, using morphological techniques and the
molecular data. Ten species are currently described from Australia but the
estimated number is much higher and up to 50 new species will be described and
named. Our study will be the first to document the full diversity and distribution
of these pseudoscorpions in Australia.
178
Chthonioid pseudoscorpions: phylogeny based on morphology
Mark S. Harvey
Department of Terrestrial Zoology, Western Australian Museum, Welshpool DC, WA,

The pseudoscorpion superfamily Chthonioidea is well defined and

seemingly monophyletic based on previously published morphological and
molecular datasets using multiple markers. However, the higher classification is
not settled with several alterations over the past two decades. To assist resolve
these issues, a phylogenetic analysis was performed using 121 chthonioid
species placed in 35 of the 48 recognised genera. Parsimony analyses under
equal weights provided little phylogenetic resolution. Implied weights analyses
drastically improved the resolution of the trees and, in all analyses,
Pseudotyrannochthoniidae were sister to the remaining chthonioids. In both
equal and implied weights analyses, the taxa currently included in the families
Tridenchthoniidae and Lechytiidae grouped strongly with the chthoniid genera
Sathrochthonius and Sathrochthoniella. Of the remaining taxa, low concavity
functions distinguished three other clades, Chthoniini, Tyrannochthoniini and
the “apochthoniines” (Apochthonius+Kleptochthonius). Higher concavity
functions retained Tyrannochthoniini and the “apochthoniines”, but divided
Chthoniini into multiple clades. Proposed changes to the classification of the
Chthonioidea are discussed.
179
Moulting in a whip scorpion

Joachim Haupt
former address: Institute of Biology, TUB, Gluckweg 6, 12247 Berlin, Germany
A thelyphonid (Ginosigma schimkewitschi (Tarnani, 1894)) was kept in

darkness while transporting from Thailand to Germany. It had prepared for
moulting, but did not dig a burrow, because it was prevented from doing so.
Thereby I could observe the moulting process.
Although the starting of the moulting could not be observed, it can be
imagined that ecdysis starts with the rupture of the peltidium which diverges in
horizontal direction. The whip scorpion sits on his three legs (the fourth is used
as tactile leg). Successively, the white next instar appears. It withdraws the
chelicerae first, then the pedipalps, next the tactile leg and the walking legs, last
the whip. The whole process needs about two hours, that's why the film is
speeding.
Even before the moulting is finished, the whip scorpion starts to move one
or the other limb.
After the whip scorpion has withdrawn from the exuvia, it will walk away.
At that time the cuticle is still white – protonymphs and deutonymphs will do the
same way.
In the discussion a comparison will be made with mesothelae and
mygalomorphae.
180
The mechanical importance of retaining the non-sticky

spiral in Nephila webs (Araneae: Nephilidae)
Thomas Hesselberg & Fritz Vollrath
Oxford Silk Group, Department of Zoology, University of Oxford, United Kingdom,

thomas.hesselberg@zoo.ox.ac.uk
The highly structured two-dimensional orb web is one of the most

recognisable traps in nature. However, despite its overall conservative design,
variations are found at all taxonomic levels. The members of the genus Nephila
in particular deviates from the norm by building large fine-meshed aerial webs
that retain the non-sticky spiral (Harvey et al. 2007). A previous study on the
transmission of prey vibrations along radii in intact Nephila webs and in webs
with the non-sticky spiral connections cut revealed that vibrations leak from the
radii into the non-sticky spiral, whose presence thus degrades the signal
(Landolfa & Barth 1996). The authors speculate that the negative contribution of
the non-sticky spiral to signal transmission efficiency might be offset by a
positive contribution to the mechanical support of the web (Landolfa & Barth
1996). In the present study we investigate this idea further by comparing the
behaviour of intact and cut webs of Nephila edulis during wind-loading and
simulated prey impact. Wind is one of the most damaging environmental forces
affecting the web. We therefore decided to subject first the intact web to
increasing windspeeds (from 0 m/s to 8.4 m/s in 11 steps) generated by a wind-
tunnel and subsequently removing the non-sticky spiral from two south-running
radii and re-testing the web in order to compare radii deflection and separation.
However, the main function of the orb is to catch prey and its geometry and
structure thus needs to be optimised for this function (Eberhard 1990, Lin et al.
1995). We therefore compared the deflection and failure force of intact parts of
the web with parts where the non-sticky spiral was removed by dropping plastic
balls from variable heights into the horizontally-held web and recording the
impact with a high-speed camera. Our preliminary results indicate that the orb
webs undergo higher deformations when the non-sticky spiral connections are
cut, which suggests that the presence of the non-sticky spiral is very important
for the ability of the large fine-meshed Nephila webs to withstand wind-loading
and to intercept and retain large prey.
References
Eberhard W.G. 1990. Function and phylogeny of spider webs. Annual Review of
Ecology and Systematics, 21: 341-372.
Harvey M.S., Austin A.D. & Adams M. 2007. The systematics and biology of
the spider genus Nephila (Araneae: Nephilidae) in the Australasian region.
Invertebrate Systematics, 21: 407-451.
181
Landolfa M.A. & Barth F.G. 1996. Vibrations in the orb web of the spider
Nephila clavipes: cues for discrimination and orientation. Journal of
Comparative Physiology, 179: 493-508.
Liao C.-P., Chi K.-J. & Tso I.-M. 2009. The effects of wind on trap structural
and material properties of a sit-and-wait predator. Behavioural Ecology,
20: 1194-1203.
Lin L.H., Edmonds D.T. & Vollrath F. 1995. Structural engineering of an orb-
spider's web. Nature, 373: 146-148.
182
Foraging in social spiders

Christina Holm & Marija Majer
Department of Biological Sciences, Aarhus University, Aarhus, Denmark,

Christina.holm@biology.au.dk, marija.majer@biology.au.dk
Web building spiders are to a certain degree polyphagous. Nevertheless

spiders discriminate between prey items, as it is not advantageous to consume all
prey items captured in the web. Spiders actively select prey that is most
beneficial for their fitness, with regards to prey size and prey species.
Stegodyphus dumicola is a social spider species found throughout southern
Africa. These spiders live their entire lives in a communal web and nest
consisting of up to a thousand individuals. They do not defend specific territories
within the web but cooperate on prey capture, feeding on prey items, and brood
care. Observations of prey capture were done directly in the field, and size of
potential prey was determined from window and sticky trap samples. Nest and
web size, and captured prey size were measured in the field as well. Nest sizes
were arbitrarily divided into small, medium, and large to see the relation
between prey acceptance and colony size. The colonies have a tendency to
accept prey in a medium to large size range, but discard small prey, which is
most abundant in the habitat of the colonies. This shows that the spiders are
selective in prey acceptance. The size of foraging groups was also noted to
ascertain the optimal prey size per spider. Moreover we noted the behaviour of
individuals while capturing and feeding. By determining the optimal prey size
range for S. dumicola, it might be possible to make a model on the distribution
of this spider species with regards to non-permanently social congeners, based
upon prey availability in their habitats.
183
Molecular approach to elucidate the hormonal regulation

of spider ecdysis
Yoshiko Honda, Mari Horigane & DeMar Taylor
Graduate School of Life and Environmental Sciences, University of Tsukuba, Japan,

yoshiko-honda@mbi.nifty.com, taylor.de.mar.ge@u.tsukuba.ac.jp
Introduction
Moulting is a common phenomenon necessary for arthropod development
and progresses with complicated sequential regulation. The mechanisms of
moulting are well-understood in insects and steroid hormone ecdysteroids are
one of the main hormones that regulate moulting in arthropods. Ecdysteroids are
secreted into the hemolymph from a synthesizing organ and interact with target
organs to prepare them for ecdysis. Active ecdysteroid is 20-hydroxyecdysone
(20E) and changes in 20E hormone titers of the hemolymph induce moulting.
Moulting consists of three phases: intermoulting, ecdysis and post ecdysis.
During the intermoulting phase, ecdysteroid titers are relatively low and
gradually increase until the next phase, ecdysis (Chapman 1998). Increases in
ecdysteroid titers induce various events necessary for moulting such as
construction of new cuticle. Ecdysteroid titers peak at moulting and immediately
decrease after ecdysis. The patterns of ecdysteroid changes differ depending on
the insect species, but changes in ecdysteroid titers are commonly important for
insect moulting.
Studies on ecdysteroids in spiders are limited to a few studies during the
1970s. Injection of 20E induced moulting of Araneus cornutus and Dugesiella
hentzi (Krishnakumaran & Schneiderman 1968, 1970). A. cornutus spiders
treated with 20E moulted 29 days after treatment and the number of moulted
spiders was fivefold higher in 20E treated spiders than ringer injected controls
(Krishnakumaran & Schneiderman 1970). However, ecdysis was abnormal and
mortality increased with injection of ecdysteroids. In addition, spiders ceased
feeding and spun a moulting pad instead of a normal orb web. In a later study by
Bonaric (1976) injection of high concentrations of 20E induced a high incidence
of moulting (81%) whereas low concentrations induced a lower incidence of
moulting (60%) in Pisaura mirabilis. However, the high doses of 20E injection
induced abnormal ecdysis with no apolysis and high mortality. The timing of
injection is another important factor. Injection of 20E into P. mirabilis during
the early stages of the 7-9th instar nymphs showed low concentrations of
ecdysteroids induce a prolonged instar, whereas injection into spiders during the
middle stages induced premature moulting (Bomaric 1976). Based on these early
studies, 20E induces spider moulting and also affects behavioural changes and
these effects are dose and time dependent. Subsequently, Bonaric and De Reggi
(1977) measured the ecdysteroid titers in 8th instar nymphs of P. mirabilis. The
8th nymphal stage lasted for 22 days and the 20E titers remained at low levels
during the intermoulting phase. However, the 20E titers rapidly peaked just
184
before ecdysis. A rapid large peak of 20E titers just before ecdysis is similar to
that seen in numerous other arthropods. In addition, Trabalon et al. (1998, 2005)
have shown that hormonal regulation is also related to secretion of pheromones,
sexual behaviour and cannibalism in T. atrica.
Understanding of the mechanisms regulating moulting in insects has
progressed in recent years through studies on ecdysteroid function at the
molecular level. These studies have established that ecdysteroids require two
nuclear receptors, the ecdysteroid receptor (EcR) and retinoid X receptor (RXR
a homologue of insect ultraspiracle) to function (Yao et al. 1992, 1993; Thomae
et al. 1993). EcR and RXR form a heterodimer and the heterodimer binds
ecdysteroids (E) to form a functional complex (E/EcR/RXR) for the induction of
transcription in target genes. E/EcR/RXR binds to the gene regulatory region,
ecdysone response element (EcRE), of the target gene to trigger gene
transcription. In addition, specific target genes called ecdysone early genes are
induced following transactivation to further regulate responses to the
E/EcR/RXR complex. The E75 gene is one ecdysone early gene that is important
in the regulation of cuticle formation (Hiruma & Riddiford 2009). Therefore,
ecdysteroid dependent regulation of moulting requires ecdysone, EcR, RXR and
early genes. Ecdysteroid dependent ecdysis is common in arthropods and
identification of EcR, RXR and early genes are reported from numerous other
arthropods (Nakagawa & Henrich 2009). In Chelicerata, EcR and RXR have
been identified from ticks (Guo et al. 1997, 1998; Horigane et al. 2007, 2008)
and scorpions (Nakagawa et al. 2007). These EcRs and RXRs from the
chelicerates show similar affinities to ecdysteroid and transcriptional activity in
insects (Guo et al. 1998, Nakagawa et. al. 2007). These results indicate ecdysis
of chelicerates is also regulated by E/EcR/RXR and early genes. However,
identification and functional analysis of ecdysteroids, EcR, RXR and early genes
have not been done in any spider. To understand spider moulting, clarification of
the regulation of ecdysis by ecdysteroids, EcR and RXR are essential. In
addition, this will contribute to understanding the hormonal regulation of other
physiological processes and behaviours such as pheromone secretion, and
feeding, web spinning and sexual behaviours of spiders. Therefore, in this study
we identified and analyzed EcR, RXR and E75 from the spider Agelena
silvatica.

A. silvatica, a common web-weaving spider in Japan, used in this study
was collected from Tsukuba, Japan and reared in our laboratory. 5’ and 3’ rapid
amplification of cDNA ends (RACE) were performed to determine the full
length of the EcR, RXR and E75 cDNA sequences as well as an actin sequence.
The degenerate primers were designed from other arthropods and the sequences
were determined by standard subcloning and sequencing techniques. Homology
analyses were performed with pairwise alignment by water program in
EMBOSS. Expression patterns of EcR and RXR were determined during
moulting by daily extraction of total RNA from the whole bodies of A. silvatica
3rd instar nymphs, synthesis of cDNA by reverse transcription and polymerase
185
chain reactions (PCR). Actin from A. silvatica also determined in this study was
used as an internal control.

In this study, full length sequences of EcR and RXR were identified from
A. silvatica (AsEcR and AsRXR). Nuclear receptor family genes have a
common structure with a DNA binding domain (DBD) and a ligand binding
domain (LBD). DBD is required to bind with the specific sequence (EcRE) of
target genes for gene transcription and is conserved among arthropod species.
LBD is necessary to bind the hormone and is also conserved among arthropods.
DBD of AsEcR showed high homology with insects, crustaceans and other
chelicerate species, while the LBD of AsEcR showed high homology to tick and
scorpion but low with insects. As observed in AsEcR, AsRXR also showed high
DBD identities with other arthropods and LBD homology was high with other
chelicerates but low with insects. The homology analysis indicates that spider
EcR and RXR can form a heterodimer and bind to EcRE, and the LBD of
AsEcR has a similar affinity to bind 20E for transactivation of target genes.
We also identified E75 from A. silvatica (AsE75) and there are at least two
isoforms with different sequences. One isoform has all E75 domains conserved,
while the other isoform lacks the DBD. AsE75 has a high homology with the
DBD and LBD of other arthropods. Therefore, the regulation of gene
transcription by ecdysteroids appears to be conserved in spiders. Subsequently,
the expression of AsEcR and AsRXR were determined for 3rd instar A. silvatica
nymphs. Both AsEcR and AsRXR mRNA expression were observed throughout
the 3rd instar nymphs indicating AsEcR and AsRXR are present to regulate
ecdysis of spiders when 20E titers increase.
In this study, we characterized EcR, RXR and E75 transcriptional factors
of A. silvatica. This is the first identification of these genes from spiders and
indicates the moulting mechanism is widely conserved in chelicerates,
crustaceans and insects. Ecdysteroids are also indicated to be important in other
physiological processes and behaviour of spiders such as pheromone secretion,
feeding behaviour, web spinning behaviour, sexual behaviour and cannibalism.
Therefore, further elucidation of the mechanisms by which ecdysteroids affect
the physiology, ecology and behaviour of spiders will greatly contribute not only
to understanding spiders themselves but also understanding the evolution of
hormonal regulatory mechanisms in arthropods.
References
Bonaric J.C. & De Reggi M. 1977. Changes in ecdysone levels in the spider Pisaura
mirabilis nymphs (Araneae, Pisauridae). Experientia, 33: 1664-1665.
Bonaric J.C. 1976. Effects of Ecdysterone on the Moulting Mechanisms and
Duration of the Intermolt Period in Pisaura mirabilis Cl. General and
Comparative Endocrinology, 30: 267-272.
Chapman R.F. 1998. The Insects: Structure and Function, 4th ed. Cambridge
University Press, Melbourne, pp. 363-412.
186
Guo X., Harmon M.A., Laudet V., Mangelsdorf D.J. & Palmer M.J. 1997.
Isolation of a functional ecdysteroid receptor homologue from the ixodid
tick Amblyomma americanum (L.). Insect Biochemistry and Molecular
Biology, 11: 945-962.
Guo X., Xu Q., Harmon M.A., Jin X., Laudet V. & Mangelsdorf D.J. 1998.
Isolation of two functional retinoid X receptor subtypes from the Ixodid
tick, Amblyomma americanum (L.). Molecular and Cellular
Endocrinology, 139: 45-60.
Hiruma K. & Riddiford L.M. 2009. The molecular mechanisms of cuticular
melanization: the ecdysone cascade leading to dopa decarboxylase
expression in Manduca sexta. Insect Biochemistry and Molecular Biology,
39: 245-253.
Horigane M., Ogihara K., Nakajima Y. & Taylor D. 2008. Isolation and
expression of the retinoid X receptor from last instar nymphs and adult
females of the soft tick Ornithodoros moubata (Acari: Argasidae). General
and Comparative Endocrinology, 156: 298-311.
Horigane M., Ogihara K., Nakajima Y., Shinoda T. & Taylor D. 2007. Cloning
and expression of the ecdysteroid receptor during ecdysis and reproduction
in females of the soft tick, Ornithodoros moubata (Acari: Argasidae).
Insect Molecular Biology, 16: 601-612.
Krishnakumaran A. & Schneiderman H.A. 1968. Chemical Control of Moulting
in Arthropods. Nature, 220: 601-603.
Krishnakumaran A., & Schneiderman H.A.1970. Control of molting in
mandibulate and chelicerate arthropods by ecdysones. The Biological
Bulletin, 139: 520-538.
Nakagawa Y., Henrich V.C. 2009. Arthropod nuclear receptors and their role in
molting. The FEBS Journal, 276: 6128-6157.
Nakagawa Y., Sakai A., Magata F., Ogura T., Miyashita M., Miyagawa H. 2007.
Molecular cloning of the ecdysone receptor and the retinoid X receptor from
the scorpion Liocheles australasiae. The FEBS Journal, 274: 6191-6203.
Thomas H.E, Stunnenberg H.G. & Stewart A.F. 1993. Heterodimerization of the
Drosophila ecdysone receptor with retinoid X receptor and ultraspiracle.
Nature, 362: 471-475.
Trabalon M., Niogret J. & Legrand-Frossi C. 2005. Effect of 20-
hydroxyecdysone on cannibalism, sexual behavior, and contact sex
pheromone in the solitary female spider, Tegenaria atrica. General and
Comparative Endocrinology, 144: 60-66.
Trabalon M., Pourié G. & Hartmann N. 1998. Relationships among cannibalism,
contact signals, ovarian development and ecdysteroid levels in Tegenaria
atrica (Araneae, Agelenidae). Insect Biochemistry and Molecular Biology,
28: 751-758.
187
Yao T.P., Forman B.M., Jiang Z., Cherbas L., Chen J.D., McKeown M.,
Cherbas P. & Evans R.M.1993. Functional ecdysone receptor is the
product of EcR and Ultraspiracle genes. Nature: 366, 476-479.
Yao T.P., Segraves W.A., Oro A.O., McKeown M. & Evans R.M. 1992.
Drosophila ultraspiracle modulates ecdysone receptor function via
heterodimer formation. Cell, 71: 63-72.
188
Phylogeny of the spider family Pimoidae (Araneoidea)

Gustavo Hormiga1* & Dimitar Dimitrov1,2
1
2
Zoological Museum, University of Copenhagen, Copenhagen, Denmark,
dimitard.gwu@gmail.com
*
Presenting author
The spider family Pimoidae comprises four genera and thirty seven extant
species. Members of this relictual lineage are known from Western North
America, Southern Europe and Asia (the Himalayas, China, Japan and the
Sakhalin islands). Six fossil species of pimoids have been described from Baltic
amber. We will present recent progress towards a phylogeny of Pimoidae based
on morphological and nucleotide sequence data.
189
Possible evolutionary causes of sexual size dimorphism

in giant wood spider Nephila pilipes (Nephilidae)
Chueh Hou
Department of Life Science, Tunghai University, Taiwan, emma200057@hotmail.com
Extreme female-biased sexual size dimorphism is generally assumed to result

from female gigantism as a result of selection for fecundity. However, the mechanisms that
maintain male’s smaller size should still be considered. Scramble and sperm competition
are considered two potential evolutionary causes of male dwarfism, driving extreme sexual
size dimorphism. Each hypothesis is supported by different studies, but relevant empirical
evidence from the field is still rare. The giant wood spider Nephila pilipes, widely
distribute in E and SE Asia, has extreme sexual size dimorphism, making it a suitable
organism for studying this topic. In this study, I investigated pre- and post-copulatory
male-male competition to realize potential selection pressures on N. pilipes males. Results
of an intensive field census revealed a low intensity of male-male competition. Female N.
pilipes changed their web locations frequently and moved long distances, which made it
difficult for small males to locate them. Low recapture of marked males also indicated high
mortality of males during mate searching. Moreover, preliminary results from laboratory
mating trials showed a tendency of first male sperm priority in N. pilipes. All these results
suggest that scramble competition and sperm competition exert selection pressure to
shorten the juvenile stage, enhancing the probability of a male being the first to mate with a
receptive female.
190
Effect of grassland burning on spider assemblages

in Terai, India
Upamanyu Hore & Virendra Prasad Uniyal
Wildlife Institute of India
Annual low-intensity fire is a conspicuous management strategy in

virtually all floodplain grassland of protected areas in India. While it is primarily
used to reduce fuel levels and to facilitate regeneration of desire species for wild
ungulate communities, little is known about the effects of its repeated use on
natural ecosystems over long periods of time. The increased use of prescribed
fire generates questions regarding the effects of burning events on spider
assemblage, and recovery of these grassland spiders following fire disturbance.
In this study we describe the ecological consequences of burning tall grass of
Terai on spider assemblages at different seasons and with different frequencies.
The study was conducted at Dudhwa National Park which represents one of
extensive tall grassland in Terai Region. Sixteen grassland sites from burnt and
unburnt areas in both grassland types were sampled from October 2006 to
August 2007, representing 4 seasons of sampling. During sampling seasons a
total of 8 sampling sites were established in each of the lowland and upland
grassland habitats; 4 on burnt areas and 4 on unburnt areas. Further, within burnt
areas we assessed two fire regimes for their impact on grassland spider
assemblage: (i) single fire, sites currently under management practices, burnt
annually early in the dry season (January - February); (ii) repeated fire, sites
burnt multiple times (as commonly occurs as uncontrolled) wildfires before the
end of the dry season (January - May). At each site, ten plots were randomly
established. Each plot consisted of a transect containing six sampling points at
approximately 10 m intervals. These six points along transect were used for both
spider sampling and grassland microhabitat assessment. Spiders were sampled
by pitfall trapping and sweep netting methods and for each fire regime spider
abundance, richness, diversity and evenness were calculated. Grassland
characteristics were also delineated to measures impacts of the prescribed fires
and to assess variability and heterogeneity of the grassland environment. A
total of 10,172 individuals were collected during the entire sampling period,
represents 98 species belonging to 58 genera and 22 families. Following
burning in this Terai grassland, we found the effects of fire on spider
assemblages varied with habitat type, frequency of burn and marginally with
seasons of burn. The differential response is likely to be related to differing
levels of habitat change in the two grassland types following fire. Spider
assemblages are mostly affected by fire because of fire-induced habitat
modification, altering microhabitats, resource availability and even
interspecific relationships. Combining habitat variables of two grassland types
to contrast similarity between patterns explained by best variables for all
spiders found highly correlated with grass cover (rho=0.78), soil moisture
191
(rho=0.65) and grass height (rho=0.64). Results from this study indicate there
can be great variation in response to fire; spider assemblages in upland grassland
habitats being less resilient to fire than those in lowland grassland habitats and
this has implications for the scale at which current fire management is
implemented. In the study we also found species strongly associated with
particular fire regime and rarefied species richness was higher at single fire sites,
represented high diversity compared to unburnt sites. This study highlights the
importance of considering habitat types and sensitivity to fire when burning for
biodiversity conservation, and cautions against applying prescribed fire in a
‘blanket-fashion’ across the conservation area. Adaptive management of
appropriate fire prescription should be taken account to provide wide range of
microhabitats that support a large proportion of species and to meet conservation
efforts for these grassland spider assemblages.
192
Effects of urbanisation on ground-dwelling spiders along a

rural-suburban-urban lowland forest gradient in Hungary
Roland Horváth1, Csaba Szinetár2, Tibor Magura3

& Béla Tóthmérész1
1
University of Debrecen, Department of Ecology, Hungary, horvathr@tigris.unideb.hu
2
University of West Hungary, Department of Zoology, Hungary
3
Hortobágy National Park Directorate, Hungary
We studied the effects of urbanisation on ground-dwelling spider

assemblages (Araneae) along a rural-suburban-urban forest gradient in Debrecen
(Hungary). We collected spider species with pitfall traps every two weeks from
the end of March to the end of November, 2001. We tested several hypotheses
which explain the effects of urbanisation (increasing disturbance hypothesis,
habitat alteration hypothesis). We also investigated the relationships between the
abundance of spiders and certain environmental variables (ground temperature at
2 cm depth, air temperature on the soil surface, relative humidity on the soil
surface and percentage cover of leaf litter, decaying wood material, herbs,
shrubs and canopy cover) along the urbanisation gradient. During the statistical
analyses we grouped the collected spider species according to their habitat
affinity (forest, generalist and open-habitat species). We found that overall
spider species richness was significantly higher in the urban sites than in the
suburban and rural ones. The higher diversity was due to the significantly more
open-habitat species in the urban sites. This result suggests that species living in
the surrounding matrix (grasslands and arable lands) penetrated the disturbed
urban sites. The ratio of forest species was significantly higher in the rural sites
compared to the suburban and urban ones, suggesting that forest species are
sensitive to the disturbance. Canonical correspondence analysis showed that the
species composition changed greatly along the urbanisation gradient. Open-
habitat species were associated with the urban sites where there was higher
ground and air temperature. Forest spiders were characteristic of the rural sites
with higher amount of decaying woods. Our findings suggest that the overall
diversity was not the most adequate indicator of disturbance; species with
different habitat affinity should be analyzed separately to get an ecologically
relevant picture of the effects of urbanisation.
193
Sampling and estimating spider diversity

in an alpine environment
Hubert Höfer, Theo Blick, Christoph Muster & Detlev Paulsch
Staatliches Museum für Naturkunde Karlsruhe, Germany, hubert.hoefer@smnk.de
We sampled spiders at an alpine meadow (1430-2000 m a.s.l.) in the

Allgäu Alps, Bavaria, Germany during a 6-year-project (2003-2009) with pitfall
traps operating during the vegetation period from early June to late September.
The area is known for its extraordinary botanical diversity originating
from the specific edaphic conditions: deep humid non-calcareous soils on
laminated Jurassic marl. These mountains attract attention due to flower-rich
mat-grass vegetation on their steep flanks up to the summits. However,
vegetation in the study area had suffered considerable alteration by long lasting
intense sheep grazing. The area along the ridge was used by sheep to lair and
therefore became strongly eutrophic and dominated by the grass Deschampsia
cespitosa. This type of land-use ended in 2000 and instead a controlled grazing
by cattle was realized with the objective to regenerate the species-rich alpine
mat-grass (Nardus stricta) vegetation.
For our study six pitfall traps were installed every year in each of 16
permanent plots in the grazed Nardetalia grassland and in 22 plots, presenting
other vegetation types.
We captured 81,700 (69,480 adults) spiders and identified 158 species.
We observed an extreme dominance (85%) of four lycosid species originating
from an extremely high activity density of the males of these species during the
first two weeks after snow melt. Species richness at single plots varied between
11 and 36 with a mean of 19 species per year. Pooled over 6 years between 22
and 54 species were captured at single plot. The most species-rich plots were in
the lower calcareous grassland with dwarf pine, in ungrazed (and thermally
favoured) plots, but also in the strongly altered (eutrophic and botanically
degraded) ridge area.
We were interested to know how often and how long sampling with pitfall
traps should be done to get a good estimate for the diversity of the spider
assemblages. Several species estimators (ICE, Chao 2, Jackknife 2, Michaelis-
Menten) were used to compare observed and estimated species richness for all
and several subsets of samples. With the total data set, the Chao 2 and ICE
estimators resulted in values very close to the observed ones, whereas Jackknife
2 seemed to overestimate and Michaelis-Menten underestimate species richness.
For assessing completeness of sampling in the main habitat type we divided the
estimated species number of subsets by the observed species number of all
samples for this habitat. Resulting values should be close to 100%. Here we
present only data estimated by Chao 2. For the characteristic vegetation type
mat-grass sward 116 spider species were observed during the whole
investigation and 138 estimated. This value is closest to the species richness
194
observed in the whole open grassland (136). Considering only the captures of the
two-week spring periods (over 6 years) 92 species (79% of all observed) were
observed and 118 estimated (102%). Considering only two spring periods 57%
of the overall observed species were captured and 72% estimated. Considering
samples of three two-week-periods from 2 consecutive years resulted in an
estimate of 91% of the observed species. Sampling one year during the whole
vegetation period from early June to end of September resulted in 96 species
(83%) and an estimate of 147 species (108% of all species observed in open land
and 127% of all species observed in all permanent plots).
195
Does equilibrium or non-equilibrium process shape

arboreal spider assemblage in the deciduous forest canopy?
Samuel Yu-Lung Hsieh & Karl Eduard Linsenmair
Department of Animal Ecology and Tropical Biology, University of Würzburg,

Germany, hsieh@biozentrum.uni-wuerzburg.de
The neutral theory and niche theory are still controversial and offer useful
perspectives regarding fundamental questions of ecology. However, we lack
even a basic understanding and universal model of how the community
organizations change within temporal and spatial resolution, even though this
information is vital for suggesting that equilibrium or non-equilibrium process
control biodiversity. Here we use null model to test whether arboreal spider
species in the European beech canopy have competitive interactions, while
controlling for temperature (seasons) and micro-environmental (canopy strata)
variables. We have also listed the co-occurrence species which have high niche
overlaps. These arboreal spider communities are assembled deterministically in
the cold season and stochastically in the warm season. The vertical stratification
of species composition between high canopies and low canopies occur only in
the warm season, however, they are competitive and do not demonstrate niche
segregation within these canopies. Using this data, we show that both
equilibrium and non-equilibrium patterns control the biodiversity in the
temperate forest and shape the arboreal spider communities which support the
predictions of the continuum hypothesis, which combine both neutral theory and
niche theory.
196
The arboreal spider diversity and community composition

change with growth of European beech: what does it mean
to forestry management and conservation?
Samuel Yu-Lung Hsieh & Karl Eduard Linsenmair
Department of Animal Ecology and Tropical Biology, University of Würzburg,

Germany, hsieh@biozentrum.uni-wuerzburg.de
European beech (Fagus sylvatica) is a deciduous tree species with high

economic value. Recently, the European system of environmental forestry
monitoring showed that the percentage of visible crown defoliation of beeches in
Germany has abruptly increased from 13% to 55% in the past 20 years, and it
caused that German forest enterprises had a low profitability due to low prices
for bad-quality timbers. The reasons are not only directly due to global warming,
but also to rising infestation by herbivorous insects. Hence an understanding of
the role of the forest canopy for diversity and community compositions of
insectivorous arthropods is urgently needed, to serve as an important reference
for further improving the forest management toward an even more nature-
friendly direction.
Würzburg University Forest (2664 ha) represents the type of a near-
primary mixed temperate forest in central Europe. Twenty-one percent of the
forest area here is covered by European beech, which is also the dominant
deciduous tree species distributed in the temperate region between southern
Europe and the British Isles. Six beeches in each of three classes categorized
according to age, height and trunk diameter at breast height respectively, have
been fogged on a monthly basis, with previously fogged trees excluded, always
sampling new trees (overall 162 trees were fogged). Arboreal spiders were
collected by insecticidal knockdown fogging from all strata of European beeches
for three consecutive summers. Analyses of species and guild composition
among various years, demonstrated significant differences. In particular, the
species composition in each summer showed a unique pattern. This reveals that
the community structures and diversities change dynamically from year to year,
therefore we made predictions based on spider composition and diversity during
different years. The Grey System helped us find the relationship between
changes of several diversity indices in consecutive years and did even forecast
the index values of the following two years. The Accumulated Generating
Operation complied with the Verhulst Grey Model even produced a high level of
accuracy for forecasting, in our study, some significant information appeared
only when the spider diversity under analysis was considered in light of different
periods of age classes. We found that, regardless of age of the canopies, the
model failed to show some crucial details: a declining trend in Fisher’s Alpha
index in the mature canopies was evident. One reason for this may be that in the
summer, the foresters carried out selective logging in mature tree areas. In some
cases there are some dominant beeches in the mature forest area, and their
197
presence oppresses the other neighbour beeches and results in bad-quality

timber. Thus, forest management policy is to cut the dominant beeches
selectively in order to let the other oppressed beeches grow straight up to ensure
better quality timber. If this policy continues, the prediction model shows that
the spider diversity will decrease in the mature forest. We suggest that mature
beeches not be logged in the summer for the purposes of natural habitat
protection. Furthermore, the arboreal spiders inhibit the population of
herbivorous insects in the forest, which is the essential way to protect not only
beech canopies, but also the forest ecosystem. Our results show that a true
picture at canopy level in the forests can only be drawn when also assessing
temporal dynamics from different years.
Currently, only 1% of Würzburg University Forest is covered by old-
growth beeches, which have a huge amount of herbivorous insects based on the
large volume of canopy particularly. However, comparisons of spider densities
per cube metre between three tree crown categories indicate that old-growth
canopy has in a significant reduction of spider abundance. It could explain why
old-growth canopy has the most serious problem of defoliation compared to the
other two age classes, if defoliation is considered being an effect of global
warming and increasing herbivorous insects. Nevertheless, old-growth canopy
has a distinctive species and guild composition of arboreal spiders. The
composition varies based on the different physical properties of the European
beeches and microclimate of the beech forests. Thus, for maintaining the unique
spider composition and abundance in higher strata of beech canopy, the old-
growth forest should not be disturbed or over-logged, so that the arboreal spiders
can inhibit the increasing herbivorous insects and reducing the beech defoliation.
198
Reconstructing the pholcid tree: a progress report

Bernhard A. Huber
Alexander Koenig Research Museum of Zoology, Bonn, Germany,

b.huber.zfmk@uni-bonn.de
Despite considerable progress during the last decade, the relationships

among the currently 86 pholcid genera are still far from settled. Some major
groups are fairly well supported both by morphological and molecular data, but
several large polytomies and doubtful nodes remain. The effort to place all
nominal genera into a cladogram must be seen before this background, as a
working hypothesis directing attention both to some major clades considered
likely to remain stable and to some of the most relevant and urging open
questions. A broad and extensive DNA sequencing effort will probably be
necessary for the next major step forward, but the lack of suitable material from
many taxa is a major obstacle to this proposition.
Beating the world record: diversity and endemism

of pholcid spiders in Brazil’s Atlantic Forest
Bernhard A. Huber
Alexander Koenig Research Museum of Zoology, Bonn, Germany,

b.huber.zfmk@uni-bonn.de
A worldwide comparison of published and unpublished species counts per

locality reveals that only 11 localities so far are known to contain more than 10
pholcid species each. Almost half of them (five) are located in the Serra do Mar
region of the Brazilian Atlantic Forest, including the current record of 15 species
at the Reserva Ecológica de Guapiaçú in the state of Rio de Janeiro. Recent
efforts to collect all species at six Atlantic Forest sites resulted in a total of 39
species, 22 of which were new, and 24 of which were found at only one locality
each. The two extreme (northern and southern) localities did not share any
species, suggesting a high level of endemism and immense unknown species
diversity. The dominant genera are Metagonia, Mesabolivar, Carapoia, and
Tupigea, with the last genus being endemic to the Atlantic Forest.
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3D data on the circulatory system in Cupiennius salei

Keyserling (Araneae: Ctenidae)
Katarina Huckstorf & Christian S. Wirkner
Allgemeine & Spezielle Zoologie, Institut für Biowissenschaften, Universität Rostock, Germany,
kati.huckstorf@gmx.de, christian.wirkner@web.de)
In terms of the central nervous system, behaviour and physiology,

Cupiennius salei is one of the best investigated spider species. However, parts of
its anatomy have only been described superficially to date. In the course of a
comparative study of the hemolymph vascular system in Araneae we
investigated the circulatory system of Cupiennius salei.
Like all other Arthropoda, spiders possess an open circulatory system. A
tubular heart lies dorsally in the opisthosoma. Hemolymph enters it via ostia,
and laterally paired cardiac arteries emanate from it. Anteriorly, the heart is
extended by an anterior aorta which runs through the petiolus and splits off into
two arteries (trunci peristomacales) within the prosoma. Each truncus branches
off into two arteries - the arteria cephalica and the arteria crassa. The arteria
crassa opens out into two sinus thoracales, from which vessels emanate which
supply the pedipalps, the four legs pairs and the central nervous system. The
arteria cephalica supplies, amongst other organs, the supraesophageal ganglion,
the eyes, the chelicerae and the mouthparts via several vessels.
Here we present the first three-dimensional data relating to the circulatory
system in Cupiennius salei, obtained by combining semi-thin sections and micro
computer tomography (microCT). Prior to microCT scanning specimens were
injected with a casting resin to enhance contrast on the hollow arteries. Data are
compared with existing literature on the circulatory system in Araneae.
200
Investigation of spiders overwintering in land-snail shells

in Southern Moravia (Czech Republic), with special regard
to importance of snail species 5
Vladimír Hula, Jana Niedobová, Václav Psota & Ondřej Košulič
Department of Zoology, Mendel University, Brno, Czech Republic, hula@mendelu.cz
Spider overwintering in land snail shells is poorly known. In Europe only

several studies from Germany, Hungary and recently from the Czech Republic
focused on this theme. On the conference in Alexandroupolis, Greece we
presented preliminary results of our investigation and now we would like show
more accurate data.
We collected more than 8000 land-snail shells in more than 50 different
xeric habitats – nature reserves, roadsides, quarries and on vineyards terraces at
southern Moravia (Czech Republic).
Collected shells we stored in controlled laboratory conditions (25°C, 50%
humidity) in plastic bags. We collected all emerging spiders. The juveniles were
kept in laboratory till maturity (mainly Heliophanus spp., Cheiracanthium spp.,
Xysticus spp. and Gnaphosidae s.l.). Particular species of land-snail shells were
investigated separately (Helicella obvia, Helix pomatia and Cepea
vindoboniensis).
Preliminary results show that different snale species host different species
of spiders, Pellenes nigrociliatus and Sitticus pennicillatus are found in Helicella
obvia; Euryopis quinquegutatta, Pellenes tripunctatus and Myrmarachne
formicaria in Cepea vindoboniensis. Several other spider species also show
similar relations.
The most common jumping spiders of genus Talavera (T. aperta, T.
aequipes, T. petrensis) do not prefer particular species. T. aequipes occupies all
three snail species, being the most frequent in Helicella. Specimens of this genus
can overwinter outside the shell.
Helix pomatia shells are not prefered by particular spider species. We have
found there mostly subrecedent common species such as Pardosa lugubris and
Diplostyla concolor.
In Cepea and Helicella shells we have found several rare species (e.g.
Cheiracanthium pennyi, Steatoda albomaculata, Micaria formicaria etc.).
5
The study supported by grants IGA-AF-MZLU-SP2100101/224 and VaV-MZP-CR-
SP/2D4/59/07.
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Subterranean arachnids of the Western Italian Alps

(Arachnida: Araneae, Opiliones, Palpigradi,
Pseudoscorpiones)
Marco Isaia
Dipartimento di Biologia Animale e dell’Uomo, Università di Torino, Italy,

marco.isaia@unito.it
The presentation shows the results of five years of work dedicated to the
Subterranean Arachnids of the Western Italian Alps. The work, that has just been
published by the Natural Museum of Torino (NW-Italy), is based on unpublished
material collected by Enrico Lana and Marco Isaia throughout an intense field work
from 2005 to 2010, on literature records and on the complete revision of the material
cited in the previous regional catalogue of the cave-dwelling spiders of Piemonte.
The work is the outcome of a fruitful collaboration of several European
arachnologists, who identified different arachnid groups. A special mention to Axel
L. Schönhofer (Germany) and Erhard Christian (Austria) for their contributions on
harvestmen and palpigrades, respectively.
Scorpions and mites have not been considered. The exclusion of scorpions is
justified by the trogloxenic life of the few species recorded in the studied area.
Concerning mites, records of an extremely specialized species of Troglocheles
(Prostigmata: Rhagidiidae) will be published separately.
The work covers 366 subterranean cavities, most of them located in province
of Cuneo (166), followed by Torino (69), Biella (38), Vercelli (30), Aosta (22),
Novara (21), Verbania (15), and Alessandria (5). We present 104 species (74
spiders, 14 harvestmen, 2 palpigrades, and 14 pseudoscorpions). Twelve (4 spiders,
2 palpigrades, and 6 pseudoscorpions) are considered as troglobiont on the basis of
obvious troglomorphy, 28 as troglophilic (20 spiders, 6 harvestmen, and 2
pseudoscorpions). For each species identification aids are provided, including
several illustrations of diagnostic features (original drawings), in situ photographs,
maps of the hypogean localities in the Western Italian Alps, large-scale distribution
and considerations from the ecological and faunistic points of view.
Spiders represent the major order of arachnids recorded in the study area, with
Meta menardi and Metellina merianae being the most abundant, followed by
Nesticus eremita, Malthonica silvestris, Pimoa rupicola and Troglohyphantes
lucifuga. The latter two are endemic to the southern and northern sectors of the
Western Alps respectively. Most remarkable are the troglobiont species of the genus
Troglohyphantes (T. konradi, T. pedemontanus, T. lanai) that also show, together
with the troglophilic T. bornensis, T. nigraerosae and T. pluto, the most restricted
distributions. Another interesting species is the troglobiont Nesticus morisii, only
known from the type locality in the Maritime Alps. Meta bourneti, a troglophilic and
markedly thermophilous species of Turanic-European-Mediterranean corotype, is
confined, in NW Italy, to one cave in the province of Cuneo and a few caves in Susa
Valley (province of Torino).
202
Among opilionids, the most interesting taxa are Holoscotolemon oreophilum,

three species of Ischyropsalis, and Leiobunum religiosum. All of these species show
restricted distribution and a strong relation with subterranean habitats.
Palpigrades represent the flagship of this work and of the entire
arachnological fauna of the Western Italian Alps. The data presented in the book are
of outstanding significance. According to current knowledge, the south-western part
of the Alpine chain houses Eukoenenia bonadonai and E. strinatii. Both species
belong to the spelaea/austriaca complex and show highly developed troglomorphic
features. Records are from three caves in the province of Cuneo.
Despite the lack of details and the difficulty in finding updated information on
the Western Alpine species, the pseudoscorpion section is mainly based on literature
data. Several species such as Pseudoblothrus peyerimhoffi, P. ellingseni, Chthonius
italicus, C. troglophilus, and Neobisium zoiai deserve special attention for their
pronounced troglomorphy and the restricted distribution.
With respect to overall arachnid species richness, the most important caves
are located in the Alpine districts of Alpi Marittime and Alpi Liguri (province of
Cuneo) which may thus be considered as a hot-spot of biodiversity. Curiously, the
most interesting assemblage is found in an artificial cave, the abandoned military
bunker of Vernante (province of Cuneo), that houses 9 species of subterranean
arachnids. Among these, at least 6 taxa are extremely specialized and some of them,
like Troglohyphantes konradi and Nesticus morisii (for which the bunker is the
locus typicus), show restricted or punctual distribution.
The current state of conservation policy in the Western Italian Alps is also
discussed. Nearly one third (122) of the recorded caves are situated in Protected
Areas, but only in five cases the cave habitat (“8310, Caves not open to public”
according to 92/43 Habitat Directive) is mentioned in the official document. Despite
the presence of extraordinary biocoenoses and the proximity to protected areas, a
number of caves are still unprotected. Examples are the above-mentioned military
bunker of Vernante, which is also known for the presence of endemic cave-dwelling
insects, and many caves in the Southern and Western Alpine districts, which
likewise harbour numerous specialized endemics. The vulnerability of several
populations of species deserving protection measures, such as Ischyropsalis carli,
Nesticus morisii, Troglohyphantes pluto, Meta bourneti, Neobisium zoiai and
Pseudoblothrus peyerimhoffi, is highlighted.
203
How many species of Sibianor/Bianor are there

in Middle Europe?
Piotr Jastrzębski1, Barbara Patoleta1 & Robert Rozwałka2
1
Department of Zoology, University of Podlasie, Siedlce, Poland, pjast@ap.siedlce.pl
2
Department of Zoology, Institute of Biology and Earth Sciences of Maria Curie-
Skłodowska University, Lublin, Poland, arachnologia@wp.pl
From Middle Europe only two species of Sibianor, (or Bianor), S.

aurocinctus and S. tantulus have been recognized so far. During the field study
in different parts of Poland we have found a number of populations showing
some variety in genitalia, body coloration, hairiness and habitat preferences.
Further research has revealed the presence of five species, 2 of them new for
science.
The results suggest that the genus is still poorly studied and some species
are not recognized and classifies as aurocinctus.
It seems likely that Middle European Sibianor (Bianor) is a good example
of local speciation, the timing of which, biogeographical aspects and
relationships between particular species should be subjected to further
(molecular) analyses. Here, we diagnose, describe and illustrate the new species
and present preliminary remarks on genus’ distribution in Middle Europe.
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Distribution patterns of Afrotropical arachnids:

staying ahead of shifting baselines
Rudy Jocqué
Royal Museum for Central Africa, Tervuren, Belgium, rudy.jocque@africamuseum.be
Astride across the equator, with almost perfectly parallel vegetation zones
in the north and a mosaic in its southern part, Africa is the dreamt continent to
study distribution patterns. Although the inventory of arachnids is far from
complete and in certain areas hardly begun, for certain taxa the amount of
information is sufficient to infer a reasonably good picture of the distribution
and its origin. Thanks to recently developed GIS technology, it is now possible
to predict distributions on the base of a restricted number of locality data. Yet,
the rapid decline of natural habitats and the global climatic changes are bound to
alter the baseline and the meaning of a concept like endemicity, for those groups
that so far have only been collected occasionally. Litter dwelling
Cyphophthalmi, canopy living spiders and cryptic soil spiders are examples of
such animal groups.
On the base of reasonably well collected taxa several distribution patterns
can be recognized although the continental dynamics on a geological time scale
highly complicate the historical reconstruction of present day distributions.
Typical distributions are those from the West Equatorial lowlands, often
with relatives in the Neotropics, and the eastern African elements often with
congenerics in South East Asia. Some ancestral taxa from southern Africa,
considered as typical Gondwanan distributions are known to have close, often
congeneric relatives, on the southern tips of other continents. These taxa should
be qualified as Afro-temperate elements. The combination of Afrotropical and
Afrotemperate faunas has lead to the idea of the pear-shaped distribution of
biodiversity on the African continent, which has come about as a result of
mixture of both elements in Afromontane areas. For some rare taxa it is possible
to demonstrate the importance of refuge areas in periods of drought. Similar
information can serve to elucidate the origin of forest faunas in East and West
Africa.
Surprisingly, only few examples of transverse distributions along parallel
vegetation zones across the northern part of the continent are known and may be
symptomatic of the climatic changes in recent geological times.
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Pseudoscorpions (Chelonethi: Neobisiidae: Neobisium)

parasitized by mites (Acari: Trombidiidae: Trombidium)
Mark L.I. Judson1 & Joanna Mąkol2
1
UMR 7205, Muséum national d’Histoire naturelle, Paris, France, judson@mnhn.fr
2
Institute of Biology and Institute of Natural Sciences, Wrocław University of
Environmental and Life Sciences, Poland, joanna.makol@up.wroc.pl
The few previous records of mites parasitizing pseudoscorpions have only

identified the host or the parasite. Here we report new host-parasite associations
from Europe. An adult male of Neobisium bernardi bernardi Vachon, 1937,
from Ariège, France, was found to be parasitized by two larvae of Trombidium
brevimanum (Berlese, 1910) (new host and country record for T. brevimanum)
An adult male of N. carcinoides (Hermann, 1804), from Morud, Norway was
found carrying ten larvae of T. mediterraneum (new host and country record for
T. mediterraneum). These are the first cases in which both the mite and the
pseudoscorpion have been identified. An additional record of a larva of
Trombidium sp. (probably T. mediterraneum) on Neobisium fuscimanum (C.L.
Koch, 1843) is given from Lower Silesia, Poland. The stylostomes (feeding
canals) formed by the mites are similar to those produced in other hosts and the
engorged state of some of the larvae suggests that Pseudoscorpions represent
viable alternative hosts for Trombidium larvae, although the successful
completion of development has yet to be demonstrated. Scars in the pleurum of
the pseudoscorpions are interpreted as the result of attempts at fixation by the
mite larvae. The low frequency of parasitism of pseudoscorpions by mites is
discussed. It is suggested that this might be due to the fact that larval
Parasitengona fall well within the size range of pseudoscorpion prey and would
have difficulty approaching without being attacked. Another factor that might
reduce the vulnerability of pseudoscorpions to mite parasitism is their strategy of
moulting within silk nests.
206
Experiences with floating traps

Béla Kancsal1, Csaba Szinetár2 & Dorottya Angyal3
1
Pannon Univesity, Keszthely, Hungary, kabakpityoka@gmail.com
2
University of West Hungary, Savaria University Center, Zoological Department,
Szombathely, Hungary, szcsaba@bdf.hu
3
Pécs, Hungary, angyal.dorottya@gmail.com
Collection of spiders in wetland habitats, especially in reeds is often

difficult. During pitfall trap monitoring high water level often causes some
troubles. We tried to eliminate this problem by developing a new type of trap
which floats at the top of water. The cups which contained killing-fluid were
sunk into a polystyrene plate. Upthrust of water was taken out by lead-ballasts
which were fixed with plaster at the bottom of the cups. Researches were carried
out from April to November of 2009 on the southern coast of Lake Velencei in
the offshore reed of Chernel István Madárvárta (47°11'24"N, 18°35'4"E) which
belongs to Agárd town. The 26 km2 Lake Velencei is Hungary’s third largest
natural stagnant water. Besides floating traps traditional pitfall traps were also
used as references to make catching results comparable. A total 3008 specimens
were collected, representing 20 families and 67 species. 422 specimen was
juvenile. With the help of floating traps 816 specimens of 17 families and 49
species, while with the standard traps 1079 specimens of 15 families and 44
species were collected in this comparison. The number of common species was
34. Although more spiders were collected by control traps, species richness and
diversity were significantly higher when floating traps were used. NMDS and
cluster analysis represent that floating traps were separated well from standard
traps. The most abundant species were Pirata latitans (Blackwall, 1841) (33%),
Pardosa prativaga (L. Koch, 1870) (13%), Trachyzelotes pedestris (C. L. Koch,
1837) (7%) and Arctosa leopardus (Sundevall, 1833) (7%). Some interesting
species were also collected e.g. Argyroneta aquatica (Clerck, 1757), Trebacosa
europaea Szinetár & Kancsal, 2007 and Philodromus pulchellus Lucas, 1846.
We conclude that collecting arthropods living on soil surface in reeds with
floating traps proved to be more successful than with the traditional traps.
207
The spider fauna (Araneae) of Maldives Islands

in Indian Ocean
Sunil Jose Kanniparambil
Deva Matha College, Kerala, India, sunil32@gmail.com
Introduction
Maldives is a small archipelago supporting a rich biodiversity of
invertebrates. Although many reports were made on the terrestrial and aquatic
animals, the spider fauna of the archipelago is poorly investigated, with only one
paper (Pocock 1904), listing 19 species.
Because of its proximity to several lands masses such as India, Seychelles,
Australia and Madagascar, the fauna shows unique features. The aim of the
present paper is to provide a preliminary spider check-list and to form a basis for
further investigations.

Check-list is based on an examination of specimens collected by the
author (January 2007 - November 2008) and on literature data (Pocock 1904,
Platnick 2006). The classification of Araneae follows Platnick (2009). For each
species, information on guild structure, distribution and affinities are given. The
specimens were preserved in 75% ethanol. The exact geographical locations
were determined with the Global Positioning System hand unit (GPS). The
identification of spiders was done following Pocock (1900, 1904), Tikader
(1970, 1977, 1980, 1982), Koh (1996), Murphy & Murphy (2000) and
Dippenaar (2002).
The specimens are deposited in the Arachnological Collections of Deva
Matha College, Kuravilangad, Kerala, India.
Study area
The Maldives make a chain of 26 coral atolls, 80-120 km wide and 860
km long (7°6'35"N-0°42'24"S, 72°33'19"E-73°46'13"E), located on the 1600 km
long Laccadives-Chagos submarine ridge extending from the south-west coast of
the Indian sub-continent to the central Indian Ocean. It is believed that the
Maldives were formed about 65-225 million years ago in the Mesozoic Era
(Maniku 1990). The atolls comprise 1192 islands (from 0.5 km2 to around 5.0
km2) varying in location, topography, form and shape (from small sandbanks
with sparse vegetation to elongated strip islands). The maximum height is
around 3 meters, while some 80% of the land area is less than 1 meter above
mean high tide level (MHAHE 1999). Out of the 1192 islands, 199 are inhabited
and 87 have been developed as tourist resorts. The largest island is Gan (1°55'N,
73°32'5''E) in Laamu Atoll, with an area 5.16 km2 and most of the present study
was conducted in this island during the period January 2007 - November 2008.
The relative humidity ranges from 73% to 85%. Daily temperatures vary little
208
throughout the year with a mean annual temperature of 28°C. Average annual
rainfall varies from 1,407 mm to 2,707 mm between different atolls.
Results
Family diversity: 16 families are recorded from Maldives during the
study. Families like Araneidae (12 species), Salticidae (10 species),
Tetragnathidae (7 species) and Sparassidae (7 species) exhibit highest species
diversity. Theridiidae (2 species), Pholicidae (2 species) are also widely present
in the islands. Families like Barychelidae, Hersilidae, Desidae, Scytodidae,
Thomisidae and Uloboridae are represented by one species only.
Generic diversity: 34 genera are found in 16 families. Maximum generic
diversity is found in families like Araneidae (7), Salticidae (5), Tetragnathidae
(2), Pholcidae (2) and Sparassidae (2). Most genera discovered show affinities
with oriental region and are widely present in the Indian mainland. Genera like
Cyclosa, Cyrtophora (Araneidae), Hersilia (Hersiliidae), Pardosa (Lycosidae),
Artema, Crossopriza (Pholcidae), Bavia, Myrmarachne, Plexippus (Salticidae,
Tylorida (Tetragnathidae) are first records from Maldives.
Species diversity: 54 species are collected from Maldives during the
study. Genera like Neoscona (6), Tetragnatha (5), Oxyopes (3), Heteropoda (3),
Olios (3) shows highest diversity of species in the collection.
New records: The most striking feature of the spider fauna of Maldives
islands is the high number of new records. About 30 species recorded during the
study are new records to Maldives. Araneidae and Salticidae exhibit highest
number of new records. Similarly 10 genera recorded during the study are also
new to Maldives.
Functional groups: The collected spiders can be divided into six
functional groups (guilds) based on their foraging behaviour in the field (Uetz et
al. 1999). The dominant guild was of the orb web builders and it comprised of
20 species of spiders. Spiders of the families Araneidae, Tetragnathidae and
Uloboridae fall under this category. Spiders of the category Stalkers formed the
next dominant guild comprising of 13 species of spiders. Ground runners (10
species), scattered line weavers (4 species), ambushers (3 species) and Foliage
runners (1 species) are the other functional groups.
Endemism: A total of 50 species are discovered from Maldives so far.
Among the collection Heteropoda atollicola is endemic to Maldives. Desis
gardineri, and Tetragnatha foveata are also restricted to Laccadive and
Srilankan region.
Affinities: The present studies conducted in Maldives revealed that the
spider fauna of this ecosystem bears affinities with Oriental (21 spp), Australian
(3 spp.), Palaearctic (4 spp.) and Nearctic (1 sp.) regions. High number of Indian
species suggests the arrival of majority of spiders here from the neighbouring
Indian mainland.
Zoogeographic analysis: About 29 species recorded in Maldives are
widely distributed in South Asia; a few of these are found only in the Indo-
Srilankan region. Most of the widely distributed species in south Asia belong to
Araneidae (11 species) and Salticidae (6). Because of bright colouration and
209
large orb webs, spiders of the above mentioned families were easily observed.
Species like Crossopriza lyoni (Blackwall, 1867); Plexippus paykulli (Audouin,
1826) are cosmopolitan in distribution; whereas species like Artema atlanta
Walckenaer, 1837; Zosis geniculata (Olivier, 1789); Heteropoda venatoria
(Linnaeus, 1767) are pantropical in distribution.
Discussion
The spider fauna of Maldives is not rich compared with many other
tropical islands. Around 1447 species are reported from the neighbouring Indian
mainland and around 354 species are reported from Sri Lanka (Siliwal 2007).
The lack of high species diversity can be attributed to the limited diversity of
habitats in these coral islands. The limited floral diversity is also a contributing
factor in reducing the number of invertebrates. A notable feature in the diversity
of spiders is the higher family and generic diversity. Except the common
families like Araneidae and Salticidae most families are represented by a few
species. Seven families are represented by only single species. Rare families like
Desidae which are not found in neighbouring mainland are also recorded from
these coral islands.
The spider fauna here is a chance assemblage of species arrived from
neighbouring lands. Most species found here are also found in Indian mainland
and Sri Lanka, which shows the primary route of spider migration. The sub order
Mygalamorphae is represented by only a Barychelid species, Sason robustum.
The scarcity of mygalmorphs can be attributed to the vast separation of these
coral islands from the neighbouring land. Legendre (1979) suggested that in the
case of Sason, its arboreal nest allowed for its transport as flotsam in ocean
currents. Another notable feature in the spider fauna is the high number of
Tetragnathid spiders of the genus Tetragnatha observed during the study. These
are common in most areas; the frequent equatorial rain also favours the
abundance of moisture loving genera.
There are many environmental factors that affect species diversity
(Rosenzweig 1995). However, when spiders were divided according to their
functional group there was a significant effect of habitat on the diversity of these
groups. The web building and foliage running spiders rely on vegetation for
some part of their lives, either for finding food, building retreats or for web
building. The structure of the vegetation is therefore expected to influence the
diversity of spiders found in the habitat. Studies have demonstrated that a
correlation exists between the structural complexity of habitats and species
diversity (Hawksworth, Kalin-Arroyo 1995). Diversity generally increases when
a greater variety of habitat types are present (Ried & Miller 1989). Uetz (1991)
suggests that structurally more complex shrubs can support a more diverse
spider community. The lack of high diversity of spiders in Maldives has to be
viewed in this context.
210
Comparative morphology of the circulatory system

in scorpions: an evolutionary character analysis
Bastian J. Klußmann & Christian S. Wirkner
Allgemeine & Spezielle Zoologie, Universität Rostock, Germany,

Bklussmann@gmx.net, christian.wirkner@uni-rostock.de
Although scorpions are one of the better-known arachnid groups,

investigation of their anatomy has been superficial. Existing comparative work
on the circulatory system, one of the major organ systems, is preliminary to say
the least. To fill this knowledge gap the aim of this study is to describe and
visualize the circulatory systems of different scorpion taxa in detail. To this end
we compared the circulatory organ structures of nearly all the major scorpion
taxa (42 species) using corrosion casting, MicroCT in combination with
computer-aided 3D reconstruction, and scanning electron microscopy. The open
circulatory system of scorpions is made up of a hemolymph vascular system (i.e.
heart and associated structures) and a lacunar system (lacunae and sinuses). The
hemolymph vascular system in scorpions comprises a dorsal tubular heart which
extends the entire length of the mesosoma and is surrounded by a pericardium.
The heart is extended posteriorly by the posterior aorta which pervades the
metasoma to the aculeus. In the prosoma, there is an anterior aorta which
supplies the appendages (e.g., the chelicerae, the pedipalps and the walking
legs). There is also a dense meshwork of small hemolymph channels which
supply the central nervous system. The ventral vessel originates from the
anterior aorta and runs through the entire opisthosoma. Our study investigates in
detail the branching pattern of the anterior aorta and the characters of the heart
and associated structures (e.g. cardiac arteries and arrangement of the ostia). Our
results enable us to conceptualize a number of phylogenetic characters. These
are mapped onto existing hypotheses of scorpion phylogeny in order to trace
their evolutionary transformations.
211
The classical taxonomic approach towards a phylogenetic

system of the Cyphophthalmi (Opiliones)
Ivo M. Karaman
University of Novi Sad, Department of Biology and Ecology, Novi Sad, Serbia,
ivo.karaman@dbe.uns.ac.rs
There are a few as good land organisms suitable for studying historical
biogeography as Cyphophthalmi. Their ancient origin (Dunlop 2007),
conservative morphoanatomic characters, distribution, criptical way of life and
low dispersal rates recommend them for study.
From among 60 scientists that have been studying this group so far,
Christian Juberthie is the most prolific author. Following Hansen and Sørensen
(1904), he was the first to use the classical taxonomic approach to set the
foundation of modern taxonomy of Cyphophthalmi. Juberthie (and co-authors)
was the first to study the biogeography of Cyphophthalmi, including the plate
tectincs aspect (Juberthie & Massoud 1976). Juberthie (1988) also illustrated the
slow evolution of the group on the example of the genus Parasiro.
Equaly significant for our understanding of the diversity and distribution
of the group are the papers by Shear. In 1980 he proposed a reclassification of
Cyphophthalmi and provided a novel opinion of their relationships, based on
cladistic analysis of morphological characters. He also raised the group to the
family rank, describing it as a new taxo (Shear 1993).
Today the Cyphophthalmi are classified within 6 families: Sironidae
(Europe, North America, Japan), Pettalidae (temperate Gondwanan distribution),
Stylocelidae (great part of Indomalaya), Neogoveidae (Tropical South America,
Florida and West Africa), Ogoveidae (West Africa) and Troglosironidae (New
Caledonia).
A new period of the complex approach to the group, with cladistic
analyses of molecular and morphological data started with the papers by
Gonzalo Giribet and colaborators. Particularly important was the work on the
reconstruction of phylogenetic relationships and biogeographical studies (Giribet
& Boyer 2002, De Bivort & Giribet 2004, Boyer et al. 2007, Murienne et al.
2010, Clouse & Giribet in press). Recently, the Giribet’s group used also a
morphometric approach in an attempt to solve phyletic relationships among
certain groups of Cyphophthalmi (Clouse et al. 2009, De Bivort et al. 2010).
The classical taxonomic approach (recognition and assessment of
characters) has recently been wrongly neglected. Based on the analysis of a
number of representatives of various genera and families of Cyphophthalmi, I
have concluded that certain characters may clearly define families and phyletic
relations within them. Based on this analysis, Ankaratra franzi Shear & Gruber,
1996 from Madagascar clearly belongs to Ogoveid - Troglosironid clade. This
species shows affinity to Troglosironidae from New Caledonia, and this suggests
a possible connection of Cyphophthalmi from New Caledonia with Ogoveidae
212
from West Africa. Consequently, the synonymy of these two families is

possible.
Manangotria taolanaro Shear & Gruber, 1996, the second species from
Madagascar, and Karripurcellia sierwaldae Giribet, 2003 from Western
Australia, are congeneric, indicating that these territories are closely related.
The characters analyzed show that the family Stylocellidae is more
diversified than it was thought. Based on the same criteria that define other
families, a new family has to be raised.
Cladistic analyses of morphological characters may lead to a somewhat
superficial approach that may result in serious oversights.
References
Boyer S.L., Clouse R.M., Benavides L.R., Sharma P., Schwendinger P.J.,
Kuranarathna I. & Giribet G. 2007. Biogeography of the World: a case
study from cyphophthalmid Opiliones, a globally distributed group of
arachnids. Journal of Biogeography, 34: 2070-2085
Clouse R.M., de Bivort B.L. & Giribet G. 2009. A phylogenetic analysis for the
Southeast Asian mite harvestman family Stylocellidae (Opiliones,
Cyphophthalmi) - a combined analysis using morphometric and molecular
data. Invertebrate Systematics, 23: 515-529.
Clouse R.M. & Giribet G. in press. When Thailand was an island - the
phylogeny and biogeography of mite harvestmen (Opiliones,
Cyphophthalmi, Stylocellidae) in Southeast Asia. Journal of
Biogeography.
De Bivort B.L. & Giribet G. 2004. A new genus of cyphophthalmid from the
Iberian Peninsula with a phylogenetic analysis of the Sironidae
(Arachnida: Opiliones: Cyphophthalmi) and a SEM database of external
morphology. Invertebrate Systematics, 18: 7-52.
De Bivort B.L., Clouse R.M. & Giribet G. 2010. A morphometrics-based
phylogeny of the temperate Gondwanan mite harvestmen (Opiliones,
Cyphophthalmi, Pettalidae). Journal of Zoological Systematics and
Evolutionary Research.
Dunlop J.A. 2007. Paleontology. In: Pinto-da-Rocha R., Machado G. & Giribet
G. (eds), Harvestmen. The Biology of Opiliones. Harvard University
Press, Cambridge, MA, pp. 247-265.
Giribet G. & Boyer S. 2002. A cladistic analysis of the cyphophthalmid genera.
Juberthie C. 1988. Les opilions cyphophthalmes biogeographie, vittesse
d’evolution, periodes de colonisation du mileu souterrain. XI
Europaisches Arachnologisches Colloquium, Berlin, pp. 303-308.
Juberthie C. & Massoud Z. 1976. Biogeographie, taxonomie et morphologie
ultrastructurale des opilions cyphophthalmes. Revue d Ecologie et de
Biologie du Sol, 13(1): 219-231.
Murienne J., Karaman I. & Giribet G. 2009. Explosive evolution of an ancient
group of Cyphophthalmi (Arachnida: Opiliones) in the Balkan Peninsula.
Journal of Biogeography.
213
Shear W.A. 1980. A review of the Cyphophthalmi of the United States and
Mexico, with a proposed reclassification of the suborder (Arachnida,
Opiliones). American Museum Novitates, 2705: 1-34.
Shear W.A. 1993.The genus Troglosiro and the new family Troglosironidae
(Opiliones, Cyphophthalmi). The Journal of Arachnology, 21: 81-90.
Shear W.A. & Gruber J. 1996. Cyphophthalmid opilionids new to Madagascar:
two new genera (Opiliones, Cyphophthalmi ? Pettalidae). Bulletin of the
214
Diversity of the spider communities on the intensively

managed grasslands of the “Medvednica” Nature Park,
Croatia
Luka Katušić
Association for Biological Research – BIOM, Croatia, luka.katusic@biom.hr
A lot of studies have been done so far dealing with the impact of different
grassland management on the occurring arthropod communities. They show that
an extensive grazing or mowing has a positive effect on the grassland
communities by preventing succession, while intensive mowing or grazing
impoverishes the spider communities through decreasing vegetation diversity
and microhabitat availability (Cattin et al. 2003, Rothenbücher 2004, Plantureux
2005, Warui et al. 2005, Pétillon et al. 2007).
Grassland habitats of the mountainous “Medvednica” Nature Park are
threatened by the loss of the traditional agricultural activities and an increasing
number of Park visitors followed by the need to reform grasslands for
recreational purposes.
From June to October 2007 a research of spider communities was
conducted on four meadows of the “Medvednica” Nature Park with a main goal
to determine the influence of intensive management of meadows used for
recreational purposes (skiing) on spider communities. The aim was to detect
differences in the assemblages of spider communities on three frequently mowed
meadows and on meadow not used for recreational purposes and mowed two
times a year.
The spiders were collected by hand, exhauster, sweeping and pitfall
trapping. Only the data sampled by pitfall traps were statistically analysed. On
each meadow nine pitfall traps were placed within a 10 x 10 m plot.
Overall 5023 spider specimens were collected, with 3384 adult specimens.
99 species were identified, out of which 25 were recorded for the first time in
Croatian spider fauna (Helsdingen 2009). As a new species also the
allochthonous spider species Mermessus trilobatus (Emerton, 1882) was
recorded. As expected, the spider community of the control plot differed
significantly from the intensively managed meadows and had the highest
biodiversity. On these three disturbed plots the most abundant species were
Oedothorax apicatus and Erigone dentipalpis, abundant and ubiquitous species
on the all kinds of grassland habitats, frequent in habitats under anthropogenic
impact (Roberts 1987, Hänggi et al. 1995, Buchar & Ružička 2002, Nentwig et
al. 2003, Schmidt et al. 2008). On the control plot, species Alopecosa cuneata
was the most abundant. On the disturbed plots the most abundant family, both in
the number of specimens and in the number of species, was family Linyphiidae.
On the undisturbed plot family Lycosidae was most abundant.
Although a further research, based on a detailed analysis on environmental
variables and habitat preferences of the recorded species, is needed, this
215
preliminary research shows that intensive management of the meadows prevents

the development of the natural grassland communities and lowers the
biodiversity on such areas with the prevalence of species with broad ecological
valence.
References
Buchar J. & Ružička V. 2002. Catalogue of spiders of the Czech Republic, Peres
Publishers, Praha.
Cattin M.F., Blandenier G., Banašek-Richter C. & Bersier L.F. 2003. The impact
of mowing as a management strategy for wet meadows on spider
(Araneae) communities. Biological Conservation, 113: 179-188.
Hänggi A., Stöckli E. & Nentwig W. 1995. Habitats of Central European
Spiders. Miscelanea Faunistica Helvetiae, 4: 1-460.
Helsdingen P.J. van 2009. Araneae. In: Fauna Europaea Database (Version
2009.2), http://www.european-arachnology.org.
Nentwig W., Hänggi A., Kropf C. & Blick T. 2003. Spinnen Mitteleuropas -
Determination Key. Ver. 8.12.2003, http://www.araneae.unibe.ch/.
Pétillon J., Georges A., Canard A. & Ysnel F. 2007. Impact of cutting and sheep
grazing on ground-active spiders and carabids in intertidal salt marshes
(Western France). Animal Biodiversity and Conservation, 30: 201-209.
Plantureux S., Peeters A. & McCracken D. 2005. Biodiversity in intensive
grasslands : Effect of management, improvement and challenges.
Agronomy research, 3(2): 153-164.
Roberts M.J. 1987. The Spiders of Great Britain and Ireland. Volume 2:
Linyphiidae. Harley Books, Colchester.
Rothenbücher J. 2004. The Impact of Mowing and Flooding on the Diversity of
Arthropods in Floodplain Grassland Habitats of the Lower Oder Valley
National Park, Germany. Dissertation.
Schmidt M.H. Rocker S., Hanafi J. & Gigon A. 2008. Rotational fallows as
overwintering habitat for grassland arthropods: the case of spiders in fen
meadows. Biodiversity and Conservation, 17: 3003-3012.
Vickery J.A., Tallowin J.R., Feber R.E., Asteraki E.J., Atkinson P.W., Fuller
R.J. & Brown V.K. 2001. The management of lowland neutral grasslands
in Britain: effects of agricultural practices on birds and their food
resources. Journal of Applied Ecology, 38: 647-664.
Warui C.M., Villet M.H., Young T.P & Jocque R. 2005. Influence of grazing by
large mammals on the spider community of a Kenyan savanna biome.
216
Ultrastructural characterization of Hexisopus spermatozoa

in comparison to other solifugid species
Anja E. Klann1 & Tharina Bird2
1
Ernst-Moritz-Arndt-University Greifswald, Institute of Legal Medicine, Greifswald,
Germany, anja.klann@uni-greifswald.de
2
Department of Arachnology and Myriapodology, National Museum of Namibia,
Windhoek, Namibia; Department of Zoology, Denver Museum of Nature and
Science, Denver, CO, USA tharinab@gmail.com
Solifugae is one of the mesodiverse arachnid orders comprising 12

families and about 1087 species. The family Hexisopodidae, which occurs in
southern Africa, represents a very peculiar group differing from all the other
solifugid species. In contrast to all other species, representatives of this family
possess fossorial legs, rather than cursorial legs. The phylogeny of Solifugae is
widely unresolved. The ultrastructure of spermatozoa has successfully been used
for phylogenetic analyses in other animal groups. Therefore the question arose
whether the morphological peculiarity of the family Hexisopodidae is also
reflected in the ultrastructure of their spermatozoa. The sperm cells of a species
of the genus Hexisopus are aflagellate, roundish and their chromatin bodies are
irregularly shaped. The sperm cells form finger-like membrane protuberances
and each sperm is surrounded by a secretion sheath. Apparently neither in the
testes nor in the Vasa deferentia the spermatozoa aggregate to groups. The
acrosomal complex is relatively small. Overall the spermatozoa of this species
resemble most spermatozoa of certain species of the families Ammotrechidae,
Daesiidae and Solpugidae to a certain degree.
217
The impact of vegetation structure on weeds, ground

beetles (Coleoptera: Carabidae), arachnid communities
(Arachnida: Araneae, Opiliones): distribution patterns and
drivers for epigeic activity
Jessika Konrad
Institute for Land Use Systems, Leibniz-Centre for Agricultural Landscape Research
(ZALF), Germany, konrad@zalf.de
The aim of the study was to investigate the impact of vegetation structure
on species composition of ground beetle and spider assemblages, respectively as
well as its function as a driver for epigeic activity patterns. In 2008, field trials
were performed to analyse the communities of weeds, ground beetles
(Carabidae) and arachnids (Araneae, Opiliones) in three different conventionally
managed crop fields in Brandenburg, Germany. Each field was divided into
eight experimental plots. The crops cultivated were maize, peas, and winter rye,
respectively. The winter rye was intercropped with Sudan grass and mustard,
respectively, at the end of June. Vegetation structure was identified by the height
and cover values of the crop plants as well as of the weed. Ground beetles and
spiders were caught with pitfall traps in intervals of 14 days from April to
September and the species were identified in the laboratory. Vegetation surveys
on 72 sample plots on each field were performed. A total of 74 weed, 72 ground
beetle and 84 spider species were found. Analyses of the data with different
multivariate methods (Cluster analysis, discriminant analysis (DA),
correspondence analysis (CA) and canonical correspondence analysis (CCA)
revealed patterns of the structural parameters, as well as of the assemblages of
these species groups characteristic for each of the fields. These patterns
represent qualitative and quantitative differences in the assemblages of these
species groups, including species composition and -diversity as well as
dominance and biomass patterns, revealed to be stable on different spatial scales
(from single traps up to the field scale). After ploughing and seeding the
intercrops in two of the fields, rigorous species-specific changes in activity
patterns occurred. Vegetation structure was identified as a crucial driver for
epigeic activity patterns. The impact of the vegetation structure on spider
coenoses as a whole as well as on etho-ecological traits is discussed in relation
to some hypotheses of other authors about vegetation structure and animal
communities.
218
Arachnology in Finland. 2
Seppo Koponen
Zoological Museum, University of Turku, Finland, sepkopo@utu.fi
The history of arachnology in Finland has been earlier briefly described by

Terhivuo (1996) and Koponen (2008). In the present report, I will deal with the
Finnish arachnologists, who have worked either at the University of Helsinki or
at the University of Oulu during the second half of the 20th century. However,
Professor Pontus Palmgren (Helsinki), who’s well-known activity in
arachnology lasted from 1932 until 1983, was presented in an earlier report, and
he is now excluded. Sixteen researchers are dealt with, and a selection of 35
publications from their production is presented here. The most productive
arachnologists of this period are (in alphabetic order) Walter Hackman, Veikko
Huhta, Juhani Itämies, Jouko Kaisila, Aarno Kleemola, Timo Pajunen and
Juhani Terhivuo.
Spider fauna (Araneae) in the southwestern

archipelago of Finland
Seppo Koponen
Zoological Museum, University of Turku, Finland, sepkopo@utu.fi
About 450 species, ca 70% of the known Finnish spider fauna, are hitherto
collected from the southwestern archipelago of Finland. This large archipelago is
situated in the Baltic Sea, between Turku (Finland) and Stockholm (Sweden). It
consists of over 40 000 islands and islets of different size. The archipelago is,
due to land rising after the Ice Age (ca 0.5 m /100 years), rather young. This
report is based on the databases of the Zoological Museum, University of Turku.
Species diversity and faunal similarity in different parts of the archipelago are
discussed, and some interesting distribution patterns are shown. For example,
some continental or northern species, found in inner parts of the archipelago, are
absent on the large-size island of Aland, e.g. Alopecosa pinetorum. On the other
hand, some species are known in Finland only from Aland, e.g. Labulla
thoracica and Hyptiotes paradoxus. Many other spiders can be found only in
Aland and other outer parts the archipelago, not in the inner, coastal zone, e.g.
Amaurobius fenestralis, Pseudicius encarpatus and Lasiargus hirsutus. Many
species included in the national Red Data Book live in Finland only or mainly in
this archipelago area.
219
Host specific manipulation of spiders by a parasitoid wasp
Stanislav Korenko & Stano Pekár
Department of Botany and Zoology, Faculty of Science, Masaryk University, Brno,

Czech Republic, korenko.stanislav@yahoo.com, pekar@sci.muni.cz
Many parasites and parasitoids control behaviour of their host to achieve

more preferable conditions for their development. Here, we show how larva of
Zatypota percontatoria Müller, a narrowly specialised wasp parasitoid of
theridiid spiders, performs species-specific interaction with their hosts, Neottiura
bimaculata (Linné) and Theridion varians Hahn. The wasp larva modified the
web architecture of parasitised spiders shortly before pupating. In N. bimaculata
spiders the web architecture was modified in 19% (N=31). In remaining cases
the change in the structure was inconspicuous. The modified web had denser silk
strands around the position of spider in the web. In T. varians the web
construction was modified in 90% of cases (N=27). Parasitised T. varians spider
built a cupola-like web around the wasp larva that was used as a shelter for the
pupa. We found that unparasitised T. varians spiders construct such cupola-like
structure during overwintering in litter which seems to serve as a protection
structure against predators.
220
Cytogenetic data do not support a close relationship

between the families Mimetidae and Palpimanidae
(Araneae: Araneomorphae)
Tereza Kořínková1, Jiří Král1, Stano Pekár2 & Charles R. Haddad3
1
Department of Genetics and Microbiology, Faculty of Science, Charles University in
Prague, Czech Republic, korinko1@natur.cuni.cz, spider@natur.cuni.cz
2
Department of Zoology and Ecology, Faculty of Science, Masaryk University, Czech
Republic, pekar@sci.muni.cz
3
Department of Zoology & Entomology, University of the Free State, Bloemfontein,
South Africa, HaddadCR@ufs.ac.za
The systematic position the araneomorph family Palpimanidae and limits

of the superfamily Palpimanoidea are a matter of controversy. Palpimanids were
traditionally considered as entelegyne araneomorph spiders with secondarily
reduced female genitalia. However, the recent study of Huber (2004) showed
that the characters of both female and male copulatory organs of Palpimanidae
rather correspond to those of haplogynes. The families Stenochilidae and
Huttoniidae possess similar copulatory organ morphology and structure to
palpimanids. These families are currently grouped with Palpimanidae into the
superfamily Palpimanoidea. Palpimanidae and some other araneomorph families
possess peculiar morphological characters related to their araneophagy.
According to a concept, which emphasizes mainly these characters, up to 10
extant families belong to the superfamily Palpimanoidea. Two of them,
Archaeidae and Huttoniidae, are among the most ancient groups of araneomorph
spiders - their fossil record is known already from the Mesozoic. Following
several recent revisions, only the clade comprising the families Huttoniidae,
Palpimanidae and Stenochilidae is considered monophyletic. The remaining
families are considered to be related to the entelegyne superfamily Araneoidea.
Considering Palpimanoidea in the broad sense, only two species of the
family Mimetidae have till now been studied cytogenetically, namely Ero
japonica (2n=24, X1X20, Suzuki 1954) and Gelanor sp. (2n=25, X0, Schneider
et al. 2008). In contrast to this, karyotypes of 138 species of six families
classified as Araneoidea (sensu Coddington 2005) have been reported. As a
contribution to the discussion on the taxonomy of the superfamily
Palpimanoidea, we provide a comparative karyotype analysis of four species of
Palpimanidae, one species of Mimetidae, and two representatives of the
superfamily Araneoidea. Male diploid numbers and sex chromosome systems
were as follows: Palpimanus gibbulus from Greece 27 (X1X2X30), Palpimanus
schmitzi from Israel 28 (X1X2X3X40), Palpimanus cyprius from Israel 28
(X1X2X3X40) (Palpimanidae; Palpimaninae); Diaphorocellus sp. from Namibia
24 (X1X20) (Palpimanidae, Chediminae); Mimetus sp. 24 (X1X20) (Mimetidae);
Robertus lividus 22, (X1X20) (Theridiidae); Meta menardi 24 (X1X20)
(Tetragnathidae). In contrast to the overwhelming majority of entelegyne spiders
221
studied so far, the palpimanid karyotypes contain both acrocentric and biarmed
autosome pairs. Male meiosis of palpimanids is probably achiasmate. This mode
of meiotic division is characterized by a lack of crossovers and consequently by
the absence of chiasmata at prophase during the first meiotic division. Due to the
lack of chiasmata, paired homologous chromosomes maintain a parallel
alignment during late prophase I (the so-called postpachytene stage). The sex
chromosomes of palpimanids are heteropycnotic and more or less associated
during most of the first meiotic division. At early prophase I they lie at the
nuclear periphery. From postpachytene to metaphase I, they form a highly
condensed unit usually situated in the centre of the plate. Unlike the autosome
bivalents, the sex chromosome unit does not decondense during a particular
stage of transitional chromatin decondensation, which was observed between
pachytene and postpachytene. In contrast to palpimanids, males of Mimetus sp.
show a typical entelegyne karyotype formed entirely of acrocentric
chromosomes, including two X chromosomes. Mimetids exhibit standard
meiosis; each of the autosome bivalents usually bears a single chiasma. By
transit from premeiotic interphase to leptotene, the two non-homologous
gonosomes form a positively heteropycnotic body usually situated at the nuclear
periphery and persisting until the zygotene – early pachytene stage. At diplotene
the gonosomes become isopycnotic with the autosomes and later negatively
heteropycnotic and less condensed than the autosomes. At the beginning of
diplotene the X chromosomes pair end-to-end without chiasmata, whereas at
metaphase I they already lie separately. The karyotypes of M. segmentata and R.
lividus are similar to mimetid representative in terms of chromosome numbers
and morphology, sex chromosome system, course of meiosis and meiotic
behaviour of the sex chromosomes. The co-occurrence of biarmed and
acrocentric automosomes in a karyotype is a very unusual phenomenon feature
amongst entelegyne spiders. Within the frame of Opisthothelae, the presence of
biarmed chromosomes is probably, as indicated by our previous data. This could
support the position of Palpimanidae as one of the basal araneomorph clades.
However, the combination of a high proportion of acrocentric chromosomes and
the presence of multiple X chromosomes has not been found in previously
karyotyped basal araneomorphs (Haplogynae, Hypochiloidea or
Austrochiliodiea). Instead, these characters are rather typical for entelegynes and
would in fact support the placement of Palpimanidae amongst entelegynes or
proto-entelegynes. Within the genus Palpimanus, systems with three or four X
chromosomes seem to be a frequent phenomenon. Such systems are regarded as
derived in spider sex chromosome evolution. Achiasmate meiosis is another
derived character that has not been confirmed in any other spider group so far
except for some diplurid mygalomorphs (see abstract of J. Král et al. for details).
We found this course of meiosis in two palpimanid subfamilies (Palpimaninae
and Chediminae). Although the third palpimanid subfamily Otiothopinae was
unfortunately not included in our analysis, the present findings indicate that
achiasmate meiosis might be a common characteristic of the whole family
Palpimanidae. Unlike palpimanids, the karyotype and course of meiosis in
Mimetus is similar to that of typical araneoids. Therefore, our data do not
222
support a phylogenetic relationship between the families Mimetidae and

Palpimanidae. Karyotype analyses of other groups considered closely related to
palpimanids (especially Huttoniidae and Stenochilidae) would be needed in
order to test for congruency of chromosome data and taxonomic hypotheses in
the superfamily Palpimanoidea.
This research was partially supported by the Grant Agency of the Czech
Academy of Sciences (project no. IAA601110808 – TK, JK) and by Czech Ministry
of Education, Youth and Sports (project no. MSM0021620828 – TK, JK).
References
Coddington J.A. 2005. Phylogeny and classification of spiders. In: Ubick D,
Paquin P, Cushing PE, Roth V, eds., Spiders of North America: an
Identification Manual. American Arachnological Society, pp. 18-24.
Huber B. 2004. Evolutionary transformation from muscular to hydraulic
movements in spider (Arachnida, Araneae) genitalia: a study based on
histological serial sections. Journal of Morphology, 261: 364-376.
Schneider M.C., Stávale L.M., Brescovit A.D., Cella D.M. 2008. Cytogenetic
information on five species of entelegyne spiders [Infirmações
citogenéticas de cinco espécies die aranhas entelegynae (Araneae:
Araneomorphae)]. Abstract, 2nd Latinoamerican Congress of Arachnology,
30th Nov.- 4th Dec. 2008, Salta, Argentina, p 152
Suzuki S. 1954. Cytological studies in spiders. III. Studies on the chromosomes
of fifty-seven species of spiders belonging to seventeen families with
general considerations on chromosomal evolution. Journal of Science of
the Hiroshima University, B, 15: 23-136.
223
Diversity and endemism of spiders (Arachnida: Araneae)

of the Crimean Peninsula, Ukraine
Mykola M. Kovblyuk
Zoology Department, V.I. Vernadsky Taurida National University, Ukraine,

kovblyuk@mail.ru
Introduction
The Crimean Peninsula is the mountainous highly isolated region, surrounded
by Black and Azov seas and connected with the mainland only by the narrow
Perekop isthmus.
The first spider record from Crimea was published by Falk (1786) while the
first description of spider species was published by Doblika (1853).
In 1875 Thorell published his two significant works (1875 a,b), describing
about 50 new species; many of them synonymised or found also in other regions
(Bulgaria, continental part of Ukraine, Rostov Area of Russia, Caucasus and
Turkey). These species were analyzed by the author (Kovblyuk 2002), who also
published Catalogue of Crimean spiders (Kovblyuk 2004). During the last years
more species and data on their distribution were found.
List of Crimean endemic spider taxa and their distribution (from published and
personal unpublished data):
Genera
Deliriosa Kovblyuk, 2009 (Lycosidae)
Spinestis Saaristo, Marusik, 2009 (Oonopidae)
Species
Agelenidae
Malthonica podoprygorai Kovblyuk, 2006 – all the mountainous part of Crimea
Tegenaria taurica Charitonov, 1947 (also recorded from Caucasus by Mkheidze
(1997), probably misidentified) – south part of the Crimean Mountains, in
caves and buildings.
Clubionidae
Clubiona mykolai Michailov, 2003 – all the mountainous part of Crimea.
Dysderidae
Harpactea doblikae (Thorell, 1875) – all the mountainous part of Crimea.
Gnaphosidae
Berlandina shumskyi Kovblyuk, 2003 – steppe in the plain part of Crimea.
Micaria blicki Kovblyuk, Nadolny, 2008 – all the mountainous part of Crimea.
M. bosmansi Kovblyuk, Nadolny, 2008 – saline lands in the plain part of Crimea,
southern coast with sub-Mediterranean vegetation.
Parasyrisca marusiki Kovblyuk, 2003 – mountain plateaus at 1000-1200 m a.s.l., in
the rocky mountain steppes.
224
Linyphiidae
Diplocephalus pseudocrassilobus Gnelitsa, 2006 – south slope of the Crimean
Mountains.
Incestophantes australis Gnelitsa, 2009 – south slope of the Crimean Mountains.
Typhochrestus longisulcus Gnelitsa, 2006 – north slope of the Crimean Mountains.
Lycosidae
Lycosidae
Deliriosa karadagensis Kovblyuk, 2009 – sub-Mediterranean steppes in the east
part of the southern coast of Crimea.
Oonopidae
Spinestis nikita Saaristo, Marusik, 2009 – southern coast of the Crimean Mountains.
Salticidae
Neon kovblyuki Logunov, 2004 – southern coast of the Crimean Mountains.
Synaphridae
Synaphris lehtineni Marusik, Gnelitsa, Kovblyuk, 2005 – southern coast of the
Crimean Mountains.
Theridiidae
"Crustulina" albovittata (Thorell, 1875) (species with unclear generic status) – the
habitat and distribution is unknown
"Dipoena" lindholmi (Strand, 1910) (the species with the unknown generic position)
– southern part of the Crimean Mountains
Discussion
In total, 525 valid species from 225 genera and 38 families were recorded, of
them 17 endemic species (3.2%) and 2 endemic genera (0.8%). Most endemic
species live in the mountainous part, especially on southern slopes, except for
Berlandina shumskyi, which inhabits the steppe plains.
Endemism in Crimea is much lower than at the Balkan Peninsula (25%
species from ~ 1500) (Deltshev 2000) and Caucasus (~22% species from ~1022)
(Marusik et al. 2006). Low endemism is probably caused by the smaller area, land
age and geographical isolation. Nevertheless, total species diversity of Crimean
spiders is only twice lower than in Caucasus and 3 times lower than in the Balkans.
With the area of about 27,000 km2 and maximum elevation of 1,545 m a.s.l.,
Crimea is similar to Sardinia (about 24,000 km2; 1,834 m) and Corsica (about 9000
km2; 2710 m). Also the number of spider species (500 from each island) is similarly,
while endemism on Sardinia (27-29 species, ~6%) and Corsica (43-51 endemic
species, ~10%) it is much higher (Wunderlich 1995). It is probably the result of
stronger isolation of both islands (190 km and for Corsica respectively).
Another distinctive feature of Crimean spider fauna is the absence of endemic
adaptive radiation. The phenomenon is well known in some genera from Sardinia
(Harpactea, Leptoneta, Tegenaria) and from Corsica (Nemesia, Lepthyphantes
sensu lato) (Wunderlich 1995); from the Balkans (Troglohyphantes, Dysdera,
Lepthyphantes sensu lato, Tegenaria) (Deltshev 2000); and from Caucasus
(Dysdera, Harpactea) (Dunin 1992).
Karst topography is very well developed in the Crimean Mountains, with
225
many caves. However, cave spider fauna is very poor (5 species only), and no
endemic spiders are found. In this aspect, Crimea is very different from adjacent
Balkans, Turkey and Caucasus.
Conclusions
1. Crimean spider species diversity is large, but endemism is low.
2. Adaptive radiation in endemic spider groups does not occur.
3. Crimean cave species diversity is low, and cave endemics are lacking.
References
Deltshev Ch. 2000. The endemic spiders (Araneae) of the Balkan Peninsula. In:
Gajdos P., Pekar S. (eds), Proceedings of the 18th European Colloquium
of Arachnology, Stara Lesna, 1999. Ecologia (Bratislava), Supplement 3,
19: 59-65.
Doblika K. 1853. Beitrag zur Monographie der Spinnengeschlechts Dysdera.
Verhandlungen der zoologisch-botanischen Vereins in Wien, 3: 115-124.
Dunin P.M. 1992. The spider family Dysderidae of the Caucasian fauna (Arachnida,
Aranei, Haplogynae). Arthropoda Selecta, 1(3): 35-76 [in Russian with
English summary].
Falk J.P. 1789. Beiträge zur topographischen Kenntnis des Russischen Reiches. Bd.
3. Beitrage zur Thierkunde und Volkerbeschreibung, pp. 283-584, 1-25
(spiders on 443-444).
Kovblyuk M.M. 2002. To question about endemism of Crimean spiders (Arachnida,
Aranei). Zapovedniki Krima. Materialy II nauchnoi konferencii (25-26 April
2002). Simferopol, pp. 103-109 [in Russian].
Kovblyuk M.M. 2004. Catalogue of the spiders (Arachnida: Aranei) of the Crimea,
South Ukraine. Points on the development of the Crimea. Analytical,
scientific and practical collected articles open to discussion. 15-th issue:
Problems of the ecology in the Crimea. Inventory animals and plants species
in the Crimea. Simferopol: Tavriya-Plus, pp. 211-262 [in Russian].
Marusik Yu.M., Mikhailov K.G. & Guseinov E.F. 2006. Advance in the study of
biodiversity of Caucasian spiders (Araneae). Acta zoologica bulgarica,
Supplement 1, pp. 259-268.
Mkheidze T.S. 1997. Spiders of Georgia (taxonomy, ecology, zoogeoraphical
review). Tbilisi, University Publishing House, 384 pp. [in Georgian].
Thorell T. 1875a. Verzeichniss Südrussischer Spinnen. Horae Societatis
Entomologicae Rossicae, 11: 39-122.
Thorell T. 1875b. Descriptions of several European and North-African spiders.
Kungl Svenska Vetenskaps-Akademiens Handlingar, 13(5): 1-204.
Wunderlich J. 1995. Zur Kenntnis der Endemiten, zur Evolution und
Biogeographie der Spinnen Korsikas und Sardiniens, mit
neubeschreibungen (Arachnida: Araneae). Beiträge zur Araneologie, 4:
353-383.
226
Evolution of the karyotype and sex chromosome systems

in mesothelid and mygalomorph spiders
Jiří Král1, Lenka Dulíková2, Tereza Kořínková1 , Jana Musilová1,

Magda Vítková3, Marshal Hedin4, Sérgio S. Henriques5
& Charles R. Haddad6
1
Faculty of Science, Charles University in Prague, Czech Republic,
spider@natur.cuni.cz, korinko1@natur.cuni.cz,
jana-musilova@seznam.cz
2
GENvia Ltd., Praha, Czech Republic, lenka_dulikova@seznam.cz
3
Department of Genetics, Institute of Entomology, Czech Academy of Sciences, České
Budějovice, Czech Republic, vitkova@entu.cas.cz
4
Department of Biology, San Diego State University, San Diego, CA, USA,
mhedin@sciences.sdsu.edu
5
Department of Entomology, School of Biology, University of Évora, Évora, Portugal,
henriquesbio@gmail.com
6
Based on the entelegyne lineage of araneomorph spiders, the typical spider

karyotype is supposed to consist of acrocentric chromosomes comprising the
X1X20 sex chromosome system. Karyotypes of other spider groups remain
poorly understood. To determine fundamental trends in karyotype evolution of
basal spiders, we studied 2 mesothelids and 22 mygalomorph species belonging
to 9 families. The 2n of male and sex chromosome systems were found as
follows: Liphistiidae: Liphistius sp. 73 (X0), Ryuthela nishihirai 69 (X0);
Antrodiaetidae: Aliatypus sp. 27 (X0); Mecicobothriidae: Megahexura sp. 43
(X0); Ctenizidae: Cyclocosmia siamensis 128 (?), Ummidia sp. 53 (X0);
Cyrtaucheniidae: Cyrtauchenius walckenaeri 66 (X1X20); Migidae:
Poecilomigas sp. 33 (X0); Hexathelidae: Macrothele gigas 85 (X1.X130);
Dipluridae: Diplura cf. petrunkevitchi 90 (X1X2X3X40), Euagrus lynceus 59
(X1X2X3X4X50), Ischnothele caudata 14 (XY); Nemesiidae: Acanthogonatus
pissii 61 (X1X2X30), Iberesia machadoi 76 (X1X20); Theraphosidae: Avicularia
minatrix 78 (X1X2X3X40), Brachypelma albopilosum 74 (X1X2X3X40),
Citharischius crawshayi 67 (X1X2X30), Grammostola rosea 72 (X1X20),
Holothele cf. longipes 73 (X1X2X30), Idiothele sp. 25 (X0), Ischnocolus
adenense 85 (X1X2X30), Poecilotheria formosa 110 (X1X2X3X40), Psalmopoeus
cambridgei 84 (X1X20), Pterinochilus murinus 43 (X0).
Our study revealed a considerable diversity of diploid chromosome
numbers, chromosome morphology, and sex chromosome systems of
mygalomorphs. Diploid numbers of males vary significantly from 14 (I.
caudata, Dipluridae) up to 128 (C. siamensis, Ctenizidae). The greatest
variability of 2n was found in the families Ctenizidae, Dipluridae, and
227
Theraphosidae. The majority of mygalomorphs exhibits higher 2n than

araneomorph spiders. Biarmed chromosomes (metacentric and submetacentric)
predominate in most mygalomorphs. Some species however display a
predominance of acrocentric chromosomes, namely the representatives of the
genera Diplura (Dipluridae), Iberesia (Nemesiidae) as well as a representative of
the family Migidae. We hypothesize that the large portion of acrocentric
chromosomes in karyotype of two representatives with the highest 2n,
Poecilotheria (Theraphosidae), and Cyclocosmia (Ctenizidae), reflects frequent
centric fissions during evolution of their genome. Mygalomorph autosomes
contain a low number of nucleolar organizer regions (NOR). One pair of NOR
can often be distinguished by its enormous size. In some diplurids, NOR was
found also on one sex chromosome. Most of the studied mygalomorphs exhibit a
standard course of meiosis. Remarkably, we found the evolutionary intermediate
between standard chiasmate and achiasmate meiosis (“cryptochiasmate
meiosis”) in males of Diplura, and achiasmate meiosis in males of another
diplurid genus, Euagrus.
Sex chromosomes of mygalomorphs exhibit usually a biarmed
morphology. The system X1X20, which is supposed to be ancestral for spiders, is
less common in mygalomorphs than in entelegyne spiders. Other multiple X
chromosome systems (X1X2X30, X1X2X3X40, X1X2X3X4X50 and X1.X130) have
evolved from the X1X20 mode probably by non-disjunctions or by non-
disjunctions and fissions. Rearrangements between autosomes and sex
chromosomes resulted into neo-sex chromosomes in Ischnothele (neo-XY
system). Multiple X chromosomes show an unusual distal end-to-end pairing
during male meiosis. Interestingly, this pairing is the same as found in the basal
araneomorph spiders with X1X2Y system but different from the distal end-to-end
pairing of basal entelegyne spiders. Presence of the same type of sex
chromosome pairing in mygalomorphs and basal araneomorphs is a challenge to
test in detail the relationship of these groups. Similarly to other spiders, we
found a unique behaviour of sex chromosomes during female meiosis in
mesothelids and mygalomorphs. In contrast to homogametic sex of other
animals, meiotic behaviour of sex chromosomes and autosomes in spider
females is different. Homologous sex chromosomes pair already in premeiotic
nuclei and are facultatively heterochromatinized until prophase of the first
meiotic division. Interestingly, X chromosomes retain unusual behaviour at
female meiosis after fissions or fusions with other sex chromosomes or
autosomes. We suggest that inactivation of sex chromosome pairs in females has
evolved to avoid negative effects of X chromosome duplications during spider
evolution, especially to restrict pairing and recombination only to pairs of
homologous sex chromosomes. The complexity of sex chromosome behaviour
in female meiosis is increased by an end-to-end association of sex chromosome
pairs in premeiotic and prophase I nuclei. We hypothesize that chromosome
ends involved in this association contain information for initial assignment of
homologous X chromosomes and are likely to promote their pairing.
Finally, our data allow us to determine specific features of mesothelid
karyotypes. Both examined genera of mesothelids show a high number of
228
predominantly acrocentric chromosomes and X0 sex chromosome system. In

contrast to mesothelids, biarmed chromosomes predominate in most
mygalomorphs as well as basal clades of araneomorph spiders. This pattern can
indicate that the predominance of biarmed chromosomes is ancestral for
opisthothele spiders. Furthermore, the karyotype diversity presented in our study
allows to distinguish three basic evolutionary lineages of mygalomorphs. The
first mygalomorph lineage involves the superfamily Atypoidea, i.e. families
Atypidae, Antrodiaetidae, and Mecicobothriidae. This group is characterized by
relatively low chromosome numbers (<50 chromosomes) and X0 system.
Providing primary absence of multiple X chromosome systems in mesothelids
and the first mygalomorph lineage, the superfamily Atypoidea could constitute a
sister clade to opisthotheles. The second mygalomorph group is formed by
Rastelloidina. These spiders often exhibit higher 2n than Atypoidea; sex
chromosome system is X0 or X1X20. The third lineage is formed by the group
Crassitarsae plus the families Hexathelidae and Dipluridae. Predominance of
karyotypes with 70-90 chromosomes suggests that ancestral diploid number of
the lineage falls into these limits. The X1X2X30 system is hypothesized to be
ancestral in families Nemesiidae and Theraphosidae. X1X20 and X0 systems
found in these families arose probably by a gradual fusion of sex chromosomes.
229
Evolution of holocentric chromosomes and X1X2Y system:

two karyotype traits found in basal clades of araneomorph
spiders (Araneae: Araneomorphae)
Jiří Král1, Markéta Pastuchová1, Jana Musilová1,

Tereza Kořínková1,Milan Řezáč2, Magda Vítková3,
Bernhard A. Huber4 & Charles R. Haddad5
1
Faculty of Science, Charles University in Prague, Czech Republic,
spider@natur.cuni.cz, marketa.pastuchova@seznam.cz,
jana-musilova@seznam.cz, korinko1@natur.cuni.cz
2
Research Institute of Crop Production, Praha, Czech Republic, rezac@vurv.cz
3
Department of Genetics, Institute of Entomology, Czech Academy of Sciences, České
Budějovice, Czech Republic, vitkova@entu.cas.cz
4
Alexander Koenig Zoological Research Museum, Bonn, Germany, b.huber.zfmk@uni-
bonn.de
5
Basal clades of araneomorph spiders show a considerable diversity of

chromosome structure and sex chromosome systems. Their karyotypes consist of
holocentric chromosomes (superfamily Dysderoidea) or standard chromosomes
with a localized centromere. Holocentric chromosomes are in general of
polyphyletic origin in animal and plant kingdoms, however, the mode of their
origin is unknown. A discussed possibility of their origin is a fusion of enormous
number of chromosomes with the expansion of kinetic activity over a large
surface area of fusion products.
To highlight the origin of holocentric chromosomes in the superfamily
Dysderoidea, we studied cytogenetics of their putative relatives, family
Caponiidae. Males of the examined species show 127 chromosomes including
11 sex chromosomes. During male meiosis, sex chromosomes of Caponia
display achiasmatic pairing. Biarmed sex chromosomes pair by both ends,
whereas the acrocentric and subtelocentric sex chromosomes pair only by their
long arms. It is tempting to speculate that holocentric chromosomes of the
superfamily Dysderoidea originated by multiple fusions from caponiid
karyotype. Our analysis indicates that ancestral male karyotype of superfamily
Dysderoidea was formed only by three autosome pairs and a single X
chromosome. In spite of the considerable karyotype diversity of Dysderoidea,
this karyotype predominates in oonopids, dysderids of the genera Kaemis,
Harpactea, and Dasumia, and it was also found in the segestriid genus Ariadna.
Further evolution of spider holocentric chromosomes has frequently included
chromosome fragmentations. On the other hand, our data show that the ancestral
diploid number was also lowered in some lineages by chromosome fusions.
230
Karyotypes of some oonopid spiders are unique among all organisms because
they consist only of a single chromosome pair, formed by sex chromosomes X
and Y.
A derived X1X2Y system with an achiasmatic sex-chromosome pairing
during meiosis was found in some basal araneomorphs with standard
chromosomes, namely in the families Drymusidae, Filistatidae, Hypochilidae,
Pholcidae, and Sicariidae, suggesting that these groups form a monophyletic
clade. The structure of the X1X2Y system is conservative. These sex
chromosomes usually exhibit a metacentric morphology including a tiny Y
chromosome. Furthermore, they show the same type of distal end-to-end pairing
as found in Caponia. In contrast to other X1X2Y spiders, the X1X2Y system of
pholcids was converted gradually into the X0 system. To trace the evolution of
pholcid karyotypes and sex chromosomes, we have studied diploid numbers and
morphology of chromosomes, sex chromosome systems, and pattern of nucleolar
organizer regions (NOR) in selected species. In general, the evolution of
pholcids is characterized by a reduction of the number of autosome pairs and sex
chromosomes. The original type of X1X2Y system is retained in Spermophora
senoculata (2n♂=25; X1X2Y) only. Our data suggest reduction of the X2
chromosome in the common ancestor of the genera Leptopholcus and Pholcus.
Reduction of the X2 chromosome was accompanied by considerable enlargement
of the Y chromosome, possibly by translocation of autosome material. This type
of X1X2Y system was found in two afrotropical taxa, L. guineensis (2n♂=17;
X1X2Y) and P. guineensis (2n♂=23; X1X2Y). In the Palaearctic species P.
opilionoides and P. phalangioides (2n♂=25; X1X2Y), the metacentric
morphology of the reduced X2 chromosome was changed to an acrocentric one.
Interestingly, changes in the structure of sex chromosomes were accompanied by
changes of the NOR pattern. In S. senoculata, NORs were detected on three
autosome pairs. The karyotype of P. phalangioides contains NORs also on the
ends of X chromosomes included into meiotic pairing. This modification of sex
chromosomes may facilitate their achiasmatic pairing.
'Holocnemine' pholcids exhibit a considerably different evolution of the
X1X2Y system. An early loss of the ancestral Y microchromosome changed the
X1X2Y mode into a secondary X1X20 system. We found the most ancestral
karyotype in Smeringopus sp. from South Africa (♂2n=28; X1X20). Both X
chromosomes of this species retain a metacentric morphology. In Smeringopus
sp. from Madagascar, Hoplopholcus cf. cecconii from Israel (♂2n=28; X1X20),
and Crossopriza cylindrogaster (♂2n=26; X1X20), the morphology of the X2
chromosome was changed to subtelocentric or acrocentric by pericentric
inversion. Finally, a fusion of X1 and X2 chromosomes produced an X0 system
in some species of Crossopriza (C. lyoni and Crossopriza sp.nov. from
Morocco; 2n♂=23; X0) and in the genus Holocnemus (H. pluchei ♂2n=27; X0,
H. hispanicus ♂2n=23; X0). In contrast to the Old World pholcids, published as
well as our data suggest that the X0 system originated early in the clade of New
World pholcids. All species studied so far show the X0 system.
Altogether, our data confirm the evolutionary plasticity of the X1X2Y
system in the family Pholcidae. Furthermore, these results indicate that the
231
X1X2Y clade can be more diversified than previously thought. It might include
also families whose extant representatives do not have the X1X2Y system. This
concerns especially the families Scytodidae (X0), Austrochilidae (XY), and
Diguetidae (XY) whose karyotypes show some features typical for the X1X2Y
clade.
232
The adaptiveness of being a spider eunuch

Simona Kralj-Fišer1*, Matjaž Gregorič1, Shichang Zhang2,
Daiqin Li2 & Matjaž Kuntner1
1
2
Department of Biological Sciences, National University of Singapore, Singapore
* Presenting author: simonakf@gmail.com
Nephilid spiders show peculiar sexual behaviours, including male genital

mutilation, the plugging of female genitals, and sexual cannibalism, where giant
females regularly devour their tiny mates. Furthermore, males of Nephilengys
and Herennia exhibit an extreme form of emasculation by removing their palpal
bulbs entirely to become "eunuchs". The behavioural pathways leading to
emasculation in nephilids are plastic and poorly understood. Thus, explanations
of the seemingly non-adaptive eunuch phenomenon along with more precise
observations are needed. We studied sexual behaviour in Nephilengys
malabarensis, a common SE Asian species, and tested four explanations for
adaptiveness of being a eunuch: 1. the broken embolic conductor (EC) plugs the
copulatory opening (CO) and thus prevents rival males' access; 2. removing the
bulb after initial EC breakage is advantageous for the sterile male in stopping
hemolymph leakage; 3. eunuch protects his parental investment by fighting off
rival males and 4. emasculation increases agility allowing the male to better fend
off rival males. First, we staged mating trials to precisely document palpal
damage and emasculation and to test whether the broken EC functions as a plug.
Second, to establish the effect of palpal loss on eunuch behaviour, we conducted
a series of male-male contests: intact vs. intact male, intact vs. half-eunuch,
intact vs. full eunuch, and half-eunuch vs. full eunuch. A palp insertion always
resulted in male genital mutilation: 90% of males broke the entire bulb and
plugged the female CO, and in 10% of males the palp was first disfigured, then
severed. Sexual cannibalism was rampant at 75%. We confirmed the plugging
function of broken male genitals with a 75% prevention of any subsequent
copulation. Full eunuchs were the most active guardians of females against rival
males; they most frequently initiated and won contests and were most aggressive
against competitors. Intact males and half-eunuchs showed similar, less
aggressive behaviours. Logically, only intact males and half-eunuchs achieved
copulation, but only when not prevented by a full eunuch. Copulation with a
female was independent from the outcome of the male – male contest.
Therefore, we found support for our first, third and fourth hypotheses, but not
the second. We conclude that being a spider eunuch is adaptive: i) palpal loss
enables higher levels of agility and aggressiveness, which help outcompete rival
males; ii) aggressiveness and functional epigynal plugs together likely increase
233
the eunuch's paternal investment. We also suggest another function of palpal

severance, iii) “remote copulation”. Due to short insertion time, and high levels
of sexual cannibalism, which males try to evade, breakage of the entire bulb in
the female CO may secure continuous sperm transfer even after the male has
been (forcefully) removed from copula.
234
Spiders have personalities: Intraspecific behavioural

variability in Nephilengys malabarensis
(Araneae: Nephilidae)
Simona Kralj-Fišer1*, Larisa Zemljič2, Urška Velikonja2,

Tadeja Papler2, Tjaša Lukanc2 & Matjaž Kuntner1
1
2
Department of Biology, University of Ljubljana, Slovenia
* Presenting author: simonakf@gmail.com
Individuals of the same sex and species commonly differ from each other
in their behavioural characteristics. Individual behavioural characteristics may
be repeatable and consistent across multiple contexts. Analogous to humans,
such characteristics are referred to as personality-related traits. Behavioural
characteristics commonly come in packages (correlate) and form personality
dimensions. Some personality dimensions, such as aggression, boldness,
sociability, exploration and general activity are universal for vertebrates,
whereas others are species-specific. Personalities are moderately heritable and
often contingent on neuroendocrinological processes. Personalities influence
individuals' ecology, survival and reproductive success. Few studies have
investigated intraspecific behavioural variation in invertebrates, although their
species diversity as well as behavioural, ecological and morphological
variability is much higher than in vertebrates. This paucity of data is probably
due to general belief that invertebrates, in contrast to behaviourally plastic
vertebrates, behave as “mini-robots”. We investigated intraspecific behavioural
differences in females Nephilengys malabarensis, a common SE Asian nephilid
spider, and their importance for web building and reproductive success. In the
first step we determined personality related behaviours. For this purpose, each
individual was repeatedly tested in a series of standardized tests, i.e. predator test
(reactivity), contest (aggressiveness to a conspecific of the same sex), mating
(aggressiveness towards a mate, sexual cannibalism) and novel environment test
(boldness and exploratory behaviour). To reveal species personality dimensions,
personality related behaviours were correlated. Repeatability was shown for
reactivity, exploration and aggressiveness towards a conspecific, suggesting
these behaviours are heritable and related to personality in N. malabarensis. The
more aggressive the individuals were, the less explorative they tended to be in a
novel environment. This is in accordance with studies in vertebrates (great tits
and rodents), suggesting a common, albeit opposite, evolutionary pathway of
these two behavioural traits. Aggressiveness towards a conspecific did not
correlate to aggressiveness towards a male during mating, implying their
different underlying mechanisms (natural vs. sexual selection). The most and the
least explorative spiders differed in their web building in captivity: more
explorative spiders built webs with retreat only, whereas non-explorative spiders
235
built finished webs. Females that were more aggressive towards their mates also
captured more prey, as in previous studies on Dolomedes and Agelenopsis.
Unexpectedly however, the females that were most aggressive towards
conspecifics of the same gender, caught least prey. Spiders of different
behavioural characteristics did not differ in their willingness to mate. In
conclusion, orb-weaving spiders N. malabarensis consistently differ individually
in reactivity, exploration and aggressiveness towards conspecifics. Individual
differences in exploration of a novel environment are reflected in web building
abilities. To date, only a handful of studies exist researching spider personality,
and these are currently too scattered taxonomically to allow generalizations and
syntheses in this promising field of research.
236
Biological hours tick for everyone - time chart of diel

activity of epigeic spiders
Zuzana Krumpálová1, Miroslav Krumpál2 & Ivan Hadrián Tuf3
1
Institute of Zoology, Slovak Academy of Sciences, Bratislava, Slovakia,
zuzana.krumpalova@savba.sk
2
Department of Zoology, Faculty of Natural Sciences, Comenius University, Bratislava,
Slovakia,
krumpal@fns.uniba.sk
3
Department of Ecology and Environmental Sciences, Palacký University, Olomouc,
Czech Republic, ivan.tuf@upol.cz
Introduction
The environment is periodic with respect to day-night; a periodical
pattern of animals synchronised with this environmental rhythm. This periodism
is specific and individual. To compete in periodic world, the organism must have
a periodic activity (Park 1941).

Spiders were caught using the pitfall traps (plastic pots) without fixative
solution. 60 traps were arranged in the forest and 40 traps in the deforested clear-
cut area in Litovelské Pomoraví Protected Landscape Area (Czech Republic,
Europe). The traps were set in a line with three meters distances. The
investigation was carried out in 2004, between 20 May – 7 June for 18 days and
23 September – 18 October for 25 days. Traps were checked once a three hour
period (i.e. at 3, 6, 12, 15, 18, 21 and 24 o’clock), i.e. eight times a day. Circular
data of diurnal activity of spiders was evaluated according to Oriana (Kovach
2009), including the circular mean, length of mean vector, circular standard
deviation 95% and 99% confidence limit. We applied the multivariate analysis
using Past (Hammer et al. 2001) to evaluate of the epigeic spiders.
Results
Of almost 12,000 specimens of ground-dwelling invertebrates, 5,817
spiders were collected and identified. Spiders represented 108 taxa from 20
families. In general, a high number of species was noticed at forest in spring (71
taxa), in autumn the coenose decreased in number of species to 42; 31 species
were identical for both seasons. In the clear-cut area the number of species
decreased from 62 to 26 taxa; 17 species were identical, only.
Considering the eudominant and dominant species, Abacoproeces
saltuum (43.8%) and Pardosa lugubris (14.6%) were eudominant species,
Coelotes terrestris (5.2%) was dominant and subdominants were Pardosa
prativaga (4.5%), Panamomops mengei (3.7%), Ceratinella brevipes (2.5%) and
juveniles of the genus Pardosa (4.6%). 101 species had recedent or subrecedent
status.
237
The constant presence and higher dominance of P. lugubris, C. terrestris,

P. prativaga, Pisaura mirabilis and juveniles of the genus Pardosa at both study
sites also characterized spider communities. Dominant species showed a high
frequency of occurrence at study sites alike. The species diversity in the spider
communities varied.
P. mengei is a diurnal diphase spider, with main activities in the morning
and in the evening. O. praticola (males and females) unambiguously
demonstrated the diurnal diphase of diel activity with two peaks at 12 and 21
o’clock. Thanks to the two different diel activities of sexes, we contribute to the
diurnal diphase of C. brevipes, D. picinus and M. herbigradus – with main
activity at the noon and high males’ activity in the evening or night. Similar
activity we observed at W. dysderoides. W. obtusa is a diurnal spider with one-
phase activity in the morning. Considering to the genus Walckenaeria, time lag
of activity of W. dysderoides was shifted to the afternoon, opposite to the
morning activity of W. obtusa. A. saltuum is a diurnal linyphiid, its high activity
was noticed early in the morning.
Diurnal activity of P. lugubris, with maximum at 15 o’clock, is uniform
for males, females and juveniles. We noticed the certain uniformity in the
population of P. prativaga with maximum in the afternoon (males, females and
juveniles) and P. amentata (confirmed males and females). Juveniles of the
genus Pardosa as the adults of this genus demonstrated typical afternoon diurnal
activity with the uniformity of examined species.
Males and females of C. terrestris seem to be a midnight spiders, while
the juveniles typical diurnal ones. This phenomenon of different activity of
adults and juveniles we found at Coelotes terrestris, only one spider species.
Thanks to the research a high nocturnal epigeic activity of P. mirabilis were
confirmed at both sexes and juveniles.
Conclusions
Spiders’ body size seems to be very closely related with their diel
activities. The body size of 108 spider species markedly differed in length; vary
in size from 1.2 mm (P. mengei) to 17.0 mm (P. mirabilis). We found four types
of diel activity of epigeic spiders: diurnal one- phase in the morning; diurnal
one-phase in the afternoon; nocturnal one-phase in the night; diurnal-nocturnal
diphase in the morning and evening (or night).
Spiders with differences in body-size of sexes (females larger by about
0.6-1.1 mm) - the smallest spiders (1.2-1.8 mm) and small spiders (2.9 - 4.0 mm)
showed the diurnal diphase activity with two peaks; in the morning were active
larger females, and in the evening or night were active males. In this category of
diurnal diphase activity (morning – evening) belong – D. picinus, C. brevipes,
W. dysderoides or P. mengei; into the diphase morning – night belong O.
praticola or M. herbigradus.
Small spiders (1.6-2.9 mm) with no differences in sexes’ body size
showed the diurnal activity with one phase in the morning and the activity of
both sexes were similar during the day. In the category of diurnal morning
activity belong – A. saltuum or W. obtusa.
238
Bigger spiders (8.0-12 mm) with sexes’ similar body size showed diurnal
activity with one phase in the afternoon. Both sexes as well as their juveniles
were active before the sun shining, nocturnal activity was not confirmed. In this
category of diurnal afternoon activity belong all species of the genus Pardosa –
P. lugubris, P. prativaga, P. amentata, P. pullata, their juveniles, and the
juveniles of C. terrestris.
Big spiders (12-17 mm) demonstrated a typical one phase of nocturnal
activity. This diel activity was confirmed at both sexes. In the category of
nocturnal activity (midnight) belong spiders – P. mirabilis or C. terrestris.
Acknowledgement
Thanks to the leaders and ardent students, the tireless collectors, this
research could be realised. We would like thanks to the Slovak Grant Agency
VEGA (No. 02/0067/08 and 01/0176/09), Ministry of Environment of the Czech
Republic (No. SP/2D3/155/08) and Czech National Research Programme II (No.
2B 06101) for support this work.
239
A molecular species level phylogeny of nephilid spiders

Matjaž Kuntner1*, Peter Trontelj2, Miquel Arnedo3,
Tjaša Lokovšek1 & Ingi Agnarsson4
1
2
Department of Biology, University of Ljubljana, Slovenia
3
Department of Animal Biology, University of Barcelona, Spain
4
* Presenting author: kuntner@gmail.com
The pantropical orbicularian spider family Nephilidae, currently with four

genera and close to 40 species, is becoming a model clade in spider evolutionary
research. Recent studies have focused on the evolution of extreme sexual size
dimorphism, sexual behaviours, and webs. All of them built on a recent species
level phylogeny based on morphological and behavioural data, which
established a fully resolved monophyletic Nephilidae as sister to all other
araneoid spiders. Our research now focuses on testing the nephilid phylogenetic
relationships by generating an extensive molecular dataset using nuclear and
mitochondrial genes for over a 100 terminals spanning nephilids and
orbicularian outgroups. Here, we report on the first analyses of the incomplete
dataset using maximum likelihood and Bayesian phylogenetic inference. All
analyses find strong support for nephilid monophyly and exclusivity, and thus
Kuntner's clade definition of Nephilidae separate from either Araneidae or
Tetragnathidae is supported. However, some nephilid genera show a surprising
phylogenetic structure strikingly differing from the current morphological
phylogeny. We contrast our preliminary phylogenetic results with published
analyses, and discuss the first reinterpretations of nephilid evolutionary history.
240
Experimental analysis of male cephalic modifications

in the dwarf spider Oedothorax retusus
(Araneae: Linyphiidae)
Katrin Kunz & Gabriele Uhl
Department General Zoology and Systematics, University of Greifswald, Germany,

katrin.kunz@uni-greifswald.de, gabriele.uhl@uni-greifswald.de
Males of many Erigoninae (dwarf spiders) have complex head structures –

pits or humps equipped with numerous pores connected to extensive glandular
tissue. During courtship and/or copulation females are in contact with these
structures. The secretions released by the male are ingested by the female and
may represent a mating effort or paternal investment. Within the genus
Oedothorax (Linyphiidae, Erigoninae) such gustatory courtship is common.
In O. retusus the secretions that are produced by the male head structure
are taken up by the female during courtship and copulation. We investigated the
influence of the head secretions during different stages of a mating bout
(courtship, copulation probability, copulation behaviour, oviposition and
paternity). To this aim, encounters between females and males with head
secretions or males with an experimentally covered secretory hump were
compared. Copulation duration was limited to one minute to keep chances equal
for both males. Virgin females preferred to copulate with males that were able to
release secretion. Males with an experimentally covered hump were not
handicapped in terms of courtship behaviour but performed significantly more
coupling trials before copulation commenced. This suggests, that our
manipulation influenced the males ability to achieve a specific mating posture
and that females were more hesitant to accept these males. However, after
successful mating paternity success of both male types was not significantly
different. Our results indicate that the head secretions in O. retusus males serve
as a mating effort, not as paternal investment.
241
Another male of thomisid spider from the Baltic amber

Janusz Kupryjanowicz
Institute of Biology, University of Białystok, Białystok, Poland, kuprzool@uwb.edu.pl
Among 13 spider species of Thomisidae family, known from the Late

Eocene Baltic amber, only 2 male specimens have been described so far. In the
work presented further one male of Thomisiraptor is described.
Discussion covers the justification for description of new species basing
on juvenile specimens.
Spiders of the genus Larinia in Poland

Janusz Kupryjanowicz1 & Robert Rozwałka2
1
Institute of Biology, University of Białystok, Białystok, Poland, kuprzool@uwb.edu.pl
2
Department of Zoology, Institute of Biology and Earth Sciences of Maria Curie-
Skłodowska University, Lublin, Poland, arachnologia@wp.pl
The data presented concern distribution of two species from genus

Larinia: Larinia jeskovi Marusik, 1986 and Larinia bonneti Spassky, 1939 in
Poland. Based on 15 years of research, the dynamics of a population size of
Larinia jeskovi in north-east Poland is demonstrated. Also an influence of
succession of vegetation of sedges marshes on size of the population is
presented. The project of active protection of these species is proposed. Habitat
preferences in Poland for both species of Larinia are presented as well as
information about their biology. For Larinia bonneti, the species reported for the
first time in Poland, redescription was made supported by SEM photographs.
242
The use of DNA barcodes to assess the biodiversity

patterns of megadiverse groups in tropical regions:
the PANCODING project
Facundo M. Labarque1, Luis N. Piacentini1, Martín J. Ramírez1,

Gustavo Hormiga2, Dimitar Dimitrov2, Ligia Benavidez2,
Miquel A. Arnedo3 & Joan Pons4
1
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires,
Argentina, facundo.labarque@macn.gov.ar, luis.piacentini@macn.gov.ar,
ramirez@macn.gov.ar
2
The George Washington University, Washington DC, USA, hormiga@gwu.edu,
dimitard@gwu.edu, ligia@gwmail.gwu.edu
3
Universitat de Barcelona, Barcelona, Spain, marnedo@ub.edu
4
Institut Mediterrani d’Estudis Avançats, Mallorca, Spain, jpons@imedea.uib-csic.es
The main goal of the PANCODING project is to evaluate the advantages

of the DNA barcoding techniques to estimate patterns of species-richness and
turnover of a megadiverse group (spiders) in a biodiversity hot-spot (the
mountain forests of Panama).
We conducted a semi-quantitative sampling of four one hectare plots
along a longitudinal transect spanning 350 km on the main cordillera of Panama.
We have sorted more than 30,000 specimens, 60% of which were immature,
nearly 25% females and 15% males. Of the 408 hypothesized morphospecies,
25% were represented by singletons and 10% by doubletons in our samples.
These percentages fall within values reported from previous biodiversity
inventories of Arthropods in Neotropical and Palearctic region. We use these
data to estimate alfa and beta diversity using species acumulation curves and
non-parametric estimators (ICE, ACE, Chao1, Chao2, Jackknife 1, Jackknife 2),
and complementarity and similarity estimators. Results indicate levels of
inventory completion of 70% for each locality and 10% of complementarity
between localities.
In parallel, we have sequenced the first half of the cox1 mitochondrial
gene of more than 2,400 specimens, including immatures, which yielded more
than 1500 unique haplotypes. Distinct genetic lineages were identified using the
GMYC method.
Here we present and contrast the results of comparing the patterns of
genetic based alfa and beta diversitity with those inferred from morphological
identification. In addition, we will discuss the effect of including DNA-aided
identified immatures, generally removed from morphology-based analyses, on
the estimates of alfa and beta diversity.
243
Phylogenetically important characters

in the spider family Scytodidae
Pekka T. Lehtinen
Zoological Museum, University of Turku, Finland, pekleh@utu.fi
Scytodidae has mostly remained almost monogeneric. The two-clawed

Dictis L. Koch, 1872 has quite seldom been used outside catalogs, in spite of
their prey catching without web. A treatment similar to that of other spider
families is now regarded necessary. Widely different genital organs were found
in the genus Stedocys Ono, 1995, which also included an Oriental species always
before catalogued as a Scytodes. Cytologically the “genus” Scytodes s. lato has
been found to include an exceptional variability of chromosomal numbers. My
work on Scytodidae has revealed a large number of morphological,
ultrastructural and ethological characters allowing the division of this family to
two subfamilies, several tribes in Scytodinae and altogether more than 20 new
genera, which are all left unnamed here and my lecture only uses species-groups
names.
Macromorphological characters, whose details are best studied with SEM
include especially the extremely variable patterns of the epiandrous spigots (=e.
fusulae) close to the gonopore in male, the exact shape of the opening of the
spitting glands in chelicerae, and several types of modified setae at the tarsal tip.
Pure ultrastructural characters include shape of epiandrous spigots and
trichobothrial base, tarsal organ, surfaces of chelicerae and different leg
segments. The phylogenetically most important macromorphological somatic
characters are: number of tarsal claws, presence and type of cheliceral
stridulatory ridges, presence of spination on different segments of legs, colour
patterns of legs, abdomen, and carapace, and modifications of sternum. All two-
clawed scytodids do not belong to the same clade. Characters in copulatory
organs include shape of male palpal tarsus, structure of tegulum and embolus,
presence and shape of the female copulatory pockets, and number of seminal
receptacula characters related to sexual dimorphism include also lower carapace
of male and similarity of male and juvenile colour patterns, as well as
concentration of leg and other modifications to males. Sufficiently studied
ethological characters include the mode of prey catching (web or hunting) as
well as the type of the extremely variable web. The presence of minute scytodids
with clavate hairs as an ecological adaptation to live in litter layer of many
tropical areas has not been known before.
A critical survey of published records of Scytodidae reveals that
previously described species have more often been wrongly than correctly
identified! Presence of infraspecific variation suggesting presence of obvious
subspecies has only been confirmed in some Oriental and Pacific species of
Dictis s. lat., but information about many Ethiopian and Neotropical taxa is too
patchy for final conclusions. The most important other type of infraspecific
244
variation is a type of colour polymorphism present in parallel in several, not

related groups.
Only the taxonomic results dealing with the type species of Scytodes are
published here. S. thoracica (Latreille, 1802) is present only in the western half
of Europe, the Western Mediterranean region, and in the area around the big
lakes in NE USA, while most populations published as Scytodes thoracica
outside these areas actually concern misidentified Scytodes tigrina C.L. Koch,
1838 (syntypes from Greece – examined). All Australian records as S. thoracica
belong to the cosmopolitan synanthropic “S.” bertheloti Lucas, 1838, which
belongs to a different, still unnamed tribe.
245
Spider assemblages as indicators of habitat conservation

value: a multi-scale approach in Western France
Boris Leroy1, Julien Pétillon2, Alain Canard1 & Frédéric Ysnel1
1
URU 420, University of Rennes I - Service du Patrimoine Naturel, Muséum National
d’Histoire Naturelle, France, leroy.boris@gmail.com
2
Evolutionary Ecology Group, Department of Biology, University of Antwerp, Belgium
Introduction
There is a general underrepresentation of invertebrates in conservation
research: only 11% of conservation articles between 1987 and 2001 involved
invertebrates despite the fact that they represent 75–79% of all described species
worldwide (Clark and May 2002). As a consequence, conservation planning
tools have been developed specifically for taxa whose distributions are well
known, such as birds, mammals and plants. However, conservation planning
based on single surrogate taxa may fail to cover at-risk species (Lawler et al.
2003), and in some cases, it has been demonstrated that although invertebrates
are strong predictors of conservation priorities for vertebrates, the reverse cannot
be said (Moritz et al. 2001). In this context, spiders have characteristics that
make them a good indicator taxon (Marc et al. 1999), and the availability of
biodiversity databases compiling previous samplings and/or species lists
available in a given region (e.g. Pétillon et al. 2007, Samu et al. 2008) make
them a good candidate to develop suitable conservation tools.
The selection of areas for biodiversity conservation is currently based on
two main methodological approaches: (i) complementarity-based approaches,
and (ii) scoring procedures. Complementarity-based approaches aim at selecting
a network of complementary sites in the context of concerted conservation plans.
Scoring procedures aim at ranking sites in order of value according to one or
several criteria, such as rarity or threat status (e.g. Fattorini 2006, Simaika &
Samways 2009). Hence, unlike the complementarity based-approaches, scoring
procedures are more suitable for spiders because they can be applied to set local
priorities, select or monitor one or a few sites, or assess the impact of different
management modes.
However, existing scoring procedures are generally not transferable
between different taxa, geographic areas and spatial scales, limiting their scope
and precluding their use in conservation strategies. Therefore, we propose a new
flexible index for assessing the conservation value of species assemblages, based
on biodiversity databases.

This index integrates two parameters: the proportion of rare species and
the intensity of their rarity. Species are weighted on the basis of their range size
as estimated by their occurrence in the reference region, with a method taking
246
into account a rarity cut-off point (i.e. the threshold of occurrence below which
species are considered rare) (Gaston 1994). This new weighting method thus
allows for calculating indices over a range of rarity cut-off points in order to
precisely analyse the patterns of rarity in assemblages. Furthermore, this method
can be applied at different geographic scales and/or areas, and to other taxa. We
developed guidelines for practical applications of the index following Gaston’s
recommendations on rarity (Gaston 1994).
We proposed two methods for ranking sites according to the index: the
first consisted in ranking the sites by their median rank over a range of rarity cut-
offs. The second method consisted in comparing sites with the known
assemblages of the reference region. The latter allowed for ranking sites
depending on their position in relation to all the known assemblages of the
reference region, as well as to characterize a particular site in relation to a
comparable pool of sites from the reference region (e.g. same biotope).
Results
At a local scale we applied the index to spider assemblages colonising
different biotopes of a National Nature Reserve in the Armorican Massif
(Western France). The analysis provided consistent results, highlighting four
main patterns of rarity (see Fig. 1):
1. First, there were assemblages having high indices whatever the rarity cut-off
(i.e. high proportions of rare species, whatever the intensity of rarity
considered).
2. Second, there were assemblages with high indices at low rarity cut-offs but
not at high rarity cut-offs (i.e. presence of very rare species despite a low
general proportion of rare species).
3. Third, there were assemblages with low indices at low rarity cut-offs, but
high indices at higher rarity cut-offs (i.e. lack of very rare species, but high
proportion of less rare species).
4. Finally, there were assemblages with low indices whatever the rarity cut-off,
(i.e. globally weak proportion of rare species).
Therefore, this analysis avoided the twofold risk of both neglecting assemblages
of potential interest (with a high proportion of rare species) and missing very
rare species (e.g. when the rest of the assemblage is composed of common
species).
At a higher scale, we applied the index to compare the conservation values
of the main habitat types of Western France (heathlands, dunes, forests,.) in
relation to the macroclimatic areas within which they are embedded. The
proposed guidelines worked well in fitting the range of rarity cut-offs to the
considered geographic scale and thus permitted an accurate emphasis on at-risk
species.
247
0.20 Type 1 assemblage

Type 2 assemblage
Type 3 assemblage
Type 4 assemblage
0.15
Conservation value (Ic Index)
0.10
0.05
0.00
2% 3% 4% 5% 6% 7% 8% 9% 10 % 11 % 12 % 13 % 14 %
Very rare species  Rare species  Uncommon species
Rarity cut-off
Fig. 1. Values of the index as a function of the rarity cut-off point (based on four
assemblages of the National Nature Reserve of Séné, Western France). The rarity cut-off
point is expressed as a percentage of the occurrence of the most widespread species.
Discussion
In conclusion, such an assessment of species assemblages provided
consistent and reliable results, and could be quickly applied to different
geographic areas and/or scales once occurrence data are available or can be
gathered. This new index is particularly hopeful for setting conservation
priorities, characterizing and/or monitoring the conservation values of one or
several sites in relation to a reference region, or assessing the impact of different
management modes.
References
Clark J.A., May R.M. 2002. Taxonomic bias in conservation research. Science,
297: 191-192.
Fattorini S. 2006. A new method to identify important conservation areas
applied to the butterflies of the Aegean Islands (Greece). Animal
Conservation, 9: 75-83.
Gaston K.J. 1994. Rarity. Chapman & Hall, London.
Lawler J.J., White D., Sifneos J.C. & Master L.L. 2003. Rare species and the use
of indicator groups for conservation planning. Conservation Biology, 17:
875-882.
248
Marc P., Canard A. & Ysnel F. 1999. Spiders (Araneae) useful for pest
limitation and bioindication. Agriculture, Ecosystems and Environment,
74: 229-273.
Moritz C., Richardson K.S., Ferrier S., Monteith G.B., Stanisic J., Williams S.E.
& Whiffin T. 2001. Biogeographical concordance and efficiency of taxon
indicators for establishing conservation priority in a tropical rainforest
biota. Proceedings of the Royal Society London B., 268: 1875-1881.
Pétillon J., Courtial C., Canard A. & Ysnel F. 2007. First assessment of spider
rarity in Western France. Revista Ibérica Aracnologia, 15: 105-113.
Samu F., Csontos P. & Szinetar C. 2008. From multi-criteria approach to simple
protocol: Assessing habitat patches for conservation value using species
rarity. Conservation Biology, 141: 1310-1320.
Simaika J.P. & Samways M.J. 2009. Reserve selection using Red Listed taxa in
three global biodiversity hotspots: Dragonflies in South Africa.
Conservation Biology, 142: 638-651.
249
The male genital system of a European mite harvestman,

Cyphophthalmus duricorius (Opiliones: Cyphophthalmi)
Elisabeth Lipke1,2, Gerd Alberti1, Melanie Gutjahr3
& Reinhart Schuster4
1
Ernst-Moritz Arndt University, Zoological Institute & Museum, Department of General
Zoology and Zoological Systematics, Greifswald, Germany,
alberti@uni-greifswald.de
2
RWTH Aachen, Institute for Biology II, Department of Developmental Biology and
Morphology of Animals, Aachen, Germany
3
Ernst Moritz Arndt University, Interfaculty Institute for Genetics and Functional
Genomics, Department of Functional Genomics, Greifswald, Germany
4
Karl-Franzens University, Institute of Zoology, Graz, Austria
Cyphophthalmi or mite harvestmen are considered to represent the sister

group of all other Opiliones. Although they are distributed worldwide and in
recent times part of many phylogenetic and phylogeographic studies only little is
known about their internal anatomy and the reproductive behaviour. Former
studies on the reproductive system of Opiliones have generally focused on the
structure of the penis (and ovipositor) for the search of characters of taxonomic
relevance. In contrast to other Opiliones, mite harvestmen transfer spermatozoa
through spermatophores and show a peculiar sperm dimorphism. The present
study describes the male genital system of Cyphpophthalmus duricorius, a
European representative of the family Sironidae using light and electron
microscopy. The male genital system can be divided into three parts: 1) the
unpaired testis, 2) two vasa deferentia which fuse to the long and thin seminal
vesicle, and 3) a short spermatopositor. The anterior part of the seminal vesicle
incorporates the ducts of two accessory glands. The epithelium of the testis
consists mainly of somatic cells and germ cells. The germ cells are arranged in
clusters (or germ-cell-cysts), and are surrounded by a thin layer of muscles.
Within these germ cell cysts two types of spermatozoa develop: the later fertile
eusperm and the supposed infertile parasperm. Each type can clearly be
distinguished by the shape and condensation of the nucleus or the number of
organelles like mitochondria. Once released into the lumen of vasa deferentia
and vesicula seminalis the sperm cells arrange in a distinct pattern. The fertile
eusperm group in the centre of the establishing sperm ball, surrounded by the
supposed non-fertile parasperm and secretions. The spermatopositor of C.
duricorius, in contrast to the homologous penis of other Opiliones, is short, flat,
undivided and equipped with long bristles. Secretions of the accessory glands
might help forming the spermatophores, which the male will affix to the genital
opening of the female. Ultrastructural details of vasa deferentia, vesicula
seminalis and spermatopositor are described.
250
Impact of land management on soil arachnofauna

(Arachnida) in the southwest Pará, Brazil 6
Nancy F. Lo-Man-Hung1*, Raphaël Marichal1,2, Alexandre B.

Bonaldo1, Leonardo S. Carvalho3, Rafael P. Indicatti4, Stéphanie
Tselouiko2, Catarina Praxedes1, Georges Brown 5, Elena Velasquez6,
Thibaud Decaёns7, Marlucia Martins1 & Patrick Lavelle2,6
1
Museu Paraense Emílio Goeldi, Coordenação de Zoologia, Belém, Pará, Brasil
2
Université Pierre et Marie Curie, Cedex, France
3
Universidade Federal do Piauí, Campus Amílcar Ferreira Sobral, Floriano, Piauí, Brazil
4
Instituto Butantan, Laboratório de Artrópodes, São Paulo, São Paulo, Brazil
5
Embrapa Florestas, Colombo, Paraná, Brazil
6
Centro Internacional de Agricultura Tropical (CIAT), Unidad Suelos, Cali, Colombia
7
Université de Rouen, UFR Sciences et Techniques, Mont Saint Aignan, France
*
Corresponding author: nancylo@terra.com.br
Much of the biodiversity found in tropical forests lives in the soil

(Decaёns et. al. 2006). Unfortunately the global demand for agricultural
commodities produced in the Amazonia is growing and this region is
increasingly affected by drought, fragmentation and forest fires (Laurance et. al.
2002). The changing of soil use in Amazonia is intense and the conversion of
native forest for agricultural practices or pasture is the principal factor pressing
the remaining native forests (Fearnside 2006). Additionally, very few
information about diversity patterns of soil arachnid is available in Amazonia
and the consequences of different land management techniques on this fauna are
virtually unknown. The soil macrofauna plays important rules regulating soil
physical properties, soil carbon storage and maintaining nutrient cycles, thus
providing a whole host of ecosystem services that helps to increase the
heterogeneity in soils and the soil ecosystem’s resilience and resistance to
ecological disturbance (Decaёns et. al. 1994).
Understanding changes that directly influence these processes are
important for local people, whose livelihoods depend on agriculture practices or
pasture areas. In fact, determining the composition of macrofauna is crucial to
understand the ecologic complex of these assemblages. Additionally, several
organisms of soil fauna can be proposed as bioindicators of soil quality and
sustainability. The AMAZ (2007-2011) is an international project carried on
Brazil and Colombia aiming to evaluate Amazonian landscapes changed by
6
Research support by: ANR (Agence Nationale de la Recherche, França), CNPq
(Conselho Nacional de Desenvolvimento Científico e Tecnológico), IRD (Institut de
Recherche pour le Développement), UFPA (Universidade Federal do Pará), UFRA
(Universidade Federal Rural da Amazônia), MPEG (Museu Paraense Emílio Goeldi).
251
familiar agriculture. The social and economic importance of this kind of

agriculture is well recognized, but its environmental impact is not sufficiently
known. Therefore, this study integrates the project AMAZ under the title
"Ecosystems Services and Agricultural Systems Support on Landscapes in
Oriental Amazonia" with the objective to evaluate socioeconomic aspects, the
impacts on the landscape structure and the biodiversity in Amazonia. The
samples were extending to sites in Colombia and Brazil but, so far, reliable
results are limited to samples taken in Brazilian Amazonia.
The experiment was established between April and June 2008, during the
dry season. Three sites (windows) were chosen on southwestern Pará state:
Palmares II in Parauapebas municipality, Maçaranduba in Nova Ipixuna
municipality and Travessão 338 Sul in Pacajá municipality. Each window was
divided in three sub-windows (total of 9), and each sub-window was divided in
three plots (total of 45). Each plot was divided in five sampling points (45 points
each window). A total of 135 spatially independent points were sampled. Five
different vegetation types (systems) under each window were identified: forest,
“capoeira” (secondary forest), “juquira” (secondary forest dominated by
herbaceous and shrubs), pasture and “roça’ (shifting cultivation). The sampling
methodology used to collect soil and litter macro-invertebrates was based on the
Tropical Soil Biology and Fertility Program (TSBF method): which was the
digging on soil one block of dimensions 25 cm long, 25 cm wide and 20 cm
deep and two blocks 25x25x10 cm on each sampled point. The macrofauna were
hand-sorted in the field and the arachnids were stored in 80% alcohol. In the
laboratory, the arachnids were counted and identified. All analyses were based
only on adults. The voucher specimens are stored in the collection of Museu
Paraense Emílio Goeldi in Belém, Pará, Brazil. Patterns of species richness
between three windows and vegetation system were analyzed by visual
inspection of 95% confidence intervals of individual-based rarefaction curves.
To compare arachnids’ communities among the various types of land use and to
try to identify their determinants, we used a principal component analysis
(PCA).
From a total of 897 arachnids (Araneae, Opiliones, Pseudoscorpiones,
Schizomida and Scorpiones), 111 species (including morphospecies) were
identified: 48 species in Palmares, 52 in Maçaranduba, and 37 in Pacajá.
Rarefaction analyses showed that Pacajá significantly harboured more species of
Arachnida than the other windows, although none of the accumulation curves
approached saturation. Forest system comprised significantly more species of
Arachnida (78 adults – 50 species) followed by juquira (35-30), capoeira (22-
14), pasture (22-13) and roça (11-9).
The abundance and diversity of arachnids communities varied
significantly in relation to the land use. The principal component analysis clearly
separated two point of forest system (in Palmares II and Maçaranduba) from the
other systems. According the Monte-Carlo test (p=0.001) there was a significant
difference of the relationships of the arachnid communities between the
windows. The PCA result suggested that this pattern can at least be partly
explained by changes in structural features of the soil, by agricultural practices
252
or pasture. Altogether, in forest systems, biological activity is concentrated in

the litter and few centimetres upper the soil. In fact, most similar studies
reported a reduction in the macrofauna density in used landscapes when
compared with the original forest (e.g. Lavelle et. al. 1992, Decaёns et.al. 1994).
Our study suggests that soil arachnid fauna can be considered as a sensitive
indicator of land management.
As mentioned, forest and juquira systems had higher abundance and
species richness than the remaining systems. Besides, these systems exhibited
more dense litter and consequently more soil biological activity. This conclusion
is supported by other studies in Amazonia where population sizes and species
richness are clear negatively influenced by human impacts on land management.
The addition of the samples obtained in Colombia and the analysis of soil and
environmental variables will be essential to understand the population dynamics
and the process of arachnofauna re-colonization in these regenerating
environments.
References
Decaёns T., Lavelle P., Jimenez Jaen J.J., Escobar G. & Rippstein G. 1994.
Impact of land management on soil macrofauna in the Oriental Llanos of
Colombia. European Journal of Soil Biology, 30(4): 157-168.
Decaёns T., Jimenez J.J., Gioia C., Measey G.J. & Lavelle P. 2006. The values
of soil animals for conservation biology. European Journal of Soil
Biology, 42: 23-38.
Fearnside P.M. 2006. Fragile soils and deforestation impacts: The rationale for
environmental services of standing forest as a development paradigm in
Latin America. In: Posey D.A. & Balick M.J. (eds.), Human Impacts on
Amazonia: The Role of Traditional Ecological Knowledge in
Conservation and Development. Columbia University Press, New York,
pp. 158-171.
Laurance W.F., Albernaz A.K.M., Schroth G., Fearnside P.M., Bergen S.,
Venticinque E.M. & Da Costa C. Predictors of deforestation in the
Brazilian Amazon. Journal of Biogeography, 29: 737-748.
Lavelle P., Blanchart E., Martin A., Spain A.V. & Martin S. 1992. The impact of
soil fauna on the properties of soils in the humid tropics. In: Sanchez P.A.
& Lal R. (eds), Myths and science of soils of the tropics. Soil Science
Society of America, 185 p.
253
Resolving the nomenclatural nightmare for tarantulas

with molecular analyses
Stuart J. Longhorn1, Steven Turner2,3 & Alfried Voger3

1
Department of Biology, National University of Ireland at Maynooth, Kildare, EIRE,
sjl197@hotmail.com
2
Department of Entomology, North Carolina State University, Raleigh, NC, USA
3
Department of Entomology, The Natural History Museum (NHM), London, UK
The tarantula family Theraphosidae is the largest mygalomorph spider

family, with about 115 valid genera and more than 900 described species to date.
The group is itself dominated by the new-world subfamily Theraphosinae, with
around 50 currently valid genera and more than 400 species. The most
prominent attribute of this diverse subfamily is the presence of urticating hairs,
with several hair subtypes that may be useful to diagnose both specimen identity
and clarify systematic relationships. Adaptation to differing environments has
had a persuasive impact on the morphology and behaviour of distinct tarantula
species, confusing traditional taxonomy through misinterpretation of
convergence. Add that much of the existing taxonomic foundation is either
unrevised Victorian work, or newer descriptions without examination of suitable
types and small notes of poor scientific rigor, and the nomenclatural nightmare
becomes apparent. Despite this, several genera have been more thoroughly
reworked using modern morphological techniques and higher-quality standards.
This has lead to significant revision and progress in recent years, and now is
more common to reinterpret morphological character congruence in a plausible
evolutionary framework. Yet here, we discuss the ability of molecular data to
provide radical new insights for the systematics and taxonomy of tarantulas, to
resolve long-standing debates and quickly revoke controversial results of
superficial amateur studies. Preliminary analyses with molecular data suggest
the doubtful monophyly of several existing genera within the Theraphosinae,
and quickly highlight priorities for more intensive revision with all available
data. Furthermore, phylogenetic analysis of molecular data can now provide an
independent scaffold of plausible generic relationships, building towards a
robust molecular phylogeny of the subfamily and beyond. When combined with
rigorous morphological analysis and revision of historical material, it should be
possible to construct a stable systematic scheme for the tarantula spiders within
the subfamily Theraphosinae, on which to base broader studies. As molecular
technologies rapidly advance in the post-genomic age, we suggest how high
profile genera and species of tarantula spiders can be use to explore new
sequencing technologies, and gain additional value to historical museum
collections.
254
Towards resolution of the arachnid phylogeny

with molecular data
Stuart J. Longhorn1 & Susan Masta2

1
Dept of Biology, National University of Ireland at Maynooth. Kildare, EIRE,
sjl197@hotmail.com
2
Dept of Biology, Portland State University, Portland, OR, USA, smasta@pdx.edu
Using genomic technologies, modern molecular analyses should be able to

provide useful insights into the ancient evolutionary history of diverse arachnid
lineages. Yet, despite recent accumulation of substantial data from complete
mitochondrial genomes, multiple orthologous nuclear genes, and rare genomic
characters, the deepest relationships among arachnids are proving difficult to
resolve. Here, we compare insights from several recent molecular analyses
across major arachnid lineages to explore their relative utility to infer such deep-
phylogenetic relationships. Importantly, we highlight how the organization and
relative arrangements of genes in mitochondrial genomes can have a persuasive
influence on the reliability of such mitogenomic data for systematic inferences.
Expanded or modified taxon sampling can similarly influence analytical results
and subsequent interpretation.
Here, we specifically introduce information from new mitochondrial
genomes of several additional acariform mites and an enigmatic opilioacarid, to
put systematic insights on these lineages in the context of other arachnid orders,
and polarize unusual features of their genome evolution. Given the current taxon
sampling, it appears that mites often have unusually fast rates of mutation
accrual that can lead to conspicuous long-branch effects in molecular analyses of
arachnids, complicating phylogenetic analyses and even misleading the inferred
topology. In contrast, when systematic relationships among major arachnid
groups can be inferred with confidence, it is possible to map on unusual aspects
such as tRNA truncation to give polarized insights about changes over deep-
time.
255
Phylogeny of the spider family Mysmenidae and evolution

of web architecture in “symphytognathoids”
(Araneae: Orbiculariae)
Lara Lopardo1,3, Gonzalo Giribet2 & Gustavo Hormiga3

1
Zoologisches Institut und Museum, Allgemeine und Systematische Zoologie, E-M-A-
University, Greifswald, Germany, laralopardo@gmail.com
2
Biology, Harvard University, Cambridge, MA, USA
3
DC, USA
One of the most distal clades within Orbiculariae is the suprafamilial

group informally known as “symphytognathoids” (sensu Griswold et al., 1998),
which comprises about 400 species of minute orb-weavers that build highly
modified orb webs. The composition and the inter-familial relationships among
symphytognathoids remain contentious and poorly studied. Mysmenidae, a small
symphytognathoid family, is one of the least studied families among all orb-
weaving spiders. Its circumscription and diagnosis have always been elusive, its
monophyly never thoroughly tested. Behavioural aspects have been documented
only for a few species. We combine nucleotide sequences from six genes with an
extensive morphological dataset in a total-evidence phylogenetic analysis (ca.
6100 characters, 109 taxa: 74 mysmenids). We explore the phylogenetic signal
of the combined dataset and of the data partitions. Mysmenidae monophyly is
supported by at least 20 morphological and about 420 molecular
synapomorphies, with relatively high support and moderate clade stability.
Mysmenidae is monophyletic under the combined partitions analysis, including
the morphological dataset. Symphytognathoids are circumscribed and
interrelated as follows: (Theridiosomatidae (Mysmenidae (Synaphridae
(Anapidae + Symphytognathidae))). We also study the evolution web
architecture in symphytognathoids. Our results suggest that within
symphytognathoids the planar orb web evolved independently twice from three-
dimensional webs. Furthermore, the orb web has been secondarily and
independently modified into a sheet or a cobweb three times. The characteristic
spherical web of Mysmeninae evolved once and has never been lost. In addition,
our preferred phylogenetic hypothesis suggests that the kleptoparasitic behaviour
of some mysmenids has a single origin and has never been lost.
References
Griswold C., Coddington J., Hormiga G. & Scharff N. 1998. Phylogeny of the
orb-web building spiders (Araneae, Orbiculariae: Deinopoidea,
Araneoidea). Zoological Journal of the Linnaean Society, 123: 1-99.
256
Take a deep breath… Diversity and evolution of the

respiratory system in “symphytognathoid” spiders
(Araneae: Araneoidea)
Lara Lopardo1,2 & Gustavo Hormiga2
1
University, Greifswald, Germany
2
DC, USA
The great diversity of respiratory arrangements in “symphytognathoids”

(i.e., Theridiosomatidae, Anapidae, Symphytognathidae, Mysmenidae, and
Synaphridae) has resulted in many hypotheses to explain its variation and
evolution. For the anterior system, a pattern of reduction of the primitive, well
developed book lungs has been proposed, with a progressive decrease in leaf
number, towards a complete replacement of book lung leaves by anterior
tracheae. For the posterior system, a pattern of two separate spiracles located
midway between the spinnerets and the epigastric furrow has been hypothesized
as the ancestral condition of Symphytognathidae s.l. (Anapidae + Mysmenidae +
Symphytognathidae), later becoming fused into a single median spiracle situated
posteriorly at the base of the spinnerets, and ultimately being lost. We present
the first comparative morphological study based on an extensive SEM survey of
respiratory structures in symphytognathoids. We explore and discuss the
evolution of such structures in a phylogenetic context, and test some of the
hypotheses on its evolution. We use our recent cladistic hypothesis of
symphytognathoid relationships as the comparative framework to explore the
evolution of these respiratory structures. We propose a new evolutionary pattern
for both the anterior and posterior respiratory systems of symphytognathoids.
Our preferred phylogenetic hypothesis suggests a striking and challenging
pattern of evolutionary changes in the anterior respiratory system, where the
anterior tracheal system evolved from fully developed book lungs and,
conversely, reduced book lungs have originated at least twice from its
homologous tracheal system. Furthermore, our data suggest that structurally
similar book lungs might have evolved from different pathways of tracheal
transformation. A tendency for the posterior tracheal system to become either a
highly complex or completely lost is suggested. The evolutionary implications of
behavioural and morphological variation within symphytognathoids are also
discussed. The driving forces that have produced this exceptional diversity of
respiratory arrangements in symphytognathoids remain unclear.
257
Evolution of head structures in the dwarf spider genus

Oedothorax (Araneae: Linyphiidae): Preliminary results
based on mitochondrial sequence data
Lara Lopardo & Gabriele Uhl

University, Greifswald, Germany, laralopardo@gmail.com
Male cephalic modifications (such as grooves, pits, bulges, lobes, humps,

spires, and sulci) are extremely common in many genera of erigonine spiders.
These dimorphic modifications and their associated arrangements of glandular
tissue seem to play a crucial role during courtship, and at least the cephalic
modifications appear to have originated independently several times across the
subfamily. Males of Oedothorax species exhibit different degrees of cephalic
modifications, glandular tissue arrangements, as well as different mating
strategies. In order to explore the evolution of these features within the genus
and preliminary asses their implications in the evolution of their associated
mating strategies, partial fragments of two mitochondrial genes (16S and CO1)
were analyzed independently and combined into a single total-evidence dataset.
The ingroup includes sequences from up to 76 specimens representing six
European Oedothorax species: O. agrestis, O. apicatus, O. fuscus, O. gibbosus,
O. gibbifer, and O. retusus. The outgroup comprises 10 non-linyphiid species
and representatives of Linyphiinae (16 species), Micronetinae (nine species),
Erigoninae (three species), Mynogleninae (three species) and two Linyphiidae
incertae sedis. Phylogenetic analyses were performed under a variety of
algorithms: parsimony analyses using dynamic and static homology approaches,
maximum likelihood and bayesian inference. We report on the preliminary
results of such analyses, the hypotheses of species phylogenetic relationships
and discuss their evolutionary implications. The phylogenetic hypotheses
resulting from the analyses of the combined dataset do not recover the
monophyly of some Oedothorax species. However, they suggest the presence of
a cephalic hump optimizing at the base of Oedothorax, being secondarily lost at
least in O. agrestis, although no trend can be perceived regarding the evolution
of its size. The sulci seem to have a single origin within the genus, in a distal
clade comprising O. gibbifer, O. retusus, and some representatives of O.
apicatus. We also report on the evolution of the complexity of cephalic
glandular arrangements. In addition, we comment on the discrepancies between
the phylogenetic signals of individual genes: surprisingly, the 16S gene fragment
seems to provide more reliable information regarding species monophyly and
identification than the traditionally used CO1 fragment.
258
Redescription and new distribution records of

Acanthoscurria paulensis (Araneae: Mygalomorphae:
Theraphosidae)
Sylvia M. Lucas1, Felipe S. Paula1, Hector M.O. Gonzalez-Filho1,2

& Antonio D. Brescovit1
1
sylvialucas@butantan.gov.br
2
Uniesp-Faculdade Renascença de São Paulo, Brazil
The genus Acanthoscurria Ausserer, 1871 currently includes 40 species,

two from Central America and 38 from South America (Platnick 2010). Twenty-
seven South American species are cited or described for Brazil (Platnick 2010).
Ten new Acanthoscurria species were described from Brazil by Mello Leitão in
1923. The author did not mention the morphology of the male palpal bulb and
seminal receptacle, important for the identification of species. The spider
collection of the Instituto Butantan holds a great number of specimens belonging
to this genus. This material enabled us to redescribe the male of Acanthoscurria
paulensis Mello Leitão 1923 a common species in state of São Paulo, Brazil and
detect and describe the conspecific female and illustrate the sexual structures of
male and female. Acanthoscurria atrox Vellard, 1924 is considered a junior
synonym of A. paulensis based on the original descriptions and figures of the
male palpal bulb, and also supported by the study of a several specimens from
both type localities, Pirassununga, São Paulo and Campo Grande, Mato Grosso
do Sul, Brazil. Acanthoscurria guaxupe Piza, 1972 based on a male from
Guaxupé, state of Minas Gerais is also considered a junior synonym of A.
paulensis based on examination of the holotype. Acanthoscurria paulensis
resembles A. chacoana Brèthes, 1909, A. juruenicola Mello-Leitão, 1923 and A.
geniculata (C. L. Koch, 1842) by the large size (over 50 mm). The male of A.
paulensis resembles that of A. chacoana, A. juruenicola and A. geniculata by the
aspect of the male palpal bulb with an embolus ending like a shell, due to the
well developed prolateral inferior and superior keels. It can be distinguished
from A. chacoana by the male palpal bulb less compact and with a longer
embolus and from A. juruenicola and A. geniculata by the absence of a third
accessory keel. The female resembles A. chacoana, A. juruenicola and A.
geniculata by the fused base of the spermathecae, and differs by the square or
slightly wider than longer base of the spermathecae and by the terminal lobes
directed towards each other. New distribution records for A. paulensis are
provided from the Brazilian states of Mato Grosso, Goiás, Minas Gerais, Mato
Grosso do Sul, Espírito Santo, Paraná and Rio Grande do Sul.
259
Contrasting phylogeographies underlay among-lineage

variation in species diversification in the spider genus
Dysdera from the Canary Islands
Nuria Macías-Hernández1,2, Leticia Bidegaray-Batista1,
Pedro Oromí2 & Miquel A. Arnedo1
1
Biodiversity Research Institute & Department of Animal Biology, Universitat de
Barcelona, Spain
2
Universidad de La Laguna, Tenerife, Canary Islands, Spain, nemacias@ull.es
Phylogenetic studies at the population/species interface combined with the

use of historical population genetics tools hold great promise for addressing key
questions concerning the geography of speciation and its association to adaptive
processes. We present a comparative phylogeographic and demographic analysis
of two lineages originated as part of the large species radiation of the
woodlouse-hunter spider genus Dysdera in the Canary Islands. Both lineages are
endemic to Tenerife, share similar within-lineage genetic divergences and
estimated time of origin. The morphological and ecological diversification
patterns, however, differ significantly between the two lineages. The first lineage
includes four nominal species, two of which have large, allopatric distributions
on Tenerife (one restricted to laurel forest and the other widespread along
different habitats) and the other two which are cave-dwelling species with
restricted geographical ranges. The second lineage includes a single, widespread
species that exhibits limited amount of phenotypic variation mainly associated to
an elevation gradient. Phylogenetic and population analyses of mitochondrial
and nuclear gene sequence data of 200 individuals confirmed lower gene flow
and deeper geographical population structure in the highly diverse lineage, and
uncovered cryptic diversity in both lineages. Common phylogeographic features
in the two widespread species suggest that the geological history of Tenerife had
left a footprint on the distribution of the genetic diversity. Our results also
indicate that demographic and phylogeographic patterns may explain phenotypic
diversification asymmetries among lineages, and demonstrate that contrasting
ecological strategies (specialist vs. generalist) play a major role on structuring
populations of these species.
260
Phylogenetic radiation of salticid spiders:

so many species, so little time
Wayne P. Maddison
University of British Columbia, Vancouver, Canada, wmaddisn@interchange.ubc.ca
The beautiful diversity of salticid spiders has long proved a challenge for
systematists, enticing but overwhelming. However, one hundred years of
phylogenetic work has yielded progress, and a surprising result: each major
region (the Americas, Afro-Eurasia, and Australasia) is dominated by different
major salticid clades, and conversely many major salticid clades are largely
restricted to a single continental area. Thus, the salticids of each area have
radiated into diverse body forms and ecologies more or less independently from
those in other areas. These replicate radiations provide an opportunity to study
general patterns in community assembly over evolutionary time. A combination
of molecular and fossil data suggests that salticids are young, diversifying during
the Tertiary after the continents had separated. Our reasonably-complete
phylogeny now allows us to consider evolutionary patterns in many features,
from chromosomes to ant mimicry.
261
Spiders (Araneae) of two areas of Caatinga under

different levels of anthropogenic habitat disturbance,
Rio Grande do Norte, Brazil
Marília G. Maia & Alexandre Vasconcellos

araneae@hotmail.com
The fauna of spiders from Estação Ecológica do Seridó (ESEC-Seridó)

was evaluated in two areas of Caatinga under different levels of anthropogenic
habitat disturbance. The ESEC-Seridó is located in the municipalities of Serra
Negra do Norte, Rio Grande do Norte State. A standardized sampling was used
in both sites, with pitfall traps, transects, entomologic-umbrella and direct
sampling in leaf-litter. A total of 942 individuals and 28 families were present in
the two sites, with 25 in the preserved site and 22 in disturbed site. Salticidae
was the family most abundant in both sites, followed by Anyphaenidae. The
pitfall traps was the method that caught the greatest richness of families and
greater abundance of individuals. About 75% of the specimens were young. This
proportion of young in the dry season, compared with the results of an
occasional sampling in the same area during the rainy season, and with the data
of the literature for fauna of spiders in areas of Cerrado and Atlantic forest,
suggests that this may be linked with a strategy theses organism to tolerate the
low supply of resources during the dry season in the Caatinga.
Keywords: araneofauna, biodiversity, semi-arid, richness of families,
sampling methods.
262
Macroecological approach to evolution of group living

in spider genus Stegodyphus (Araneae: Eresidae)
Marija Majer*, Jens-Christian Svenning & Trine Bilde
Department of Biological Sciences, Ecology and Genetics, Aarhus University, Denmark

* Corresponding author: marija.majer@biology.au.dk
Social spiders occur in tropical and subtropical habitats worldwide.

Sociality (non-territorial permanently social behaviour sensu Avilés 1997) in
spiders is rare, it is found in c. 25 species (out of 41.000 described species
(Platnick 2009)), but is widespread across the spider phylogeny (Lubin & Bilde
2007). Evolution of sociality could involve similar ecological factors, as the
phylogenetic relationships among social spiders suggest several independent
origins (Agnarsson et al. 2006).
Genus Stegodyphus contains three independently derived permanent social
species and 15 ancestral subsocial species, distributed across arid and semi arid
habitats in Africa, around the Mediterranean basin, the Middle East and Asia
(Kraus & Kraus 1988). The aim of this study is to perform species distribution
modelling (SDM) on collected presence data of all Stegodyphus species to
identify and test potential ecological or environmental factors that drive
Stegodyphus diversity gradients. Within the same genus, spiders may vary
between permanent social living and periodic‐social living depending on
altitudinal and latitudinal gradients (Powers & Aviles 2007). This geographical
distribution may be related to the size distribution of available prey (Guevara &
Aviles 2007). Moreover, social spider prey capture and nest sites are highly
related to habitat structure which shapes their niche within habitats. In order to
estimate the relative importance of hypothesized environmental predictor
variables, niche utilisation of each species is estimated through correlates of
their ranges with the following variables: annual and seasonal temperature and
precipitation gradients, vegetation index (NDVI from remote-sensed data),
elevation range and slope aspects. These variables have so far been shown to be
correlated to species richness and niche width of European spiders (Entling et al.
2007, Finch et al. 2008).
The understanding of functional relationships shaping Stegodyphus
species niche is relevant for gaining insight to the evolution of group living in
spiders (Downes 1994). My study aims to reveal how diversification of niche
utilisation among species varies in relation to their level of sociality.
References
Agnarsson I., Avilés L., Coddington J. & Maddison W. 2006. Sociality in
Theridiid spiders: repeated origins of an evolutionary dead end. Evolution,
60: 2342-2351.
263
Downes M.F. 1994. Nest of the Social Spider Phryganoporus candidus

(Araneae, Desidae) - Structure, Annual Growth-Cycle and Host-Plant
Relationships. Australian Journal of Zoology, 42: 237-259.
Entling W., Schmidt M.H., Bacher S., Brandl R. & Nentwig W. 2007. Niche
properties of Central European spiders: shading, moisture and the evolution
of the habitat niche. Global Ecology and Biogeography, 16: 440-448.
Finch O.D., Blick T., Schuldt A. 2008. Macroecological patterns of spider species
richness across Europe. Biodiversity and Conservation, 17: 2849-2868.
Guevara J. & Aviles L. 2007. Multiple techniques confirm elevational differences
in insect size that maz influence spider sociality. Ecology, 88: 2015-2023.
Kraus O. & Kraus M. 1988. The genus Stegodyphus (Arachnida, Araneae) sibling
species, species groups, and parallel evolution of social living. Verhandlungen
des Naturwissenschaftlichen Vereins in Hamburg, 30: 151-254.
Lubin Y & Bilde T. 2007. The evolution of sociality in spiders. Advances in the
Study of Behavior, 37: 83-145.
Platnick N.I. 2009. The world spider catalog, version 10.0. AMNHistory, online at:
http://research.amnh.org/entomology/spiders/catalog/index.html
Powers K.S. & Aviles L. 2007. The role of prey size and abundance in the
geographical distribution of spider sociality. Journal of Animal Ecology, 76:
995-1003.
264
Effects of vegetation structure and fire on spider diversity

in New Zealand tussocklands
Jagoba Malumbres-Olarte1, Adrian M. Paterson1,

Rob H. Cruickshank1 & Cor J. Vink2,3
1
Ecology Department, Agricultural and Life Science Faculty, Lincoln University, New
Zealand, Jagoba.malumbres.olarte@gmail.com
2
Biosecurity Group, AgResearch, New Zealand, Cor.Vink@agresearch.co.nz
3
Entomology Research Museum, Lincoln University, New Zealand,
New Zealand native ecosystems have been modified and reduced since the
arrival of humans. Tussock grasslands or, which once covered large parts of the
South Island, have been transformed into pasture for livestock through clearing,
burning and over sowing in the last 150 years. Considered ecologically distinctive
because of their large levels of endemicity and economically important for their
utility as pasture for livestock, tussock grasslands have been the subject of
relatively extensive botanical research. However, their invertebrate, and more
particularly spider fauna has been overlooked and little ecological research has
been carried out on them.
Our study aims to answer some of the questions about the arachnofauna of
these ecosystems. First, we conducted an evaluation of the efficiency of spider
sampling methods in narrow leaved snow tussock (Chionochloa rigida) grasslands
in order to determine which one(s) should be given preference and included in
monitoring protocols or ecological studies, and which conditions they should be
used in. Next, we analysed and identified the biotic and abiotic factors that drive
diversity in tussock covered areas, giving special attention to the families and
species with potential as ecological indicators of changes in the ecosystem.
Finally, we investigated the effects of controlled fire in semi-modified
tussocklands through a medium-long term experimental study, where we assessed
the changes in the spider communities over a period of eight years, covering pre-
and post-fire years. The extent of the presence and impact of exotic species was
also assessed. In addition, the potential of molecular techniques to complete
ecological data and help answer ecological questions was evaluated.

Pitfall traps were identified as the most effective and efficient method to
collect spiders in tussock grasslands. The physical characteristics of tussocks were
determined as the main cause of the limited efficiency of other sampling
techniques, such as suction sampling. The predominantly vertical structure of
tussock plants constrains the living space available to spiders; the thin stems and
tips are too exposed to the wind and severe climatic conditions. Thus, most spiders
are found on the lowest layers of vegetation where the grass blades at the base of
the plant are densely packed. Therefore, methods that target these layers will be
the most successful for collecting a large proportion of the spider fauna. However,
265
unique species were found through other collecting methods. Pitfall trapping
should be combined with other techniques when the objective is to carry out an
exhaustive spider survey or a complete ecological study of the spider community.
A number of biotic and abiotic factors were combined through data
reduction in order to create surrogate explanatory variables that explained the
variation in spider assemblages in space. Such surrogate variables were related to
environmental gradients between areas with different types of vegetation that
indicated differing soil conditions or the three dimensional structure of the plants.
These gradients in vegetation were matched by gradient in spider communities,
with families like Orsolobidae favouring areas with marshland vegetation and
aerial web building families, such as Linyphiidae, preferring sparsely distributed
patches of shrubby plant species. Therefore, and after the confirmation of such
relationships by individual variables, it was concluded that environmental factors,
such as soil moisture affects the plant composition and structure in tussock
ecosystems, which in turn determines the spider composition and the relative
abundances of the families and species present. The information obtained allowed
the identification of certain families and species as potential ecological indicators
of the structure and conditions of a tussock area, which could be included in
monitoring programs for conservation management.
A larger project looking at the changes in invertebrate diversity of native
tussock grasslands of New Zealand provided the opportunity to study and
demonstrate the profound effects of fire in an ecosystem not adapted to regular
burning. This experiment, run for eight years, consisted of plots representing
spring and summer burnt treatments as well as a control treatment with samples
collected before and after a controlled fire. Spring and summer burnt treatments
showed significant differences in spider diversity with respect to the control, with
number of families, species and values of other diversity indicators decreasing
drastically after the fire and remaining significantly different for four years.
Although the overall trend was a decrease in the abundance of most spider
families, the family Linyphiidae showed a large increase in the years following the
fire. This can be explained by their efficient dispersal and ability to colonise new
habitats. An increase in the number of exotic linyphiid species, and most
particularly the species Diplocephalus cristatus, a European species well
established in New Zealand, may be behind this trend. This shows the importance
of addressing the question of the effects of disturbances on the interactions
between native – mostly endemic – and exotic species in New Zealand ecosystems
and the dangers that they may pose to native biodiversity.
Mitochondrial DNA sequences of the species present in the study areas
allowed the separation and identification of undescribed species, which was
necessary to complete the data prior to ecological analyses. In addition, it helped
with the identification of undescribed species, which would have been classified as
exotic species using morphological criteria.
Conclusions
This study shows the complex and combined effects that different
environmental variables have on spider assemblages in tussock grasslands. The
266
abundance of tussocks determines spider diversity at both general and specific

level through their physical structure, which also affects the performance of
collection methods. We also demonstrate that fire as a disturbance has a drastic
effect on spider assemblages, probably in part through changes caused to the
vegetation, which last for several years and may be beneficial for exotic species.
More research is necessary to elucidate the interactions between spiders and
other invertebrates at different trophic levels and thus understand the specific
mechanisms that drive the changes in tussock ecosystems. The use of key or
indicator species or groups like the ones identified in our study have the potential
to help with this task. Furthermore, indicator species may be used in simple
protocols designed for monitoring natural or human caused changes in this
ecosystem. In this context, molecular techniques provide valuable information that
can complete ecological data by identifying cryptic species that otherwise would
not be considered. Molecular techniques were also useful to discern whether some
species were native or exotic.
267
A revision of Gandanameno (Araneae, Eresidae)

and the promise of cybertaxonomy
Mohammad Marhabaie1 & Jeremy Miller2
1
Isfahan University, Isfahan, Iran, marhabaie@gmail.com
2
Nationaal Natuurhistorisch Museum Naturalis, Leiden, The Netherlands,
miller@naturalis.nl
Taxonomic research benefits from easy exchange of people, specimens,

and data around the world. For the most part, experts and libraries are
concentrated in developed countries while biodiversity is concentrated in
developing countries. To ameliorate this disparity, knowledge in the form of
training and publications must be transferred from the developed world to the
developing world. For training, students or experts must travel and participate in
courses or collaborative projects. For taxonomic information, the internet offers
the potential for building a profoundly democratizing resource.
As a student from a developing country (Iran) with virtually no endemic
expertise in arachnology, I (MM) found it very challenging to access basic
information on spider taxonomy. No identification key for Iranian spiders exists.
Libraries featuring good coverage of publications in arachnology do not exist in
Iran. Thus, I turned to the internet and international contacts.
Cybertaxonomy, or eTaxonomy, is a process that involves the use of
standardized electronic tools to access information (databases, e-publications)
and/or to generate knowledge bases such as identification keys (Malte C. Ebach,
BGBM) (http://wp5.e-taxonomy.eu/blog/2007/04/11/what-is-cybertaxonomy/).
While cybertaxonomic content is currently too sparse to be very practical, basic
infrastructure is in place and new content is being added all the time. By contrast
with traditional libraries, online digital information is relatively easy to access in
Iran and throughout most of the developing world.
I recently travelled to Leiden (The Netherlands) for three months of
training in revisionary taxonomy and arachnology. The subject was the small
eresid genus Gandanameno (five described species as of the start of the project),
endemic to southern and eastern Africa. In addition to a standard journal
publication, I will contribute content to cybertaxonomic resources including the
Encyclopedia of Life, the Global Biodiversity Information Facility, and
ZooBank. In the future, I plan to apply what I have learned to develop
knowledge of the spider fauna of Iran and to train new colleagues.
268
How many scuta do the Linyphiidae have?

Yuri M. Marusik
Institute for Biological Problems of the North, Russian Academy of Sciences, Magadan,
Russia, yurmar@mail.ru
Introduction
Scutum is defined as a sclerotized abdominal plate of some spiders
(Dippenaar-Schoeman & Jocqué 1997). The scutum has been described in many
spider families belonging to different superfamilies and even suborders. In the
Holarctic region, spiders with the scutum have been reported from Anapidae
(Comaroma), Araneidae (Cercidia), Corinnidae (♂, Castianeira, Trachelas,
Phrurolithus), Gnaphosidae (males of Zelotini), Linyphiidae (several genera:
Ceratinella, Pelecopsis, Styloctetor, Ceraticellus), Oonopidae (several genera),
Palpimanidae (all species), Salticidae (males of Chalcoscirtus, Euophrys
frontalis, some Aelurillus), Theridiidae (some Euryopis s. l., Pholcomma, etc.),
Zodariidae (several genera) and others. Outside of the Holarctic spiders scutum
has been described in Clubiona s. l. and in several other families (e.g.,
Tetrablemmidae).
Most of arachnologists are familiar with the dorsal abdominal scutum.
However, besides a single dorsal abdominal scutum (rarely two in Oonopidae),
there can be more scuta, from one to six pairs. With regard to the number of
scuta, the world champions are Tetrablemmidae. They possess the dorsal
scutum, up to four ventral unpaired scuta, and up to four pairs of lateral stripe-
like scuta.
Abdominal scuta can be divided into three types according to their
position: dorsal, lateral and ventral. As the dorsal scutum is most common, no
attention is usually paid to the ventral scutum (scuta). Thus, the aim of this
presentation is to describe different types of the ventral scuta and to demonstrate
their importance in the identification and classification of spiders.
In addition to the dorsal scutum, the majority of scutate spiders have one
or more ventral scuta. Ventral scuta have been reported for such genera as
Castianeira (Corinnidae), Palpimanus (Palpimanidae), the male of Euophrys
frontalis (Salticidae), Comaroma (Anapidae), Silhouettella (Oonopidae),
Linyphiidae (several genera), and Zodariidae.
Ventral scuta can be grouped into three types: 1) epigastral (anterior part
of abdomen, in front of the epigastral furrow, 2) inframammillary (close or
around the spinnerets) [the terminology follows Crosby & Bishop 1925] and 3)
intermediate (between the epigastral furrow and spinnerets). All three types can
be represented by a single solid scutum or by several scuta.
Scuta in Linyphiidae
My attention was attracted to ventral scuta when I had found a heavily
scutate Erigoninae spider from the Caucasus. Later, it was described as a new
species in a new genus Scutpelecopsis wunderlichi (Marusik & Gnelitsa, 2009).
269
The male abdomen of this species is almost a capsule, with large dorsal and
ventral scuta leaving two orifices: for the pedicel and for the spinnerets. The
female of this species possesses 5 ventral scuta! The question arose: how could
these numerous scuta have originated, and whether such a scuta are present in
other Erigoninae, or whether S. wunderlichi is unique in this respect.
Dorsal scutum is known in several Erigoninae: Pelecopsis s.l. (several
species), Ceratinella,
Ceraticelus, Idionella, Styloctetor, etc. In most of these taxa only males
possess the scutum. Dorsal scutum usually covers a part of the dorsum, except
for some species of Ceraticelus s.l., Pelecopsis bishopi Kaston, 1945, and
Ceratinella spp.
Ventral scuta: When I had compared S. wunderlichi with other scutate
erigonids, I recognized that many of them have several ventral scuta (maximum
of 5), some of which could be fused: 1) a pair of the infrapetiolar scuta (ip); 2) a
pair of the book-lung scuta (bl); 3) an unpaired inframammillary (im). Epigastral
scuta can be fused to various extend or can even form a single solid scutum. A
fusion level of the epigastric scuta in males is higher than in females. In the male
of S. wunderlichi, all the ventral scuta are fused, which leads to a unique
conformation of the genital area. In the literature, I have found only one
mentioning of the paired epigastral scuta.
Fig. 1. Ventral scuta in some erigonids, showing different levels of the sclerotization and
fusion of scuta. Males have larger scuta than females, ip - infrapetiolar scutum; bl -
book-lung scutum; im – inframammillary scutum.
270
While studying scutate erigonids, I have found two interesting structures

connected to the ventral scuta. The tracheal spiracle is always situated on the
inframammillary scutum. Such the spiracle is displaced and wider compared to
the species lacking the inframammillary scutum.
The anterior part of the male scutum of S. wunderlichi or the infrapetiolar scuta
in females seem to have stridulatory organs (setae with enlarged bases). Perhaps
similar structures are present in other armored linyphiids (Ceraticelus and/or
Idionella). I have not studied other scutate taxa by means of SEM.
Taxonomy
A comparison of different populations of widespread Ceratinella wideri
has revealed differences in the size of ventral scuta. This fact may indicate that
the Far Eastern populations may belong to a different species or subspecies. A
comparison of different species of Ceratinella or scutate Pelecopsis from
Finland has demonstrated that the shape of epigastral scuta can be used as an
additional character for species identification.
Acknowledgements
This project was supported in part by the Russian Foundation for Basic
Research grants: 08-04-92230-ГФЕН, 09-04-01365 and 10-04-91225.
References
Crosby C.R. & Bishop S.C. 1925. Studies in New York spiders; Genera: Ceratinella
and Ceraticelus. New York State Museum Bulletin, 264: 1-71.
Dippenaar-Schoeman A.S. & Jocqué R. 1997. African Spiders: An Identification
Manual. Plant Protection Res. Inst. Handbook, no. 9, Pretoria, 392 pp.
Marusik Y.M. & Gnelitsa V.A. 2009. Description of a new genus of spiders from
the eastern Mediterranean and the most armored erigonid species from the
western Caucasus (Aranei: Linyphiidae: Erigoninae). Arthropoda Selecta, 18:
57-68.
271
Conformation of the male palp in some spiders belonging

to the RTA-clade and problems in taxonomy
Yuri M. Marusik1 & David Penney2
1
Institute for Biological Problems of the North, Russian Academy of Sciences,
Magadan, Russia, yurmar@mail.ru
2
Faculty of Life Sciences, The University of Manchester, United Kingdom,
david.penney@manchester.ac.uk
Introduction
For a long time the suprageneric classification of spiders was based
exclusively on somatic characters (as in entomology). Lehtinen (1967) was the
first to clearly demonstrate the importance of copulatory organs in spider
classification. He significantly changed the whole principles of spider taxonomy
and made many changes. He especially made many changes in the classification
of families within the RTA-clade.
While working with different families in the RTA-clade we recognized
that many supraspecific taxa (genera, tribes, subfamilies) have an identical
conformation of the male palp (we call this conformation the Dictyna-type palp).
If the Dictyna-type palp is plesiomorphic, then the occurrence of this palpal
conformation in different families can be expected. What surprised us however,
was that several families have very different types of male palp, despite their
somatic morphology being more or less uniform. For example all Dictynidae or
Cybaeidae have the same type of palp, while in Agelenidae, one subfamily
(Ageleninae) includes groups with completely different palps (Agelenini vs
Tegenariini & Textrixini). A similar situation occurs in Desidae and Hahniidae.
Terminology
Before further discussing morphology and possible relationships between
taxa with similar palps or similar somatic morphologies, we should consider
some terminological issues.
The current use of terminology is rather chaotic: 1) In some instances
different names are used for homologous structures, whereas in others the same
name is applied to non-homologous structures. 2) The etymology of terms is
often based on very different principles, and same term can be applied for
different structures on the same palp. We recognized at least six different
etymological backgrounds for palpal structures:
1) Topographical/position: median, terminal, subterminal, sub(tegulum),
supra(tegulum), basal, retrolateral, palea (if derived from ‘upper’).
2) Deviating: (belonging, derivative): tegular, tibial, embolic, radial,
paracymbium.
3) Shape: lamella characteristica, palea (if derived from ‘membranous’).
4) Functional: Conductor, embolus.
5) Constructional: radix, psembolus, embolic division.
272
6) Patronymic: Fickert’s gland.

The same is true for epigynal structures. Examples of confusables include:
Spermatheca: in Hersiliidae used in addition to receptaculum.
Receptaculum (seminis): in Hersiliidae used in addition to spermatheca.
Ducts (insemination): furrows vs true ducts (many groups have closed furrows
(Araneidae, Linyphiidae, Tegenaria, etc; while many groups have true
ducts Hahniidae, Lathys, Theridiidae).
Septum: applied for non-homologous parts.
Scapus (scape): applied for nonhomologous, even for non-morphologically or
non-functionally similar things.
It may be impossible to standardize terminology for all spiders, but an
effort to use same term for homologous structures should be made, at least in
sister groups.
Always remember in morphological analysis/comparison that the same
term is often used for non-homologous characters; and that different names can
be used for homologous characters in different groups.
Priority is good (great) in the ICZN because it serves for the stabilization
of taxonomic nomenclature, but this idea is not applicable to morphology.
An unlimited number of names are available in taxonomy, but in
comparison very few terms can be used in descriptive morphology. Species
names are more or less fixed, while the interpretation of morphological terms
tend to be broader or thinner.
Suggestion: if an author is not sure what to call a specific structure, it is
better to give it a neutral term, and stress that it is not a real establishment of a
new term, but just a term used in that specific publication for a character of
unclear status with regard to origin, homology etc.
The Dictyna-type of palp

In this presentation we concentrate on the Dictyna-type of palp as it
occurs in different families, in taxa currently attributed to Agelenidae,
Argyronetidae, Cybaeidae, Desidae, Dictynidae, Hahniidae and possibly
Amaurobiidae. In Agelenidae it occurs in Ageleninae: Textricini and all
Tegenariini.
Such a pattern of distribution can be explained by at least two alternative
hypotheses:
1) The same type of the male palp evolved independently in somatically
different families, even independently within different tribes of the same
subfamily.
or
2) The different somatic characters, such as spinneret structure, position of
tracheal spiracle, etc. are a result of divergence from a common ancestral stock
with a Dictyna-type of palp.
But how can we explain why Agelenidae and Hahniidae have at least two
different types of palp, one of which is the Dictyna-type and another, which is
totally different (Agelena versus Textrix or Hahnia versus Cryphoeca)? The
most parsimonious explanation is that the current systematic classification of
273
these groups is poor, because it is based only on somatic characters and totally
ignores the copulatory organs.
How we define the Dictyna-type of palp.
1) Fungus-like (two-armed) conductor
2) Round and large base of embolus (if short)
3) Conductor originates near the base of the embolus
4) Flat tegulum, with seminal duct following a circular course
5) Filamentous embolus
6) Modified (complicated) tip (retrolateral tip) of conductor.
7) Conductor (retrolateral part) terminates on the retrolateral side of the bulbus.
Fig. 1. Examples of Dictyna- and Tegenaria- subtypes of the male palp.
The upper (prolateral) part of the conductor can be extraordinarily

enlarged (Lathys, see Marusik et al. 2006) or almost reduced (Cicurina).
This basic type can be split in two easily recognizable subtypes: Dictyna
s. str. type & Tegenaria-type.
Tegenaria-type is easily recognizable due to 1) presence of median
(tegular) apophysis, starting near the base of the conductor; 2) in most genera the
prolateral part of the conductor terminates at the level of the mid-part of the bulb
and is bifid (exception Azeriscape).
274
In all Dictyna s. str. type the retrolateral part of the conductor terminates,
or at least passes through (nearby) the base of the bulbus (cymbial-tibial
connection).
Dictyna s. str. type occurs in:
Cybaeidae: Cybaeus s. l., (treated by Lehtinen (1967) in Dictynidae, as a
subfamily).
Argyronetidae: Argyroneta (treated by Lehtinen (1967) in Dictynidae, as a
subfamily).
Desidae: Paratheuma, Corthesia.
Dictynidae: Dictyninae, Tricholathysinae (Devade, Argenna, Saltonia
(previously placed in Agelenidae), etc); Cicurininae (Yorima, Cicurina,
Blabomma, Brommella).
Agelenidae: Agelininae: Textricini (Textrix).
“Hahniidae”: Cryphoeca (?C. montana).
Mastigusa – currently in Dictynidae, but palp not typical, terminal arm
terminates in the wrong place, placed by Lehtinen (1967) in Tuberta.
Tuberta – currently in Hahniidae:+Cryphoecinae, but the palp is more similar to
the Dictyna-type.
Tegenaria-type:
Agelenidae/Ageleninae: Tegenariini.
“Hahniidae”: Cryphoeca silvicola and related taxa
Fig. 2. Distribution of the Dictyna- and Tegenaria- subtypes among different taxa.
Note: Tegenaria-subtype is plesiomorphic in comparison to Dictyna (apomorphy

– reduction of median (=tegular) apophysis).
Why Dictyna-type but not Tegenaria-type if Tegenaria has plesiomorphic state?
Tegenaria-type is known in one subfamily of Agelenidae, and one current
subfamily in Hahniidae), but Dictyna-type is known in at least five families, in at
least seven subfamilies or tribes.=> Dictyna-type widespread, better known and
causes more ‘problems’ (open questions) in comparison to the Tegenaria-type.
275
Conclusion
What is wrong (or what seems less probable than others): 1) taxonomy, 2)
somatic morphology, or 3) morphology of the copulatory organs? In our opinion
taxonomy and somatic morphology are causing the problems. It is not difficult to
test what is more correct, just apply DNA analysis for a selected group. It may
be more appropriate to test simple, but clear hypotheses rather than attempting to
make a standard cladogram in the first instance. For example we suggest to
compare two absolutely different types (Agelena – Tegenaria) and two subtypes
of Dictyna, for example: Agelena (not a Dictyna-type) – Tegenaria (Tegenaria
subtype) – Dictyna or Agelena – Tegenaria – Cryphoeca – Dictyna or Tegenaria
– Dictyna – Cybaeus – Textrix.
Acknowledgements
This project was supported in part by the Russian Foundation for Basic
Research grants: 08-04-92230-ГФЕН, 09-04-01365 and 10-04-91225.
References
Lehtinen P.T. 1967. Classification of the cribellate spiders and some allied
families, with notes on the evolution of the suborder Araneomorpha.
Annales Zoologici Fennici, 4: 199-468.
Marusik Yu. M., Ovchinnikov S.V. & Koponen S. 2006. Uncommon
conformation of the male palp in common Holarctic species belonging to
the Lathys stigmatisata group (Araneae, Dictynidae). Bulletin of the
British Arachnological Society, 13(9): 353-360.
276
Chelicerate relationships as inferred by mitogenomic data

Susan Masta
Portland State University, Portland, OR, USA, smasta@pdx.edu
Relationships among the lineages of chelicerates have proved difficult to

infer with confidence. Without well-supported phylogenetic hypotheses of
relationships, we can gain only a limited understanding of how features of these
organisms have evolved. My lab has addressed this challenge by sequencing
mitochondrial genomes from divergent lineages of arachnids and pycnogonids.
We use the sequence data from mitochondrial genomes, in conjunction with the
rare structural changes we have discovered in these genomes, to help infer
systematic relationships. Together, these different types of data are yielding
surprising information about the evolution of chelicerates.
Ancient gene rearrangements have reshaped the mitochondrial genome in
multiple lineages of chelicerates. For example, within spiders, numerous gene
rearrangements occurred after opistothele spiders diverged from mesothele
spiders, but before the opistothele lineages diversified. I map these gene
rearrangements onto a phylogeny of chelicerates inferred from amino acid
sequence data from the 13 mitochondrial protein-coding genes. Additionally, the
sizes and structures of some mitochondrial genes differ drastically within certain
lineages of arachnids, despite their strong conservation in other metazoans.
These rare changes in genes that are otherwise highly conserved can also be used
in phylogenetic inference. In general, rare genomic changes show little to no
homoplasy within chelicerates, and can thus be used as informative characters
for phylogeny reconstruction to help us understand aspects of chelicerate
evolution.
277
Vertical stratification of spiders

in Kuttanad rice agroecosystem, India
Elizabeth V. Mathew*, Ambalaparambil Vasu Sudhikumar

& Pothalil Antony Sebastian
Division of Arachnology, Department of Zoology, Sacred Heart College, India

*
Corresponding author: elizabethvmathew.here@gmail.com
Introduction
Spiders are of economic value to mankind because of their ability to
suppress pest abundance in agroecosystems. The diversity and density of spiders
are important in any kind of attempt for the implementation of integrated pest
management (IPM). They exhibit extremely high diversity and are the dominant
insectivores in many terrestrial ecosystems. Faced with the need to reduce
pesticide usage on crops and optimize natural biological control, full
investigation of the means by which spiders influence pest abundance is long
overdue.
Rice agroecosystems demand greater attention as rice is the staple food in
developing countries. Asia accounts for more than 90% of the world’s rice
production and consumption. Being a wetland agroecosystem, paddy cultivation
is different from other agricultural productions. The biodiversity of wetland rice
ecosystem is highly varied than that of many natural ecosystems (Schoenly et al.
1996). Consequently, there are a growing number of investigations in which
spiders in agroecosystems are used as tools to gain insights into the role of
generalist predators in community and ecosystem function (Sunderland 1999).
Studies of the rice field spider fauna of India are limited to the identification of
spiders and investigation of the dominant spider species (Sebastian et al. 2005).
It is less common for workers to analyze the vertical stratification of spider
community in the field (Anbalagan & Narayanasamy 1999). In this context, a
preliminary study was conducted on vertical stratification of spiders in the
Kuttanad rice agroecosystem to analyze the microhabitat association of spiders
in paddy fields.

Kuttanad (9°17'N-9°40'N, 76°19'E-76°33'E), one of the “Rice bowls of
Kerala”, is the region selected for the study. It contributes nearly 20% of the
total rice production of the state. The investigation was carried out for a period
of 6 months, from July 2009 to January 2010. Sampling was conducted during
two seasons viz., Rabi 1 (Monsoon crop: July 2009 to September 2009) and
Kharif 1 (Winter crop: November 2009 to January 2010). The Rabi season is
characterized by heavy rain (Southwest monsoon) and high humidity while the
Kharif season is characterized by low rainfall and dry weather (Menon et al.
2000).
278
Spiders were collected by handpicking. For the present study, five main
functional groups were recognized based on the activity and foraging behaviour
related to average height of the rice plant, namely <20 cm from water/soil
surface, 20-40 cm, 40-60 cm, 60-80 cm and >80 cm (Lee & Kim 2001). The
final growth stage of each plant was thoroughly examined from top to bottom,
on leaf blades, flowers, dry leaves and ground stratum. The ground area near
plants was also searched. The location where the spiders were spotted was taken
note of. Spiders were collected by leading them into glass vials from the ground
stratum and terminals of the plants. The collected specimens were preserved in
70% ethyl alcohol, properly labelled with collection locality, date and other
significant notes. They were then identified with the help of available literature
(Barrion & Litsinger 1999).
Results
A total of 1632 individuals from 70 species, 45 genera and 15 families
were collected during the study period. The most species rich family was
Salticidae (15 species) followed by Tetragnathidae (12 species) and Araneidae
(7 species). Spiders were divided into five strata based on the activity and
foraging behaviour related to average height of the rice plant. The spiders which
build perfect orb-webs were mainly present at the canopy level of the crop.
Hence Araneidae and Tetragnathidae were found foraging mainly at the top
layer of the rice plants. This stratum provides sufficient area for the construction
of web and increases the chance of prey entanglement in the webs. There is very
little chance of locating ground dwelling spiders at the canopy level of the plant.
Ground dwellers such as lycosids were mainly present at the bottom level of the
field, although there is possibility of these spiders coming up for pursuing the
insect prey. The spiders which build irregular cob-webs were also present near
the bottom of the field or below half level of the average plant height.
The spiders collected during the study were classified into 7 ecological
guilds based on the foraging mode of the spiders. The spider guild classification
was composed according to the families collected during the study. Designation
of the spider guild was based on the ecological characteristic known for the
family (Young & Edwards 1990). Among the 70 species of spiders collected
from Kuttanad, 60% belongs to stalkers (30%) and orb weavers (30%). The
second dominant guilds are the ground runners and space web builders (11%
each). Ambushers (10%), sheet web builders (5%) and foliage runners (3%) are
the other ecological guilds of these spiders.
Spiders under the category stalkers actively jump over the prey for
feeding. Spiders of the family Salticidae and Oxyopidae show this type of
feeding behaviour. Oxyopidae represented by 6 species of a single genus and
Salticidae consisted of 15 species coming under 12 genera. Spiders of the orb
weavers guild construct perfect orb webs for prey capture. Families Araneidae (7
species), Tetragnathidae (12 species) and Uloboridae (2 species) constitute this
category.
Ground running spiders mainly feed on ground layer of the field and rarely
come to the foliage or canopy of the plant for prey capture. The families
279
Lycosidae (6 species) and Scytodidae (2 species) come under this guild. Spiders
of the guild space web builders construct irregularly spaced webs for prey
capture. Belonging to this category are the families Pholcidae (2 species) and
Theridiidae (6 species). Ambushers show a “sit-and-wait” type of behaviour for
prey capture. Spiders of the families Philodromidae (1 species), Pisauridae (1
species) and Thomisidae (5 species) are members of this guild.
Foliage runners hunt on foliage for phytophagous insect pests. This guild
is formed of 2 families viz., Clubionidae and Miturgidae with one species each.
Spiders of the guild sheet web builders construct sheet like web for capture.
Only one family of paddy field spiders belong to this category namely,
Linyphiidae, with 3 species.
Discussion
The variation in spider diversity between the base of the plant and its
canopy was primarily due to the difference in position of their habitation in the
paddy field. The most common explanation for the observed pattern of spider
guild structure accounts for the effects of host crop, including its structural
diversity, micro environment and the level of disturbance (Young & Edwards
1990). The web building and plant wandering spiders rely on vegetation for
some part of their lives. The structure of the vegetation is therefore expected to
influence the diversity of spiders found in the habitat.
Habitat structure maintains diverse spider assemblages (Uetz 1991, Wise
1993) and may be critical to successful insect suppression (Riechert & Bishop
1990). Uetz (1991) suggested that structurally more complex shrubs can support
more diverse spider community. The physical structure of the environment has
an important influence on the habitat preferences of the spider species especially
web-building species. Choice of foraging habitat has been recognized as one of
primary importance through its effect on feeding rates, with derived benefits to
growth and reproduction.
Once in feeding patch, the spider is confronted with an array of potential
prey species. Since more than half of the predatory fauna in rice agroecosystem
are spiders and it is known that changes in spider density can impact pest
populations (Nyefeller et al. 1994), it would seem logical that the spider
community would be a key component of integrated pest management strategies.
In order to increase the emphasis on spiders as agents of biological control, it is
imperative to decipher exactly how crop growth stages that change the habitat
structure within relatively homogenous fields influence the density, diversity and
foraging behaviour of spiders. This will ultimately lead to the path of increased
food security.
280
Molecular and morphological analysis of Sicarius

systematics across a wide geographic range
Elise N. Maxwell1, Jeremy Miller2 & Greta J. Binford1
1
Department of Biology, Lewis & Clark College, Portland, OR, USA
2
National Museum of Natural History, Leiden, The Netherlands
The taxonomy of the Haplogyne spider genus Siciarus has been piecemeal
with no unified systematic revision. We present a standardized analysis of
Sicarius systematics across a wide geographic range of taxon sampling from
Africa, Central and South America. We analyzed genetic structure in the lineage
using molecular phylogenetics of 28S, CO1, 16S and ND1. These analyses show
consistent support for many terminal clades, but little consistency of patterns of
relationships at deeper levels. We also investigated a wide range of somatic and
genitalic morphological characters. Out of all characters examined, genitalic
characters are the most helpful in differentiating species. We confidently assign
species names to most of the South American taxa, but are not confident about
assigning species names to any of our African taxa given the current
nomenclature. This work provides the basis for a future thorough systematic
revision.
281
Araneofauna of the Western Ghats of India

Mundackatharappel J. Mathew1, Pothalil A. Sebastian2
& John Joseph2
1
IT@School Project, Kochi, India, mathewmj@asianetindia.com
2
Sacred Heart College, Kochi, India
Introduction
The Western Ghats is a chain of mountains running parallel to India’s
western coast, between the latitudes 8ºN to 20ºN and longitudes 73ºE to 77ºE.
They cover an area of about 160,000 km² and stretch for 1,600 km from the
country’s southern tip. Western Ghats supports some of the pristine forest
patches. The average height of the Ghats is 1500 m above sea level, but in the
southern reaches, it rises up to 2000 m and to exceptionally higher peaks of 2500
m and above. This area is one of the world's ten "Hottest biodiversity hotspots"
and at least 325 globally threatened species occur here (Myers et al. 2000). This
region exhibits very high level of endemism, which ranges from 75% in
amphibians to 40% in insects and hence, the Western Ghats is listed among the
34 biodiversity hotspots of the world (Mittermeier et al. 2005).
Although around 15% of the Western Ghats is protected in 20 national
parks and 68 sanctuaries, extremely high population pressure has seriously
stressed the region's biodiversity. Nearly 40 million people inhabit this hotspot,
at a density of 260 people/km² (one of the highest in hotspots) (Jha et al. 2000).
The forests of the Western Ghats have been selectively logged and highly
fragmented throughout the entire range. Forests have been converted to
agricultural lands for monoculture plantations and are also cleared for building
reservoirs, roads, and railways. Much of the remaining forest cover consists of
timber plantations or disturbed secondary growth. Due to the paucity of workers
and funds, a large share of spider diversity in the Western Ghats still remains
unexplored. As a result, the disappearance of many species is left
undocumented.
Studies on patterns of species richness, diversity and their spatial
distribution has widely been acknowledged as sine qua non for effective
conservation planning (Cushman & Mc Garigal 2003, 2004). Against this
backdrop, a study was conducted to document the araneofuana of the Kerala
region of Western Ghats, as well as to analyse their diversity, abundance and
ecology.

The study area consisted of Kerala Region of Western Ghats. In order to
conduct exploratory surveys of spider species richness and diversity, the
different study sites corresponding to the various geographic locations of the
Western Ghats of Kerala region were identified. The collection sites selected for
the study include Agasthyamalai Biosphere Reserve, Periyar Tiger Reserve
282
(West Division), Periyar Tiger Reserve (East Division), Idukki Wildlife

Sanctuary, Chimmony Wildlife Sanctuary, Chinnar Wildlife Sanctuary, Silent
Valley National Park and Wynad Wildlife Sanctuary. The random transect
method using the technique adopted by Aiken and Coyle (2000) was used for
spider sampling. This technique involved a combination of four collection
methods to assess the diversity of spider fauna namely, ground hand collection,
aerial hand collection, beating and sweeping. Time was used as a as a measure
of sampling effort to make the methods comparable. One sample unit equalled
one hour of uninterrupted time during which all spiders encountered were
collected (Sebastian et al. 2005). Sampling was conducted monthly in the
selected study sites for a period of three years from February 2006 to February
2009. The diversity, richness, and evenness indices of spider communities were
computed using the software SPDIVERS.BAS of Ludwig and Reynolds (1998).
A detailed analysis was also made on the taxonomy, endemism, affinities and
conservation status. Habitat associations of spiders were observed and recorded.
The sampling data was also utilized to analyse the guild composition as well as
seasonality.

The study documented 173 species of spiders belonging to 85 genera and
25 families from Western Ghats of Kerala including 6 new species. At the
generic level, 4 genera and at the species level, 6 species were new records to
India. Discovery of new species, as well as sighting of a number of species and
genera for the first time from India indicates the biological wealth of this region
and further points out the necessity of more detailed exploration in order to
comprehensively understand the biodiversity of the country. A total of 6 species
recorded from the study area endemic to Kerala and 29 species endemic to Indo-
Sri Lankan region. It has been observed that the araneofauna of Western Ghats
of Kerala bear affinities mainly to oriental and Palearctic regions, as well as to
the fauna of Sri Lanka. Analysis of the faunal composition revealed that
Araneidae was the taxonomically dominant family. Abundance data revealed
that Pardosa pseudoannulata (Family Lycosidae) was the most abundant
species. Analysis of the guild structure revealed that orb weavers were the
dominant feeding guild. Analysis of the various diversity indices revealed that
Periyar Tiger Reserve East Division recorded the highest values for most of the
diversity indices computed. The overall trends in abundance of spider
community showed correlation with seasons. The highest species occurrence
was recorded in the post-monsoon months followed by pre-monsoon months and
the least occurrence was during the monsoon period. The 2009 IUCN Red Data
Book (IUCN 2009) lists 26 species of spiders as threatened. Among them, 3 spp.
were recorded from Western Ghats of Kerala during this investigation. These are
Haploclastus kayi (endangered), Poecilotheria rufilata (endangered) and P.
striata (vulnerable). Nonetheless, these small numbers reflect how little we
actually know about spider populations in general, let alone the real extent of
endangered species. Many species may be threatened, endangered or extinct, but
research on them is lacking. Unfortunately, due to this limited information on
283
distribution, population and conservation status of spiders, it is difficult to get

them listed under national or international legislation. Conserving biological
wealth requires action in both areas of endemic species and areas of high
biological diversity. The uniqueness of species compositions, as indicated by
levels of endemism and habitat specialization, is more important in establishing
regional conservation priorities (Platnick 1991). Threatened centres of endemism
are major biodiversity hotspots (Roberts et al. 2002), and conservation efforts
targeted toward them could help to avert the loss of biodiversity. The present
investigation could serve as a baseline for future study of spiders of Western
Ghats.
References
Aiken M. & Coyle F.A. 2000. Habitat distribution, life history and behavior of
Tetragnatha spider species in the Great Smoky Mountains National Park.
Cushman S.A. & McGarigal K. 2003. Landscape-level patterns of avian diversity in
the Oregon Coast Range. Ecological Monographs, 73: 259-281.
Cushman S.A. & McGarigal. K. 2004. Hierarchical analysis of forest bird species
environment relationships in the Oregon Coast Range. Ecological
Applications, 14: 1090-1105.
IUCN 2009. IUCN Red List of Threatened Species. Version 2009.1.
www.iucnredlist.org. Downloaded on 18 September 2009.
Jha C.S., Dutt C.B.S. & Bawa K.S. 2000. Deforestation and land use changes in
Western Ghats, India. Current Science, 79: 231-238.
Ludwig J.A. & Reynolds J.F. 1998. Statistical Ecology: A primer on methods in
computing. John Wiley & Sons, New York, 352 pp.
Mittermeier R.A., Patricio R.G., Hoffman M., Pilgrim J., Brooks T., Mittermeier
C.G., Lamoreux J. &. Fonseca G.A.B. 2005. Hotspots revisited: Earth's
biologically richest and most endangered terrestrial ecoregions.
Conservation International, pp. 1-432.
Myers N., Mittermeier R.A., Mittermeier C.G., Da Fonseca G.A.B. & Kent J. 2000.
Biodiversity Hotspots for Conservation Priorities. Nature, 403: 853-858.
Platnick N.I. 1991. Patterns of biodiversity: tropical vs. temperate. Journal of
Natural History, 25: 1083-1088.
Roberts C.M., Colin J.M. & John E.N., et al. 2002. Marine Biodiversity
Hotspots and Conservation Priorities for Tropical Reefs. Science, 295
(5558): 1280-1284.
Sebastian P.A., Mathew M.J., Pathummal Beevi S., Joseph J. & Biju C.R. 2005.
The Spider Fauna of the Irrigated Rice Ecosystem in Central Kerala, India
across Different Elevational Ranges. Journal of Arachnology, 33: 247-255.
284
Cytogenetical characterization of five orb-weaver species

(Araneae: Araneidae) with special regard to the diploid
number polymorphism in Parawixia velutina
(Taczanowski, 1878)
Viviane F. Mattos1, Marcia G. Kraeski1, Leonardo S. Carvalho2,

Antonio D. Brescovit3 & Douglas Araujo1
1
Universidade Estadual de Mato Grosso do Sul, Unidade de Mundo Novo, Mato Grosso
do Sul, Brazil, vivianefagundesmn@hotmail.com, marcia_kraeski@hotmail.com,
daraujo@uems.br
2
Universidade Federal do Piauí, Campus Amílcar Ferreira Sobral, Piauí, Brazil,
carvalho@ufpi.edu.br
3
adbresc@terra.com.br
Araneidae comprises 168 genera and 2992 species taxonomically

described, with about 65 species from 18 genera cytogenetically characterized.
Thus, represents the third largest family concerning the number of described
species as well as chromosomal data. From the 65 species karyotyped, more than
50 revealed a diploid complement composed by 2n♂=24, with 22 autosomal
elements and a sex chromosome system of the type X1X2. On other studied
species the diploid number varied from 2n♂=13 to 2n♂=49. Besides the sex
chromosome system of the type X1X2, the most common within the family, the
type X occurred in a smaller frequency, and the X1X2X3 and XY type were
registered only once for each species.
In Araneidae, the chromosomal morphology is acro/telocentric on most
species. The aim of this work is to characterize Alpaida truncata (Keyserling,
1865), Alpaida veniliae (Keyserling, 1865), Parawixia kochi (Taczanowski,
1873), Parawixia velutina (Taczanowski, 1878), and Wagneriana sp. in relation
to the diploid number, chromosomal morphology, and sex chromosome system
type.
The spiders were collected at Parque Nacional de Ilha Grande, in the
boundary between the states of Mato Grosso do Sul and Paraná, Brazil. After the
gonads dissection, the specimens were deposited in the arachnological collection
of Instituto Butantan, municipality of São Paulo, State of São Paulo. All
individuals were collected in São Francisco Island, with the exception of A.
veniliae, which was collected at the margins of Xambrê lake, both in the State of
Paraná, Brazil. The chromosomal preparations were obtained from testis and
ovaries of two male specimens of A. truncata, one male and one female of A.
veniliae, one male of P. kochi, four males and three females of P. velutina, and
one male of Wagneriana sp. The gonads were dissected in physiologic solution
(7.5 g of NaCl, 2.38 g of Na2HPO4, 2.72 g of KH2PO4, diluted in 1 litre of
distilled water) and submitted to a 0.16% colchicine solution (prepared with
285
physiologic solution) during 2 hours. The hypotonic treatment consisted in

immersion of the gonads in tap water during 20 minutes, and the fixation is
performed with Methanol: Acetic Acid solution (3:1). The microscopy slides
containing the preparations were stained with 3% Giemsa solution. The mitotic
and meiotic cells were photographed using a digital capture system coupled to a
light microscope.
The analyses of spermatogonial diplotenes in A. truncata and A. veniliae
showed meiotic formulae composed by 10 autosomal bivalents and two sexual
univalents (10II+X1X2) for both species, indicating a diploid number of 2n♂=22,
composed by 20 autosomes and the X1X2 sex chromosomes. In A. truncata the
X1 and X2 sex univalents appears always associated side by side. On both
species, the sex chromosomes are heteropycnotic in a number of diplotene
nuclei. The majority of autosomal bivalents presented only one terminal chiasm,
but bivalents with an interstitial chiasm also occurred. A female pachytene cell
of A. veniliae revealed 12 chromosomal bivalents, indicating possibly the
occurrence of 2n♀=24, composed by 20 autosomes and the sex chromosome
system of the type X1X1X2X2. The chromosomal morphology was not
established in both species, due to the lack of mitotic or metaphase II cells. The
observation of spermatogonial diplotenes of P. kochi and P. velutina showed 10
and 11 autosomal bivalents, respectively. Both species presented two sexual
univalents that always appear side by side and correspond to the X1 and X2
chromosomes. Therefore, the meiotic formulae of P. kochi is 10II+X1X2,
corresponding to a diploid number of 2n♂=22. In P. velutina the meiotic
formulae is 11II+X1X2, indicating a diploid complement of 2n♂=24. This
complement was confirmed by the presence of 2n♀=26 in oogonial metaphases
of two females in P. velutina. Mitotic metaphases of one female individual of P.
velutina presented an astonishing characteristic, the diploid number is composed
by 2n♀=25, with a large metacentric chromosome, without pair, contrasting with
the telocentric elements of the karyotype. In both species, the autosomal
bivalents presented only one interstitial or terminal chiasma. Differential
pycnosis was not noticed in any chromosome of the complement in Parawixia
species. Only one metaphase II nucleus was found in P. kochi, evidencing n=10
chromosomes.
The analyses of male diplotene cells in Wagneriana sp. revealed 10
autosomal bivalents and two sexual univalents (10II+X1X2), suggesting a diploid
number of 2n♂=22. The autosomal bivalents presented only one terminal or
interstitial chiasma. The sex chromosomes were heteropycnotic in relation to the
autosomes and appeared always closely associated. It was not possible to
identify the chromosomal morphology due to the lack of mitotic metaphases of
metaphases II. The differences regarding male diploid number between P. kochi
(2n=22) and P. velutina (2n=24) could be useful as a cytotaxonomical character,
together with the external genital characteristics described on a taxonomical
review of the genus, taking into account that both species were collected in the
same area.
Among all araneids, the diploid number of 2n♂=22 was registered for the
first time for P. kochi, A. truncata, A. veniliae, and Wagneriana sp., all studied
286
in this work. It is interesting to notice that according to a taxonomical review,

the genus Parawixia, Alpaida, and Wagneriana shared some external
morphological putative synapomorphies. The presence of species with 2n=22
exclusively in these genera among araneids could be a reinforcement of their
close relationship. Unfortunately, there is no cytogenetical data on other genera
considered closely related to Parawixia, such as Eriophora and Acanthepeira.
Parawixia velutina with 2n♀=25 represents a case of numerical
polymorphism within a population. Taken into account that the other specimens
of P. velutina, as well as the other araneids in general, presents exclusively
acro/telocentric chromosomes, the metacentric element observed in the
polymorphic specimen has arisen through a Robertsonian fusion (centric fusion),
in which the centromeric regions of two acro/telocentric chromosomes fuse to
form a single meta/submetacentric chromosome, generating a heterozygosis
condition. This type of rearrangement is a very common evolutionary change
and has been reported in most groups of organisms. In many cases, chromosome
changes produce reproductive barriers when they cause problems at meiosis in
heterozygotes, leading to reduced fertility, due to the formation of a trivalent
during meiosis and unbalanced segregation. However, there are a number of
ways in which a trivalent can generate balanced gametes after segregation.
Unfortunately, as the polymorphic individual was a female, it was not
possible to observe the pairing, because it is rare the presence of certain meiotic
stages. However, it can be supposed that the fusioned metacentric pairs with
their two homologous telocentric elements, formed a trivalent. Centric fusions
have been of major importance in the karyotype evolution of many groups of
animals. This type of rearrangement can be observed in the heterozygous
condition in some population, as presently verified in the cytotype of P. velutina
with 2n♀=25, but in many cases the homozygous condition rapidly reached
fixation, with the original telo/acrocentric elements disappearing from the
population.
It is not possible to determine if the chromosome number polymorphism
found is presented over a wide geographic area or if there are two chromosome
cytotypes, one with 2n♀=26 and the other with 2n♀=24, each cytologically
monomorphic throughout most of its range, but with a zone of overlap in which
chromosome number heterozygotes with 2n♀=25 occurs. The distinction will be
clear only after an extensive investigation involving a study of many
populations.
In this study we found four species with 2n♂=22, suggesting that this
diploid number could be more common in araneids than anyone could imagine.
This diploid number is also very frequent in Theridiidae, another araneoid
family. We also show that even in some genera, as Parawixia and Alpaida, some
species presents 2n♂=24 while others presents 2n♂=22, which suggests that the
rearrangements involved in the conversion of 24 to 22 chromosomes, or 22 to 24
chromosomes are relatively common among araneoid spiders. With some degree
of chance the intermediary or heterozygotic condition can also be found, as in P.
velutina with 2n♀=25. A similar case of heterozygosis in spiders was registered
in Evarcha hoyi (Peckham & Peckham, 1883; Salticidae).
287
Some araneids with diploid numbers smaller than 2n=24, such as some
Neoscona species with 2n=14, presented predominantly biarmed chromosomes
in the karyotype, evidencing the occurrence of centric fusions. On the other
hand, Gasteracantha possess 2n=16 exclusively acro/telocentric chromosomes.
According with some literature proposals, tandem/centric fusion followed by
pericentric inversion from a karyotype with 2n♂=24 acro/telocentric
chromosomes are involved in the origin of the karyotype with 2n=16. Due to the
pericentric inversions, there is no metacentric chromosomes in the karyotype
with 2n=16, despite the chromosome number reduction.
288
Host - parasite relations between spiders (Araneae)

and terrestrial Parasitengona mites
(Acari: Actinotrichida, Prostigmata)
Joanna Mąkol1,2 & Magdalena Felska1
1
Institute of Biology, Department of Invertebrate Systematics and Ecology, Wrocław
University of Environmental and Life Sciences, Poland,
joanna.makol@up.wroc.pl
2
Institute of Natural Sciences, Wrocław University of Environmental and Life Sciences,
Poland
Terrestrial Parasitengona mites are known as protelean parasites of

arthropods and vertebrates, the only exception being the record pertaining to
molluscs. Data on host–parasite relations between spiders and Parasitengona
mites are scattered in the literature. In many cases the host identification is
limited to higher taxonomic ranks (e.g. order or family), this being a serious
limitation in studies on the host spectrum of parasitic species. The summaries
concerning the present topic have been provided by Welbourn and Young (1988)
and Fain and Jocque (1996a). Baker and Selden (1997) have summarised the
records on Araneae, parasitised by larvae of Leptus (Parasitengona:
Erythraeidae).
The aim of this work is to summarise all the existing data on larvae of
Parasitengona terrestria, which exploit spiders as hosts. The list of taxa is based
on the literature and results of the recent survey of alcohol-preserved material
originating from Poland, Russia and France.
Larvae of four, out of 14, Parasitengona families: Erythraeidae (eight
species), Trombidiidae (seven species), Microtrombidiidae (one species) and
Eutrombidiidae (two species) were recorded as parasites of spiders representing
19 families. Eight records apply to new, hitherto unpublished data. Achaearanea
lunata, Theridion varians (Theridiidae), Bathyphantes alticeps, Helophora
insignis, Oedothorax retusus, Prinerigone vagans (Linyphiidae), Pachygnatha
clercki and Tetragnatha montana (Tetragnathidae) were recorded as hosts of
terrestrial Parasitengona for the first time. Out of 28 species of Araneae recorded
as hosts, only two, i.e. Pachygnatha clercki Sundevall, 1823 and Nuctenea
umbratica (Clerck, 1758) were parasitised, though not simultaneously, by larvae
of two different Parasitengona species.
All new records involve the interaction with the representative of one mite
species, i.e. Trombidium brevimanum (Berlese, 1910). For the time being T.
brevimanum remains the only member of terrestrial Parasitengona, whose wide
host spectrum points to affiliation with Arachnida, with particular reference to
spiders as regular hosts of these mites.
289
Metropolitan spider fauna in the Brazilian Atlantic Forest

remnants under urban pressure
Tércio S. Melo, Marcelo Cesar L. Peres, Marcelo A. Dias,

Katia R. Benati, Alessandra R.S. Andrade, Marcos Vinicius A.
Guimarães & Sheila Luzia de Santana Varjão
Catholic University of Salvador, Centro de Ecologia e Conservação Animal, Brazil,

terciosilvamelo@hotmail.com
In urban influenced landscapes such as those found in Salvador, the

heterogenity of habitats may influence the local biodiversity. The habitats are
highly modified by human occupation, bringing a variety of new elements, such
as building sites, motorways, shopping centres and industries (Pickett et al.
2006). The forest remnants created by fragmentation represent important refugia
for a high number of species (Rodrigues et al. 1993, Brown Jr & Freitas 2003).
The spider distribution patterns in these rainforest landscapes is still
poorly known (Brescovit 2009). Therefore, this study objective was to build a
spider database for the fauna of Salvador city Atlantic Forest remnants and to
compare the species composition for seven most important localities in order to
undertake the future conservation actions.

The data came from the arachnological collection of the Centro de
Ecologia e Conservação Animal at the Universidade Católica do Salvador, and
from surveys carried out in seven Atlantic Forest remnants in the city of
Salvador, Bahia, Brazil between 2000 and 2009. The surveyed remnants were
the 19º Batalhão de Caçadores de Salvador – 19 BC – with an area of 166 ha; the
4ª Companhia de Guardas de Salvador – 4 CGS - 60 ha; the Área de Proteção
Ambiental Lagoas e Dunas do Abaeté, 1800 ha; the Grande Moinho Aratu
remnant – GMA - 5 ha; the Jardim Botânico de Salvador – JBS - 18 ha; the
Parque Joventino Silva park – PJS - 72 ha; and the Parque Metropolitano de
Pituaçu – PMP - 425 ha. For all of these Forest remnants the urban matrix was
made, showing low connectivity and mostly composed of motorways, residential
zones and business areas.
The spiders were sampled with pitfall traps (19 BC, GMA, Abaeté, PJS e
PMP), direct searching (4ªCGS, JBS, PJS, PMP), Berlesse funnels, canopy traps
(PMP), beating trays (Abaeté, JBS, PJS,PMP) and Winkler extractors (19 BC e
GM). In addition, 39 samples of one meter square leaf litter were taken for
sampling in the Winkler extractor for 24 hours each.
Spiders were identified to the species or genus level, otherwise classified
as morpho-species. Specimens were deposited at the Coleção Aracnológica in
290
the Laboratório de Artrópodes Peçonhentos do Instituto Butantan (curated by Dr.

Antônio Brescovit).
Presence (1) and absence (0) matrix of the recorded species were created
for fauna composition analysis in each remnant, using data collection from non
standardized techniques during surveys. We also used three replicated remnants
to compare species composition (4ªCGS, JBS e PJS), once they had the same
sample design applied during the study effort. The multiple response
permutation procedure - MRPP (PcOrd ©: McCune and Mefford, 1999),
applying the Bray Curtis distance, was then used to compare these remnants
composition. This method does not have the assumption of Gausian distribution,
therefore was considered the most suitable for this study.

We registered 7,182 spiders representing 44 families. Araneidae,
Corinnidae, Ctenidae, Oonopidae, Scytodidae and Theridiidae were the most
frequent. The families Clubionidae, Diguetidae, Hahniidae, Mysmenidae,
Pisauridae were represented only by immature individuals in all sample sites and
surveys.
Among the recorded species, we identified 3,201 adults from 39 families,
143 genera, 170 species and morpho species. A study conducted in 1,390 ha
Atlantic Forest remnants, distant 100 km from Salvador, the authors found 130
species (Pinto-Leite et al. 2008). Peres et al. (2006) registered 117 species in an
urban matrix remnant in the Brazilian north-eastern region. The information
found in literature suggests that the richness in Salvador can be considered low,
if compared with other similar regions and landscapes. Halaj et al. (1998) stated
that changes in the structural complexity and the spatial heterogeneity may result
in the loss of richness of spider species in time.
Some recorded families presented higher richness: Salticidae (n=28;
16,47%), Theridiidae (n=26; 15,29%), Araneidae (n=16; 9,41%), Corinnidae
(n=10; 5,88%) and Oonopidae (n=10; 5,88%). The most frequent species were
Scytodes fusca Walckenaer, 1837, showing 100% frequency, and Alpaida sp.,
Corinna sp., Mesabovilar sp., Tenedos sp., presenting 85,71%. Higher frequency
of Araneidae, Salticidae and Theridiidae, may be related to the environmental
structure - secondary Forest formations promoting web builders and foraging
behaviour mode of Salticidae (Wise 1993).
We also found a significant difference for richness, when remnants were
compared (MRPP: p=0.0000, A=0.05803102, T=-6.8678563). When post hoc
comparison were performed, it was found significant differences between 4ª
CGS and JBS (MRPP: p=0.0003, A=0.04575580, T=-4.9105393), between
4ªCGS and PJS (MRPP: p=0.00562091, A=0.02763257, T=-3.0660198), and
JBS and PJS (MRPP: p=0.0000, A=0.06077587, T=-6.6071755). These may
suggest that events such as, fragmentation (Brazil et al. 2005), fragment size and
age (Bolger et al. 2000, Laurence & Vasconcelos 2009), the form and distance
291
between remnants (Steffan-Dewenter & Tscharntke 2002, Mcintyre 2000), as

well as the landscape matrix where these fragments are immerged, may directly
influence the species composition (Laurence & Vasconcelos 2009). However,
these structural differences in association with the loss of connectivity, which
was consequently promoted by the urban growth along the century, may have
contributed to the differentiation among these remnants as observed by Gibb and
Hochuli (2002) in Sydney, Australia.
The spider species composition recorded in the Atlantic Forest remnants in
Salvador city was considered low. However, and even though they are located in
the same urban matrix, this composition was different, indicating that remnant
size and shape is important.
References
Brandão C.R.F., Cancello E.M. & Yamamoto C.I. 2000. Avalização do estado
do conhecimento da diversidade biológica do Brasil - invertebrados
terrestre - USP, Museu de Zoologia.
Brescovit A. 2009. Aranhas e araneísmo: uma curta historia das araneomorphae
peçonhentas da bahia. Gazeta Medica da Bahia, 79, 78.
Gibb H. & Hochuli D.F. 2002. Habitat fragmentation in an urban environment:
large and small fragments support different arthropod assemblages.
Biological Conservation, 106: 91-100.
Halaj J., Ross D.W. & Moldenke A.R. 1998. Habitat structure and prey
availability as predictors of the abundance and community organization of
spiders in western Oregon forest canopies. The Journal of Arachnology,
26:203-220.
Laurence W.F. & Vasconcelos H.L. 2009. Conseqüências ecológicas da
fragmentação florestal na Amazônia. Oecologia Brasiliensis, 13(3): 434-451.
Mcintyre N.E. 2000. Ecology of Urban Arthropods: A Review and a Call to
Action. Annals of the entomological society of America, 93.
Mcintyre N.E., Rang J., Fagan W.F. & Faeth S.H. 2001. Ground arthropod
community structure in a heterogeneous urban environment. Landscape
and Urban Planning, 52: 257-274.
Pickett S.T.A. & Cadenasso M.L. 2006. Advancing urban ecological studies:
frameworks, concepts, and results from the Baltimore ecosystem study.
Australian Ecology, 31(2): 114-125.
Pinto-Leite C.M., Guerrero A.C. & Brazil T.K. 2008. Non-random patterns of
spider species composition in an Atlantic rainforest. The Journal of
Wise D.H. 1993. Spiders in Ecological Webs. Cambridge University,
Cambridge.
292
Spider fauna of coastal and floodplain meadows in Matsalu

(Estonia), influence of flooding and management
Mart Meriste
Tartu Collage of Tallinn University of Technology, Estonia, mart.meriste@ttu.ee
Spiders play an essential role in grassland ecosystems, as they are among

most important predators and form an important link between grass layer and soil
food chains. According to the environmental conditions and food availability, spider
community composition can be very variable in different grasslands habitats.
Due to more or less regular floods, coastal and floodplain meadows are rather
uncomfortable habitat for spiders. Flooding is lethal for most spiders, although some
species can survive shorter flooding either by climbing on the higher vegetation or
even sunken under water for few days. Due to the harsh environmental conditions,
spider fauna of flooded habitats is similar to other disturbed sites and consist mostly
of euryoecious species with good dispersal ability. When comparing habitats where
flooding occurs – coastal and floodplain meadows – there are differences in the
duration of flooding as well as origin (and salinity) of flooding water. Salinity of
water determines the differences the plant community composition, but how the
different flooding regimes (and salinity of water) are affecting spiders, is largely
unknown.
Here I studied the spider community composition and dynamics in semi-
natural grasslands with different flooding regime. Studied communities situate in
historical river delta area. In this delta, floodplain meadows turn slowly to coastal
meadows on the coasts of long and narrow Matsalu bay. Sites on floodplain are
influenced by fresh water from rivers (mainly Kasari river), while coastal sites are
influenced by brackish water from the sea. Thus, a salinity gradient forms from
floodplain to the coast of open sea. Intermediate areas are usually flooded by rivers
but with strong western winds the sea water rises in the narrow bay and flow to these
grasslands. Some of the grasslands in the region are currently managed, whereas
others are abandoned already years ago. Hence, semi-natural grasslands in this
region provide good system for studying the influence of flooding regime and
management on spider communities.
Spider fauna of the floodplain meadows in Estonia has earlier been described
by A. Vilbaste 1964 but on coastal meadows the knowledge is very scarce.
However, information about fauna of seminatural grasslands is certainly needed due
to their possibly high and characteristic diversity and occurrence of rare species. In
addition, spiders particularly are considered to be precise indicators of the
environmental conditions in communities.
The aim of this study was to:
1. investigate the abundance and diversity of spider fauna in flooded
seminatural grasslands in Western Estonia;
293
2. study environmental conditions that affect spider community composition,

especially flooding in both coastal and floodplain meadows and in sites with
intermediate origin;
3. study differences in spider fauna in managed and unmanaged sites.

Study is based on the material collected from Matsalu National Park in
western Estonia in 2006. Spiders were collected with pitfall traps from ten
different wet meadow sites which formed a gradient from typical floodplain
meadow to the typical coastal meadow. The salinity of habitats in Matsalu bay
are dependent on the balance between the fresh water from river Kasari and the
brackish water (around 6‰) from the sea. Thus the salinity of the flooding water
is different in different meadows. The area, duration and depth of inundation are
dependent on height from the sea level which on average was between 0.3 and
3.6m in studied sites. In three locations (two coastal and one floodplain
meadow), the pairs of managed and unmanaged sites for comparison were
chosen. Organic matter content, soil moisture, K and Na content were measured
from soil samples.
Results
Altogether 1460 adult and 170 juvenile spider specimens were collected,
but also other invertebrates that fell into pitfall traps were recorded. Total spider
abundance and diversity did not differ according to the height from sea level.
The abundance of Lycosid spiders decreased significantly from floodplain
meadows towards coastal ones. This is probably not because the saltwater
influence but more likely due the different flooding regime and duration. The
abundance and diversity of spiders were not dependent on the abundances of
other arthropods as potential pray or other predators as potential competitors.
Significant differences occurred between managed and unmanaged sites, but the
pattern differed between meadow types. It is noteworthy that considering
different ecotypes of spiders in coastal meadows (eurytoecious, hygrophilous,
xerophilous etc.), the faunas of both unmanaged sites seems to be quite similar
whereas grazed sites were very different. This is probably because higher
vegetation in unmanaged sites that creates stable condition for spider community
and protects it against the floods. Vegetation in managed sites is very low and
the structure of spider fauna is more random.
In conclusion, spider fauna on flooded grasslands in Estonia consists
mainly of euryoecious species, but the community structure is dependent on the
characteristics of the inundation. The differences in spider diversity and
abundance between managed and unmanaged sites can be explained by different
vegetation structure.
294
Impact of local prey abundance on a colonial spider,

Cyrtophora citricola (Araneidae) in a hyper-arid
environment
Laia Mestre1,2 & Yael Lubin3

1
Departament de Biologia Animal, Biologia Vegetal i Ecologia (BABVE). Centre de
Recerca Ecològica i Aplicacions Forestals (CREAF). Facultat de Biociències,
Universitat Autònoma de Barcelona, Barcelona, Spain, laia.mestre@uab.cat
2
Centre de Recerca Ecològica i Aplicacions Forestals (CREAF). Facultat de
Biociències. Universitat Autònoma de Barcelona, Barcelona, Spain
3
Mitrani Department of Desert Ecology. Blaustein Institutes for Desert Research, Ben-
Gurion University of the Negev, Sede Boqer Campus, Israel, lubin@bgu.ac.il
Several studies on colonial spiders have pointed at prey abundance as the

main cause of grouping tendency. However, the link between group formation
due to higher food availability and colony success in a given site remains largely
unexplored, with most work focusing either on group formation or on the
advantages of coloniality. We investigated the effect of prey availability on
juvenile site tenacity and on colony success in the spider Cyrtophora citricola
(Araneidae).
We carried out two field surveys on spider colonies found in Acacia trees
in the Arava valley, a hyper-arid region in Israel. Some trees were near an
organic fertilizer depot that attracted large numbers of insects (“compost sites”)
and others were at increasing distances from it (“non-compost sites”). In each
survey, we counted the number of spiders and eggs and measured colony volume
in each tree, as well as sampling flying insects with sticky traps. We further
conducted two experiments inside net houses with Acacia trees, where we left
females to build webs and released juveniles on these webs after a few days. We
tested the effect of prey remains in the female’s webs on the establishment
decisions of the juveniles (first experiment), and the effect of food supply on
their tendency to remain in the web (second experiment).
Colonies in the compost sites were more successful both in numbers of
individuals per colony and fecundity, but colony success and prey abundance
were not correlated. The field experiments showed that juvenile spiders are able
to detect prey remains in a female web: they settled with lower probability when
they were released on empty webs in comparison with webs containing prey
remains. However, once established, differences in food supply only slightly
affected the number of juveniles that remained in the female web, which
suggests that individuals are reluctant to disperse after an establishment decision
has been made on the basis of indirect cues.
295
An unseen world… First phylogenetic analysis of spiders

(Araneae) based on sperm morphology and male genital
system
Peter Michalik1 & Martín J. Ramírez2

1
Zoologisches Institut und Museum, Ernst-Moritz-Arndt Universität Greifswald,
Germany, michalik@uni-greifswald.de
2
Museo Argentino de Ciencias Naturales „Bernadino Rivadavia“ - CONICET, Buenos
Aires, Argentina
Spiders are characterized by an astonishing diversity in sperm morphology,

seminal fluids and organization of the male genital system (Michalik 2009). Despite
the high amount of potential phylogenetic information that sperm characters might
offer, they have never been included in phylogenetic analyses to date. Here, we
present the first approach to incorporate such characters in previous analyses of
spider relationships. In addition, we explore character transformations based on the
resulting phylogenetic hypothesis in order to elucidate the evolution of spider
spermatozoa and male genital system. We defined 45 sperm characters at different
organizational levels (cellular to aggregation levels). Most sperm characters are
related to the level of sperm cell organelles, e.g., shape of nucleus and acrosomal
complex, axonemal patterns, or centriole-associated structures. More than 90 species
representing 38 families of spiders were investigated by means of transmission
electron microscopy. The taxon sampling includes representatives of all main spider
lineages, i.e. Mesothelae, Mygalomorphae (5 families), Haplogynae (10 families)
and Entelegynae (23 families). Characters were added to previous and ongoing
morphological phylogenetic matrices and analyzed using parsimony. Based on the
results of our analyses spermatozoal synapomorphies can be proposed for the
following taxa: Mesothelae, Mygalomorphae, Araneomorphae, Liphistiidae,
Heptathelidae, Atypidae, Dipluridae, Haplogynae (excl. Filistatidae), Filistatidae,
Segestriidae, Dysderidae, Oonopidae, Pholcidae, Ochyroceratidae, Sicariidae und
Scytodidae, Mimetidae, Araneidae, Mysmenidae, Anapidae, Linyphiidae,
Nesticidae and Theridiidae, Synotaxidae, the „RTA clade“, Philodromidae,
Dictynidae, Anyphaenidae, Corinnidae, Sparassidae and Selenopidae. Our results
further revealed insights in the evolution of spermatozoa, sperm aggregates (transfer
forms) and the male genital system which are also discussed.
References
Michalik P. 2009. The male genital system of spiders (Araneae) with notes on
the fine structure of seminal secretions. Contributions to Natural History,
12: 959-972.
296
Permanent sperm depletion in spiders (Araneae)

Peter Michalik1 & Clare C. Rittschof2
1
Zoologisches Institut und Museum, Ernst-Moritz-Arndt-Universität, Greifswald,
Germany, michalik@uni-greifswald.de
2
Department of Biology, University of Florida, Gainesville, FL, USA, critter@ufl.edu
Sperm is a costly and limited resource for males. In some spider species,
there is behavioural documentation that males permanently deplete all of their
lifetime sperm after a single mating. No study, however, has confirmed this
phenomenon by addressing the mechanistic bases of sperm depletion in spiders.
Here we evaluate permanent sperm depletion in the spider Nephila clavipes
using gross measurements and light and transmission electron microscopy of the
male genital system. Male testis size decreases as males approach their
maturation moult and reaches its lowest point after the first sperm induction. The
decrease in testis size corresponds to the maturation process of the sperm and a
subsequent absence of testis tissue devoted to sperm production as males mature
and age. Spermatogenesis occurs only during the young male sub-adult stage,
and no male testes produce sperm after the terminal moult. Permanent sperm
depletion has implications for the evolution of male mating strategies and male
terminal investment strategies in spiders.
297
Latradectism in Iran and laboratory scale antivenom

production against Latrodectus tredicimguttatus
Abbas Zare Mirakabadi, Hamid Reza Goudarzi,

Housein Zolfagharian & Naser Mohammadpur
Department of venomous animals and antivenom production, Razi Vaccine and Serum
Research Institute, Karaj, Iran, abbas.zare8@gmail.com
Hundreds of people suffer from spider bite yearly in Iran. Reports from
hospitals in Khorasan province indicate the serious cases of spider bites which
some cases lead to death in children. The duration of treatment varies from few
days to two weeks in hospital. In the present work the venom of spider was
extracted by removing the venom glands from spider Latrodectus
tredecimguttatus and dissolved in distilled water. LD50 of the venom was
determined and found to be 18.5 when the venom extraction was carried out in
4C. Two healthy horses were selected for immunization and the antigen was
prepared using complete and non complete Frunds adjuvant. Immunization of
horses were carried out for the first time in 9 injection protocol and the titer of
antivenom was found to be 315 LD50 while when second time immunization
was carried out in 5 series of injection the titer of antivenom was found to be
762 LD50. The specific tests for standardization of antivenom was carried
according to WHO recommended parameters. The total protein of product was
estimated to be 2.76% and dry weight was 4.09%. PH and phenol content of
product was adjusted to 6.75 and 1.5g/l respectively. Pyrogeny test indicated that
the product is free of endotoxins and pyrogenic factors. In conclusion the
laboratory scale of spider antivenom production showed that there is possibility
of producing safe and effective anivenom against medically important spiders of
Iran.
298
A new anophthalmous species of the genus

Paranemastoma Redikorzev, 1936 from Bulgaria
(Opiliones: Nemastomatidae)
Plamen G. Mitov
Department of Zoology and Anthropology, Faculty of Biology, Sofia University, Sofia,

Bulgaria, pl_mitov@yahoo.com
In this paper a new eyeless representative of Paranemastoma, collected

from the Stojkova Dupka 1 cave in the Slavyanka Mountains (SW Bulgaria) is
described and illustrated based on a single male specimen and two juveniles. The
penis, chelicerae and form of the pedipalp, as well as the absence of scutum
armament, clearly separates this new species from any other nemastomatid. The
closest species morphologically, and the only other eyeless example, is the
troglobiont Paranemastoma (Buresiola) bureschi, which is known from
numerous caves in the Balkan mountain range (Stara Planina Mts.).
299
Nemastomatid harvestman (Opiliones) in Baltic amber

Plamen G. Mitov1 & Jason A. Dunlop2
1
Department of Zoology and Anthropology, Faculty of Biology, Sofia University, Sofia,
Bulgaria, pl_mitov@yahoo.com
2
Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity
at the Humboldt University Berlin, Berlin, Germany, jason.dunlop@mfn-berlin.de
Nematostomatid harvestmen (Arachnida: Opiliones: Dyspnoi:

Nematosomatidae) are described from Palaeogene (Eocene) Baltic amber. The
fauna includes the first fossil example of the subfamily Ortholasmatinae. This is
a significant find for the Palaeogene of Europe, given that modern
ortholasmatines are restricted to East Asia and North/Central America. Within
the other subfamily, Nematostominae, three new species, each in a new genus,
can be recognised. Considering species previously described in the literature,
Mitostoma gruberi Dunlop and Mitov, 2009 – previously known from the
German Bitterfeld amber – is now recorded for the first time in Baltic amber.
Generally, the absence of (male) genital characters makes unequivocal
comparisons with extant genera difficult, but both ?Histricostoma tuberculatum
(Koch and Berendt, 1854) and ?Mitostoma denticulatum (Koch and Berendt,
1854) show subtle somatic differences compared to living representatives of
these groups; in particular the legs and pedipalps of the fossils appear to be
longer. These species probably belong to new (fossil) genera. Key pedipalp
characters cannot be resolved in the poorly-preserved holotype of Nemastoma
incertum Koch and Berendt, 1854, which is here treated as a nomen dubium.
300
High species richness and abundance of grassland dwelling

spiders in the landscape with an intermediate level
of forest surroundings
Tadashi Miyashita & Yuki Chishiki

Laboratory of Biodiversity Science, School of Agriculture and Life Sciences, University
of Tokyo, Japan, tmiya@es.a.u-tokyo.ac.jp
Spiders are one of the most diverse and abundant arthropod predators in
terrestrial ecosystems, and they often determine population and community
structures at lower trophic levels. In particular, insect pests for agricultural crops
are believed to be kept at low densities by spiders. These characteristics led
researchers to examine the effects of environmental factors on species richness
and abundance of spiders in various ecosystems. However, most studies were
conducted either in small-scale experimental settings, or by using descriptive
statistics that examined the relationship between spider richness/abundance and
local environmental measures. Here we investigated how species richness and
abundance of grassland-dwelling spiders are determined in multi-scale contexts
in a typical agricultural landscape in Japan, which consists mostly of rice fields
and secondary forests. Because mosaic structure of the landscape is considered
to increase species richness, we expected that spiders are more diverse in the
grasslands with an intermediate level of forest surroundings.
We have chosen 35 sampling sites on Sado island, the largest island of
Japan; each site being located at least 500 m apart from another. We collected
spiders inhabiting grasslands by sweeping, and recorded local vegetation
structure (plant height, plant species composition). To identify factors
determining species richness and abundance of spiders, we used GLIM (error
distribution was either Poisson or negative binomial) that includes two local
variables (plant height and PCA of plant composition) and three landscape
variables (forest cover, square of forest cover, and residual of the forest-edge
length from forest cover). Because it was not possible to determine the spatial
scale in advance, we searched for the appropriate scale by 1) generating different
sizes of buffer circles around the focal site, 2) calculating percentage of forest
cover within a buffer circle, and 3) selected the best model on the basis of AIC
(Akaike’s information criterion).
Species richness of spiders exhibited the lowest AIC with a 400m buffer
radius, and the best model at this scale included forest cover, the square of forest
cover, and residual of forest-edge length. Species richness showed a peak with
around 60% forest cover, and increased with increasing forest-edge length. Total
abundance of spiders showed a similar pattern. When analyzed separately for
each species, 6 out of 8 species exhibited a peak at intermediate levels of forest
cover, and one species each showed a monotonic increase and decrease with
301
increasing forest cover, respectively. Local factors were mostly unimportant for
determining species richness and abundance of spiders. The reason why
intermediate levels of forest surroundings favoured species richness and
abundance is not clear, but we speculate that prey abundance is higher or
temporarily more stable where two ecosystems (rice fields and forests) meet
with a moderately ratio, thereby enhancing local density of each species and
species richness as a result.
In summary, species richness and abundance of grass-dwelling spiders are
largely determined by landscape-level factors, and mosaic structures of a
landscape seemed to have an important role.
302
Liochelid scorpions of the Indo-Pacific:

systematics and biogeography
Lionel Monod1,2 & Lorenzo Prendini2
1
Muséum d’histoire naturelle, Route de Malagnou 1, Genève, Switzerland,
lionel.monod@ville-ge.ch
2
Scorpion Systematics Research Group, Division of Invertebrate Zoology, American
Museum of Natural History, New York, NY, USA, lorenzo@amnh.org
Phylogenetic relationships in the scorpion family Liochelidae Fet &

Bechly, 2001 are poorly understood. The taxonomic validity of several currently
recognised genera remains uncertain. A phylogenetic analysis of relationships
among the Indo-Pacific liochelid genera, based on 358 morphological characters
and ca. 4 kb of DNA sequence from four loci in the nuclear (18S rDNA and 28S
rDNA) and mitochondrial (12S rDNA and 16S rDNA) genomes, was conducted
to test hypotheses concerning their current pattern of distribution. Based on the
phylogeny and known distributions, a revised biogeography of Indo-Pacific
Liochelidae is presented. Further remarks are given concerning the
diversification of Australian liochelids which appears to be the result of the
fragmentation of a once widespread rainforest habitat due to aridisation in the
Middle Miocene.
Financial support: AMNH Graduate Student Fellowship; National Science
Foundation, USA; Richard Lounsbery Foundation.
303
On three new species of the genus Spariolenus Simon,

1880 (Sparassidae: Heteropodinae) from Iran,
with comments on taxonomy and zoogeography
Majid Moradmand & Peter Jäger
Research Institute Senckenberg, Frankfurt am Main, Germany,

majid.moradmand@senckenberg.de, peter.jaeger@senckenberg.de
Spariolenus Simon, 1880 is one of the rarest genera of the spider family
Sparassidae with just few species described so far. Currently, six nominal
species of the genus are reported from Asia.
Sparassids of Iran have been poorly investigated with just three recorded
species. During surveys in semiarid parts of Iran (caves as well as river banks),
three morphologically different specimens of the genus were encountered.
Results from investigations of somatical and copulatory characters as well as
analyzing CO-I sequences will be presented as well as a discussion about the
species status of the new forms.
The cave-dwelling species, Spariolenus sp. 1, are impressive giant spiders
and have leg spans up to 15 cm. The other two species were caught from
crevices in rocks near river banks. In this study, the subfamily Heteropodinae is
recorded for the first time from Iran.
Representatives of Heteropodinae are common inhabitants of subtropical
and tropical forests of Africa (Barylestis), Asia (Barylestis, Bhutaniella,
Heteropoda, Martensopoda, Pandercetes, Pseudopoda, Sinopoda, Spariolenus)
and Australia (Heteropoda, Pandercetes, Yiinthi). Occurring of the members of
Heteropodinae in the current arid and semiarid areas suggests that the region
used to be humid in former times. After vanishing of the ancient tropical forest
in the territory of the today’s Iran, the relict populations retreated into places like
caves as remaining suitable (=humid) habitats. Taxonomy and zoogeography of
the current species in relation to other species of the genus are discussed.
304
Taxonomic revision of the genus Eusparassus Simon, 1903

(Araneae: Sparassidae), Part 1: finding diagnostic characters
Majid Moradmand & Peter Jäger
Research Institute Senckenberg, Frankfurt am Main, Germany,

majid.moradmand@senckenberg.de, peter.jaeger@senckenberg.de
The genus Eusparassus Simon, 1903 comprises 29 nominal species

described from Southern Africa to Mediterranean Europe and through the
Middle East toward Central Asia. These medium to large Sparassidae are among
the foremost arthropod predators of deserts and semiarid areas. Despite
representatives of the genus are distributed through the vast part of the Old
World, they show a strange type of uniformity in somatical and copulatory
characters. These similarities in traits as well as some interspecific variations
have challenged a discrimination of species.
The aim of our study is to explore the diagnostic characters which can be
used to recognize species boundaries and define and describe the species within
the genus.
305
Phylogeny and biogeography of Ricinulei

Jerome Murienne1, Ligia R. Benavides2, Gustavo Hormiga2
& Gonzalo Giribet1
1
Biology, Harvard University, 26 Oxford Street, Cambridge, MA, USA
2
DC, USA
Very little has been published about the interrelationships of the ca. 60
species of Ricinulei. Here we present a molecular data set based on multiple
markers for the entire distribution range of the group to generate the first
molecular phylogeny of Ricinulei in order to test the monophyly of the three
genera, Ricinoides in west Africa, and Cryptocellus – Pseudocellus in the
Neotropical region. We use this phylogeny to discuss biogeographic aspects of
this group of arachnids with low vagility and compare it to other studies of
arachnids that overlap with Ricinulei in their distribution range.
306
Differentiation of Nurscia albomaculata and

N. albosignata (Araneae: Titanoecidae): two sibling
and sympatric species from Crimea, Ukraine
Anton A. Nadolny & Mykola M. Kovblyuk
Zoology Department of V.I. Vernadsky Taurida National University, Ukraine,

nadolnyanton@mail.ru, kovblyuk@mail.ru
Introduction
The genus Nurscia Simon, 1874 contains only 4 described species
distributed in Palaearctic (Platnick 2010). Two species of Nurscia sympatrically
occur in Crimea: Nurscia albomaculata (Lucas, 1846) and N. albosignata
Simon, 1874. N. albomaculata is distributed from Europe to Central Asia, N.
albosignata – from Bulgaria and Cyprus to Central Asia. Males of these species
well differ by length of palps. However, identification of the females is very
difficult, because their epigynes are similar. Comparative identification drawings
for these species were never published.

29 specimens of N. albomaculata and 69 specimens of N. albosignata
collected mostly by pitfall traps in Crimea in 1995-2007 have been studied. In
order to found differences in females, they were studied from samples
containing both males and females. Then the identified differences were checked
in females from samples without males.
Results
Males of these species well differ by the length of the palp (shorter than
carapace in N. albomaculata and longer than carapace in N. albosignata), by the
teeth on hooked tibial apophysis (long in N. albomaculata and short in N.
albosignata), and by the shape of embolus (straight in N. albomaculata and
curved in N. albosignata).
Females are well distinguished by the shape of epigynal median plate,
narrow in N. albomaculata and wide in N. albosignata. Septum in N.
albomaculata is three times longer then wide; septum in N. albosignata is as
long as wide in posterior part. Margins of epigynal septum are parallel in N.
albomaculata, and curved and not parallel in N. albosignata.
These species do not differ by habitats and are frequently found in the
same biotopes. Both species are found in some kinds of steppe and in salt
marshes. However, these two species differ by phenology. The peaks of
abundance for both species are in July, but males and females of N.
albomaculata were collected since May, and males of N. albosignata – since
June, females – from July. So, activity of N. albomaculata starts one month
earlier than of N. albosignata. The latest males of both species were collected in
307
August. The latest females of N. albomaculata were collected also in August,

and these of N. albosignata – in October.
Discussion
It is interesting when two related species, occurring sympatrically, do not
differ in geography and habitat, but only in phenology.
References
Platnick N.I. 2010. The world spider catalog, version 11.0. American Museum of
Natural History, http://research.amnh.org/entomology/spiders/catalog/.
308
Identification of Wolbachia in the Iranian scorpions

Shahrokh Navidpour1 & Jelodar Zadeh Abbas2
1
Razi Reference Laboratory of Scorpion Research, Venomous Animals Department,
Razi Vaccine and Serum Research Institute, Karaj, Tehran, Iran,
Navid1038@hotmail.com
2
Veterinary Medicine School, Chamran University, Ahvaz, Iran
Bacteria of the genus Wolbachia are intracellular parasites of insects,

filarial nematodes, crustaceans, and mites. The successful spread of Wolbachia
is partly due to their extraordinary ability to alter host development, sex
determination, and reproduction to their own advantage. Similarly to other
bacteria in the order Rickettsiales, Wolbachia are obligate intracellular
endosymbionts of eukaryotes.
The founding member of this genus, Wolbachia pipentis, was first
observed by Hertig and Wolbach in the ovaries of the mosquito Culex pipiens.
Since then, Wolbachia have been detected in most orders of insects and are
believed to be the most pervasive endosymbiont, infecting 15–76% of all insect
species worldwide.
Here we report the Wolbachia in the Iranian scorpions for first time. Four
species of Iran scorpions were studied, Androctonus crassicauda, Mesobuthus
eupeus, Scorpio murus townsendi and Hemiscorpius lepturus, and the bacteria
was found in H. lepturus by Multilocus Sequence Typing and by Wolbachia
Surface Protein.
309
Faunistic study on scorpions of the Province Lorestan, Iran

Hassan Nayebzadeh1*, Shahrokh Navidpour2
& Mohammad Hassan Kayedi3
1
Department of Parasitology, School of Veterinary Medicine, Lorestan University,
Khorramabad, Iran, hassannayeb@yahoo.com
2
Department of Venomous Animals & Toxin, Razi Reference Laboratory of Scorpion
Research, Razi Vaccine and Serum Research Institute, Hesarak, Karaj, Iran
3
Department of Parasitology, School of Medicine, Lorestan University of Medical
Sciences, Khorramabad, Iran
*
Corresponding author
Lorestan is a mountainous province located to the south-west of Iran

between eastern 46º51' to 50°3', and northern 32°37' to 34°32'; surrounded by
the mountains of the Zagros range. In localities of this province there are two
scorpion taxa, Mesobuthus eupeus phillipsii (Pocock, 1889) and Hottentotta
saulcyi (Simon, 1880). Additionally, Androctonus crassicauda (Olivier, 1807),
Compsobuthus matthiesseni (Birula, 1905), Hemiscorpius lepturus (Peters,
1862); Scorpio maurus townsendi (Pocock, 1900) and Orthochirus iranus
(Kovařík, 2004) have been seen in the province. Researchers from Razi
Reference Laboratory of Scorpion Research (RRLS); Lorestan University of
Medical Sciences, and Lorestan University have identified three other species in
the province for the first time: Hottentotta zagrosensis (Kovařík, 1997);
Razianus zarudnyi (Birula, 1903); Hemiscorpius lepturus (Peters, 1862). In
addition, a new species of Hottentotta was described.
310
Grazed versus ungrazed – case study of one xerothermic

slope in Moravian Karst (Czech Republic) 7
Jana Niedobová1, Vladimír Hula1 & Fric Z. Faltýnek2
1
Department of Zoology, Mendel University, Brno, Czech Republic,
Naaudia@seznam.cz
2
Department of Ecology and Conservation Biology, Institute of Entomology, Biology
Centre, ASCR, České Budějovice, Czech Republic, fric@entu.cas.cz
Grazing undoubtedly influences grassland ecosystems and belongs to the

widely discussed questions of present natural and landscape protection. It is
necessary to maintain some kinds of grasslands by grazing especially in the
landscape protected areas because most of critically endangered species are
depended on grazing ecosystems. Grazing helps to keep biotopes on the some
succession stage and it leads to enrichment ecosystem by endangered and
critically endangered species (mainly in case of plants). But grazing can destroy
some kinds of organisms too. It happens especially by trampling and pasturing.
We tried to find influence of grazing on invertebrates in Moravian Carst.
There were two transects in the Macošská stráň slope. First of them was
ungrazed and second transect was grazed. Our research took place from April to
November 2008 and samples were taken in a month or two-week intervals
respectively. To be able to find the most species of invertebrates we used three
samplings methods: pitfall traps, yellow Möerick cups and sweeping. We used
two groups (Araneae and Carabidae) of invertebrates for evaluating grazing
influence. And than we evaluated data with the programme CANOCO.
DCA analyses didn’t showed significant affinity to grazed or ungrazed
transect. We found only one difference between transects – t-test showed that
localities under grazing are inhabit by ubiquitous species of spiders (t=-2.89,
df=6, p=0.027). It means that this habitat has little bit richer fauna of spiders, but
only the common ones.
7
The study was supported by grants IGA-AF-MZLU-SP2100101/224 and VaV-MZP-
CR-SP/2D4/59/07.
311
Spiders from the Pico da Neblina: species diversity

and distribution along an altitudinal gradient
in Brazilian Amazonia (Arachnida: Araneae)
André do Amaral Nogueira1,3, Eduardo Martins Venticinque1,2

& Antonio Domingos Brescovit3
1
Pós-Graduação em Ecologia, INPA - Instituto Nacional de Pesquisas da Amazônia,
Brazil, andrearanhas@gmail.com
2
Universidade Federal do Amazonas (UFAM), Departamento de Biologia, Brazil,
eventicinque@wcs.org,
3
Laboratório de Artrópodes, Instituto Butantan, São Paulo, SP, Brazil,
adbresc@terra.com.br
Montane ecosystems have always drawn the attention of biologists. The

conspicuous changes observed in biotic communities along a mountain have
always interested ecologists and biogeographers, while their usually high
diversity and endemism level also attracts zoologists and botanists. One of the
most studied features about mountain biota is the species richness distribution
patterns along the altitudinal gradient. Early surveys, as well as the resemblance
in some aspects with the latitudinal gradient of species richness, led to the belief
that richness always decrease with increasing altitude. However, several recent
works, usually based on standardized sampling, have revealed other patterns,
especially the occurrence of richness peaks at intermediate altitudes. The reasons
responsible for this pattern have been intensively debated in recent years, and
explanations are usually based either on ecological/environmental factors, or in a
hypothesis known as geometric constraints. This hypothesis, also known as the
mid-domain effect (MDE), sustains that the disposition of species ranges in a
bounded domain tend to lead to a greater richness near the centre of this domain,
due to the overlap of ranges, suggesting thus that intermediate richness peak
occurs regardless of environmental or ecological gradients.
In this work, we investigate the spider community of the Pico da Neblina,
a mountain located in Brazilian Amazonia. Our goal was to assess the patterns of
richness distribution along the gradient, and to test the MDE hypotheses, using it
as a null model to generate an expected richness pattern and compare it with the
observed pattern. We also test another hypothesis concerning gradients, the
Rapoport Effect, which claims that there is positive relation between the size of
the range of a specie and the altitude (or latitude) where it lives, i.e., species
from higher altitudes would also be expected to have larger ranges. Finally, we
also investigated other parameters of the community, as structure and
composition, discussing the changes observed along the gradient.
Our study site was the Pico da Neblina (0°15'N, 66º10'W), the highest
Brazilian mountain, with 2.994 m. a.s.l. The Pico da Neblina is located on the
northern border of the Brazilian state of Amazonas, a mountain region that
represents the boundary between Brazil and Venezuela, as well as the watershed
312
between the Orinoco and Amazonas basins. The climate of the region can be
classified as tropical humid, with an annual average rainfall of 2.500-3.000
mm/year, an average temperature of 25° and 85-90% of humidity, with little
variation through the year. With increasing elevation, there is an increase in
rainfall and humidity and a decrease in temperature, and, in the highlands (>
1.800 m) of the region, the rainfall decreases and is replaced by a constant mist,
with humidity reaching almost 100%. In those altitudes, temperature can drop to
an average of 10° in the coolest month. Vegetation of the lowlands is mainly
composed by a tall, evergreen forest, being gradually replaced by submontane
(400 to 800 m), montane (800 to 1.500 m) and upper-montane forest (1.500 to
2.000 m) (Huber 1995). Higher altitudes present more open types of vegetation,
such as high-altitude meadows and grasslands.
Spiders were collected in September/October 2007, with a beating tray,
during the day, and through manual active searching, during the night. Sample
unit of the first method correspond to the investigation, with a beating tray, of 20
small trees, or shrubs, or other components of the vegetation. Sample unit of the
second method correspond to one hour of search along a 30 m long transect. All
spiders obtained with those two methods were conserved in alcohol at 70%.
Adult spiders were sorted in morphospecies and identified to the most accurate
taxonomic level possible. We sampled at six different altitudes, 100, 400, 860,
1.550, 2.000 and 2.400 m.a.s.l., and in each altitude we investigated three
different sites. In each site the sampling effort corresponded to 9 diurnal and 9
nocturnal samples, which lead us to an amount of 54 samples by altitude (27
diurnal and 27 nocturnal), and to the final count of 324 samples (172 of each
method). We also measured the temperature at each sampled site.
We used a Monte Carlo simulation to generate the richness pattern
expected by MDE predictions, and tested the Rapoport effect with a simple
linear regression, relating for each species the size of their range with their
weighted altitudinal midpoint (calculated as the number of individuals in each
altitude, multiplied by the altitude value, and divided by the total abundance of
the species, a.k.a. specimen method). Finally, we performed an NMDS to verify
the similarity of spider communities from all the sampled sites.
We obtained 2.950 adult spiders (1.896 females and 1.054 males), divided
into 469 species. Since the match of males and females was very hazardous in
most cases, results presented below refer only to females, which accounted for
almost two thirds of adult individuals and were sorted to 349 species. The mean
richness (i.e. the mean richness of the three sites sampled by altitude) and
standard deviation by altitude are presented below, with the total richness by
altitude in brackets: 100 m, 62,6±7 (141); 400 m, 51,3±6 (121); 860 m, 54,3±5
(119); 1.550 m, 42±6 (79); 2.000 m, 14,3±4 (37); 2.400, 9,6±4 (18). Thus,
richness decreased along the gradient, and presented a very poor fit with the
richness predicted by the MDE (R2=0,2497; p>0,1). The Rapoport Effect
couldn’t be observed either, since the relation between the size of the range and
the weighted average midpoint was also very weak and non-significant (R2=2.7,
p >0.01). The richness distribution pattern exhibited by the community is clearly
non-random, and seems to be very influenced by environmental factors, notably
313
temperature. Our gradient presents a plateau of high diversity in the three first
altitudes, with a small decrease in the fourth altitude and a steeper decrease in
the fifth altitude. Several studies indicates that the structural complexity of the
environment have a positive relationship with the diversity of spiders, which
would explain the large number of species found at the first altitudes. So, the
lower richness of the last two altitudes would be a consequence of a less
complex environment, with open types of vegetation, and also of a harsher
climate. We believe that this environmental disruption in our gradient was also
one of the causes of the absence of a Rapoport Effect in our community, since
the species from forested sites would have difficulties to expand their ranges
upward, at non-forested and cooler sites, while the species adapted to those high-
altitude environments couldn’t expand their range downward. So, species
distributed at higher altitudes would have shorter ranges, while species from
forested sites could have larger ranges, but at lower altitudes, the opposite of
what is predicted by the Rapoport Effect.
The changes observed in the composition, however, revealed an
unexpected pattern, since the fourth altitude (1.550 m) was associated with the
higher sites, despite the structural difference between them. It indicates that the
temperature may have an important role as limiting factor for many forest
species, and the decrease in richness observed at 1.550 when compared with the
other forested sites seems to confirm that. Anyway, the similarity between the
faunas from the three last altitudes, revealed by the NMDS (Stress=0.17), seems
to be very influenced by a few dominant species.
Finally, the structure of the community, defined here as the dominance
and proportion of rare species (singletons and doubletons), goes through very
important changes along the gradient. Community from the first altitude
presented a very low dominance and a very large proportion of rare species (the
dominant specie represented 4% of the total abundance, while the rare species
represented 24%), and, as altitude increase, there is an increase in the dominance
and a decrease in the importance of rare species. As a consequence, the last
altitude presented a very different community, where the dominant specie
accounted for 55% of the total abundance, while the rare species only 9%. So, at
least some of the changes observed in the spider community of the Pico da
Neblina along the gradient, notably the richness pattern and the structure of the
community, seems to be in agreement with the changes observed in spiders
communities along the latitudinal gradient.
314
Does seasonal isolation influence on relations between

saprophagous and predatory invertebrates,
Collembola and Araneae?
Izabella Olejniczak1, Paweł Boniecki1, Marzena Stańska2

& Izabela Hajdamowicz2
1
Centre for Ecological Research PAS, Dziekanów Leśny, 05-092 Łomianki, Poland, iza-
olejniczak@wp.pl
2
University of Podlasie, Department of Zoology, Siedlce, Poland
Relations between saprophagous and predatory invertebrates are one of

the most important questions in soil biology, especially in seasonal flooded
environments. Both groups of invertebrates may indirectly affect mineralization
and humification of organic matter. Moreover they are known as indicators of
soil properties.
Activity of epigeic saprophagous Collembola and predatory Araneae were
studied in alder forest-Alnus glutinosa-Carex elata association. There were
chosen 25 alder tussocks that average surface was 0,50m2 and the high was
0,40m. Distances between particular tussocks ranged from 1m to 11m. Carex
elongata was the dominant plant in all alder tussocks.
Samples were taken twice in the season: during the spring, when tussocks
were washed by water and during the autumn when tussocks were not washed by
water. Three plastic tubes of 1 cm diameter and 6 cm high were placed in each
tussock. Tubes were filled with 96% ethanol and emptied every seven days in
June and September.
Activities of Collembola and Araneae were higher in the spring than in
the autumn (P<0,05), but there were no significant differences between
particular tussocks as well in the spring as in the autumn. Size of the alder
tussocks did not influence on invertebrates’ activity. The contribution of
collembolans in invertebrates communities was higher in the autumn – 54% than
in the spring – 18%. The contribution of spiders in invertebrates communities
was nearly the same in the spring and autumn and ranged from 20% to 11%.
The impact of predatory invertebrates on collembolans activity was differ
in relation to degree of isolation of alder tussocks. Spiders’ activity influenced
on collembolan activity only in the autumn (P=0,05).
315
Local and landscape scale factors influencing

spider density and diversity in crop fields
Maria Oleszczuk
Institute for Agricultural and Forest Environment, Polish Academy of Sciences, Poznań,
Poland, oleszczukm@vp.pl
The studies of local (distance from forest belt and crop type) and
landscape (heterogeneous and homogenous landscape) factor’s influence on
spider density and diversity in the crop fields of winter cereals, sugar beet and
alfalfa are presented. The samples were taken with biocenometer from the area
of 0.25 m2 (10 samples in one site) twice a year for three consecutive years.
The spider density were between 0.4 to 5.2 ind./1 m2. Statistically
important difference in spider density and number of species were noted
between winter cereal crop in homogenous landscape and sugar beet crop in
homogenous landscape and sugar beet crop in heterogeneous landscape.
The proximity of forest belts positively influenced spider species diversity
in the studied crop fields. In the distance of 10 m from forest belts 14 spider
species in cereal crops and 8 species in sugar beet crops were noted. In the
distance of 50 m nine and five spider species respectively were recorded. The
exception was the alfalfa crop, were the number of spider species has not been
changed with the distance for the forest belts. However, the above differences
were not statistically important.
The influence of landscape type and the distance from forest belts on the
share of ecological groups of spiders was analyzed. In the landscape with forest
belts larger part of foliage-dwelling spiders than aeronautic spider species was
noted, and in the homogenous landscape the tendency was opposite. The
distance from forest belts has not changed essentially the share of ecological
groups of spiders.
316
Mortal combat and nuptial display of the loser

in Cheiracanthium japonicum Bösenberg & Strand, 1906
(Araneae: Cheiracanthiidae) 8
Mikhail M. Omelko
Mountain-taiga station of Russian Academy of Sciences, Russia, omelkom@gmail.com
Nuptial prey gifts are known in various groups of spiders and insects. For
example in Pisaura mirabilis (Clerck, 1757) the prey gift works as a sensory
trap where the male exploits the female's foraging motivation in a sexual context
(Stálhandske 2001, Bilde et al. 2007).
At present time only two species of Cheiracanthidae (Cheiracanthium
japonicum Bösenberg & Strand, 1906 and C. virescens (Sundevall, 1833)) are
known from continental Southern Far East of Russia (Mikhailov 1997). Our
observations were carried out in Maritime Province near the Gornotaezhnoe
biological station of the Russian Academy of Science (43º41'42.77"N,
132º09'25.48"E) in June of 2004.
During mating season the males of C. japonicum wander on upper leaves
of herbaceous plants, low trees and bushes and try to found females. The females
make their shelters on grasses. They roll part of the leaf into a short tube and
cover openings by the web. When a male find out the female’s shelter it settles
there and constructs its own refuge. Male’s shelter is a light semi-transparent
marquee located above female’s shelter.
The observations of the mortal combat and nuptial display are based on
single case by this time. First spider detected and met newcomer one rather far
from the female’s shelter. When the spiders approached to each other they had
menacing poses and started to spin around the stem with spread first pair of legs
and chelicerae. A demonstration of menacing poses was going on a little less 1
minute. During this time each spider both spiders were trying to near to its rival
from one side. When the males approached so mach close to each other that
were almost touching by its chelicerae the first one very fast rushed in attack and
pierced the second spider’s cephalothorax. Hereon the fighting was finished
instantly and the killed one became rigid at once. The victorious spider was
standing at that place and holding the dead motionless rival. Then it lifted the
corpse up and brought to the shelter where the female was hiding. There male
stayed approximately 4 minutes as though making a demonstration of the dead
body to female. Afterwards it hung the corpse at the side of the shelter on a short
thread for approximately 5 minutes. Then it threw it down and had gone to its
own shelter. The all event since meeting with the newcomer spider up to the
moment when the corpse was thrown took 15 minutes.
8
The work supported by Russian Foundation for Basic Research (grant #10-04-01424-а).
317
At present time it isn’t clear for me which function the display of the
corpse is performing. The female doesn’t neither eat the loser’s body nor change
its behaviour in any way so this nuptial gift cannot be something like a sensory
trap in other groups of spiders. Conducting an investigation in nature and
laboratory to solve this question of behaviour of C. japonicum is planned.
References
Bilde T., Tuni C., Elsayed R., Pekar S. & Toft S. 2007. Nuptial gifts of male
spiders: sensory exploitation of the female's maternal care instinct or
foraging motivation? Animal Behaviour, 73: 267-273.
Mikhailov K.G. 1997. Catalogue of the spiders of the territories of the former
Soviet Union (Arachnida, Aranei). Trudy Zoologicheskogo Muzeya
MGU, 37: 1-416.
Stálhandske P. 2002. Nuptial gifts of male spiders function as sensory traps.
Proceedings of the Royal Society, Biological Sciences, 269: 905-908.
318
The spider fauna of Japan, with a brief view of the

geological history traced back to the Mesozoic
Hirotsugu Ono
National Museum of Nature and Science, Tokyo, Japan, ono@kahaku.go.jp
After the opening of the Samurai country to the West urged by the Meiji
Restoration around 1868, European researchers were active in spider collecting
in Japan, such as Albrecht von Roretz (1846-1884) from Austria (the length of
his stay in Japan: eight years between 1874 and 1882), Franz Hilgendorf (1839-
1904) from Germany (three years in Japan, 1873-1876), A. (?Anatole) Mellottée
from France (1881-1882 in Japan) and Friedrich Karl Wilhelm Dönitz (1838-
1912) from Estonia (1873-1886 in Japan). Having described thirty-one species
based on Roretz Collection in Wien, Ludwig Koch (1878) took the lead in
reporting Japanese spiders. Since then, about 1500 species of 64 families were
recorded from this country [Karsch (1879, 1881), Simon (1886, 1889),
Bösenberg and Strand (1906), Kishida (1909, 1914), Saito (1934, 1939),
Nakatsudi (1937, 1942), Oi (1960), Yaginuma (1960), Komatsu (1961), Okuma
(1988) and further works of many arachnologists]. The spider fauna of Japan is
biogeographically surveyed here on the basis of results of these taxonomical
studies continuing for 130 years, which are integrated into a volume of ‘The
Spiders of Japan’ recently published (Ono 2009). Some general and extreme
examples of distribution are shown. A difference of species diversity between
the Ryukyu Islands (Southwest) and the Izu Ogasawara Islands (Southeast) is
made clear based on some aspects referring to the geological history of the
Japanese islands.
319
Origins and phylogeography of the European protected

funnel-web Macrothele spiders (Araneae: Hexathelidae)
Věra Opatová1, Pedro Cardoso2,3, Miguel-Angel Fernández4

& Miquel A. Arnedo1
1
Department of Animal Biology, University of Barcelona, Spain,
Vera.Opatova@seznam.cz, marnedo@ub.edu
2
Smithsonian Institution, National Museum of Natural History, Washington DC, USA,
pcardoso@ennor.org
3
Azorean Biodiversity Group (CITA-A), Departamento de Ciências Agrárias,
Universidade dos Açores, Portugal
4
Sociedad para el Estudio y la Conservación de las Arañas, Spain, harpactea@yahoo.es
The funnel web spider genus Macrothele includes 26 species with a

disjunct distribution. Most of them are found across South-eastern Asia, few
have been reported from Central Africa, and only two are known from Europe:
Macrothele calpeiana (Walckenaer, 1805) from the southern Iberian Peninsula
and Macrothele cretica Kulczyński, 1903 endemic to Crete. Little is known on
the origins of the European Macrothele spiders. The genus belongs to the family
Hexathelidae, which is particularly diverse in Australia and New Zealand. Some
authors have suggested that European Macrothele originated in Asia and
subsequently colonized Europe following a west-wise pattern, while others
proposed that, at least M. calpeiana, would have colonized Iberia from northern
Africa. It has also been suggested that M. calpeaina is a recent introduction from
China.
The two European Macrothele species have received conservation
attention. M. cretica is included in the IUCN red list under the data deficient
category. Macrothele calpeiana is the only spider species protected by European
Union legislation. Its inclusion in the Bern Convention listings and the Habitats
Directive was motivated by its putative role as quality bio indicator of the
diminishing cork forest areas. Nevertheless, a better understanding of the species
distribution revealed that the species was not restricted to cork forest and that
climatic variables, such as hot, dry summers and mild, humid winters are better
predictors of its presence. In addition, some population of M. calpeiana far from
being threatened seem to be expanding their range. As an example, previously
unknown localities have been recently reported from southeast Spain, southern
Portugal or even northern Italy. These observations led some authors to propose
that the species should be removed from the Bern convention and Habitats
Directive. Nevertheless, molecular analyses have revealed that M. calpeiana
includes distinct genetic lineages, some with restricted distribution only
explained by the long-term presence of the species in the Iberian Peninsula, and
ancient separation of populations, which probably dates back to the Pliocene.
To investigate the origins of European Macrothele species and the genetic
affinities of the newly found populations of M. calpeiana, we have expanded
320
former phylogenetic analyses to include for the first time samples of M. cretica
and Macrothele from China and Central Africa, as well as new samples of M.
calpeiana from Italy and Portugal. Phylogenetic relationships of Macrothele
were inferred from a combined data set including the nuclear ribosomal genes
18S and 28S and the protein gene EF1gamma. Population analyses were based
on the mitochondrial genes cytochrome oxidase I and a fragment expanding the
ribosomal 16S, the tRNA leucine and the NADH dehydrogenase subunit I.
Results of these analyses bring new light on the origins and evolutionary
relationships of European Macrothele and provide essential information for
conservation planning and management of these unique elements of the
European fauna.
321
Molecular tracking of the role of agrobiont and immigrant

spiders in Negev (Israel) wheat fields: trophic delineation
of biological control potential
Itai Opatovsky1, Eric G. Chapman2, Phyllis G. Weintraub3,

Yael Lubin1 & James D. Harwood2
1
Mitrani Department of Desert Ecology. Blaustein Institutes for Desert Research, Ben-
Gurion University of the Negev, Sede Boqer Campus, Israel, lubin@bgu.ac.il
2
Department of Entomology, University of Kentucky, Agricultural Science Center
North, Lexington, KY, USA
3
Agricultural Research Organization, Department of Entomology, Gilat Research
Center, Israel
Natural enemy populations are important in suppressing outbreaks of pest

populations. However, disturbances in agricultural fields due to crop
management do not allow maintenance of sustainable predator populations.
Therefore, alternative habitats, which provide reproduction sites, shelter and
alternative prey, are important for natural enemies that immigrate into the
agricultural fields during the crop season. In the desert agroecosystems of Israel
there are large ecological differences between natural habitats and crop fields.
As a result, the higher-productivity agricultural fields attract predators from the
surrounding natural desert environment during the crop season. However, it is
not known whether these immigrant predators consume pests or alternative prey
after colonizing the fields. Using DNA based gut-content analysis, the
consumption of pests (aphids and hessian flies) or alternative prey (Collembola)
by spiders inhabiting wheat fields was examined. The two web building spiders
that are common during the wheat season, share the same niche, and may be
competing for the same resources: an agrobiont species, Alioranus pastoralis
(Linyphiidae), and an immigrant species, Enoplognatha sp. (Theridiidae), were
examined. The agrobiont species showed feeding preference towards the more
nutritious prey, the Collembola, while the immigrant spiders consumed the less
nutritious prey, the aphids. These results clearly demonstrate the potential role of
immigrant spiders in biological control of pest species in winter wheat fields in
Israel.
322
Morphological organization of male palpal organ of the

Australian ground spiders (Araneae: Gnaphosidae)
Vladimir I. Ovtcharenko & Boris Zakharov
Department of Natural Sciences, Hostos Community College of the City University of New
York, New York, NY, USA, vio@hostos.cuny.edu, zakharov@amnh.org
A detailed morphology of the male palpal organs of Australian ground

spiders of genera Encoptarthria Main, 1954 and Taieria Foster, 1979 is
presented. The genera Encoptarthria and Taieria are the sister groups. The
genus Encoptarthria is endemic although widely distributed all over the
Australia. The genus Taieria was known previously only from New Zealand,
and was also widely distributed all over Australia. Comparative analysis shows
that Encoptarthria males have more complicated palps and differ significantly
from other Australian gnaphosid spiders by having a very complex organization
of the terminal part of the bulbus. Besides a distal sclerotized tube, the terminal
part of bulbus poses a prominent distal apophysis and a closely spaced embolus
terminal apophysis and fulcrum. The sister genus Taieria has simpler palps with
a fulcrum only. Moreover, Taieria has an exceptional structure that does not
occur in other Australian gnaphosid spiders – the terminal membrane. The
genera Intruda and Anzacia have the simplest male palps of the Australian
gnaphosids. The palpal organization is a tripartite structure that consists of a
subtegulum, a tegulum and an embolus, which includes the conductor and
median apophysis. Other additional structures are missing from the palps of
these spiders. In addition, Anzacia male palps have unique shape of the
conductor, which is divided on three parts: the distal, median and proximal
lobes. Analysis of the male palpal organs is presented and its implication for
classification of these groups is discussed.
323
Arachnid cave-dwelling fauna on Biokovo Mt.,

Central Dalmatia, Croatia
Roman Ozimec
Croatian Biospeleological Society (CBSS), Demetrova 1, 10000 Zagreb, Croatia,

roman.ozimec@hbsd.hr
Biokovo Mt. (1762 m), with relatively small surface of 200 km2, is
situated in Central Dalmatia (Croatia) and belongs to the Dinaride Mountains.
Very similar to Velebit Mt. and Orjen Mt., Biokovo was under continental
glacial and Mediterranean influence. Due to extremely karstification, unique
geomorphologic features, biodiversity and endemicity, Biokovo was declared as
Nature Park in 1981.
Biospeleological research in region began in the first decades of 20th
century but most intensive systematic research was performed between 2002-
2006 with cooperation of Biokovo Nature Park and Croatian Biospeleological
Society. During that period 115 speleological objects have been researched
through 192 visits. Five biogeographical zones are recognized on Biokovo Mt.,
with many different cave habitats. A total of 186 different taxa have been
recorded that show some cave-dwelling affinities. Endemism of cave-dwelling
fauna is extremely high, even 65 taxa are endemic for Biokovo Mt. and further
47 taxa are endemic for Dinarides. Until now, 44 taxa new for science have been
recognized.
Among them, 57 cave-dwelling taxa belong to Arachnids: Acari (7),
Palpigradi (1), Pseudoscorpiones (23), Opiliones (4) and Araneae (22). All cave-
dwelling arachnid taxa are endemic for Dinarides, 23 taxa are endemic for
Biokovo Mt. with at least 20 new for science. Most representative genus are:
Rhagidia, Opilioacaris, Eukoenenia, Chthonius (Chthonius), Chthonius
(Globochthonius), Chthonius (n. subg.), Troglochthonius, Protoneobisium,
Neobisium (Neobisium), Neobisium (Blothrus), Neobisium (Ommatoblothrus),
Roncus, Cyphophthalmus, Folkia, Stalagtia, Mesostalita, Barusia, Sulcia,
Stygopholcus, Centromerus, Typhlonyphia, Histopona.
It seems that Biokovo Mt. is a hot spot of arachnid cave-dwelling fauna,
but also development centre for some phylletic lines of families Chthoniidae and
Neobisiidae (Pseudoscorpiones), same as Dysderidae and Leptonetidae
(Araneae). Further systematic research on Biokovo Mt. will continue on cave-
dwelling, but also on soil and surface arachnid fauna.
324
Spiders and wetlands conservation: a new challenge

at regional scale in Piedmont (NW Italy)
Mauro Paschetta & Marco Isaia
Dipartimento di Biologia Animale e dell’Uomo, Università di Torino, Italy,

mauro.paschetta@gmail.com, marco.isaia@unito.it
The Ramsar Convention, signed in 1971, recognized for the first time, at
international level, the ecological values of wetland habitats and the importance
of their conservation. These habitats are characterized by the presence of high
bio-diverse ecosystems that harbour species with relevant conservation value,
especially among birds, amphibians and plants. Studies aiming to monitor the
present state of conservation of these habitats are essential to evaluate their on-
going degradation, strictly related to agricultural intensification, land
reclamation, urbanization, pollution and global warming.
Despite the huge diversity of spiders and their great potential for bio-
diagnostic purposes in wetland biodiversity assessment (Cristofoli et al. 2010),
studies on spider communities are generally lacking. To give two examples of
the extraordinary spider biodiversity of wetland ecosystems, Villepoux (1993)
recorded 177 species in wetlands of the Rhine valley and Patocchi (1993) 171
species in riparian habitats in Switzerland. When considering the Italian fauna,
the lack of studies on this topic is particularly striking. At regional level for
example, the only contributions given on this topic in Piedmont are the work of
Isaia et al. (2005) and two more works on riparian habitats by Arnò (2001) and
Beikes et al. (2004). Furthermore, wetlands may harbour peculiar species whose
conservation is closely related to the survival of such ecosystems. An example is
provided by the fen raft spider (Dolomedes plantarius), whose presence in Italy
is only recorded from a few localities (Pavesi 1873, 1879: Lombardia;
Pesarini1994: Sardinia; Hansen 2002, 2007: Veneto), without any information
on the state of conservation of the habitat. The importance of this species is
underlined by the fact that Dolomedes plantarius is among the few species of
spiders cited in the IUCN European Red List (“vulnerable”), for which
conservation measures are required, especially considering the decline in area of
occupancy, the extent of occurrence and the quality of habitat and the effects of
introduced taxa, hybridisation, pathogens, pollutants, competitors or parasites.
Our recent finding of this species in the Natural Reserve of Fondotoce
(North-Eastern Piedmont) in the framework of a preliminary research founded
by the Reserve itself, induced us to develop an extensive three-years project in
collaboration with the Regional Museum of Natural Science of Turin. The
research started with a preliminary ecological analysis of the Natural Reserve of
Fondotoce, focusing especially on spiders, with special reference to the
environmental evaluation of the habitat, on the presence of stenoecious species,
on the relation species-habitat, on the structure of spider communities and on
specific turnover across different habitats. Spiders were collected by means of
325
standardized pitfall transects placed in five different habitats: a bed of reeds, a

riparian wood, a mown wet meadow, an anthropogenic heath colonized by an
invasive allochtonous plant (Solidago sp.) and a transition zone at the border of
the bed of reeds. Random hand-collecting was also performed in order to
maximise species richness and to provide new faunistic data. We recorded 37
species by pitfall traps and 40 by hand-collecting. From the ecological point of
view the bed of reeds stood out for the presence of several interesting
stenoecious species, like the previously cited Dolomedes plantarius and
Donacochara speciosa, Pachygnatha terilis, Mendoza canestrinii and Pirata
hygrophilus that also represent new records for the regional fauna. The habitats
with higher species richness (anthropogenic heath and mown meadow) were
dominated by euriecious species like Trochosa ruricola, Pardosa prativaga and
P. proxima. Indicator species analysis on pitfall data revealed statistically the
close relation between hygrophilous species like Bathyphantes gracilis, Pirata
piraticus, Gnathonarium dentatum, Antistea elegans and the bed of reeds
ecosystem. On the other hand, the mown meadow was characterized by semi-
hygrophilous species like Arctosa leopardus and Pachygnatha terilis and also by
generalist species like Pardosa proxima. The riparian wood was characterized
by several generalist species, like Trochosa hispanica, Pardosa alacris and
Ozyptila praticola, attesting a certain degree of disturbance (the undergrowth
was, in fact, dominated by the allochtonous Phytolacca and Impatiens shrubs).
We performed the analysis of specific turnover by means of MRPP (Multi
Response Permutation Procedure), a class of multivariate permutation tests of
group-differences that offers vast improvement over the traditional analysis of
variance-based tests in terms of robustness under non-normal data and
heterogeneous error variances. The result of this test is a numerical value (T with
associated p value) that quantifies in statistical terms the distance between two
or more matrixes. By combining average distances between each habitat, MRPP
demonstrated the uniqueness of spider assemblages found in the bed of reeds
and in the riparian wood (high specific turnover). According to the classical
trend of ecotonal habitats, the lowest mean T was found in the transition area
that was characterized by low specific turnover. The structure of the spider
assemblage found in each habitat was tested in terms of rank-abundance
modelling, highlighting that the distribution of the spider assemblage in the
anthropogenic heath followed a logarithmic distribution, confirming the high
degree of disturbance.
Our work underlines the vulnerability of the bed of reeds, especially
considering its peculiar spider assemblage (presence of Dolomedes plantarius),
the general low ecological quality of nearby habitats (dominance of
euriecious/invasive species) and the high environmental pressures around it
(rapid process of urbanization taking place around the Reserve).
The research carried out at the Reserve of Fondotoce is part of a three-
years PhD project that is going to involve some other wetlands in Piedmont.
Analogous ecological analysis are going to be performed with the main purpose
to assess the environmental quality of wetlands using spiders as bioindicators
326
and to outline the current state of conservation and distribution of Dolomedes

plantarius in Piedmont.
References
Arnò C. 2001. Ragni dell’area protetta “Fascia fluviale del Po”: nota preliminare
su tre specie nuove per l’Italia e una nuova per il Piemonte (Arachnida,
Araneae). Rivista piemontese di storia naturale, 22: 155-164.
Beikes S., Isaia M., Bona F. & Badino G. 2004. Analisi ecologica
dell'araneofauna di fasce riparie di tratti fluviali a differente livello di
funzionalità. XIV Convegno Nazionale della Società Ecologica Italiana,
Siena, 4-6 ottobre 2003, p. 45.
Cristofoli S., Mahya G., Kekenbosch R. & Lambeets K. 2010. Spider
communities as evaluation tools for wet heathland restoration. Ecological
Indicators, 10: 773-780.
Hansen H. 2002. Segnalazioni 21 – Scytodes velutina, …, Segnalazioni 40 –
Philaeus chrysops. Bollettino del Museo civico di Storia Naturale di
Venezia, 53: 268-272.
Hansen H. 2007. Stato attuale della conoscenza della fauna dei ragni presente
nel territorio della laguna di Venezia e nelle aree limitrofe (Arachnida:
Araneae). Bollettino del Museo civico di Storia Naturale di Venezia, 58:
11-82.
Isaia M., Bonelli S., Montani M., Badino G. & Balletto E. 2005. Spiders
(Arachnida, Araneae) of a Maculinea alcon - M. teleius pSCI in NW Italy.
In: Settele J., Kuhn E. & Thomas J.A. (eds.), Studies on the Ecology and
conservation of Butterflies in Europe, vol. 2: Species ecology along a
European Gradient: Maculinea Butterflies as a model, 9-15. Pensoft
Publishers, Sofia-Moscow.
Patocchi N. 1993. I ragni della Valle Maggia: studio faunistico ecologico delle zone
alluvionali. Memorie Società Ticinese di Scienze Naturali, 3: 209-267.
Pavesi P. 1873. Enumerazione dei ragni dei dintorni di Pavia. Atti della Società
Italiana di scienze naturali, 16: 68-78.
Pavesi P. 1879. Saggio di una fauna aracnologica del Varesotto. Atti della
Società Italiana di scienze naturali, 21: 789-817.
Pesarini C. 1994. Arachnida Araneae. In: Minelli A., Ruffo S. & La Posta S.
(eds.), Check-list delle specie della fauna italiana 23. Calderini, Bologna.
Villepoux O. 1993. Etude de la rèpartition des araignèes d’une zone humide.
Proc. 14th Eur. Colloq. of Arachnology. Catania, Bollettino
dell’Accademia Gioenia di Scienze Naturali, 26: 361-370.
327
The New Caledonian Salticids: uniqueness and diversity

– zoogeographical account
Barbara M. Patoleta
Department of Zoology, University of Podlasie, Siedlce, Poland, patoleta@ap.siedlce.pl
The salticid spider fauna of the Pacific Islands is mixture of Asian,

Australian/New Guinean and wide spread taxa, with contribution of local
endemics (Prószyński 1996). The analysis for particular islands shows
distinctive differences is genus and species composition, depending on island
origin, age, size, biota and distance from other faunistic sources.
My study shows that New Caledonia has unique salticid fauna, with 25%
endemic genera and over 70% endemic species.
The uniqueness of New Caledonian fauna is related to its 65-70 MY long
post-Gondwanan isolation, accompanying climatic changes and evolution of
biota.
The work is part of a long-term project aimed to study salticids of SW
Pacific region in order to describe entire systematic composition and to analyze
the origin, evolution and relationships of faunas.
Salticidae (Arachnida, Araneae) of Fiji. List of species

with notes on their zoogeography
Barbara M. Patoleta
Department of Zoology, University of Podlasie, Siedlce, Poland, patoleta@ap.siedlce.pl
Before this project started, 36 salticid species were recorded from Fiji
(Karsch 1878, 1879, Roewer 1944, Prószyński 1984, Żabka 1988, Berry et al.
1996, 1997, 1998). As the result of my study (Patoleta 2002 unpubished PhD
thesis, 2008a, 2008b), 49 species were recorded from the archipelago.
The Fiji archipelago is of volcanic origin and its fauna is largely the result
of influence of Indopacific (54%) and widespread (36%) genera. The most
interesting is high endemism of species (53%), being the result of geological
history and isolation. The archipelago seems to be the centre of speciation for
same genera e.g. Xenocytaea, Cytaea, Sobasina and Palpelius.
328
Evolution of stenophagy in spiders (Araneae): evidence

based on the comparative analysis of spider diets
Stano Pekár1, Todd Blackledge2 & Jonathan Coddington3
1
Department of Botany and Zoology, Masaryk University, Brno, Czech republic,
pekar@sci.muni.cz
2
Department of Biology, University of Akron, Akron, OH, USA
3
Smithsonian Institution, National Museum of Natural History, Washington DC, USA
Most studies centred upon the evolution of stenophagy, i.e. utilisation of a

narrow trophic niche, were done on herbivores or parasites. Therefore, little is
known about forces that drive evolution of stenophagy in carnivores. We used
spiders (Araneae), as the most diversified group of terrestrial predators, to test
whether stenophagy can be explained by phylogeny. Utilising published data on
the prey of almost 600 species of spiders and phylogenetic comparative methods
we aim to find whether stenophagy (a low diversity of prey) is a derived state,
whether it promoted diversification, and whether it is determined either by a
foraging guild or a geographical zone. Six categories of stenophagy were found
among spiders, with myrmecophagy being most frequent followed by
araneophagy, lepidopterophagy, termitophagy, dipterophagy and
crustaceophagy. At the family level, stenophagy has evolved repeatedly and
independently in several families at various positions of the tree. Specifically,
Ammoxenidae, Archaeidae, Caponiidae, Palpimanidae, Mimetidae, Zodariidae
and Oecobiidae are considered stenophagous on the family level. Within
families some types of stenophagy are clearly derived, while others are
plesiomorphic. Myrmecophagy was found to be derived in Zodariidae,
Salticidae, and Thomisidae. Lepidopterophagy was derived in Araneidae. But
araneophagy turned out to be a primitive in Salticidae, Zodariidae, and
Araneidae. Termitophagy and crustacophagy also appear to be derived on the
family level, whereas dipterophagy was most frequent in species positioned at
the middle of the family tree. As concerns diversification we found that
euryphagous families or genera have about 4-times higher species diversity than
those in stenophagous families or genera. The diet breadth of species from the
tropics and subtropics was on average less diverse than that from the temperate
zone. And the diet breadth was lower in the cursorial than in the web-building
species. Future studies are needed to bring stronger evidence on the effect of
phylogeny once better resolution of spider phylogeny and more prey data is
available (now it is only 0.1% of known species). We conclude that at present
the evolution of stenophagy in spiders appears to be governed by several
determinants including phylogeny.
329
Mercy landlord or good lodgers? A form of interaction

between kleptoparasitic and host spiders
Po Peng
Department of life science, Tunghai university, Taichung 40704, Taiwan,

g98232001@thu.edu.tw
The interactions between different species of spiders are usually described

as competition, predation, or parasitism. Mutualistic interactions involving
different species of spiders have rarely been reported. In this study, I examined
the nature of interactions between the kleptoparasitic spiders, Argyrodes
fissifrons, and their Cyrtophora host spiders. The body colouration of A.
fissifrons is very conspicuous, while that of the Cyrtophora hosts is dark brown.
Previous studies showed that conspicuous body colourations of spiders may
function as visual lure to prey. In this study I tested whether the conspicuous
body colouration of A. fissifrons functioned as a visual lure to attract prey to the
webs of Cytophora hosts, and consequently benefited the hosts. I performed a
field experiment, in which the presence of kleptoparasites was manipulated and
prey catching rates of host webs were monitored by infrared video cameras. The
presence of A. fissifrons increased the prey catching rates of Cytophora webs,
whether or not the host spiders were present. The interaction between these two
spiders may therefore be mutualistic.
Keywords: kleptoparasitic spider, mutualism, body colouration.
330
X-ray computed tomography of a new Craspedisia

(Araneae: Theridiidae) in Miocene Dominican amber
David Penney1,5, Yuri M. Marusik2,5, David I. Green3,

Andrew McNeil4, Robert Bradley4, Philip J. Withers4
& Richard F. Preziosi1
1
Faculty of Life Sciences, The University of Manchester, United Kingdom,
david.penney@manchester.ac.uk
2
Institute for Biological Problems of the North, Magadan, Russia, yurmar@mail.ru
3
Manchester Museum, The University of Manchester, United Kingdom
4
School of Materials, The University of Manchester, United Kingdom
5
Corresponding authors
Introduction
Comb-footed spiders (Araneae: Theridiidae) rank fifth (out of 109
families) in terms of extant spider diversity, with 2297 species in 112 genera
(Platnick 2010) and form the most species rich spider family in the fossil record
(Dunlop et al. 2010). This is due to their regular occurrence in Cenozoic ambers
(Marusik & Penney 2004, Penney 2008), but they are unknown from the
Mesozoic (Selden & Penney 2010). They represent the most diverse spider
family in Miocene Dominican amber, with 38 named species (Penney 2008),
most of which were described by Wunderlich (1988), who also described a
mature male specimen of Craspedisia Simon. He considered it too poorly
preserved to diagnose sufficiently to assign a species name. Only three extant
Craspedisia have been described: C. cornuta (Keyserling) from Brazil; C.
spatulata Bryant from Hispaniola and C. longioembolia Yin et al. from China
(Levi 1963, Yin et al. 2003). They are easily identified by their small size,
prominent clypeal projection and their heavily sclerotized epigastric region.
However, their pedipalps are rather variable. The species described from China
by Yin et al. (2003) differs to such an extent from the generotype, including in
the structure and location of the clypeal projection, that it is probably misplaced
in Craspedisia.

The proholotype is well preserved in a clear, oval piece of yellow
Dominican amber with two small insect syninclusions and numerous soil
particles. It was excavated in the La Bucara amber mine. La Bucara is close to
the La Toca group of mines, but has only been worked for the last several years,
so the actual mine tunnels are fairly shallow as compared to the deep La Toca
mines. The La Bucara mines tend to be lower on the mountain slopes compared
to the La Toca mines, which are close to the ridgeline. The proparatype is
preserved in a clear piece of amber-orange Dominican amber and belongs in the
collections of the Senckenberg Museum, Frankfurt (SMF); mine provenance
unknown. There are no syninclusions. The proparatype is poorly preserved and
331
may have been subject to diagenetic processes, presumably overburden pressure

and increased temperature, which have distorted and modified the cuticle of the
legs and body. However, the features important for identification are clearly
visible and it is possible to assign both specimens to the same species.
The recent application of VHR-computed tomography to amber inclusions
has produced remarkable results (Penney et al. 2007). It was used here to reveal
characters that would otherwise have been inaccessible. The specimen was
scanned using the Xradia MicroXCT at the University of Manchester's Henry
Moseley X-ray Imaging Facility. The 3D virtual volume was reconstructed from
1531 16-bit radiographs (projections) acquired with a significant sample to
detector distance (180mm) such that phase contrast was visible. This step was
necessary because such samples show low levels of attenuation contrast. These
conditions resulted in a pixel size of 2.0µm and a FOV of 4.1mm. Following the
application of a phase-retrieval algorithm, tomographic reconstruction employed
the TXMReconstructor software by Xradia. Each slice was exported as a 16-bit
TIFF file and imported into Avizo 6.1 for visualisation, using a combination of
automatic isosurface extraction via thresholding, and manual segmentation.
Light microphotographs were assembled from a stacked series of digital
images recorded by a Nikon Coolpix 4500 camera mounted on a Leica M10
stereomicroscope with 0.63 and 1.6 planapochromatic objectives, following
the technique described by Green (2005).
Taxonomy
Theridiidae Sundevall, 1833
Pholcommatinae Simon, 1894
Craspedisia Simon, 1894
Craspedisia sp. (to be diagnosed and named)
Craspedisia sp. Wunderlich, 1988: 147, fig. 343.
Description
Proholotype male. Total length 2.0 mm, carapace punctate, raised in the
ocular area. Eight eyes: AME largest, lateral eyes contiguous. Clypeus twice
diameter of AME, with a highly distinctive projection, covered with short setae
(not resolved by computed tomography). The clypeal projection does not have a
downwards-curved tip as in the extant species. Sternum punctate, maxillae not
converging, chelicerae unmodified, dentition not visible. Legs without
modifications, relatively short for a theridiid spider, covered with fine setae and
with a long proximal and distal spine dorsally on each patella and tibia (tibia of
third leg with one spine); tarsi with three claws, tarsal comb on fourth leg
indistinct. Pedipalp structure not clearly visible, nor was it possible to
reconstruct sufficiently well with computed tomography. However, there is a
distinct, long, thick embolus arising retrolaterally (clearly visible in the
proparatype). Abdomen globular, with four heavily sclerotized dorsal sigillae
and many tiny scerotized dots (possibly pits bearing setae) over entire surface.
Epigastric region with heavily sclerotized covering extending around pedicel.
332
Spinnerets without modifications, but with distinct colulus and anal tubercle.
Female unknown.
Distribution. Miocene Dominican amber forest.
Discussion
Based on both pedipalp and somatic morphology the new fossil species is
most closely related to C. spatulata, which occurs on Hispaniola today, and so
makes sense in terms of geography. C. cornuta differs from both the
aforementioned species by having a shorter embolus, the carapace raised in the
posterior region, and in possessing a longer and more curved clypeal projection.
The discovery of yet another fossil species from an extant genus in the fossil
Dominican amber assemblage that has its closest relative in the extant Hispaniolan
fauna, demonstrates the great antiquity of components of at least this neotropical
ecosystem. Many lineages appear to have survived with little change despite
proposed glacial episodes in the region (although the degree to which these
affected the Greater Antilles is unclear). While the fossil and extant spider faunas
are extremely similar at family (and in large part genus) level (Penney & Pérez-
Gelabert 2002, Penney 2008), there are some differences in other groups. Poinar
(1999) proposed that the demise of certain groups known from Dominican amber
but absent from the Greater Antilles today resulted from a cool period associated
with increased aridity during the Plio-Pleistocene, although Peñalver & Grimaldi
(2006) suggested that the insularization of Hispaniola was probably more
important. Assuming that this neotropical assemblage has remained relatively little
changed for the majority of the last 16 million years (and possibly much longer),
the rapid rate by which humankind is depleting tropical forest cover globally is
cause for immediate and great concern.
Acknowledgements
Peter Jäger (SMF) is thanked for the loan of the proparatype. Dmitri
Logunov (Manchester Museum) is thanked for access to resources and we are
grateful to Dr Chris Martin (University of Manchester) for X-ray imaging
support.
References
Dunlop J.A., Penney D. & Jekel D. 2010. A summary list of fossil spiders. In:
Platnick N.I. (2010): The world spider catalog, version 10.5. AMNH,
online at: http://research.amnh.org/entomology/spiders/catalog/index.html.
Green D.I. 2005. Digital combination photography: A technique for producing
improved images of microscopic minerals. Australian Journal of
Mineralogy, 11: 13-24.
Levi H.W. 1963. The spider genera Cerocida, Hetschkia, Wirada and
Craspedisia (Araneae: Theridiidae). Psyche, 70: 170-179.
Marusik Y.M., Penney D. 2004. A survey of Baltic amber Theridiidae (Araneae)
inclusions, with descriptions of six new species. Arthropoda Selecta,
Special Issue, 1: 201-218.
333
Peñalver E. & Grimaldi D. 2006. New data on Miocene butterflies in Dominican

amber (Lepidoptera: Riodinidae and Nymphalidae) with the description of
a new nymphalid. American Museum Novitates, 3519: 1-17.
Penney D. 2008. Dominican Amber Spiders: a comparative palaeontological-
neontological approach to identification, faunistics, ecology and
biogeography. Siri Scientific Press, Manchester.
Penney D. & Pérez-Gelabert D.E. 2002. Comparison of the Recent and Miocene
Hispaniolan spider faunas. Revista Ibérica de Aracnología, 6: 203-223.
Penney D., Dierick M., Cnudde V., Masschaele B., Vlasssenbroeck J., Van
Hoorebeke L. & Jacobs P. 2007. First fossil Micropholcommatidae
(Araneae), imaged in Eocene Paris amber using X-Ray Computed
Tomography. Zootaxa, 1623: 47-53.
Platnick N.I. 2010. The world spider catalog, version 10.5. AMNH, online at:
http://research.amnh.org/entomology/spiders/catalog/index.html.
Poinar G.O. Jr. 1999. Extinction of tropical insect lineages in Dominican amber
from Plio-Pleistocene cooling events. Russian Entomological Journal, 8:
1-4.
Selden P.A., Penney D. 2010. Fossil spiders. Biological Reviews, 85: 171-206.
Wunderlich J. 1988. Die fossilen Spinnen im dominikanischen Bernstein.
Beiträge Araneologie, 2: 1-378.
Yin C.M., Griswold C.E., Bao Y.H. & Xu X. 2003. A new species of the spider
genus Craspedisia from the Gaoligong Mountains, Yunnan, China.
Bulletin of the British Arachnological Society, 12: 383-384.
334
Reproductive behaviour and the role of the nuptial gift

for Thaumasia sp. nov. (Pisauridae) in central Brazil
Rommel B. Pereira & Paulo C. Motta
Universidade de Brasília, Brazil, bio_aracnideos@yahoo.com.br, mottapc@unb.br
Introduction
In order to achieve reproductive success, animals use courtship to
establish intersexual communication (Robinson 1982, Huber 2005, Peretti et al.
2006). Spiders’ diverse mating systems can be seen as a mechanism of
reproductive isolation (Foelix 1996). Courtship in spiders is formed by ritualized
patterns that are important to reduce female aggressiveness and stimulate sexual
excitement (Robinson 1982, Schneider & Lubin 1998).
Nuptial gifts consist on any form of nutrient transference by the male to
the female during any stage of the copula and it is best known in insects (Vahed
1998). The occurrence in spiders is rare and has been studied almost exclusively
in Pisaura mirabilis (Pisauridae) (Austad & Thornhill 1986, Drengsgaard & Toft
1999, Stålhandske 2001, Bilde et al. 2007). More recently it was also reported
for Paratrechalea ornata and Paratrechalea galianoae (Trechaleidae) by Costa-
Schmidt et al. (2008). Nuptial gifts have been thought as male mating effort,
insurance against cannibalism and parental investment (Stålhandske 2001, Bilde
et al. 2007), but these hypotheses are not excluded (Stålhandske 2001).
Thaumasia sp. nov. represents the first record for nuptial gift in Brazilian
Pisauridae, and the aims of this study are: describe the reproductive behaviour of
Thaumasia sp. nov.; analyze the importance of the nuptial gift for the courtship
and mating; and verify whether the size of the nuptial gift has any influence on
the duration of the copula and in the spiderling production.
Methods
Thaumasia sp. nov. has body length from 7 to 11mm, brownish colour
with lateral white stripes surrounding the body. White spots are found in the
abdomen and the ventral part of the abdomen is white. The sexes are similar in
size and colouration. It is semi-aquatic species and although has nocturnal
habits, it can be found on aquatic plants during the daylight. This species is
being described by Estevam L.C. Silva and Arno A. Lise.
The specimens were collected in pounds of Brasília, central Brazil, in
January and March 2007 and February and June, 2008. The juvenile spiders
were reared in laboratory (20-22ºC) separately in plastic vials, and fed ad libitum
with Leurolestes circunvagans.
Having reached the sexual maturity, the spiders were separated into three
groups of 30 individuals each (15 couples by group) according to the prey size
(Leurolestes circunvagans) to be used by the male as nuptial gift. All prey items
were measured. The males from group A was deprived from prey items, group B
received a small prey item (body length 1-7 mm) and males from group C
335
received large preys (8 - 24 mm). As a courting arena we used a glass tank (50
cm x 30 cm x 40 cm) with water (15 cm depth) and two pieces of Styrofoam
glued to each end of the tank were used as solid substrate. Females were
introduced first at one end of the tank and the males were introduced 24 hours
after the females at the opposite end. Ten minutes after introducing the males,
the prey item was offered. The experiments had duration of 1 hour per time
started to be counted right after the prey introduction. At the end of each
experiment the arena was completely cleaned, the water changed and the
Styrofoam pieces washed. The experiments were held in the evening at the same
time (07.00 pm) and were filmed. The individuals were used only once, and then
preserved and measured.
Results
Reproductive behaviour
We did not observe the sperm induction or the males touching the
females’ draglines. The courtship started as soon as the male captured the prey.
Once this happened he began to shake the body and this movement produced
concentric waves. Then the male went to the solid substrate with the prey in its
chelicerae and started moving the body up and down and, at the same time,
moved around the female in little jumps. It also shook its abdomen alternating
this movement with taps against the substrate with the legs I and II. He also
rubbed the legs I and II or II and III. Then the male placed itself in front of the
female with legs I raised up and flexed back. The female responded rubbing its
legs II. This movement was copied by the male. It, then, moved towards the
female touching her with his first legs. Female remained with legs I raised up
and flexed back.
The next step was the making of the nuptial gift. The male put the prey on
the substrate and started to cover it with layers of silk while doing this the male
never lost contact with the female touching her with his legs. While covering the
prey with silk the male would press it against the substrate with his palps. Then,
he plucked the gift out, placed himself in front of the female assuming a
hyperflexed position exposing the gift in its chelicerae. The female would grab
the gift and the copula started. The couple assumed a lateral position where the
male in the first palpal insertion placed its prosoma right beside the female
opisthosoma and she did just the opposite. This position was inverted for the
second papal insertion. The first separation was done by the male, but the end of
the copula was always the female prerogative. The female kept the present
during the whole copula.
Experiments on the role of nuptial gift
Males from the group A (without preys) do not show courtship or copula
behaviours. Males from the groups B and C that have lost their prey for the
females did not court them or interrupted the courtship (n=3).
There were nine successful copulas in the group B resulting in five egg
sacs, the average copula duration was 55.9 s and the total number of spider lings
was 6914. For the group C, four copulas were successful resulting in four egg
336
sacs, the average copula duration was 80.8 s and the total number of spiderlings
was 6201.
The size of the nuptial gift has significantly influenced the duration of the
copula (n=13, Spearman correlation r2=0.77, p=0.002). In each group, the copula
duration was different according the prey size (n=13, Mann-Whitney U=45,
p=0.003). When analyze the results between treatments the number of offspring
produced did not vary significantly (ANCOVA, n=9, F2,10=2.32, p=0.14).
However, when we analyze the size of the nuptial gift regardless the groups,
then we have a significant difference in the offspring production and the size of
the gift (Multiple Steps Regression F1,11=6.75, β=0.62±0.24, R2=32.40%,
p=0.02). The body size of the females does not have influence in the number of
spiderlings (ANCOVA, n=9, β=0.30±0.39, p=0.39).
Discussion
There are some important differences between the courtship pattern of this
new species and its most famous relative P. mirabilis and the two Trechaleidae
reported by Costa-Schmidt (2008). The nuptial gift is mandatory for the
courtship and copula since nothing happened with the couples deprived from
prey items, differently of P. mirabilis whose males have obtained success in
mating without nuptial gifts (Stålhandske 2001).
The second important aspect observed was the moment the males made
the nuptial gift. In P. mirabilis and the two Trechaleidae they made the gift
before meeting the females. Thaumasia sp. nov. starts to make it in the course of
the courtship. One possible explanation for this may reside in the fact that both
males and females live close together in their natural habitat sharing, most of the
time the same plant leaf while waiting for a prey to be trapped in the water film
so the males do not have to wander searching for the females.
Only males with gift presented some courtship display and copula, so
establishing the importance of the nuptial gift in reproduction. The nuptial gift
did not work as a protection against sexual cannibalism since none of the 45
males were attacked by any females.
The size of the nuptial gift has significantly influenced the duration of the
copula and the offspring production, indicating that a nuptial gift can work as a
parental investment. This study shows clearly that a lot more is yet to be done
with this new species in order to fully understand their behavioural patterns and
to establish evolutionary connections between the spiders’ families.
References
Austad S.N. & Thornhill R. 1986. Female reproductive variation in a nuptial-
feeding spider, P. mirabilis. Bulletin of the British Arachnological
Society, 7: 48-52.
Bilde T., Tuni C., Elsayed R., Pekár S. & Toft S. 2007. Nuptial gifts of male
spiders: sensory exploitation of the female’s maternal care instinct or
foraging motivation? Animal Behaviour, 73: 267-273.
337
Costa-Schmidt L.E., Carico J.E. & Araújo A.M. 2008. Nuptial gifts and sexual
behavior in two species of spider (Araneae, Trechaleidae, Paratrechalea).
Naturwissenschaften, 95: 731-739.
Drengsgaard I.L. & Toft S. 1999. Sperm competition in a nuptial feeding spider,
Pisaura mirabilis. Behaviour, 136: 877-897.
Foelix R.F. 1996. Biology of spiders. Harvard University Press, Cambridge,
Mass.
Huber B.A. 2005. Sexual selection research on spiders: progress and biases.
Biological Reviews, 80: 363-385.
Peretti A., Eberhard W.G. & Briceño R.D. 2006. Copulatory dialogue: female
spiders sing during copulation to influence male genitalic movements.
Animal Behaviour, 72: 413-421.
Robinson M.H. 1982. Courtship and mating behavior in spiders. Annual Review
of Entomology, 27: 1-20.
Schneider J.M. & Lubin Y. 1998. Intersexual conflict in spiders. Oikos, 83: 469-506.
Stålhandske P. 2001. Nuptial gift in spider Pisaura mirabilis maintained by
sexual selection. Behavioural Ecology, 12: 691-697.
Vahed K. 1998. The function of nuptial feeding in insects: a review of empirical
studies. Biological Reviews, 73: 3-78.
338
The fall of spiderman: silk production from tarantula

tarsus experimentally questioned (Araneae:
Theraphosidae)
Fernando Pérez-Miles, Alejandra Panzera, David Ortiz-Villatoro

& Cintya Perdomo
Sección Entomología, Facultad de Ciencias, Montevideo, Uruguay, myga@fcien.edu.uy,

alecita2007@gmail.com, theraphosidae@gmail.com, cuquis266@hotmail.com
The abdominal spinnerets are specialized structures of the spiders,

considered as key features unique to the group. As all other spiders, theraphosid
tarantulas spin silk from abdominal spinnerets. Recently Gorb et al. (2006)
reported that the zebra tarantula, Aphonopelma seemanni (F.O.P.-Cambridge,
1897) also can secret silk from their tarsi. This amazing discovery was
interpreted as a mechanism to improve the ability of tarantula to climb on
vertical surfaces. We tested that when the abdominal spinnerets are
experimentally sealed, the zebra tarantula cannot secret silk or similar threads
(Pérez-Miles et al. 2009), disagreeing with previous reports. Gorb et al. (2009)
gave some additional elements to support their hypothesis of tarsal silk
production, which are also discussed here.

We used four females of A. seemanni collected in Guatemala, Escuintla,
Torremolinos on August 15, 2008. All individuals were kept in glass terrariums
fed with insects (Blaptica dubia, Blattaria, Blaberidae) and provided with water.
We carried out two series of experiments. In the first series we placed each
specimen of A. seemanni in a glass container (10 x 33 cm of base, and 15 cm
height) which had the floor and two vertical walls covered with 20 microscope
slides. Tarantulas remained 14 hours (overnight) in the container and after that,
the slides were carefully examined. In several instances of spider walking
behaviour (mainly vertical walking) they were photographed. The second series
was performed seventy- two hours later; we completely sealed the spinnerets of
the same individuals with paraffin (60ºC) and repeated the experiments in the
same conditions. We observed spider behaviour for the first and last two hours
of the experiment. After the experiments, slides were carefully examined with
optical microscope to determine the presence of silk threads.
Leg tarsi of other preserved specimens were included in paraffin, and
transverse cuts were done with a microtome, in sections of 5 micrometers. The
histological analysis was made with standard hematoxiline-eosine technique. All
cuts were observed to determine the presence of gland tissue or gland conduits
and compared with images of spider silk glands from the literature. The internal
surface of tarsi exuviae of A. seemanni and Grammostola mollicoma (Ausserer,
1875) were also studied by observation with a scanning electron microscope, to
determine if gland conduits were present.
339

In the first series of experiments, with free spinnerets, most slides (24
out of 40 in horizontal, and 16 out of 40 in vertical slides) showed silk
threads together with dislodged abdominal urticating setae. None of the
slides in the second series with sealed spinnerets showed silk threads and we
found only urticating setae. In both series of experiments we observed the
spiders walking on horizontal and vertical surfaces.
For additional evidences we made transverse cuts of A. seemanni tarsi
and no structures interpretable as silk glands or silk conduits were observed.
When we examined the internal face of the tarsal exuvia of A. seemanni and
G. mollicoma by SEM, we found orifices without traces of silk or other
substances inside. A morphological reconstruction from several views led us
to interpret these orifices as the basis of macrosetae. No evidences of gland
conduits were found.
We discovered that tarantula entangles silk threads from the spinnerets
with their tarsi. This could explain the presence of thread footprints
photographed by Gorb et al. (2006). The tarantula usually use hind legs to
entangle silk, but we also observed A. seemanni entangling with their other
legs which answer one of the points of the reply of Gorb et al. (2009). The
other point is that tarantula silk released from spinnerets involve multiple
parallel silk threads which are better observed when magnified. This pattern
could be maintained once the silk line is adhered to the substrate, although
combed or stepped by tarsal scopulae. Gorb et al. (2009) found by SEM
examination of a silk line, a distinct broad area at the beginning of the fiber
that they interpret as an initial fluid. They propose that presumably any fluid
phase adhesive would be lost in the transference from abdomen to tarsi. We
agree with this interpretation, but a question that remains is: how could Gorb
et al. (2009) discard if it was a silk line directly adhered from abdominal
spinneret?
Tarsal structures interpreted as spigots by Gorb et al. (2006) are very
different from diverse kind of spigots reported for spiders (Bond 1994,
Coddington 1989, Griswold et al. 2005). We found that such structures are
very similar in morphology and size to fragments of tarsal thermo-sensory
setae reported for other tarantulas (Raven 2005). As well, in insects, a
common origin or convergence was proposed for silk spigots and sensory
setae (Ross 1991, Merrit 2007). The presence of a developed dense scopulae
brush with spatulated setae on the ventral surface of tarsi and metatarsi of all
Theraphosidae seems rather incompatible with the presence of spinning
spigots in a lower layer. Scopulae setae could interfere with silk release.
Additionally, Niederegger & Gorb (2006), explained that in A. seemanni the
friction with scopulae setae is enough to guarantee spider climbing.
Summarizing, we did not find evidences of silk-like secretion through leg
structures in tarantulas.
340
References
Bond J.E. 1994. Setae-spigot homology and silk production in first instar of
Antrodiaetus unicolor spiderlings (Araneae: Antrodiaetidae). Journal of
Coddington J.A. 1989. Spinneret silk spigot morphology: evidence for the
monophyly of orb weaving spiders, Cyrtophorinae (Araneae), and the
group Theridiidae plus Nesticidae. Journal of Arachnology, 17: 71-95.
Gorb S.N., Niederegger S., Hayashi C.Y., Summers A.P., Vötsch W. & Walther
P. 2006. Silk like secretion from tarantula feet. Nature, 443: 407.
Gorb S.N., Niederegger S., Hayashi C.Y., Summers A.P., Vötsch W. & Walther
P. 2009. Reply. Nature: doi:101038/nature08405.
Griswold C.E., Ramírez M.E., Coddington J.A. & Platnick N.I. 2005. Atlas of
phylogenetic data for entelegyne spiders. Proceedings of the California
Academy of Sciences, 56: 1-124.
Merrit D.J. 2007. The organule concept of insect sense organs: Sensory
transduction and organule evolution. Advances in Insect Physiology, 33:
192-241.
Niederegger S. & Gorb S.N. 2006. Friction and adhesion in the tarsal and
metatarsal scopulae of spiders. Journal of Comparative Physiology, A,
192: 1223-1232.
Pérez-Miles F., Panzera A., Ortiz-Villatoro D. & Perdomo C. 2009. Silk
production from tarantula feet questioned. Nature, 461:
oi:101038/nature8404.
Raven R.J. 2005. A new tarantula species from northern Australia (Araneae,
Theraphosidae). Zootaxa, 1004: 15-28.
Ross E.S. 1991. In: The insects of Australia 405-409, Canberra & Melbourne
Univ. Press.
341
Condition- and phenotype-dependent dispersal

in a spider with a clustered distribution
Julien Pétillon1, David Deruytter2, Arthur Decae2 & Dries Bonte2
1
Evolutionary Ecology Group, University of Antwerp, Antwerpen, Belgium,
julien.petillon@ua.ac.be
2
Terrestrial Ecology Unit, Ghent University, Gent, Belgium
Dispersal plays a central role in the dynamics and evolution of spatially

structured populations (Kokko & López-Sepulcre 2006) and is involved in
processes like colonization, gene flow, genetic drift, local adaptation, inbreeding
and long-time survival of populations and species (for reviews see Bowler &
Benton 2005, Ronce 2007). Further, the presence of heritable variation and
strong selection pressures related to e.g. landscape composition may induce fast
evolution in dispersal traits. It is recognized to be both a conditional and
phenotype-dependent process dependent on factors related to density and body
condition. Atypus affinis (Mygalomorpha, Atypidae) populations often reach
high densities within clusters under natural conditions. Therefore, we expect
natal dispersal through ballooning to be positively related to local densities of
spiderlings and to increased starvation. In first instance, we assessed the
relationship between density (assessed in the field, in Flanders, Belgium) and
individual body condition (estimated by sugars and lipids measurements). In
second instance, we confirm our specific hypotheses on ballooning dispersal by
means of laboratory experiments and additionally demonstrate that emigration
decision making is determined by preceding ballooning events.
References
Bowler D.E. & Benton T.G. 2005. Causes and consequences of animal dispersal
strategies: relating individual behaviour to spatial dynamics. Biological
Reviews, 80: 205-225.
Kokko H. & Lopez-Sepulcre A. 2006. From individual dispersal to species
ranges: Perspectives for a changing world. Science, 313: 789-791.
Ronce O. 2007. How does it feel to be like a rolling stone? Ten questions about
dispersal evolution. Annual Review of Ecology, Evolution and
Systematics, 38: 231-253.
342
Ground-dwelling spiders of Lake Elton area

(Caspian Sea Lowland, Russia)
Tatyana V. Piterkina
Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia,

piterkina@yandex.ru
The purpose of the research was investigation of the araneofauna and

seasonal dynamics of spider population structure in the Lake Elton area. Elton is
a biter lake located in the NW Caspian Sea Lowland, at the contact of steppe and
semi-desert zones (49°12.47'N, 46°39.75'E). Desert steppe associations with
different wormwoods (Artemisia lerchiana, A. pauciflora, A. austriaca) as well
as Kochia prostrata, Agropyron desertorum and Festuca valesiaca form the
main part of landscapes around the lake. Hyper-halophytic plant communities
occupy salt-marshes. Tangles of reed and cattail grow on the banks of small
rivers. Gullies running into the lake are occupied with arboreal vegetation. Ten
habitats were chosen as model ones; among them are six zonal, one intrazonal
and three extrazonal biotopes. Material was collected using pitfall traps. Lines of
traps were functioning all year round, from 10 May 2006 till 10 May 2007. The
material is at the stage of initial processing. The seasonal dynamics of total
abundant, ratio of families, sex and age structure of population is revealed.
343
Not so banal after all; latest evidence on Loxosceles

diversity in Canary Islands and Northwestern Africa
Enric Planas & Carles Ribera
Institut de Recerca de la Biodiversitat (IRBio) and Dept. Biologia Animal, Universitat de

Barcelona, Spain, enplanas@gmail.com, cribera@ub.edu
The spider genus Loxosceles has a widespread distribution, and the vast
majority of its species are concentrated in North and South Americas. The
cosmopolitan Loxosceles rufescens (Dufour, 1820), has its original distribution
range in the Mediterranean basin, and has been the only representative accepted
of the genus in this area until the recently discovered Loxosceles mrazig Ribera
& Planas, 2009 from Tunisia.
The evidence of the diversity in the Loxosceles lineage in NW Africa has
been pointed out by Duncan et al. (2010). In this paper, the authors indicated the
existence of a NW African clade, which includes the widespread L. rufescens,
the new recently described L. mrazig and some evolutionary lineages present in
the Maghreb and Canary Islands. This clade also includes Loxosceles
foutadjalloni Millot, 1941 from Guinea and the South American Loxosceles
amazonica Gertsch, 1967.
Recent collecting trips carried out in Morocco and Canaries provided us
with abundant material of Loxosceles. In this contribution we present the
preliminary results on taxonomy and phylogeny of this species from Canary
Islands and Morocco.
Canary Islands harbour an endemic group of Loxosceles species clearly
distinctive by morphology and molecular characters. Until now we have
identified 6 species: 1 in Fuerteventura and Lanzarote, 2 in Tenerife, 2 in Gran
Canaria plus the cosmopolitan L. rufescens. Our data suggest a single
colonization event to the Eastern Canary Islands (Fuerteventura and Lanzarote)
and a posterior interinsular colonization to Gran Canaria and Tenerife.
Several individuals from SW Morocco, morphologically consistent with
L. rufescens lineage are placed as the sister group of the L. rufescens clade
showing a high genetic divergence.
Within the L. rufescens lineage several independent evolutionary lineages
are discussed. We provide individuals from the type locality, Sagunt, and a wide
representation of specimens from the Mediterranean Basin. The lack of
geographical structure due to human-mediated dispersal impedes the
clarification of the dispersal patterns, but the genetic distance among groups and
the fact that populations of geographical proximity don't have genetic flux could
suggest a cryptic speciation.
344
Development and life cycles of erigonid and wolf spiders

(Araneae: Linyphiidae: Lycosidae) in arable fields:
differences due to crop type and region
Ralph Platen
Institute for Land Use Systems, Leibniz-Centre for Agricultural Landscape Research
(ZALF), Germany, platen@zalf.de
Different live stages of development (egg-cocoons, instars, juveniles,

subadults and adults) were identified for wolf spider species (e.g. Pardosa
agrestis agrestis, P. palustris, P. prativaga and Trochosa ruricola) and the
juvenile and adult stages of an erigonid spider (Oedothorax apicatus) were
identified from field material, collected with pitfall traps on arable land,
collected during three years (2005-2007) a wide spectrum of agricultural
landscapes in Germany (Federal States of Bavaria, Brandenburg, Mecklenburg-
Vorpommern, Schleswig Holstein and Thuringia). In each year five pitfall traps
were used in each field from April to July in legumes, summer as well as winter
cereals, and from April to September in maize and perennial grassland. At first,
the adults of the spiders were determined. Then the fields were identified where
one of the species of the (genera), mentioned above, was caught exclusively. We
concluded that the subadults and juvenile stages caught in these fields belong to
these species. These stages were identified and their individual lengths were
measured. The time of development was determined and compared for each
crops, year of investigation and location (Federal State), respectively. The
possibility to complete a life cycle within the field is discussed for each of the
species as well as differences in the time of development together with regional
and meteorological aspects.
345
Araneofauna of chalk lands of Eastern Ukraine

Nina Yu. Polchaninova
Kharkov National University, Ukraine, polchaninova@mail.ru
Introduction
Underlying bedrock of chalk creates specific conditions for the forming of
a chalk biota. Geological history of the landscape, structure features of the chalk,
high albedo, low daily moisture and temperature fluctuation of the upper layers
promote development of extrazonal communities, where a great number of relic
and endemic species are presented, as well as thermophilous and xerophilous
ones that enrich local fauna with southern elements. Spiders of chalk lands of
Eastern Europe have not been studied purposely. In literature, there are only a
short remark of the species richness for South-Eastern Ukraine (Polchaninova &
Prokopenko 2003), and records in the lists of local faunas of North-Eastern
Ukraine and Southern Russia (Kulczyński 1913, Prisny 2003, Ponomarev &
Polchaninova 2006, Polchaninova & Procopenko 2007, Polchaninova 2009).

In the South of Central Russian Upland, chalk lands, with chalk bedrock
on the upper interfluves and right high river banks of the river Don basin,
occupy a vast region. The altitude of hills does not exceed 300 meters. In the
forest-steppe natural zone, as well as on the Donetsky Ridge, that is regarded as
an island of the forest-steppe in the steppe zone, the upland and right river banks
are partly covered with oak forests. Among them, relic tertiary forests of Pinus
cretaceus are scattered as small patches. Grasslands on the northern slopes are
represented by a meadow steppe. In the steppe natural zone, these slopes are
occupied by a forbgrass-feathergrass-fescue steppe, while on the upper part and
on the southern slopes in both zones, specific calcareous vegetation forms an
azonal chalky steppe.
Our investigations were carried out in 12 localities of Eastern Ukraine
within the three administrative regions.

In total, 179 spider species were registered on chalk lands of Eastern
Ukraine; among them, 100 species in steppe communities, and 134 species in the
forest ones. In chalk grasslands, as in all steppes types, Gnaphosidae is the
richest family of the fauna (Tab. 1), being the most numerous in granite and
forbgrass steppes. Excluding Linyphiidae, the other six main families are equally
represented in the chalk community.
In the grass layer of chalk habitats, common steppe dominants as
Theridion impressum L.K., Agalenatea redii (Scop.) Argiope bruennichi (Scop.)
Neoscona adiantum (Walck.), Dictyna arundinacea (L.), Tibellus oblongus
(Walck.), Xysticus cristatus (Cl.), X. striatipes L.K., Heliophanus flavipes
(Hahn) prevail. Becides, Mangora acalypha (Walck.) and Cheiracanthium
346
pennyi O.P.-C. are numerous in the North of the area in question, with Dictyna
latens (Fabr.), Tibellus macellus Sim., Thomisus onustus Walck., Phylaeus
chrysops (Poda) dominating in the South. Linyphia triangularis (Cl.), L.
tenuipalpis Sim., and Philodromus cespitum (Walck.) densely settle low shrub
thickets. In the litter, a bulk of common species Meioneta rurestris (C.L.K.),
Alopecosa sulzeri (Pav.), Berlandina cinerea (Mg), Drassodes pubescens
(Thor.), Thanatus arenarius L.K., Xysticus kochi Thor. is typical; in addition
to them, local dominants Trochosa robusta (Sim.), T. ruricola (De Geer), T.
terricola Thor., Gnaphosa taurica Thor., and less numerous Atypus muralis
Bertk., Eresus cinnaberinus (Oliver), Stemonyphantes lineatus (L.), Asianellus
festivus (C.L.K.), Phlegra fasciata (Hahn) were found. Only 5 species were
recorded exclusively in chalk grasslands. Uloborus walckenaerius Latr., a
typical dweller of sand habitats, settles Artemisia thickets; the other 4 species
were rare. In the forest-steppe zone of Ukraine, some polytopic mesophilous
species (Micrommata virescens (Cl.), Pisaura mirabilis (Cl.), Phrurolithus
festivus (C.L.K.), Alopecosa cuneata (Cl.) settle northern chalk slopes.
Northwards, in Central Forest-Steppe, the tendency is more evident; a lot of
species never occurring in this habitat southwards appear on chalk hills
(Robertus lividus (Bl.), Microlinyphia pusilla (Sund.), Neriene clathrata
(Sund.), Araniella cucrbitina (Cl.), Araneus marmoreus Cl., Zora nemoralis
(Bl.), etc).
There are no specific spider species characteristics of calcareous
grasslands. Uniqueness of the spider complex consists in presence of the
southern and eastern elements which have an area disjunction and sometimes
northern- or westernmost bounds of their distribution, penetrating to other
natural zones along the chain of chalk lands. As an example, the following
species can be referred: Pardosa italica Torng., Alopecosa cursor (Hahn),
Gnaphosa licenti Schenkel, G. steppica (Ovtsh.), G. mongolica Sim., G.
taurica Thor., Micaria rossica Thor., Zora pardalis Sim., Xysticus ninni Thor.
In a pine forest on chalk hills on the Donetsky Ridge, 112 spider species
were registered. Two families (Linyphiidae, 17%, and Theridiidae, 13% of the
fauna) were dominants, the other main families (see table 1), including
Philodoromidae, made up from 7 to 8,8%. More than a half species (62) was
presented by forest or polytopic mesophiles, typical of forests of the forest-
steppe and steppe zones. Photophilous dwellers of grasslands penetrate under
the canopy and/or occupy glades and cuttings (Enolpognatha thoracica
(Hahn), Simitidion simile (C.L.K.), Theridiom impressum, Th. nigrovariegatum
Sim., Meioneta rurestris, Stemonyphantes lineatus, Cyclosa oculata (Walck.),
Hypsosinga sanguinea (C.L.K.), Neoscona adianta, Alopecosa trabalis (Cl.),
Dictyna latens, Tibellus macellus, Aelurillus festivus, Philaeus chrysops and
others, 20 species altogether), 30 species are rare or sporadic. The bulk of
dominants of the grass and lower shrub layer was composed of Mangora
acalypha, Araneus diadematus Cl., Zilla diodia (Walck.), Gibbaranea
bituberculata (Walck.), Tetragnatha pinicola L.K., Linyphia hortensis, L.
triangularis, Philodromus cespitum; in the litter, it included Atypus muralis,
Alopecosa sulzeri, Arctosa lutetiana (Sim,), Haplodrassus umbratilis (L.K.),
347
Thanatus sabulosus (Mg), Asianellus festivus in pitfall traps and Tenuiphantes

flavipes (Bl.) in quadrate samples. On the whole, the spider fauna of calcareous
pine forest is richer than that of pine forests on sand soil in river valleys (we
collected in the three local fauna in Eastern Europe 72, 77 and 89 species,
respectively). However, in spite of its ancient age, we have not found any
spider species preferring or typical of this very habitat.
In the neighbouring oak forest, 129 spider species were found.
Linyphiidae was the most numerous – 24% of the fauna, the second group was
composed of Gnaphosidae, Theridiidae and Lycosidae (11-13,5%), the third
one was formed of Araneidae, Thomisidae and Salticidae (7% each). Unlike to
pine forest, the family Philodromidae was very poor (3%). In this habitat, a
half of species also belonged to common forest dwellers; a guild of photo- and
thermophiles enriched the spider complex with atypical elements (Ero aphana
(Walck.), Microlinyphia pusilla, Pardosa palustris (L.), Trochosa robusta,
Titanoeca schineri L.K., Cheiracanthium erraticum (Walck.), Zelotes electus
(C.L.K.), Xysticus robustus (Hahn) etc, 24 species altogether).
During the Quaternary Period, the Donetsk Ridge was not covered with
ice, suggesting that representatives of nemoral biota might have survived here.
Gnaphosa montana (L.K.), G. bicolor (Hahn), Thanatus sabulosus, Ozyptila
brevipes (Hahn), and Xerolycosa nemoralis (Westr.) have been found far away
from their main area in the forest zone o (for the last one), in the north of the
forest-steppe zone. The six species, typical of the forests (Dipoena erythropus
(Sim.), D. melanogaster (C.L.K.), Ceratinella scabrosa (O.P.-C.), Tapinopa
longidens (Wid.), Ozyptila claveata (Walck.), and Cozyptila blackwalli (Sim.)
do not advance southward in the steppe zone. Zelotes aurantiacus Mill.,
Gnaphosa taurica, Trachyzelotes pedestris (C.L.K.) are characteristics of
steppe communities and/or polytopic on the North coast on the Black see.
There is the sole example of their finds in oak forests of East Europe.
A comparison of spider fauna of chalk oak forests with that of the three local
plane forests in the South of the Forest-Steppe, shows a gradual decrease of the
Linyphiidae ratio from 40 to 24%, a slight increase of the Lycosidae (4-8%)
and Theridiidae (9-12%) families, and a leap in the Gnaphosidae richness from
5 to 10%. Many close species display specific patterns of distribution in these
forests. Atypus piceus (Sulz.) occurs in plane forests in the driest places. In the
chalky forest it was found on the North wet slope, while on the South one, only
A. muralis was numerous. Phrurolitus festivus is common in the first case and
very rare in the second one. Pachygantha listeri Sund., Agroeca brunnea
(Walck.), Clubiona caerulescens L.K., Zelotes aszheganovae Esyunin, Efimik,
Z. laterillei (Sim.), Xysticus ulmii (Hahn), X. luctator L.K. do not occur in the
chalky forest, where P. degeeri Sund., A. cuprea Mg., C. pseudoneglecta
Wund., Z. electus, X. robustus are present in single finds, and Z. kukushkini
Kovblyuk and Z. pedestris are numerous. So, the araneofauna of chalky oak
forests is heterogeneous, constituted of common forest species with an addition
of northern and southern elements.
348
Table 1. Species composition of the main families in the spider fauna of different steppe
types of Eastern Ukraine (number of species and %).
Steppe type
Families
Chalk Limestone Granite Forbgrass Total
N % N % N % N % N %
Theridiidae 12 12,0 6 7,5 12 9,4 20 10,6 22 9,6
Linyphiidae 5 5,0 5 6,3 10 7,8 27 14,3 28 12,3
Araneidae 10 10,0 6 7,5 7 5,5 13 6,9 15 6,6
Lycosidae 14 14,0 8 10,0 12 9,4 19 10,1 21 9,2
Gnaphosidae 17 17,0 17 21,3 27 21,1 25 13,2 41 18,0
Thomisidae 11 11,0 9 11,3 14 10,9 17 9,0 18 7,9
Salticidae 11 11,0 14 17,5 16 12,5 29 15,3 36 15,8
Others 20 20,0 15 18,8 30 23,4 39 20,6 47 20,6
Total 100 100 80 100 128 100 189 100 228 100
349
Taxonomy, phylogeny and biogeography of the Neotropical

spider genus Celaetycheus Simon (Araneae: Ctenidae)
Daniele Polotow & Antonio D. Brescovit

1
Laboratório de Artrópodes, Instituto Butantan, São Paulo, SP, Brazil; Departamento de
Zoologia, Instituto de Biociências, Universidade de São Paulo, São Paulo, SP,
Brazil, danielepolotow@gmail.com
2
Laboratório de Artrópodes, Instituto Butantan, Av. Vital Brasil 1500, CEP 05503-900,
São Paulo, SP, Brazil, adbresc@terra.com.br
Introduction
The spider genus Celaetycheus was proposed by Simon (1897) to include the
type species C. flavostriatus Simon, 1897, described based on a single female
collected at Salobro, Bahia, Brazil. Currently, the genus is monotypic (Polotow &
Brescovit, 2009). The genus can be distinguished from the remaining Ctenidae
genera by the presence of several small ventral spines on the endites, coxae, femur
and trochanter (Polotow & Brescovit, 2009: figs 12C, 13A–C) and the presence of a
cymbial retroventral process (Polotow & Brescovit, 2009: figs 11A–B) in the male
palp and short lateral spurs, with the tip below the median plate, and the
spermathecae divided in two parts, with a round head and large and curved base
(Polotow & Brescovit, 2009: figs 11C–D) in the epigynum.
The aim of this paper is to describe six new species of Celaetycheus and apply
a cladistic analysis based on parsimony to test the phylogenetic relationships of the
species of the genus. In addition, the pattern of distribution of the species is
discussed.

A total of 15 terminals composed the taxonomic sample used in the cladistic
analysis. The data matrix comprised 30 characters. Mesquite, version 2.5 (Maddison
& Maddison 2008) was used to build and edit the character matrix. Nonapplicable
and unknown data are presented as “–” and “?”, respectively. All characters were
equally weighted and all multistate characters were coded as nonadditive. The
parsimony analysis was performed using the computer program TNT 1.0 (Goloboff
et al 2003-2004).
All measurements are in millimeters. Morphological observations and
illustrations were made using a Leica MZ12 stereomicroscope with camera lucida.
The epigynum was detached from the abdomen and submerged in clove oil to clear
the internal structures.
Results
Nine new species are described to the genus Celaetycheus, all recorded to
northeast of Brazil. The genus Celaetycheus arises as a monophyletic group and
350
it is supported by four non-ambiguous synapomorphies: basal projection of the

cymbium, rounded-shaped cymbium, spermathecae with a rounded projection
and spines in the endites and labium of the male. The genus is also supported by
two ambiguous synapomorphies: elongated conductor and five pairs of spines in
tibia I and II.
Celaetycheus appears as sister group of the Calocteninae genera used in
this analysis: (Celaetycheus (Arctenus gen. nov. (Caloctenus (Toca +
Gephyroctenus)))). The genus is transferred to Calocteninae. The subfamily
Calocteninae is supported by three non-ambiguous synapomorphies: short
labium, posterior medium spinnerets reduced and tick and elongated anal setae.
The subfamily is also supported by one ambiguous synapomorphies: reduced
number of cylindrical glands.
References
Goloboff P.A., Farris J.S. & Nixon K. 2003-2004. TNT: Tree analysis using new
technology. Version 1.0. Program and documentation available from the
authors at http://www.zmuk.dk/public/phylogeny.
Maddison W.P. & Maddison D.R. 2008. Mesquite: a modular system for
evolutionary analysis. Version 2.5. Program and documentation available
at http://mesquiteproject.org
Polotow D & Brescovit A.D. 2009. Revision of the new wandering spider genus
Ohvida and taxonomy remarks on Celaetycheus Simon, 1897 (Araneae,
Ctenidae). Zootaxa, 2115: 1-20.
351
The effects of rubber plantation-forest edges on the

distribution of web-building spider composition at Khuan
Khao Wang Forest Park, Songkhla Province, Thailand
Booppa Ponksee1*, Sunthorn Sotthibhundhu1, Sara Bumrungsri1

& George A. Gale2
1
Prince of Songkla University, Songkhla, Thailand
2
King Mongkut’s University of Technology Thonburi, Bangkok, Thailand
* Corresponding author: zigzagargiope@yahoo.com
The edges have been shown to produce both positive and negative effects
upon their inhabitants. In addition, edge preferences of predators have been
supported in some ecosystems and not in the others. A gap that makes edge
effects difficult to create any generalization is a lack of knowledge in several
aspects and the inconsistency of research results. Accordingly, the gaps, several
groups of organisms including spiders, ants, butterflies, lizards, etc. and several
types of adjacent areas namely plantations adjacent to forests, etc., especially in
tropical areas, await further studies in order that certain principles and theories
on the edge effects can be formulated.
Web-building spider assemblage is reasonably suitable for assessing their
responses to the edges since they live in a particular habitat; quickly respond to
environmental changes and they are complete predators thus they are highly
satisfactory to test the hypothesis of preference of predators to edge habitats. In
addition, being generalist predators make them directly and indirectly correlated
with many prey species and their species richness and abundance may reflect the
abundance of their prey, mainly arthropods, in the communities. Besides,
because of various life histories (web design, web site, time to prey) among species
so they are appropriate for assessing which type of life history is the most sensitive
to edge effects.
Although rubber plantation-forest edges are commonly found in many
parts of Southern Thailand, studies on their effects are not known. In the present
study, web-building spiders will be surveyed along transects across the rubber
plantation-forest edges. The prediction of the current research is that species
richness of web-building spiders at the edge will be higher than the adjacent
habitats. Since the edges are a transitional zone of neighbouring habitats,
therefore the spider species of both neighbouring habitats will be found at the
edges. The roughly result analyses is different from prediction. Species richness
of spiders at the rubber plantations is highest. Their richness at the edges is
higher than forests. Totally, 1753 individuals of spider were found. They are 917
juveniles and 836 adults. 67 morphospecies of spiders were found from the
adults. Certain species are found interesting to study in depth to find for
underlying mechanism responding to edge effects.
352
Are Salticidae (Araneae) of Indonesia exceptional?
Jerzy Prószyński
Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw,

jerzy.proszynski@wp.pl
Describing fauna of islands we should be prepared for island speciation

pattern, which surprises us by morphological similarity of prolific species within
some genera, and by body diversity of other. In each case species are
recognizable and placed only by internal structure of epigyne, and by palps.
With a ban on collecting in many countries around the world, the study of
Salticidae fauna can further develop using the passions and skills of a new
generation of naturalists, using digital photography. The challenge is to bridge
the gap between validity of deteriorated preserved specimens with beauty of
photographs of alive Salticidae.
353
The dark and salty identity of Pardosa purbeckensis

(Araneae: Lycosidae)
Charlène Puzin1, Frédéric Ysnel1, Alain Canard1 & Julien Pétillon2
1
URU 420, University of Rennes I, France, charlenepuzin@gmail.com,
frederic.ysnel@univ-rennes1.f, alain.canard@univ-rennes1.f
2
University of Antwerp, Belgium, julien.petillon@ua.ac.be
Introduction
Identifying sympatric speciation is currently a hot and growing topic in
modern ecology and taxonomy (Elias et al. 2008). Among spiders, Lycosidae
received a special attention as they include closely related species with similar
foraging strategies and overlapping patterns of microhabitat use (DeVito et al.
2004). As an example, Barthel and von Helversen (1990) considered that
Pardosa wagleri and P. saturatior are two different species because of their
differences in habitat preference, morphology and phenology. Kronestedt (1990)
reached to the same conclusion with the pulverulenta group using ecological,
morphological and sexual behaviour characters. In this study, we compared two
sibling species of Pardosa, P. agrestis and P. purbeckensis. The species status of
P. purbeckensis – P. agrestis still remains uncertain, P. purbeckensis being
frequently considered as a (junior) synonym of P. agrestis due to their high
morphological similarities (e.g. Platnick 2009). We more precisely tested the
hypothesis that both P. agrestis and P. purbeckensis are two different species, as
would reveal a cladistic analysis of different populations. As the two species can
be found in closed stations, we suggest that they have been submitted to a
sympatric speciation, ecological differences can indeed lead to reproductive
barriers between two species of invertebrate in only dozens of generations
(Hendry et al. 2007).
Model species
Pardosa agrestis (Westring, 1861) is a mainland species widely
distributed in Europe, more especially in the south and east, mainly found in
drier banks adjoining the flats, chalk pits or clay soils (Harvey et al. 2002). This
species is undistinguishable from the variety P. purbeckensis (F.O. P.-
Cambridge, 1895) regarding genitalia (Roberts 1995) and thus they are not
considered as two different species, despite a high variety of epigynes in P.
purbeckensis (Roberts 1987). But the last, exclusively found in salt marshes or
polders of the north west of Europe (Heydemann 1970, Harvey et al. 2002,
Pétillon et al. 2005), presents some differences in its habitus (median band of
cephalothorax not dilated, males’ metatarsi and tarsi I with many long hairs) and
its size (both male and female are larger than P. agrestis’ ones) (Locket &
Millidge 1951). Pétillon et al. (submitted) showed a higher salinity tolerance for
P. purbeckensis by exposing the two varieties to a gradient of salinity which is
relevant with their habitats preferences. Furthermore, populations of P. agrestis
354
exhibit two cohorts (Samu et al. 1998) whereas only one cohort has been found
for populations of P. purbeckensis (Puzin et al. submitted).
Methods and expected results

To assess the differences between the two species, several populations
from different parts of Europe were studied to eliminate variations of size due to
unsuitable local conditions (e.g. body condition and physiological tolerance may
vary with latitude: Castañeda et al. (2004). Populations of P. agrestis were
collected in France (2 populations in Brittany; N-W France), Belgium,
Switzerland and Italy (1 population in each country). Populations of P.
purbeckensis were collected in France ((2 populations in Brittany and in
Normandy), Belgium, United Kingdom (Essex) and Netherlands (1 population in
each country). Habitus of females and males of the two varieties were compared
(bands of cephalothorax, colouration,…) and several measurements were done:
body: width and length of cephalothorax, length of tibia I, genitalia: width and
length of epigyne, width and length of male palp and length of median
apophysis.
Further developments are also planned in population genetic (DNA
barcoding technique, i.e. sequencing of mitochondrial DNA and more
specifically the gene coding for the Cytochrome C Oxydase subunit I) as well as
different courtship behaviours (Chiarle et al 2009) and cross breeding
experiments if no pre-zygotic isolation occurs.
References
Barthel J. & von Helversen O. 1990. Pardosa wagleri (Hahn 1822) and Pardosa
saturatior (Simon 1937), a pair of sibling species (Araneae, Lycosidae).
In: Célérier M.L., Heurtault J. & Rollard C. (eds), Proceedings of the 12th
European Colloquium of Arachnology, Paris, 2-4 July 1990, pp. 17-23.
Castañeda L.E., Lardies M.A. & Bozinovic F. 2004. Adaptative latitudinal shifts
in the thermal physiology of terrestrial isopod. Evolutionary Ecology
Research, 6: 579-593.
Chiarle A., Isaia M. & Castellano S. 2009. Courtship behaviour in syntopic and
cryptic species: the case of Pardosa vljimi (Araneae, Lycosidae), a new
record for the Italian fauna. In: Chatzaki M., Stathi I., Spiridopoulou K.
(eds), Abstracts of the 25th European Colloquium of Arachnology,
Alexandroupolis, 16-21 August 2009.
DeVito J., Meik J.M., Gerson M.M. & Formanowicz Jr D.R. 2004.
Physiological tolerances of three sympatric riparian wolf spiders (Araneae:
Lycosidae) correspond with microhabitat distributions. Canadian Journal
of Zoology, 8: 1119-1125.
Elias M., Gompert Z., Jiggins C. & Willmott K. 2008. Mutualistic interactions
drive ecological niche convergence in a diverse butterfly community.
PLoS Biology, 6: 2642-2649.
Harvey P.R., Nellist D.R. & Telfer M.G. 2002. Provisional atlas of British
spiders (Arachnida, Araneae). Biological Records Centre, Huntington 2.
355
Heydemann B. 1970. Ökologische Untersuchungen zum Problem der halophilen

und haloresistenten Spinnen. Bulletin du Muséum d‘Histoire naturelle de
Paris, 41: 226-232.
Hendry A.P., Nosil P. & Rieseberg L.H. 2007. The speed of ecological
speciation. Functional Ecology, 21: 455-464.
Kronestedt T. 1990. Separation of two species standing as Alopecosa aculeata
(Clerck) by morphological, behavioural and ecological characters, with
remarks on related species in the pulverulenta group (Araneae,
Lycosidae). Zoologica Scripta, 19: 203-225.
Locket G. H. & Millidge A.F. 1951. British spiders. Ray Society, London 1, 310 p.
Pétillon J., Ysnel F., Lefeuvre J.-C. & Canard A. 2005. Are salt marsh invasions
by the grass Elymus athericus a threat for two dominant halophilic wolf
spiders? Journal of Arachnology, 33: 236-242.
Pétillon J., Lambeets K., Ract-Madoux B., Vernon P. & Renault D. (Submitted
to Zoology). Relationship between habitat preference and saline stress
tolerance in ground-living spiders.
Platnick N.I. 2009. The world spider catalog, version 9.5. American Museum of
Natural History, http://research.amnh.org/entomology/spiders/catalog/.
Puzin C., Acou A., Bonte D. & Pétillon J. (Submitted to Animal Biology).
Comparison of reproductive traits between two salt-marsh wolf spiders
(Araneae, Lycosidae) under different habitat suitability conditions.
Roberts M.J. 1995. Spiders of Britain and Northern Europe. HarperCollins
Publishers, London, 383 pp.
Roberts M.J. 1987. The Spiders of Great Britain and Ireland (volume 1:
Atypidae - Theridiosomatidae). Harley Books, Colchester, 229 pp.
Samu F., Németh J., Tóth F., Szita É., Kiss B. & Szinetár C. 1998. Are two
cohorts responsible for the bimodal life-history pattern in the wolf spider
Pardosa agrestis in Hungary? In: Selden P.A. (ed.), Proceedings of the
17th European Colloquium of Arachnology, Edinburgh, 14-18 July 1997,
pp. 215-221.
356
Spider (Araneae) composition structure in the chir pine

(Pinus roxburghii) forest habitat, Nanda Devi Biosphere
Reserve, Western Himalayas, India
Shazia Quasin & Virendra Prasad Uniyal*
Wildlife Institute of India, Chandrabani, Dehradun, India

*
Corresponding author: uniyalvp@wii.gov.in
Introduction
Spiders are ubiquitous in nature, being both diverse and abundant in
almost all habitats, exploiting a wide variety of niches in virtually all the earth's
biomes and play vital role in sustaining the ecosystem. In India, information on
spider assemblages in the Himalayan region is scanty in comparison to other
regions of the country because of its tough terrain and climatic conditions. For
the first time diversity of spiders has been investigated in Chir Pine (Pinus
roxburghii) forest in Western Himalayan region of India near Joshimath (1500-
2000 m) area of Chamoli district of Garhwal Himalaya, adjoining to Nanda Devi
Biosphere Reserve (NDBR).

Spider samples were collected during February 2009 to November 2009.
Sampling was carried out in select plots (10m × 10m) in the forest area. In order
to collect the spider samples from all the niches, sampling was carried out
following five semi quantitative methods viz., pitfall trapping, ground hand
collection, aerial hand collection, sweep netting and litter sampling. Pitfall
trapping, ground hand collection and litter sampling methods were employed to
collect wandering spiders while sweep netting and aerial hand collection was
carried out to capture web builders and ambushers. The spider specimens after
collection were preserved in 70% ethanol in glass vials. Identification was then
carried out in the laboratory using a binocular microscope. All voucher
specimens were deposited at the Wildlife Institute of India, Dehradun.
Results
A total of 96 species/morphospecies of spiders belonging to 52 genera and
24 families was reported during the study period. Of all the species collected,
38.5% of the specimens were identified up to species level and 61.5% were
identified up to genus level. The specimens that were immature were identified
only up to genus level. Araneidae and Salticidae were represented by 11.5% of
the 52 genus recorded followed by Theridiidae and Thomisidae (9.6%). Species
richness within families decreased as follows- Araneidae with 16 species (16.7%
of the total species), followed by Linyphiidae 8 species (8.3%); Thomisidae and
Salticidae, 11 species (11.5%) each; Theridiidae with 7 species (7.3%);
Gnaphosidae and Lycosidae with 5 species each (5.2%); Tetragnathidae and
357
Uloboridae with 4 species (4.2%); while the other families were rare and
composed of 26% of the total species with 25 species.
Discussion
The 24 families reported from this area represent 40% of the total families
reported from India. The initial result suggests that Araneidae, Thomisidae and
Salticidae are the most speciose families in this area. The genus Episinus
(Theridiidae) was documented for the first time in India. The area has rich
diversity of spider, further investigation will help to understand spider diversity
of Western Himalayan Ecosystem.
358
Does the whip spider genus Stygophrynus (Amblypygi:

Charontidae) extend its distribution eastward
to the Solomon Islands?
Cahyo Rahmadi1,3, Mark S. Harvey2 & Jun-ichi Kojima3

1
Museum Zoologicum Bogoriense, Indonesia, cahyo.rahmadi@lipi.go.id
2
Department of Terrestrial Zoology, Western Australian Museum, Welshpool, Western
3
Natural History Laboratory, Ibaraki University, Japan, jkrte@mx.ibaraki.ac.jp
Whip spiders are bizarre arachnids characterized by the strong raptorial

pedipalps equipped with sharp spines, flattened body and extremely elongate
antenniform first legs. They are nocturnal and live under stones or within tree
buttresses or they are often found in caves. The family Charontidae is one of the
five families in the Amblypygi and is restricted to the Australasian region. The
family has two genera, Charon Karsch, 1879 and Stygophrynus Kraepelin, 1895.
They are distinguished by the morphology of the pedipalpal patella spination;
two spines about equal in size in Charon and three spines in Stygophrynus and
pedipalpal tarsus; that is, the tarsus is divided in Stygophrynus and undivided in
Charon.
Nine species are currently placed in the genus Stygophrynus (Harvey 2003,
Weygoldt 2002, Rahmadi & Harvey 2008), of which the following seven are in the
nominotypical subgenus and distributed from Myanmar, Thailand, Vietnam, Malay
Peninsula to Java: S. cavernicola (Thorell, 1889); S. cerberus Simon, 1901; S.
berkeleyi Gravely 1915; S. longispina Gravely, 1915; S. dammermani Roewer,
1928; S. brevispina Weygoldt, 2002; and S. sunda Rahmadi & Harvey, 2008. The
remaining two species, S. moultoni Gravely, 1915 and S. forsteri Dunn, 1949, are
placed in the subgenus Neocharon Dunn, 1949. The two Neocharon species show
unusually disjunct distribution pattern; S. moultoni is known only from western
Borneo whereas the record from Sumatra (Quintero 1986, Harvey 2003) has been
clarified by Rahmadi and Harvey (2008) and S. forsteri only from Solomon Islands.
The holotype of Stygophrynus forsteri Dunn, 1949 has an undivided
pedipalpal tarsus, the state found in Charon, suggesting that the specimen is a
juvenile of a Charon species (Rahmadi 2009). Specimens recognized as S. moultoni
have the pedipalpal tibial spination (Gravely 1915; fig. 9) that is characteristic in the
family Charinidae. Recently collected specimens from East Kalimantan which have
similar morphological characters to S. moultoni confirm that the specimens might
prove to be members of family Charinidae since the pedipalpal tarsus lacks a basal
row of setae, a typical character for Charontidae (Quintero 1986, Weygoldt 1996).
In conclusion, whip spiders in the genus Stygophrynus could be restricted to
Myanmar, Thailand, Vietnam, Malay Peninsula, Sumatra and Java. An extensive
study on the genus is required to confirm the record and taxonomic status of the
species in Borneo and Solomon Islands.
359
References
Gravely F.H. 1915. A revision of the Oriental subfamilies of Tarantulidae order
Pedipalpi. Records of the Indian Museum, 11: 433-455.
Harvey M.S. 2003. Catalogue of the smaller arachnid orders of the world:
Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae.
CSIRO Publishing, Melbourne.
Quintero D. Jr. 1986. Revision de la classification de Amblypygidos pulvinados:
creacion de subordenes, una neuva familia y un nuevo genero con tres
nuevas especies (Arachnida: Amblypygi). In: Eberhard W.G., Lubin Y.D.
& Robinson B.C. (eds), Proceedings of the Ninth International Congress
of Arachnology, Panama 1983: 203-212, Smithsonian Institution Press,
Washington D.C.
Rahmadi C. 2009. A review of the family Charontidae (Archnida: Amblypygi)
with notes on the systematic and distribution [Abstract]. 41st Annual
Meeting of the Arachonological Society of Japan. Acta Arachnologica,
58(2): 117.
Rahmadi C. & Harvey M.S. 2008. The first epigean species of Stygophrynus
(Amblypygi: Charontidae) from Java and adjacent Islands with notes on S.
dammermani Roewer, 1928. Raffles Bulletin of Zoology, 56(2): 281-288.
Weygoldt P. 1996. Evolutionary morphology of whip spiders: towards a
phylogenetic system (Chelicerata: Arachnida: Amblypygi). Journal of
Zoological Systematics and Evolution Research, 34: 185-202.
Weygoldt P. 2002. Sperm transfer and spermatophore morphology of the whip
spiders Sarax buxtoni, S. brachydactylus (Charinidae), Charon cf. grayi,
and Stygophrynus brevispina nov. spec. (Charontidea). Zoologischer
Anzeiger, 241: 131-148.
Weygoldt P. 2000. Whip spiders: Their biology, morphology and systematics.
Apollo Books: Stenstrup.
360
Guild structures and biomass of spiders in forests

and agroforestry systems in central Amazonia, Brazil
Florian Raub & Hubert Höfer
Institution: State Museum of Natural History Karlsruhe, Germany,

florian.raub@smnk.de, hubert.hoefer@smnk.de
Data were collected during the German-Brazilian research program SHIFT

(Studies on Human Impact on Forests and Floodplains in the Tropics). The studied
sites are situated in Central Amazonia, 30 km outside the city of Manaus, within the
experimental area of the Brazilian research institute Embrapa - Amazonia Ocidental
(02°53'S, 59°59'W).
Soil/litter fauna has first been collected at two sites of a polyculture
agroforestry system, one site of secondary forest, and one site of primary forest by
repeated sampling between July 1997 and March 1999 with soil cores extracted by a
Berlese apparatus. From June to August 2001 spiders were then sampled with
pitfalls during 4 weeks in 16 replicate plots of another polyculture system, another
natural forest and a rubber tree monoculture. We analysed the data for expected
differences between natural and anthropogenic sites in abundance, body size
distribution, biomass and guild structure of spiders.
Within the first study 3073 spiders were extracted from soil cores: 1203 from
primary forest, 937 from secondary forest and 933 from the polyculture. From the
pitfall study resulted 1001 spiders (686 adults) sorted to 27 families.
Spiders accounted for 3.7 to 4.7% (abundance) resp. 0.6 to 3.4% (biomass) of
the arthropod fauna in soil cores and the respective litter fraction. Web builders were
less abundant than stalking, ambushing and running ground spiders. In pitfall traps
spiders accounted for 4.6-8.7% of all arthropods. In the activity based pitfall
samples, in contrast to the abundance based soil core sampling, slightly more web
builders than hunters or stalkers were captured. An analysis of the size distribution
of adult spiders in pitfall samples showed a pattern distinctly differing from a normal
distribution in the primary forest, with small spiders much most frequent, whereas in
the agroforestry systems the size distributions were equally normal.
The biomass of spiders was distinctly lower in the disturbed areas than in the
reference sites (primary forest, both in quadrat-based samples and pitfall samples. In
the Berlese samples of the secondary forest the biomass reached 62% and in the two
polyculture plots 5% and 27% of the biomass observed in the primary forest. The
differences in biomass between primary forest and the anthropogenic sites were
even greater in pitfall data: biomass in the polyculture reached 36% and in the
monoculture only 5% of the primary forest.
In the Berlese study the spider diversity was higher in the natural forest, but
not in the pitfall study, where the highest diversity could be measured in the
polyculture. Guild structure differences (due to the lack of some guilds and a
dominance of others) occurred in the disturbed sites and abundance as well as
biomass decreased by anthropogenic alteration of the natural habitat.
361
Scent gland chemistry in harvestmen (Opiliones)

Günther Raspotnig1, Petra Föttinger1, Miriam Schaider1
& Ivo Karaman2
1
Institute of Zoology, Karl-Franzens University Graz, Austria,
guenther.raspotnig@uni-graz.at, miriamschaider@hotmail.com
2
Department of Biology and Ecology, University Novi Sad, Serbia
Large, prosomal sac-like “scent glands” (syn. defensive glands) are an

important synapomorphic character of all Opiliones. Following disturbance,
many species discharge distinctly scented, repellent secretions from scent
gland orifices (ozopores) which typically are situated near coxae I
(Palpatores), near coxae II (Laniatores) or atop latero-dorsally protruding
prosomal tubercles (Cyphophthalmi).
The chemistry of opilionid scent gland secretions has been investigated
since the 1950s, and meanwhile, published chemical data are available for
about 40 species of Laniatores, for 12 species of Palpatores and for 3 species
of Cyphophthalmi. Scent gland secretions are a source of unique natural
products. Secretions may contain only one component (e.g. some phalangodid
Laniatores), but usually constitute more complex mixtures, even comprising up
to more than 20 components such as in some Cyphophthalmi (Raspotnig et al.
2005). Moreover, secretions exhibit a characteristic group- and even species-
specific chemical composition, clearly suggesting their potential use as novel
phylogenetically important characters. In the present contribution, the current
state of knowledge of opilionid scent gland chemistry is reviewed, with focus
on recent developments and some recently accumulated data.
Results from early opilionid scent gland research were strongly biased
and restricted to some gonyleptoid laniatoreans (Eisner et al. 1977, 1994) and
to a small group of Palpatores, namely to some North American sclerosomatid
Eupnoi (Ekpa et al. 1985). These data suggested a “chemical dichotomy”
between phenol- and benzoquinone-rich secretions of Laniatores and acyclic
compounds-dominated secretions in Palpatores. Indications for a much more
complex situation, however, have arisen since the late 1970s, beginning with
the finding of distinct chemical equipments (naphthoquinones) in secretions of
phalangiid Eupnoi (Palpatores) (Wiemer et al. 1978). Also for laniatoreans
outside the Gonyleptoidea (e.g., in a representative of Insidiatores) a
completely distinct set of compounds was detected, including bornyl esters,
terpenes, and nitrogen containing compounds (Ekpa et al. 1984).
Not until recently, some large gaps in the knowledge of opilionid gland
chemistry could be closed: In pioneering studies on secretions of
Cyphophthalmi, complex patterns of methyl ketones and naphthoquinones
(including unique chloro-naphthoquinones) were found (Raspotnig et al. 2005,
Jones et al. 2009), and naphthoquinones and unusual anthraquinones have just
recently been reported for a first representative of Dyspnoi (Raspotnig et al.
362
2010). Nitrogen-containing compounds may be very characteristic for the

Insidiatores (or a group of these) and, e.g., dominate the secretions of
Cladonychiidae (Raspotnig, unpublished). A first study on Phalangodidae is
meanwhile also available (Shear et al. 2010), demonstrating that certain
phenols (but no benzoquinones) had already been developed in (presumably)
basal Grassatores. Benzoquinones, in their turn, may have evolved in more-
derivative Laniatores since they seem to be restricted to the Gonyleptoidea
(Föttinger et al. 2010).
Despite these recent advances mentioned above, the overall-picture of
scent gland chemistry in harvestmen is still highly fragmentary. Chemical data
that possibly contribute to solve important questions on the gross phylogeny of
Opiliones are still missing: so far, no chemical synapomorphies have been
found to justify a sister-group relationship between Eupnoi and Dsypnoi, or (as
more recently proposed) between Dypnoi and Laniatores. By contrast, as
outlined above, certain compounds or classes of components seem to
characterize groups of Opiliones on superfamily and family levels.
Chemosystematics using scent gland secretion profiles may also be applied to
the problem of discrimination of cryptic or morphologically very similar
opilionid species as just successfully demonstrated for a number Balkan
Sironidae (Raspotnig & Karaman, in preparation).
References
Eisner T., Jones T.H, Hicks K., Silberglied R.E. & Meinwald J. 1977. Quinones
and phenols in the defensive secretions of neotropical opilionids. Journal
of Chemical Ecology, 3: 321-329.
Eisner T., Rossini C., González A. & Eisner M. 2004. Chemical defence of an
opilionid (Acanthopachylus aculeatus). Journal of Experimental Biology,
207: 1313-1321.
Ekpa O., Wheeler J.W., Cokendolpher J.C. & Duffield R.M. 1984. N,N-
Dimethyl-ß-phenylethylamine and bornyl esters from the harvestman
Sclerobunus robustus (Arachnida: Opiliones). Tetrahedron Letters, 25:
1315-1318.
Ekpa O., Wheeler J.W., Cokendolpher J.C. & Duffield R.M. 1985. Ketones and
alcohols in the defensive secretion of Leiobunum townsendi Weed and a
review of the known exocrine secretions of Palpatores (Arachnida:
Opiliones). Comparative Biochemistry and Physiology, 81B: 555-557.
Föttinger P., Acosta L.E., Leis H.J. & Raspotnig G. 2010. Benzoquinone-rich
exudates from the harvestman, Pachylus paessleri (Opliliones,
Gonyleptidae, Pachylinae) with comments on the phylogenetic
significance of secretion compounds in the Laniatores. Journal of
Arachnology, under review.
Jones T.H., Shear W.A. & Giribet G. 2009. The chemical defences of a stylocellid
(Arachnida, Opiliones, Stylocellidae) from Sulawesi with comparisons to
other Cyphophthalmi. Journal of Arachnology, 37: 147-150.
363
Raspotnig G., Fauler G., Leis M. & Leis H.-J. 2005. Chemical profiles of scent
glands secretions in the cyphophthalmid opilionid harvestmen, Siro
duricorius and S. exilis. Journal of Chemical Ecology, 31: 1353-1368.
Raspotnig G., Leutgeb V., Schaider M. & Komposch Ch. 2010.
Naphthoquinones and anthraquinones from scent glands of a dyspnoid
harvestman, Paranemastoma quadripunctatum. Journal of Chemical
Ecology, 36: 158-162.
Shear W.A., Jones T.H. & Snyder A.J. 2010. Chemical defence of phalangodid
harvestmen: Bishopella laciniosa (Crosby & Bishop) and Texella
bifurcata (Briggs) produce 2-methyl-5-ethylphenol (Opiliones:
Grassatores: Phalangodidae). Bulletin of the British Arachnological
Society, 15: 27-28.
364
A review of the Mygalomorphae: biology, morphology

and systematics
Robert J. Raven
Queensland Museum, South Brisbane, Qld, Australia, Robert.Raven@qm.qld.gov.au
Mygalomorph spiders, with their plesiomorphic morphology, rest uneasily

as sister group to the Araneomorphae. Their phylogeny, though much contented,
resists change from earlier hypotheses. Their species concepts are conflated by
uninformative and highly homoplasious characters and significant bilateral
assymetry but challeneged severely by the resolution provided by venom
differences at the population level.
In habitat, they range from areas snow covered in winter, through deserts
and down to the intertidal zone. In river beds, they endure weeks of inundation
during extensive and often flash flooding. In behaviour, most are conservative
burrow dwellers. A few make extensive aerial webs. Many build trapdoors
which usually work to their advantage but are sometimes exploited by birds.
Although many are the prey of mammals, some are also mammal predators.
Attacked by a diversity of organisms, often incongruent in the slowness of the
attack, they are the longest lived of spiders and some persist in suburban areas.
Challenges abound as putative neotony conflates the cladistic use of their
morphology. As with the repeated loss of the cribellum in araneomorphs, some
closely related barychelids show a tendency to lose the second pair of spinnerets.
Molecular studies are needed to better illuminate this dilemmas but so far
acceptable overarching proposals have not been forthcoming.
Research opportunities are many as still the value of their silk in
phylogeny remains untested. Controversies over the present of sil glands in
theraphosid scopula hairs rage. And their conservation status begs careful
attention to their biologies.
Theraphosids remain a dominant focus both in diversity and phylogeny
and a review of South American genera is desparately needed, especially with
the tarantulas ascending further as preferred pets. Uninformative monotypic
genera abound. Preliminary studies on the Australian fauna indicate that
ontogenetic changes in the growing animal contradict the expected development
sequence (Haeckel’s Law).
In contrast, many other mygalomorph groups remain seriously unknown.
No doubt that is often driven by unstable political issues in understudied regions,
like parts of Africa. However, the decline of taxonomy worsens as the quality of
submitted papers belies a lack of understanding of critical processes in the
science.
Biogeographically, they seem simple and informative: animals with poor
vagility, sedentary habits, isolated on continents by continental drift, restricted
by glaciation, orogenic activity and habitat fragmentation through progressive
drying. Many groups are Gondwanan in distribution and origin, or at least
365
appear so. In New Caledonia, the mygalomorph fauna challenges all

preconceptions. In south-western Australia, several species persist in close
allopatry with no evident barrier.
Biologically, mygalomorphs are surrogate “super-mammals”: long-lived
and univoltine but with high local endemicity. As such, they are among the frst
spiders to be listed as threatened but often their low incidence, cryptic burrows
and burrowing behaviour severely limits useful knowledge of their biology.
Mygalomorphs includes the most venomous spiders in the Australian
Funnel-webs (Hexathelidae) responsible for 14 known deaths. However, their
venom, of considerable interest to pharmaceutical companies is neutralised
quickly by dogs, cats, mice and guinea-pigs and the spiders themselves are the
prey of large lizards. In contrast, the unrelated Mouse Spiders (Actinopodidae)
with massive chelicerae and diagonally acting fans that “lock” on the prey,
seemed incongrously harmless until it was understood that the spider rarely
engages its reduced venom glands.
366
Social evolution in huntsman spiders: behavioural,

ecological, and physiological strategies
Linda S. Rayor
Department of Entomology, Cornell University, Ithaca, NY, USA; Research School of

Biology, Australian National University, Canberra, Australia, LSR1@cornell.edu
Why do some animals live in social groups while others are solitary?
Throughout Animalia are examples of closely related taxa in which the majority
of species live essentially solitary lives while a few species have evolved to live
in cooperative social groups. What factors make cooperation beneficial although
closely related species succeed without those benefits? How do physiological
and ecological constraints interact so that group-living becomes more beneficial
than a solitary life style? Why do young adults remain with their parents when
they could have better opportunities to breed if they left home? These questions
are at the essence of understanding the costs and benefits of group-living in
animals. Evolution of sociality must involve adaptive benefits to offset the costs
of living closely with competitors, especially among potentially cannibalistic
predators such as spiders. These adaptations may involve behavioural,
ecological, and physiological strategies for living in groups.
In this presentation, I will discuss some of these differences in
behavioural, ecological, and physiological strategies influencing social evolution
in endemic social and solitary huntsman spiders from Australia. The Australian
endemic subfamily Deleninae is monophyletic with relatively little
morphological diversity (Rheims 2007) containing ten genera with ~99 species
(Platnick 2010, Hirst per. comm.). These endemic delenine spiders share a
number of key traits that make them particularly valuable for comparative
studies: The social and solitary species overlap geographically, in body size,
general biology, and habitat preferences; many live in retreats under bark, often
in neighbouring trees. The social and solitary species appear to be very similar in
most superficial traits except that two huntsman species from different genera
live in social groups for most of their lives. To aide in determining whether there
are phylogenetically independent characters associated with social evolution in
these species, there are solitary congeners for comparison. Here I will present
data on comparative patterns of group-living, prey sharing, dispersion, life-
history, and metabolic rate in 15 species from 7 genera of delenine huntsman
spiders. I will discuss some of the factors that effectively reduce the costs of
group-living for the social species, which are not seen in the solitary huntsman
species.
367
Insect form vision as a potential shaping force

of spider web decoration design
Cheng Ren-Chung
Tunghai University, Taiwan, bolasargiope@gmail.com
Properties of prey sensory systems are important factors shaping the

design of signals generated by organisms exploiting them. In this study it was
assessed how prey sensory preference affected the exploiter signal design, by
investigating the evolutionary relationship and relative attractiveness of linear
and cruciate form web decorations built by Argiope spiders. Since insects have
innate preference for bilaterally symmetric patterns, we hypothesized that
cruciate form decorations were evolved from liner form due to their higher
visual attractiveness to insects. We first reconstructed a molecular phylogeny of
the genus Argiope and results of ancestral character state reconstruction showed
that the linear form was ancestral and the cruciate form derived. We then
performed field experiments in which the number and orientation of decoration
bands were manipulated. Decoration bands arranged in a cruciate from was
significantly more attractive to insects than those arranged in a linear form.
Moreover, dummy decoration bands arranged in a cruciate form attracted
significantly more insects than those arranged in a vertical/horizontal form. Such
results suggest that pollinator insects’ innate preference for bilateral or radial
symmetric patterns might be one of the driving forces shaping the arrangement
pattern of spider web decorations.
368
Untangling the ambiguous: taxonomy and biogeography

of Western Mediterranean Lycosa (Araneae: Lycosidae)
using molecular and morphological data
Carles Ribera1, Enric Planas1 & Carmen Fernández-Montraveta2

1
Institut de Recerca de la Biodiversitat (IRBio) and Dept. Biologia Animal. Universitat
de Barcelona, Spain, enplanas@gmail.com, cribera@ub.edu
2
Dept. Psicología Biológica y de la Salud, Universidad Autónoma de Madrid, Spain,
carmen.montraveta@uam.es
The genus Lycosa Latreille, 1804 is one of the most diverse genera of
spiders. Its worldwide distribution and its wandering life style place some
Lycosa species among the best-known and widely studied spider species. Lycosa
groups 240 species, 17 of which are recorded from Western Mediterranean, and
some of them are among the largest representatives of the spider fauna of this
area, a fact that has also contributed to the extensive number of studies
concerning their systematics, behaviour and ecology. Despite of the numerous
studies, the taxonomical status of the species of Lycosa from Western
Mediterranean remains controversial.
The fact that original descriptions are very old (18th, 19th and the first half
of the 20th century) and the majority of the type material has been lost, impedes
or seriously obstructs the identification to the specific level. Moreover, the
taxonomic status of most of these species is confused, mainly after the Roewer’s
revision (1955), which transferred many “true” Lycosa species from western
Mediterranean to the genus Allocosa Banks, 1900. There are only few species
redescriptions and no recent revision at the genus level has been carried out in
this area.
In an attempt to clarify the situation we undertook a series of collecting
trips in Western Mediterranean with a main objective: to make the redescription
of the Lycosa species from this area and to nominate the neotypes of the species
that have been lost. In this contribution we present the preliminary results on
taxonomy and phylogeny of these species.
Phylogenetic analysis using information from mitochondrial and
molecular genes allows us to examine the phylogenetic relationships and
diversification patterns of the genus in the Western Mediterranean. Results show
a basal group of species (maghrebi group) located in the South of Morocco and
Tunisia, although, some of them also colonize the northern areas near the
Mediterranean and some islands (Corsica and Sardinia). In addition there are
two independent evolutionary lineages with European representatives: the L.
tarantula Group and the L. subhirsuta Group.
The tarantula group contains the Tunisian L. bedeli and the Europeans L.
tarantula and L. hispanica. The phylogenetic analysis suggests a single
colonization event from Tunisia to Europe through the Sicilian island. Posterior
glacial periods separate the Iberian and Italic representatives.
369
Within the subhirsuta group appear L. subhirsuta (spread in Balearic

Island, Tunisia, Sardinia and Corsica) and L. munieri (from Iberian Peninsula,
Balearic Islands, Morocco and Tunisia).
370
Spider composition in less diverse sites in Brazilian

Amazonia and its importance for conservation
Janael Ricetti1 & Alexandre Bonaldo2
1
Departamento de Zoologia, Universidade Federal do Paraná, Brazil, jricetti@gmail.com
2
Laboratório de Aracnologia, Museu Paraense Emílio Goeldi, Belém, Pará, Brazil
Introduction
The white sand vegetation, also called “campina”, is known by its low plant
diversity and high endemism rates (Anderson 1981, Frasier 2008), when compared
with other Amazonian ecosystems. This vegetation grows over sandy soil and
occurs in patches throughout Amazonia, as a distinctive vegetal formation
presenting from open fields to low-canopy forests (Anderson 1981).
The Southeastern Amazonia, presenting extensive white sand areas, is
nowadays endangered mainly due to the irregular land usage for monoculture,
cattle grazing and inadequate wood extraction practices.
Several questions about species distribution in Amazonia still remain
unanswered and some megadiverse animal groups, as spiders, haven’t received
well-deserved attention. The lack of faunistic studies in this particular Amazonian
landscape leaded us to survey the spider fauna of selected sites at Serra do
Cachimbo region. The main objective of this study was to analyze how the species
composition associated with white sand vegetation ecosystem contributes to the
Amazonian spider diversity.

Serra do Cachimbo is located in Southeastern Brazilian Amazonia,
presenting a peculiar vegetation mosaic of Rain Forests (FO), savannah-like
vegetation or Cerrado (CE) and white sand vegetation (WS), with scattered
riparian forests (RP) along watercourses. The sampling was carried out in these
four vegetations types, at the “Campo de Provas Brigadeiro Velloso”, Novo
Progresso, State of Pará. Two expeditions were made, during both dry (August
and September 2003) and wet (March and April, 2004) seasons. The sampling
effort, represented by 60 samples per site, was obtained using beating tray and
sweeping net; nocturnal manual search and Winkler extractors. The similarity
among the spider assemblages in each vegetation type was calculated with
Morisita coefficient. An Analysis of Similarity (ANOSIM) were performed, using
the Bray-Curtis index (Clarke 1993), to verify the significant differences between
the spider fauna of the four vegetations types. Samples similarity were represented
in a two axis ordination generated by a non-metric multidimensional scaling
(NMDS) (Bray-Curtis index) (Legendre & Legendre 1998). Analysis were
performed with PAST v.1.94b (Hammer et al. 2009).

The WS spider fauna is different from the other vegetations in Serra do
Cachimbo. WS had the lowest species richness per sample (Ricetti & Boanldo
2008). This may be due to the structural homogeneity and low productivity of
371
this vegetation type (Mcnett & Rypstra 2000, Ysnel & Canard 2000) compared
with the Rain Forest. WS shared less then 15% with the other vegetations, while
CE, FO and RP shared more than 70% between them.
More than a half (55%) of WS species occurred only in this vegetation
type, with average of 10% more exclusive species than the other vegetations.
The spider species composition was significantly different between the areas
(ANOSIM: r=0,17; p=0,0001). The ordination showed a visible proximity of
WS samples, in contrast with the other vegetations samples, which were less
aggregated. Some recent data about avian community also demonstrate that
white sand ecosystems may harbour several species differentiated of more
complex Amazonian ecosystems. Borges (2004) discussed the association of a
distinct bird assemblage with white sand vegetation of western Amazonia, as
well Polleto & Aleixo (2005) stressed the importance of intensifying faunistic
surveys in white sand patches, aiming to add more data on endemism and
biogeography. Even with the need of more complete observations, these results
suggests that, as well for plants and birds, spider species endemism may occur in
the white sand ecosystems, highlighting the conservation importance of this
environment.
References
Anderson A. 1981. White-sand vegetation of Brasilian Amazonia. Biotropica,
13: 199-210.
Borges S.H. 2004. Species poor but distinct: bird assemblages in white sand
vegetation in Jaú National Park, Amazonian Brazil. Ibis, 146: 114-124.
Clarke K.R. 1993. Non-parametric multivariate analysis of changes in
community structure. Australian Journal of Ecology, 18(1):117-143.
Frasier C.L., Albert V.A. & Struwe L. 2008. Amazonian lowland, white sand
areas as ancestral regions for South American biodiversity: biogeographic
and phylogenetic patterns in Potalia (Angiospermae: Gentianaceae).
Organisms Diversity and Evolution, 8: 144-157.
Hammer O, Harper D.T.A. & Ryan P.D. 2009. PAST: Palaeontological Statistics
software package for and data analysis. Paleontologia Eletronica, 4(1): 9.
Legendre P. & Legendre L. 1998. Numerical Ecology. 2. ed. Elsevier,
Amsterdam.
Macnett B.J., Rypstra A.L. 2000. Habitat selection in a large orb-weaving
spider: vegetational complexity determines site selection and distribution.
Ecological Entomology, 25: 423-432.
Poletto F. & Aleixo A. 2005. Biogeographic implications of new avian records
from a patch of white-sand forest in southwestern Brazilian Amazonia.
Revista Brasileira de Zoologia [online], 22(4): 1196-1200.
Ricetti J. & Bonaldo A.B. 2008. Diversidade e estimativas de riqueza de aranhas
em quatro fitofisionomias na Serra do Cachimbo, Pará, Brasil. Iheringia,
Sér. Zool., 98: 88-99.
Ysnel F. & Canard A. 2000. Spider biodiversity in connection with the
vegetation structure and the foliage orientation of hedges. Journal of
372
Making the most of limited locality data: predicting the

distribution of jumping spider (Salticidae: Araneae)
faunas in Australia
Barry J. Richardson
CSIRO Division of Entomology, Canberra, ACT, Australia, barry.richardson@csiro.au
In much of the world the information available for making management

decisions is very limited for groups like the jumping spiders. In Australia for
example, jumping spiders are a diverse component of the Australian fauna, with
over 350 species described and possibly a further 1000 species present
(Richardson and Zabka, 2008). A recent study in the Murchison Region (Harvey
et al. 2000) found 37 genera of which 18 were undescribed and a brief survey in
south western Queensland (Richardson 2009) found 15 species of which 13 were
undescribed. Usually the only distribution information available is point data
associated with taxonomic revisions or museum collections. Innovative
approaches need to be developed and used for making the best use of these data.
Linnaean hierarchies, unlike cladistic trees, are intended to represent a
natural biological hierarchy of functional entities at each level. For example, a
genus can be considered “a species or group of species of common ancestry that
occupies an ecological situation, or adaptive zone, that is different from that
occupied by the species of another genus” (Wood and Collard 1999). Such
genera provide entities suitable for analysis using ecological attributes, for
example the estimation of geographical distribution based on bioclimatic profiles
using modelling programs. Such generic distributions can be of great use in
countries such as Australia where most species are undescribed. In taxonomic
work they allow relevant geographical areas to be identified when planning
fieldwork (i.e. areas predicted to support members of the genus even though no
specimens known) and for management decisions as to which habitats or areas
need to be surveyed or given priority protection.
Such an approach was explored using a data set of point records for
Australian jumping spider genera. The data for 4500 locality records was stored
in BioLink and a map of the predicted distribution of each of 51 genera was
generated on the basis of its bioclimatic profile, using the BIOCLIM package as
compiled in BioLink. The predictions were tested by comparing them with
independent data sets for four landscape regions in Australia (Bridgewater
1987). As well, a table was prepared allowing a list of the likely genera in any
given area to be obtained before field work is undertaken or management
decisions made.
Test 1: Eyre region in South Australia. Seven genera were predicted for
Eyre though no actual locality records in Eyre were available. Specimens of six
of these genera were found from there in the collection of the South Australian
Museum.
373
Test 2: South Western Australia. Similarly, of 24 genera predicted for SW

Western Australia for which no specimens from the area were used in making
the prediction, specimens of twelve were found from the region in the
collections of the Western Australian Museum. The remainder are largely genera
represented in eastern Australia as outliers of cosmopolitan or oriental genera
that presumably have been unable to make their way across the much drier
Nullabor region.
Test 3: Murchison Region, WA. Of 21 predicted genera, 19 were collected
during a detailed biodiversity survey carried out in the Region.
Test 4: North-eastern Simpson Desert. Seven of the eleven genera
predicted to be present were found, while one genus of the eight predicted to be
present in the general region but not on the study area was also found. The four
genera predicted but not collected were all likely to be ground dwellers and their
apparent absence, given that heavy flooding that had recently occurred, was not
surprising.
In summary, the approach gave surprisingly good predictions of the
genera likely to be present in areas where no data were available for many
hundred of miles.
Several limitations need to be noted however.
Firstly, unknown or very poorly known genera cannot be considered. In
the Murchison study, the survey also collected examples of a further 18
undescribed genera. In the Simpson study a genus known from only several
specimens elsewhere in Queensland was found, though it had not been modelled
due to lack of data.
Secondly, areas that lie beyond the range of bioclimatic zones included in
the training set cannot give proper predictions. Very dry landscape regions such
as Simpson or Cooper for example, have no salticid genera predicted. Such a
situation is difficult to imagine, especially as large parts of these areas support
vegetation. Elsewhere some genera such as Grayenula, Afraflacilla or Pellenes
are known to occur in such habitats. Few genera were also predicted for the
Australian Alps and Barrington Tops, areas that regularly receive snow.
Presumably the climate profile for these areas is outside predicted ranges based
on specimens collected at lower altitudes.
The predicted distribution of a genus can also be compared with the
combined predicted distributions of the known species in a genus to identify
geographical areas that may contain new species. This will be demonstrated for
the genus Prostheclina.
These results highlight the shortcomings of past fieldwork in Australia,
which has concentrated on the higher rainfall areas. Such systematic bias cannot
be offset in an approach such as this and it seems likely for example that inland
Australia will support a large, highly endemic, but presently unknown fauna
adapted to the region.
374
Genes, trees and tiny spiders: progress and prospects

in micropholcommatid systematics
Michael G. Rix* & Mark S. Harvey
Department of Terrestrial Zoology, Western Australian Museum, Welshpool DC, Perth, WA,
Australia, michael.rix@museum.wa.gov.au, mark.harvey@museum.wa.gov.au
*
Presenting author
The Micropholcommatidae are a family of tiny, distinctive araneoid

spiders, known from southern-temperate habitats throughout Australasia and
Chile. The greatest abundance of individuals and the largest diversity of taxa
occur in the cool-temperate rainforests of south-eastern Australia and New
Zealand, where micropholcommatid spiders can be very common within moss
and leaf litter microhabitats. Although poorly studied biologically and largely
neglected taxonomically, the Micropholcommatidae are a diverse lineage, with a
significant, and previously unrecognised, generic diversity.
Considerable recent progress has been made in micropholcommatid
systematics, at multiple taxonomic levels. Molecular and morphological data
have been used to test the phylogenetic position and phylogeny of the
micropholcommatid taxa, and now inform a comprehensive generic
classification of the family. Three subfamilies are proposed, and 18 genera are
recognised from Australia, New Zealand, Papua New Guinea, New Caledonia
and Chile. A Gondwanan biogeography is suggested for the
Micropholcommatidae, and explicitly tested in the new tribe Textricellini.
Textricellin biogeography is further explored in the south-west of Western
Australia, where phylogeographic research has revealed a rich and locally
endemic fauna. Most importantly, a taxonomic framework and a phylogenetic
foundation now exist for the Micropholcommatidae: a template by which new
species can be described and existing species can be identified, and a valuable
dataset for exploring phylogenetic hypotheses.
375
Cladistic analysis of the South American genus

Polybetes Simon (Araneae: Sparassidae),
with notes on the subfamily Sparassinae Bertkau
Cristina Anne Rheims

cris.rheims@butantan.gov.br
The genus Polybetes was originally proposed by Simon (1897a) to include

the type species, Olios martius Nicolet, from Chile, and Voconia maculata
Keyserling [=P. pythagoricus (Holmberg)], from Argentina. It was characterized
by having anterior and posterior eye rows slightly procurved, rarely straight, and
only two promarginal teeth on the chelicerae. In the same year, Simon (1897b)
included Polybetes germaini, from Paraguay and P. obnutptus Simon, from
Bolivia, and Polybetes delfini Simon, from Chile. Mello-Leitão described an
additional nine species: Polybetes vittatus and P. semivittatus from Buenos
Aires, Argentina (Mello-Leitão 1938); P. humeratus and P. pallidus from Santa
Fé, Argentina (Mello-Leitão 1941a); P. tucumanus from Tucumán, Argentina
(Mello-Leitão 1941b); P. proximus from Paraíba, Brazil (Mello-Leitão 1943a);
P. rubrosignatus from Rio Grande do Sul, Brazil (Mello-Leitão 1943b); P.
punctulatus, from Buenos Aires, Argentina (Mello-Leitão 1944); and P.
cancroides from Corrientes, Argentina (Mello-Leitão 1945), thus increasing the
genus composition to 14 species.
The genus was revised by Gerschman & Schiapelli (1965) who considered
it to be a senior synonym of Streptaedoae Järvi, with three species, and
Leptosparassus Järvi, with three species. This revision resulted in the synonymy
of 11 species restricting the genus composition to the thirteen species considered
valid to date: Polybetes delfini Simon, P. germaini Simon, P. martius (Nicolet),
P. obnuptus Simon, P. pallidus Mello-Leitão, P. parvus (Järvi), P. proximus
Mello-Leitão, P. punctulatus Mello-Leitão, P. pythagoricus (Holmberg), P.
quadrifoveatus (Järvi), P. rapidus (Keyserling), P. rubrosignatus Mello-Leitão
and P. trifoveatus (Järvi) (Platnick 2010).
Little is known on the relationships between Polybetes and the remaining
sparassid genera. At the time of it’s description, Simon (1897a) included it in
Deleneae, together with Cebrennus Simon, Cerbalus Simon, Cercetius Simon,
Damastes Simon, Delena Walckenaer, Olios Walckenaer, Origes Simon,
Paenula Simon, Pediana Simon Eusparassus Simon, Holconia Thorell, Isopeda
L. Koch, Megaloremmius Simon, Nisueta Simon, Nonianus Simon, Remmius
Simon, Rhitymna Simon, Sarotesius Pocock, Spatala Simon, Typostola Simon
and Zachria L. Koch. A few years later, Järvi (1912) removed Polybetes from
Deleneae and proposed Polybeteae based solely on genital characters, to include
Polybetes, Origes, Paenula and the new genera Leptosparassus Järvi and
Streptaedoea Järvi. This classification was not accepted by Petrunkevitch (1928)
who presented a classification with similar divisions to those proposed by Simon
376
(1897a). Järvi’s Polybetinae was placed together with Delena, Eusparassus,

Isopeda, Pediana, Pseudomicrommata Järvi, Rhytimna, Typostola and Zachria in
Eusparassinae. The most recent catalog (Roewer, 1954) lists Polybetes in
Sparassinae together with Adcatomus Karsch, Cebrennus, Cerbalus, Cercetius,
Damastes, Macrinus Simon, Megaloremmius, Micrommata Latreille, Nisueta,
Nonianus, Olios, Origes, Paenula, Remmius, Sarotesius and Vindullus Simon. No
synapomorphic characters have been found to support the monophyly of the
subfamily and nothing is known on the relationships between it’s genera and the
remaining sparassids. Mostly, the species present only two promarginal teeth in
the chelicerae and a straight or slightly procurved eye row (Jäger 1998).
Here I present a taxonomic revision of the genus Polybetes and, based on
the examination of the type specimens, propose eight synonymies: Olios vitiosus
Vellard 1924 with Polybetes germaini Simon 1897; Polybetes pallidus Mello-
Leitão 1941 with Polybetes quadrifoveatus Järvi 1914; Polybetes punctulatus
Mello-Leitão 1944 with Olios fasciatus (Keyserling 1880); Polybetes delfini
Simon 1904 with Olios bombilius (F.O.P.-Cambridge 1899); Polybetes obnuptus
Simon 1897 with Polybetes pythagoricus (Holmberg, 1875); Polybetes proximus
Mello-Leitão 1943, Olios fuscovariatus Mello-Leitão 1943 and Stasina koluene
Mello-Leitão 1949 with Olios niveomaculatus Mello-Leitão 1941. In addition, I
present a cladistic analysis of the genus, including representatives of most
Sparassinae genera and other sparassid subfamilies. The data matrix comprised 35
terminal taxa scored for 48 morphological characters. The analysis yielded three
most parsimonious tree with 131 steps and shows that Polybetes as currently
defined, does not form a monophyletic unit. Thus, the genus is redefined to
include only seven species (Polybetes martius, P. pythagoricus, P. quadrifoveatus,
P. bombilius, P. fasciatus, P. parvus and P. trifoveatus (Järvi)) based on the
presence of a modified tegular grove that acts as a conductor for the embolus of
the male palp and by the widening of the first winding of the female copulatory
ducts. Streptaedoea is revalidated to include P. rapidus, P. germaini and Olios
hyeroglyphicus Mello-Leitão based on the double helix type embolus on the male
palp and on the anterior part of the copulatory duct looped around the copulatory
openings of the female epigyne. Both genera arise as sister groups within a large
clade that includes all Sparassinae genera, except Adcatomus Karsch.
Two new genera are proposed, one to include O. niveomaculatus and Olios
subadultus Mello-Leitão, supported by the presence of two retrolateral spines on
the male palpal tibia and bell shaped pockets on the median septum of the female
epigynum, and one to include P. rubrosignatus, based on the presence of three
prolateral and four retrolateral spines on metatarsi IV and the subdistal origin of
the RTA on the male palpal tibia. Both genera arise within a clade formed by
genera currently included in Heteropodinae Thorell. Although this clearly shows
that their species are not congeneric with either Polybetes nor Streptaedoea and
are not related to the remaining Sparassinae, their placement cannot be confirmed
without a much broader analysis of the family as a whole.
377
References
Gerschman de P., Schiapelli B.S. & Schiapelli R.D. 1965. El género Polybetes
Simon, 1897, en la Argentina (Araneae-Sparassidae). Revista do Museo
Argentino de Ciencias Naturais Bernardino Rivadavia (Ent.), 1: 313-339.
Jäger P. 1998. First results of a taxonomic revision of the SE Asian Sparassidae
(Araneae). In: Selden P.A. (ed.), Proceedings of the 17th European
Colloquium of Arachnology, Edinburgh, pp. 53-59.
Järvi T.H. 1912. Das Vaginalsystem der Sparassiden. I. Allgemeiner Teil.
Annales Academiae Scientiarus Fennicae, 4: 1-131.
Mello-Leitão C.F. de 1938. Algunas arañas nuevas de la Argentina. Revista del
Museo de La Plata (N.S), 1: 89-118.
Mello-Leitão C.F. de 1944. Arañas de la provincia de Buenos Aires. Revista del
Museo de La Plata (N.S., Zool.), 3: 311-393.
Mello-Leitão C.F. de 1941a. Las arañas de la provincia de Santa Fe colectadas
por el Profesor Birabén. Revista del Museo de La Plata (N.S., Zool.), 2:
199-225.
Mello-Leitão C.F. de 1941b. Las arañas de Córdoba, La Rioja, Catamarca,
Tucumán, Salta y Jujuy colectadas por los Profesores Birabén. Revista del
Museo de La Plata (N.S., Zool.), 2: 99-198.
Mello-Leitão C.F. de 1943a. Araneologica varia brasiliana. Anais da Academia
Brasileira de Ciências, 15: 255-265.
Mello-Leitão C.F. de 1943b. Catálogo das aranhas do Rio Grande do Sul.
Archos del Museo Nacional Rio de Janeiro, 37:147-245.
Mello-Leitão C.F. de 1945. Arañas de Misiones, Corrientes y Entre Ríos. Revta.
Revista del Museo de La Plata (N.S., Zool.), 4: 213-302.
Petrunkevitch A. 1928. Systema Aranearum. Transactions of the Connecticut
Academy of Arts and Sciences, 29: 1-270.
Platnick N.I. 2010. The world spider catalog, version 9.5. AMNH,
Roewer C.F. 1954. Katalog der Araneae von 1758 bis 1940. Bruxelles, 1: 1-1040.
Simon E. 1897a. Histoire naturelle des araignées. Paris, 2: 1-192.
Simon E. 1897b. Etudes arachnologiques. 27e Mémoire. XLII. Descriptions
d'espèces nouvelles de l'ordre des Araneae. Annales de la Société
Entomologique de France, 65: 465-510.
378
On the Nearctic species of the family Sparassidae (Araneae)

Cristina Anne Rheims

cris.rheims@butantan.gov.br
The Nearctic region comprises most of the North American continent,

including Greenland and the northern highlands of Mexico. To date, only thirteen
species have been described from this region (eight from the USA and five from
Northern Mexico), all assigned to the genus Olios Walckenaer: Olios franklinus
Walckenaer, O. concolor Keyserling, O. giganteus Keyserling, O. peninsulanus
Banks, O. schistus Chamberlin, O. naturalisticus Chamberlin, O. positivus
Chamberlin, O. scepticus Chamberlin, O. pragmaticus Chamberlin, O. albinus Fox,
O. bibranchiatus Fox, O. mojavensis Fox and O foxi Roewer (Platnick 2010).
Olios franklinus, the first species to be described from the Nearctic, was
proposed by Walckenaer in 1837, from the USA. The female type was never located
and the identity of the species is considered doubtful.
Olios fasciculatus was described by Simon (1880), on both sexes from
Mariposa Co., USA. Roth (1988) examined the type series and, based on Simon’s
description, designated a male lectotype. Nevertheless, no females fitting the
original description were found amongst those in the type series suggesting that
these were added posteriorly and were not conspecific with the described male.
Also, since no other specimens were found in the collections of Olios from
California, Roth raises the possibility that the vial was actually mislabeled. Jäger &
Kunz (2005) confirmed Roth’s suspicions and matched the lectotype to specimens
from Tanzania. Thus, the species is not native and does not occur in the Nearctic
region.
Keyserling (1884) described O. giganteus, O. concolor and O. abnormis from
New Mexico, USA. The name abnormis was preoccupied by Sparassus abnormis
Blackwall 1866 and the species was given the new name Olios foxi by Roewer in
1951. Olios concolor and O. giganteus were synonymized with O. fasciculatus by
Banks (1893) and the latter removed from this synonymy by Roth (1988).
Olios peninsulanus was described by Banks (1898), based on a male and a
female from San Jose del Cabo, Baja California, Mexico. Chamberlin (1919)
described O. schistus from Claremont, Los Angeles, USA, and a few years later
(1924), O. naturalisticus, O. positivus, O. scepticus and O. pragmaticus from the
Gulf of California (Tiburon Is., San Francisco Is., Ceralba Is,. and San Lorenzo Is.,
respectively), Mexico. Olios pragmaticus was synonymized to O. fasciculatus by
Fox (1937) who revised the Nearctic fauna of Sparassidae and described three new
species: Olios albinos, from Phoenix, Arizona, O. bibranchiatus, from Madera
Canyon, Arizona, and Olios mohavensis, from the Mojave desert, California. He
also recorded the presence of Heteropoda venatoria (Linnaeus), a widely distributed
pantropical species, and Pseudosparianthis cubana Banks, originally described from
Cuba, in SW USA and Florida respectively.
379
Here I present the taxonomic revision of the Nearctic fauna of Sparassidae

based on the examination of the available type specimens. O. concolor and O.
pragmaticus are removed from the synonymy of O. fasciculatus and considered
junior synonyms of O. giganteus. O. schistus, O. scepticus and O. positivus are
synonymized with O. peninsulanus and O. albinus and O. foxi with O.
naturalisticus. All species are redescribed and illustrated. Comparisons between
these species and the type species of the genus Olios, Olios argelasius Walckenaer,
shows that none of them are congeneric and true Olios does not occur in the
Nearctic region. Nevertheless, the correct placement of these species in new genera
will only be possible after a more thorough revision of the Nearctic and Neotropical
fauna, especially that of Mexico and Central America.
References
Banks N. 1893. Notes on spiders. Journal of the New York Entomological
Society, 1: 123-134.
Banks N. 1898. Arachnida from Baja California and other parts of Mexico.
Proceedings of the California Academy of Sciences, 1: 205-308.
Blackwall J. 1866. A list of spiders captured in the southeast region of equatorial
Africa, with descriptions of such species as appear to be new to
arachnologists. Annals and Magazine of Natural History, 18: 451-468.
Chamberlin R.V. 1919. New Californian spiders. Journal of Entomology and
Zoology Claremont, 12: 1-17.
Chamberlin R.V. 1924. The spider fauna of the shores and islands of the Gulf of
California. Proceedings of the California Academy of Sciences, 12: 561-
694.
Fox I. 1937. The Nearctic spiders of the family Heteropodidae. Journal of the
Washington Academy of Sciences, 27: 461-474.
Jäger P. & Kunz D. 2005. An illustrated key to genera of African huntsman
spiders (Arachnida, Araneae, Sparassidae). Senckenbergiana biologica,
85: 163-213.
Keyserling E. 1884. Neue Spinnen aus America. Verhandlungen der Zoologisch-
Botanischen Gesellschaft Wien, 33: 649-684.
Roewer C.F. 1951. Neue Namen einiger Araneen-Arten. Abhandlungen des
Nturwissenschaftlichen Vereins zu Bremen, 32: 437-456.
Roth V.D. 1988. American Agelenidae and some misidentified spiders
(Clubionidae, Oonopidae and Sparassidae) of E. Simon in the Muséum
national d'Histoire naturelle. Bulletin du Museum d'Histoire Naturelle,
Paris, 10(A): 25-37.
Simon, E. 1880. Révision de la famille des Sparassidae (Arachnides). Actes de
la Société Linnéenne de Bordeaux, 34: 223-351.
380
Molecular evidence for the placement of Breda Peckham &

Peckham within the amycoids and proposal of two new
genera (Salticidae: Amycoida)
Gustavo R.S. Ruiz
Instituto Butantan, São Paulo, Brazil, gustavoruiz86@hotmail.com
The genus Breda Peckham & Peckham was proposed in 1894 to include
two South American species, Marpissa milvina C.L. Koch, 1846, from Bahia,
Brazil, designated as type species, and Marpissa lubomirskii Taczanowski, 1878,
from Peru. Since then, 12 other species have been described or placed in the
genus, composing the current list of 14 species in Platnick´s catalog. The genus
Breda was revised in 2005 (Ruiz & Brescovit unpublished data) and ten species
were considered valid. Also, three new species were found and described. A new
genus was also proposed to include Breda flavostriata Simon, 1901, a species
closely related to Breda that missed states considered synapomorphies for that
genus.
The exam of recently collected material from areas in Brazil revealed the
existence of two new lineages supposed to be closely related to Breda. Two
other new genera will be proposed to include these species.
The first new genus includes only one species with very low, long, dark
carapace, leg I without spines on femur or tibia, very reduced chelicerae, male
palp with three retrolateral tibial apophyses, an extra loop on the sperm duct and
a thin, long embolus; epigynum with anterior, small croissant-like atrium and a
median posterior depression; internally with short copulation ducts and small,
anterior spermathecae. This species is based on a single male and a single female
from Central Brazil (Goiás and Piauí).
The second new genus includes only one species with a shorter and higher
carapace, male palp with a single, short RTA, very long embolus, reaching two
and a half circles around the tegulum; epigynum has an anterior, almost
triangular atrium; internally with coiled copulation ducts and small, anterior
spermathecae. This species is based on several males and several females from
Alagoas, NE Brazil.
The genus Breda was proposed as belonging in the “Marptusa group”
(Marptusa is currently a junior synonym of Marpissa). The list of genera of the
Marptusa group as presented by Peckham & Peckham included all the salticid
species with low carapace known until that time. This group has been refined by
other authors, being currently known as the Marpissinae.
The subfamily Marpissinae Simon includes about nine genera, most from
the New World. The genus Breda was considered as belonging in the
Marpissinae by Maddison & Hedin, Edwards and Ruiz & Brescovit, based on
the general appearance of its flat species. Recently acquired DNA sequences
(both 28S and Actin genes), conversely, place Breda within the Amycoida.
381
The clade Amycoida was discovered by Maddison & Hedin during a study
on molecular data of salticids. The group includes a great portion of the
Neotropical diversity of jumping spiders, being composed by about 60 genera
and 420 described species. The main groups within the amycoids are the
Amycinae, Huriinae, Hyetussinae, Sitticinae, Synemosyninae and Thiodininae.
Besides, there are some incertae sedis genera, such as Agelista Simon, Asaracus
C.L. Koch, Fluda Peckham & Peckham, Scopocira Simon e Titanattus Peckham
& Peckham. The study by Maddison & Hedin did not include any member of
Breda.
Despite Breda and the related new genera have the same pattern of
cheliceral teeth as the species of the Hurieae group (Huriinae), they do not group
with one species of Hurius sampled. Instead, the genus Breda shows up close to
the base of Amycoida, sometimes in a polytomy with the Thiodininae, Sitticinae
and the rest of the group.
The amycoids will receive a better sampling and their morphology will
also be studied to help us reconstruct their phylogeny, clarifying the position of
Breda and related groups, which may be an important clade for the amycoids
due to their early branching, as shown by our preliminary results.
382
Phylogeny of dendryphantines (Araneae: Salticidae)

Gustavo R.S. Ruiz1 & Wayne P. Maddison2
1
Instituto Butantan, São Paulo, Brazil, gustavoruiz86@hotmail.com
2
University of British Columbia, Vancouver, Canada, wmaddisn@interchange.ubc.ca
The subfamily Dendryphantinae is one of the few groups of jumping spiders

with well established boundaries. It includes many hundreds of species, mostly from
the New World, whose males have a carina on the retrolateral surface of the
chelicerae and palps with a reduced spiral embolus.
Despite being one of the largest subfamilies of jumping spiders,
Dendryphantinae lineages present an unusual morphological homogeneity across the
entire group, comparable only to that of the subfamily Euophryinae. The lack of
diagnostic characters for genera caused the indiscriminate proposal, by earlier
arachnologists, of many genera with loose boundaries, sometimes resulting in small
groups of unrelated species. In part, this can be explained by the frequent homoplasy
of single male secondary sexual characters (for instance, male chelicerae are often
stout and elongated, as is the first pair of legs). All these facts have contributed to the
current chaotic classification of the group, which needed to be revised under a
phylogenetic scope.
Since male palp morphology is especially conservative in Dendryphantinae,
molecular data are of extreme importance for phylogenetic reconstructions in the
group. The main goal of this work was to sequence DNA regions of dendryphantine
terminals in order to reconstruct the phylogeny of the entire subfamily, allowing the
review of its systematics and the understanding of such large biodiversity.
During this study, we tried to sample the largest possible diversity of
dendryphantine lineages, including almost all described genera, some problematic
incertae sedis species and some lineages of undescribed species. In total, we
sampled over 100 species of Dendryphantinae, of which more than 85 were sampled
for the first time in molecular studies. In addition to the dendryphantine terminals,
we used 40 terminals as outgroups, limiting sampling to the large clade which
includes the Astioida, Baviinae, Ballinae and Marpissoida. To root the trees, we
used terminals of Astioida and Baviinae, in this order of preference, due to their
close relationship to the Marpissoida+Ballinae clade.
As an attempt to solve deeper level and shallower level divergences within the
group, four gene fragments – the nuclear 28S and Actin and the mitochondrial 16S
and ND1 – were amplified and sequenced in this study.
Bayesian analyses were done using MrBayes, using the GTR invariant-
gamma model. Model parameters were permitted to differ among data partitions.
We had two sets of partitions, one set with 8 and one set with 4 partitions. For the
first set, we treated nuclear data independently from mitochondrial. For the second
set of partitions, similar DNA types, nuclear or mitochondrial, were treated as
having similar evolution patterns.
Despite the notorious “long branch attraction” caused by Maximum
Parsimony, which can lead to wrong tree estimation especially when dealing with
383
large molecular data matrices, we also inferred trees using this method for all the
genes independently and for the same complete matrix used in the Bayesian
analysis. Parsimony analyses were done using TNT treating character states as
unordered and gaps as missing data.
As the result of the Bayesian analyses, the clades Marpissoida, Ballinae,
Marpissinae, Synagelinae and Dendryphantinae were recovered as monophyletic
(the clade Marpissoida includes the Marpissinae, the Synagelinae, the
Dendryphantinae and undetermined genera such as Itata Peckham & Peckham). The
general topology found under parsimony for the All-Genes matrix agrees with the
Bayesian analyses: (Astioida (Baviinae (Ballinae (Synagelinae (Marpissinae
(Dendryphantinae)))))). The position of the genus Itata and of a poorly understood
species from Costa Rica is still considered doubtful within the Marpissoida.
Within the Dendryphantinae, the genus Hentzia Marx seems to be sister to the
rest of the subfamily. Species of this genus, along with those of Macaroeris
Wunderlich, Phanias F.P.-Cambridge, Homalattus White, Mabellina Chickering
and Rudra Peckham & Peckham, are the only groups of dendryphantines who have
epiandrous fusules. The loss of these fusules characterizes a large clade which
includes most dendryphantine lineages. The clade of the infusulate dendryphantines
includes three major lineages. The first is the genus Zygoballus Peckham &
Peckham; the second is a huge clade including Metaphidippus F.O.P.-Cambridge
and several South American genera, such as Chirothecia Taczanowski, Ashtabula
Peckham & Peckham, Lurio Simon, Alcmena C.L. Koch and Admirala Peckham &
Peckham; the third lineage includes the genera Ramboia Mello-Leitão, Parnaenus
Peckham & Peckham, the Bagheera group (Bagheera Peckham & Peckham,
Gastromicans Mello-Leitão, Selimus Peckham & Peckham and species of the
“Messua” limbata group), the Bellota group (Bellota Peckham & Peckham,
Paradamoetas Simon and Sassacus Peckham & Peckham, among other genera) and
a huge clade from North America. The North American clade includes Ghelna
Maddison, Terralonus Maddison, Dendryphantes C.L. Koch, Tutelina Simon,
Phidippus C.L. Koch, Paraphidippus F.O.P.-Cambridge, Beata Peckham &
Peckham, Eris C.L. Koch and Pelegrina Franganillo.
Despite the fact that the subfamily seems to be a group originated in the
American continent, where it presents a fantastic diversification, the
Dendryphantinae has reached the Old World at least three times independently. Two
of these invasions are probably older and were carried out by lineages with
epiandrous fusules, namely Macaroeris and Homalattus (former Rhene Thorell),
while the third seems to be a very recent invasion by an infusulate group, the genus
Dendryphantes, who still has members in North America. Besides biogeographical
studies, many other aspects of their evolution can be inferred based on this topology,
such as the evolution of sexual secondary dimorphism (chelicerae and legs),
mimicry (beetles/ants) or the evolution of male palp structures.
Based on this phylogenetic hypothesis, natural groups of genera will be
revised with criticism, allowing their taxonomic simplification and easier
recognition. We hope that this will increase the interest of younger arachnologists on
the group and propitiate the description of the several hundred new species that
remain unknown and unnamed.
384
Survey of the araneofauna (Araneae) of Ibiza

(Balearic Islands)
Jens Runge1 & Carsten H.G. Müller2
1
Rostock, Germany, rung-gee@gmx.de
2
Ernst Moritz Arndt Universität Greifswald, Zoologisches Institut und Museum, Greifswald,
Germany
The araneofauna of the Balearic Islands has been the focus of a number of
faunistic studies. However, significantly less attention has been paid to the island
of Ibiza than to those of Majorca and Minorca. From the end of September to the
middle of October 2008 and 2009, spiders were collected at different locations
on Ibiza. To ensure the sample covered a broad faunistic spectrum, thirteen
locations were chosen which differed in vegetation type, grade of agricultural
use or building density. Due to differences in the diurnal activity of different
spider taxa, some locations were sampled at different times of day and night.
Use of a dip net and exhauster and collection by hand allowed us to draw
qualitative conclusions about the composition of the araneofauna. Specimens
from the following taxa were collected: Agelenidae, Araneidae, Dysderidae,
Gnaphosidae, Linyphiidae, Lycosidae, Miturgidae, Nemesiidae, Oecobiidae,
Oxyopidae, Philodromidae, Pholcidae, Pisauridae, Salticidae, Segestriidae,
Sicariidae, Tetragnathidae, Theridiidae, Thomisidae, Uloboridae, Zodariidae and
Zoropsidae. Over a dozen species were revealed to be new to Ibiza. Thanks to
the sampling techniques used, this survey showed the aranaeofauna of Ibiza to
be much more diverse then previously thought.
385
Soil spiders?
Vlastimil Růžička1, Vratislav Laška2, Jan Mikula2 & Ivan H. Tuf2
1
Institute of Entomology, Biology Centre, České Budějovice, Czech Republic,
vruz@entu.cas.cz
2
Department of Ecology and Environmental Sciences, Faculty of Science, Palacký
University, Olomouc, Czech Republic, vratislav.laska@email.cz,
jmikula@email.cz; ivan.tuf@upol.cz
Introduction
Spiders are among the most common and ubiquitous of animals; they are
indeed found everywhere over the life-supporting land masses of the world.
Where any form of terrestrial life exists, it is safe to assume there will also be
spiders living close by. Spiders have conquered all of the possible ecological
niches upon the land.
A wide spectrum of spider underground habitats also exists. Numerous
studies have been devoted to the study of cave spiders. We have a quantity of
knowledge on the vertical distribution of spiders in scree fields. Very little still
seems to be known about those invertebrates inhabiting the shallow void systems
within the bedrock, under the soil cover. Thus, the question arises: what about
“soil spiders”?
The objective of our study was to search for permanent soil spiders, i.e.
spiders inhabiting the deeper soil profiles for their entire life cycle. We extended
our previous study, bring new data, and prepared a mini-review.

Spiders were collected using subterranean traps. The collected animals
enter the tube through holes situated at 10 cm interval, and at different depths of
from 5 to 135 cm. The material was collected in the soil layers at two localities
in the Czech Republic: a beech forest growing on arenaceous marl bedrock, and
a floodplain forest growing on a gravel bank.

Altogether, 61 individuals from 11 spider species were trapped. In beech
woods, Cicurina cicur and Porrhomma microps were the most numerous. C.
cicur was found at a depth of 5–115 cm. P. microps exclusively inhabits the
deeper soil layers; it was found at depths of 65–135 cm. In the floodplain
forest, Porrhomma microps and Porrhomma myops were the most numerous.
P. microps was found at depths of 5–45 cm; whereas P. myops exclusively
inhabits the deeper soil layers, where it was found at depths of 35–95 cm.
Porrhomma microps is known as an inhabitant of the leaf-litter in deciduous
forests and also of caves. Porrhomma myops is known as an inhabitant of
screes and caves. In this study, both species were found to inhabit the deeper
soil layers.
386
Modes of speciation
We have evaluated our findings from the perspective of the development
of the adaptations of arthropods to subterranean life. We distinguished between
those adaptations to life in the soil environment (edaphomorphisms), and those
adaptations to life within the cave environment (troglomorphisms).
Depigmentation, desclerotization, as well as the atrophy or even loss of eyes are
common adaptations. The shortening of the appendages is a typical
edaphomorphism; while the elongation of the appendages is a typical
troglomorphism. The size ranges of soil inhabitants are generally smaller than
those of related epigeous species; however, among cave inhabitants both
gigantism and dwarfism are known.
The specimens of P. microps from the soil profile exhibit similar body
dimensions and proportions as those specimens from the leaf litter in South
Moravia, Czech Republic. The specimens of P. myops from the soil profile
exhibit relatively shorter legs than do the specimens from caves. We assume that
the ancient epigean ancestor of the contemporary P. myops exhibited similar
body proportions as did the epigean P. pygmaeum (similar to the other epigeic
species, e.g. P. microphthalmum). Compared with their close relative P.
pygmeum, the scree and cave populations of P. myops exhibit leg elongations (a
troglomorphism); whereas the soil population exhibits cephalothorax
diminutions (an edaphomorphism).
We recorded similar morphological adaptations to subterranean life in
several spider species. In the case of P. myops, we registered two principal
different routes of incursion into the undergroud realm. We registered two
different originator populations, which colonized two different shallow
underground habitats, and could have evolved into separate species when
colonizing the deep underground habitats.
The ‘hotspots’ of subterranean biodiversity in Dinaric Karst harbour
highly specialized forms at the end of a long-term underground evolution. Any
such previous specialized cave fauna at higher temperate latitudes must have
been eliminated by the rigorous periglacial climate; therefore the territory today
lying in the Pleistocene periglacial zones harbour invertebrates at the beginning
of their underground evolution.
Soil spiders?
Searching for the true edaphobionts, we evaluated the set of obvious
microphthalmous Central European spiders, by the exclusion of specialized
myrmecophilous species; supplemented by Wiehlea calcarifera (not
microphthalmous, but generally known as a soil inhabitant). The large species of
the genus Porrhomma are prevalent; they usually inhabit caves. P. microps also
inhabits the leaf litter, and the habitat of P. microcavense remains unknown. The
four smallest species i.e. Wiehlea calcarifera, Hahnia microphthalma,
Porrhomma cambridgei, and Pseudomaro aenigmaticus could be candidates for
soil spiders.
387
Condition dependent mate choice affects clutch size

and offspring sex ratio in the cellar spider
Ann L. Rypstra1, Ashley Conner2, Ashley Ziegler3, Leighton Fain2

& Chad D. Hoefler3
1
Department of Zoology, Miami University, Hamilton, OH, USA, rypstral@muohio.edu
2
Department of Zoology, Miami University, Oxford, OH, USA
3
Department of Biology, Arcadia University, Glenside, PA, USA
In species with low sexual dimorphism, we predict that there would be

less distinction in sex roles. Males of the cellar spider, Pholcus phalangioides
(Pholcidae), are large, by some measures larger than females, and reluctant to
mate in the 24 hrs after an initial copulation event. We tested for environmental
effects on mate selection and copulation as well as the sex ratio of the clutch
produced. We set up a series of laboratory experiments involving males and /or
females who had been maintained on a high or low quantity of prey. The
condition of both males and females affected both courtship and mating;
courtship commenced sooner and was most likely to end in mating with well fed
females; however, when paired with food limited females, males in good
condition started courtship sooner and were more likely to mate than their food
limited counterparts. Surprisingly the condition of both males and females
affected the number of eggs produced and the sex ratio of the offspring. Overall
the sex ratio was female biased, however clutches produced by well fed females
contained more males than the clutches of food limited females. Although male
feeding status had no influence on sex ratio when the female was food limited,
clutches of well fed females mated with well fed males contained more females
than clutches produced by well fed females mated to food limited males. Taken
together, these results suggest that the environment experienced by both males
and females affect the outcome of mating interactions. Although large egg sacs
produced by females make their choosiness within the traditional sex role
framework, the effects of male condition on mating behaviour, clutch size and
courtship suggests that they may be sperm limited and that they play are larger
role in the mating dynamic than is typically predicted by traditional sex role
theory.
388
Functional morphology of social groups

of web-building spiders
Maxence Salomon
Department of Zoology, University of British Columbia, Vancouver, B.C., Canada,

salomon@zoology.ubc.ca
For group-living species, maximizing individual fitness during activities

such as foraging may depend on a combination of individual- and group-level
characteristics. Species that live communally in built structures may
concurrently rely on the architectural properties of the group and the behaviours
of individual group members to achieve maximal prey capture success or
protection from predators. In this study, we examined how group architecture
and individual behaviour determine the functional morphology of social groups
of web-building spiders in the context of foraging. We conducted a field
experiment with two social species from the genus Anelosimus that differ in
individual and group morphology, and show that spiders achieve foraging
success by combining different functional properties of their communal webs
and individual cooperative behaviours. We discuss how the functional
morphology of a group may contribute to the overall success of a social lifestyle.
389
Scale and intensity of interaction between meadow

and arable field spider assemblages in a Hungarian
agricultural landscape
Ferenc Samu1, Dóra Neidert1, Éva Szita1, Kinga Fetykó1,

Zoltán Botta-Dukát2 & András Horváth2
1
Plant Protection Institute, HAS, Budapest, Hungary, samu@julia-nki.hu
2
Institute of Ecology and Botany, HAS, Vácrátót, Hungary
The Mezőföld region in Hungary typically consists of intensive arable

land which dominates large areas on the loess plateaus of the region, and
meadows in mosaic with wooded areas which are typical for the incised loess
valleys. We carried out a landscape experiment in seven 5x5 km quadrates in the
Mezőföld region of Hungary. The quadrates contained different proportions of
arable land and meadow areas, thus represented a land-use intensity gradient.
We studied the interaction between the spider assemblages of these habitat types
at a series of spatial scales. Samples were taken for three years in two cereal
field plots and in one meadow plot per landscape quadrate. We studied the effect
of the presence of different habitat types in the landscape neighbourhood of the
plots on spider species richness, abundance and community composition. For
spiders in the meadow plots the presence of arable habitats had in general a
negative effect, while for spiders in the cereal plots the presence of meadows
had a positive effect after controlling for local environmental variables and
taking into account spatial effect. This analysis had been performed five times
for each plot to take into account habitat types within five different radii between
50-1000 m. Strongest effects were observed for habitats within the 250 m and
600 m circles both for meadow and cereal plot spider assemblages. Cereal fields
which had at least 5% of meadow in their 250 m neighbourhood had nearly
twice the spider abundance than field with less than 1% of meadows in the same
neighbourhood. These and further studies into the effective distances of
interacting habitat types may help to optimize the spatial distribution of semi-
natural areas in a landscape in order to maximize landscape-level biodiversity.
390
Inventory of soil spiders (Arachnida: Araneae) in the

UFRRJ Campus, Seropédica, Rio de Janeiro, Brazil
Rita de Cássia Santos de Souza1, Rodrigo Santana de Sá1

& Ronald Bastos Freire2
1
Department of Animal Biology, Institute of Biology, Universidade Federal Rural do
Rio de Janeiro (UFRRJ), Brazil, * latrodectus_ufrrj@yahoo.com.br
2
Associated Professor, Department of Animal Biology, Institute of Biology, UFRRJ,
Brazil
The researches with spiders suggested that the richness of species and its
dominancy present a tendency to be highly related to the spatial heterogeneity,
determined by the plants community where they occur, their height, and by the
structure and composition of the litter. All these factors have a direct influence
on availability of refuges, structures to support the webs, places for
reproduction and deposition of the eggs (Rypstra et al. 1999) and are indirect
related to the diversity and abundance of preys (Souza 2007). In this study, a
mapping of the araneofauna (Arachnida, Araneae) from the campus of
Universidade Federal Rural do Rio de Janeiro (UFRRJ), in the city of
Seropédica was made, purposing to characterize and identify the spider fauna,
investigating the effects of environmental complexity on spiders’ assemblages.

The campus of UFRRJ presents an area about 3.024 ha and it is situated
at 22°45'S, 43°41'W. The climate presents the AW in Köppen classification,
being hot and humid, without apparent winter. There is a wet period in the
summer and a non-rigorous dryness period in winter. The vegetation is
dominated by grasses of the species Melinis minutiflora, Panicum maximum
and Imperata brasiliensis. The highest areas were covered with arbustive
vegetation, popularly called “capoeira” (Canellas et al. 2000).
The spiders were sampled (January - June, 2008, 9 times) by 30 pitfall-
trapps in each of the four different locations: in the campus of the Instituto de
Florestas (22°45'26,6"S, 43°41'43,2"W), in the housing sector (22°46'2,4"S,
43°41'24,7"W), in Lago Açu (22°45'40,0"S, 43°41'32,9"W) and in pines grove
(22°45'57,3"S, 43°41'13,9"W). The traps were filled with 300 ml of
hiperconcentrated saline solution (3 kg of salt+10 litres of water+detergent).
Such a solution has a reduced environmental impact (Moreira 2006).
The specimens were sorted out and identified at the Laboratório de
Artrópodes of Instituto Butantan, in São Paulo, Brazil under the guidance of
the Prof. Dr. Antonio Domingos Brescovit and his research group. The spiders
have been deposited in the collection of Instituto Butantan.
391

The vegetation complexity from each area was evaluated. In the forest the
vegetation was the most dense and diverse, following by the pines grove,
housing sector and the Lago Açu. Consequently, there were distinctive
differences in araneofauna between the locations. In house sector and Lago Açu
many specimens of Lycosidae of considerable size were found. In the pines
grove there was abundance of Salticidae and Corinnidae in the litter. At the
Instituto de Florestas the specimens of Mygalomorphae (Nemesiidae and
Theraphosidae) were found.
Altogether 910 spiders were collected, 285 immatures. At the Instituto de
Florestas the web spiders, especially Pholcidae predominated due to high
vegetation density, supporting the webs. Plantations and open areas have a
tendency to present high abundance of soil spiders and few orb-weavers (Hore &
Uniyal 2008). These spiders are more tolerant to to environmental disturbance
(Tsai et al. 2006).
The partial identification of the material collected shows abundance of
three dominant families, Linyphiidae, Corinnidae and Salticidae in the pines
grove. The family Corinnidae is predominated by Ianduba varia Keyserling,
1891, a typical synanthropic species. In Salticidae, the presence of the genus
Hasarius Simon, 1871, originated from Africa and also synanthropic was very
significant. This genus has a wide distribution. Members of the Lycosidae were
abundant in Lago Açu and the housing sector and the Pholcidae was
predominant in Instituto de Florestas. The highest variety of morphospecies
showed Linyphiidae (7), followed by Theridiidae (6 sp.).
The preliminary analysis has shown the relation between environment
characteristics and soil spiders assemblages.
Theridula gonygaster Simon, 1873 (Araneae: Theridiidae) was especially
unexpected from synsntripos area. The species is recorded only from few
localities in Brazil.
Acknowledgements
The authors are grateful to the Prof. Dr. Ildemar Ferreira, Prof. Dr. Marilia
Carvalho de Brasil Sato and Dr. Elaine Folly Ramos for the relevant presence in
this project and the encouragement. Thanks to Prof. Dr. Antonio Brescovit, for
important support in the identification of spiders. We also thank the CNPq and
the Universidade Federal Rural do Rio de Janeiro for the financial support. Rita
de Cássia Santos de Souza was supported by PIBIC UFRRJ/CNPq.
References
Canellas L.P., Berner P.G., Silva S.G., Silva M.B. & Santos G.A. 2000. Frações
da matéria orgânica em seis solos de uma toposseqüência no estado do rio
de janeiro. Pesquisa Agropecuária Brasileira, 35: 133-143.
Hore U. & Uniyal V.P. 2008. Diversity and composition of spider assemblages
in five vegetation types of the Terai Conservation Area, India. Journal of
392
Moreira T.S. 2006. Levantamento da Araneofauna (Arachnida, Araneae) do

Parque Nacional da Tijuca. Monografia (Bacharelado modalidade
Zoologia). Rio de Janeiro, UFRRJ/ Instituto de Biologia, 60 pp.
Rypstra A.L., Carter P.E., Balfour R.A. & Marschall S.D. 1999. Architectural
features of agricultural habitats and their impact on the spider inhabitants.
Souza L.T. 2007. Influência da Estrutura do habitat na abundância e diversidade
de aranhas. In: Gonzaga M.O., Santos A.J. & Japyassú H.F. (eds),
Ecologia e Comportamento de Aranhas. Rio de Janeiro, Editora
Interciência, pp. 25-43.
393
Spider webs in science, art and space

– an interdisciplinary project on 3D-visualisation
Tomas Saraceno1, Christof Wulff2, Dieter Steineck2,
Peter Jäger3 & Samuel Zschokke4
1
Studio Saraceno, Frankfurt am Main, Germany, tomas@t-saraceno.org
² Institut für Photogrammetrie und Kartographie, Technische Universität Darmstadt,
Germany, wulff@geod.tu-darmstadt.de, dieter@gauss.phgr.verm.tu-darmstadt.de
3
Senckenberg Research Institute, Frankfurt am Main, Germany,
peter.jaeger@senckenberg.de
4
Section of Conservation Biology, University of Basel, Switzerland,
samuel.zschokke@unibas.ch
How can we measure, analyze and transfer an entire three-dimensional

spider web electronically? This question was raised by a team of an art studio in
Frankfurt, Germany. A multidisciplinary team of scientists, artists and architects
worked for two years to find a solution. Driving force were ideas of Tomas
Saraceno to build fragile artistic web-structures inspired by architecture as well
as philosophy. The comb footed spider species Latrodectus mactans was chosen,
as its web is large enough and forms a three-dimensional construction. Results
will be presented including a stereoscopic image processing method of a laser
illuminated spider web. Properties of 3D-webs and their design can be analyzed
and compared for phylogenetic purposes. Subsequently, the web was transferred
from the electronic data set into a hand-knotted model of 30 times the original
size. It was pre-installed in a hangar close to Frankfurt and will be exhibited in
the Bonniers Konsthall in Stockholm in the first half of 2010. Another aspect for
the future work will be a photogrammetric procedure of a web built in the space
lab. The proposal passed the first evaluation by the NASA ISS (International
Space Station) within the Call “Research Opportunities for Flight Experiments
in Space Life Sciences on the ISS (ILSRA-2009)”. It should be investigated
whether a higher or lesser degree of freedom in building a spider web in
microgravity will result in different structural diversity, which, on the other
hand, could be useful for different disciplines.
394
A new species of Araneus Clerck, 1757 (Araneae:

Araneidae) from University Campus, Amravati, India
Akarte Satish1, Kondulkar Sunil2 & Milind Shirbhate3
1
Department of Zoology, Vidyabharati Mahavidyalaya, Amravati, India,
srakarte@rediffmail.com
2
MF Mahavidyalaya, Warud, Distt. Amravati, India, Sunil_kondulkar@rediffmail.com
3
Shankarlal Khandelwal College, Akola, Distt. Akola, India,
m_shirbhate@rediffmail.com
Araneus is a genus of orb-weaving spiders including the European garden

spider and the barn spider. Araneidae includes 168 genera with 2992 species.
The genus has more than 650 species. New species of Araneus, both male and
female from a single orb web is recorded from Amravati University Campus,
India.
Keywords: new species, taxonomy, Araneus, India.
395
Scent gland morphology in dyspnoid harvestmen

(Arachnida: Opiliones)
Miriam Schaider & Günther Raspotnig
Institute of Zoology, Karl-Franzens University Graz, Austria,

miriamschaider@hotmail.com, guenther.raspotnig@uni-graz.at
Introduction
Harvestmen are characterized by a pair of large prosomal scent glands.
These glands are most conspicuously developed in the suborders Laniatores and
Cyphophthalmi, but less developed in the third classical opilionid suborder, the
Palpatores. So far, data concerning scent gland morphology of Palpatores are
scarce and incomplete. Especially the Dyspnoi seem to exhibit aberrant scent
gland constructions and obviously complex, so far largely unknown secretion
modes (Juberthie et al. 1991, Schaider & Raspotnig 2009; Raspotnig et al.
2010). This study aims to shed light on the scent gland morphology of some
selected dyspnoid species.

Representatives of the two major Austrian families of Dyspnoi,
Nemastomatidae and Trogulidae, were collected from soil samples at different
locations in Upper Austria, Styria and Carinthia. The internal anatomy of scent
glands was studied by semithin-histological sectioning and subsequent 3d-
modelling of the scent glands with the help of reconstruction-software. The
external morphology of glandular openings (ozopores) was investigated by
scanning electron microscopy. Specimens of the eupnoid species Amilenus
aurantiacus were studied for comparison.
Results
Several different glandular types in the dyspnoid species herein studied
could be distinguished: Scent glands of Nemastomatidae, such as found in
Paranemastoma quadripunctatum or Nemastoma lugubre, exhibit some
characteristic features of typical opilionid defensive glands, e.g. large-scale
reservoirs internally covered by an (extensively folded) intima. Scent gland
openings, leading to the ventral side of the body, are strikingly hidden and can
hardly be detected from the outside. In Trogulidae two types could be described.
Scent glands in Anelasmocephalus hadzii are filled with cobweb-like projections
of the glandular epithelium. The glands open into a kind of external secretion
atrium, bordered dorsally by an integumental fold, ventrally by the coxa of leg I
and laterally by a loose layer of cuticular papillae. Ozopores are completely
covered and cannot be seen from the outside. Scent glands of the Trogulus-type,
such as found in Trogulus tricarinatus, are also characterized by an external
secretion-atrium with a massive lateral layer of cuticular papillae and by the
presence of solid secretion balls in the scent gland reservoirs. By contrast, scent
396
glands of Amilenus aurantiacus, a representative of Palpatores-Eupnoi, show

similar features as previously described for the eupnoid species Leiobunum
vittatum and L. flavum (Clawson 1988). Unlike the hidden ozopores of all soil-
dwelling Dyspnoi herein investigated, the slit-shaped scent gland openings of
Amilenus aurantiacus are embedded in a pore that is exposed on a cuticular
protrusion dorsal to legs I.
Discussion
Scent gland constructions of dyspnoid Troguloidea conspicuously differ
from the so far described defensive glands of other Opiliones as reported from
some Laniatores (Juberthie 1976), Cyphophthalmi (Gutjahr et al. 2005) and
palpatorid Eupnoi (Clawson 1988). Scent gland features like the cobweb-like
structures in the internal, glandular cavities of Anelasmocephalus or solid boli in
the reservoirs of Trogulus are not known from the latter groups and may
represent characteristics of the Dyspnoi. This assumption is underlined by the
fact that a similar situation with solid glandular contents in scent gland
reservoirs has also been described in ischyropsalids (Dyspnoi) (Juberthie et al.
1991). With respect to external features, a trend to hide ozopores – instead of
exposing them – can be observed: In particular the development of a secondary
atrium covering the ozopores may even be unique within the Troguloidea, and
has not been reported from the sister group Ischyropsalidoidea (Juberthie et al.
1991, Lopez et al. 1980).
The presence of distinct scent gland types especially in soil-dwelling
Dyspnoi may be indicative of multiple evolutionary traits that possibly resulted
in scent gland functions apart from typical chemical defence.
Acknowledgements
This study was supported by a DOC-fFORTE-fellowship of the Austrian
Academy of Sciences at the Institute of Zoology, Karl-Franzens-University Graz
(project number 22852).
References
Clawson R.L. 1988. Morphology of defence glands of the opilionids (Daddy
Longlegs) Leiobunum vittatum and L. flavum (Arachnida: Opiliones:
Palpatores: Phalangiidae). Journal of Morphology, 196:363-381.
Giribet G. & Kury A.B. 2007. Phylogeny and biogeography. Pp 62-87. In: Pinto-
da-Rocha R., Machado G. & Giribet G. (eds.), Harvestmen. The biology
of Opiliones. Harvard University Press, Cambridge, Mass.
Gutjahr M., Schuster R. & Alberti G. 2005. Ultrastructure of dermal and defence
glands in Cyphophthalmus duricorius Joseph, 1868 (Opiliones: Sironidae).
In: Deltshev C. & Stoev P. (eds), European Arachnology 2005 Acta
Zoologica Bulgarica, Suppl. 1: 41-48.
Juberthie C. 1976. Chemical defence in soil Opiliones. Revue d Ecologie et de
Biologie du Sol, 13: 155-160.
397
Juberthie C., Lopez A. & Juberthie-Jupeau L. 1991. Les glandes odorants des
Ischyropsalidae souterrains (Opilions): ultrastructure et role. Mémoires de
Biospéologie, 18: 39-46.
Lopez A., Emerit M. & Rambla M. 1980. Contribution a l´étude de Sabacon
paradoxum Simon, 1879 (Opiliones, Palpatores, Ischyropsalididae). Station
nouvelles, particularites électromicroscopiques du prosoma et de ses
appendices. C.R.Vé. Colloque Arach. IX, 1979 Barcelone, pp. 147-161.
Raspotnig G., Leutgeb V., Schaider M. & Komposch C. 2010. Naphthoquinones and
anthraquinones from scent glands of a dyspnoid harvestman, Paranemastoma
quadripunctatum. Journal of Chemical Ecology, 36: 158-162.
Schaider M. & Raspotnig G. 2009. Unusual morphology of scent glands in
Trogulus tricarinatus (Opiliones, Trogulidae): evidence for a non-
defensive role. Journal of Arachnology, 37: 78-83.
398
Monitoring spider diversity to assess the potential of

secondary forests for biodiversity conservation
in the southern Atlantic Rainforest of Brazil
Ludger Scheuermann, Florian Raub & Hubert Höfer
State Museum of Natural History, Karlsruhe, Germany, ludger.scheuermann@smnk.de,
florian.raub@smnk.de, hubert.hoefer@smnk.de
Spider assemblages in forests of the southern Mata Atlântica were studied

to enhance knowledge of the diversity of these forests and as part of a larger
project (www.inbioveritas.net). The aim of the study was the assessment of the
potential of secondary forests for the conservation of biodiversity in a situation
where altered forests dominate a fragmented landscape. The study was realized
between 2005 and 2008 in forests inside two private protection areas of the
Brazilian NGO SPVS “Reserva do Rio Cachoeira” and “Reserva Serra do
Itaqui” situated within the Environmental Protection Area Guaraquecaba in
Paraná. We analysed which portion of the spider assemblage observed in “old
growth” forests can exist in younger, naturally re-grown secondary forests and if
these forests can be classified by means of their spider species composition.
Sampling of spiders was carried out in three succession stages of
secondary forests (H - herbaceous, A - arboreal, M - median stage) and old
growth forest (F - forest) serving as a reference. Each stage was represented by
three replicate plots, so that in total 2x12 sites were sampled. We used three
collection methods following widely recognized standard protocols adapted to
our needs: time-based “beating” of spiders from lower vegetation during
daytime, time-based nocturnal hand sampling and pitfall trapping with an
exposition time of 1 week.
In total we collected 11,362 individuals of which 40% were adults (4,564),
which were identified to 37 families and 170 genera. Most individuals belonged
to the cob-weavers (Theridiidae), followed by zorid wandering spiders
(Zoridae), orb weavers (Araneidae) and jumping spiders (Salticidae). To
compare the stages in their genera richness the rarefaction method was used. The
forest stages showed an inconsistent image concerning genera richness. In
Cachoeira the M stage showed the highest genera number, whereas in Itaqui a
younger stage (A) was richest in genera. We used Chao 2 to estimate the genera
richness of each stage: in Itaqui the highest genera number was estimated for the
old growth (119.7+/-21.5), followed by A stage. For the M stage 90+/-10.5
genera were estimated. This stage however was the richest in Cachoeira
(111.1+/-16.5). Here for the A stage only 82.1+/-8.43 genera were estimated. An
ordination analysis of the genera composition by PCA revealed a distinct
differentiation of the two younger stages from the old secondary stage and the
old growth. Some genera only appeared in the two younger forest types (e.g.
Anodoration, Linyphiidae), other genera we found only in older forests (e.g.
Mesobolivar, Pholcidae). Spider genera of the Ctenidae family as typical forest
399
dwellers increased in abundance from young to old forest. Typical open-land

spiders like the Lycosidae were decreasing along the forest succession.
Lycosidae occurring in old growth forests belong to different genera.
We conclude that secondary forests older than 50 years can serve as a
surrogate habitat for some but not all genera (species) of the old growth. Some
genera (species) can serve as indicator taxa for forest age stages. This is an
important contribution to the multi-taxon approach to develop a biological
classification system for secondary forests in the Mata Atlântica as a knowledge
basis for future land management decision.
400
Species concepts and cryptic diversity in harvestmen

(Arachnida: Opiliones) - a short overview on methods
and perspectives
Axel L. Schönhofer
Johannes Gutenberg University of Mainz, Germany, Axel.Schoenhofer@gmx.net
The "species" is the basic unit of systematic and taxonomy and thereby
one of the most important units in the science of biology. But unlike in other
natural sciences, like physics or chemistry where units demand a clear definition,
the definition of a species naturally remains as variable as diversity itself and is a
constant issue of debate. At present about 26 definitions of “species” do exist.
They enable to define species in a variety of organisms that do not match other
species concepts or do extend species definition from present-day to future and
past. Despite these difficulties in definition, the “species” is the basis for any
political debate concerning diversity or conservation management. And while
the framework of species definitions in general deals with the human need to
classify diversity, species descriptions that meet the demand of more than one
concept appear more acceptable to satisfy modern needs.
Diversity is defined within a given frame, for example a group of
organisms or an ecosystem. The delineation of cryptic diversity and species
demands thorough knowledge of the system and its underlying mechanisms.
Only now, the next step of validation of diversity upon an appropriate species
concept is possible. So we can distinguish between variability and “true”
characters to delineate species. But how do we identify diversity? And which
characters for species delineation are useful within a set of variables? This
speech will give an overview on species concepts and methods used in
harvestmen systematic to deal with this central problem in biology.
Within arthropods, as well as in most other animals described in the last
centuries, the typological species concept, based on morphological characters,
prevailed. A morphospecies has the great advantage that, once a good
discriminating character is recognised, it can be identified by anyone without
involving expensive or difficult-to-access laboratory techniques. On the other
hand a lot of experience and knowledge is needed to identify intraspecific
variation compared to species delineating characters and it is often reasonable to
investigate other traits to discriminate the two.
For species described upon morphological characters alone the importance
of discriminating characters underwent a remarkable change when Silhavy
described several Opilio-species in Central Europe upon male genital
morphology. Albeit the size and colouration of the species varied, genital
morphology appeared constant, suggesting its high usage for species
identification, as known for many other groups of arthropods. Although this
dogmatic change first remained unacknowledged by the dominating scientist in
this field, C.F. Roewer, most of his species have or will face a revalidation of
401
their genital morphology. This was the case in the European genus Ischyropsalis
thoroughly revised by Martens.
Martens not only supported male genital morphology as key character in
many groups of Opiliones but showed combination with other morphological
features in Ischyropsalis. As in many Dyspnoi Ischyropsalis exhibits special
glands on the first cheliceral segment. Martens showed these glands to play an
important role in the mating process and their form and position on the
chelicerae to be species specific. He thereby showed mating behaviour to isolate
species and validated the connected morphological characters as useful to
identify biospecies.
The biological species concept, assuming reproductive isolation of a
species, is currently the most widely accepted species hypothesis. Nevertheless it
is upon the most difficult to substantiate. Direct confirmation is only possible
upon breeding experiments or indirectly by sympatric occurrence of species that
thereby obviously do not interbreed. Allopatric populations have to be
supported, for example, by ethological or morphological traits. The biospecies
concept is not applicable to a range of species for example fossils or
parthenogenetic organisms.
Speaking of allopatric populations, allopatry is known as one of the main
processes to induce speciation. It is common sense, despite being a “true”
definition, to include biogeographical information in the description and
delineation of species. A negative example is Roewer, who frequently misused
geographic information to describe species upon “remarkable” new records.
European Ischyropsalis showed an interesting distribution until Martens revised
the genus. Nevertheless, properly interpreted, biogeographic information is
important considering the isolation of populations that caused speciation as we
can show in several groups of the genus Trogulus.
Today, methods investigating genetic markers have strong influence on
many parts of biological science. Molecular analyses yield comparable and
quantifiable data that enable statistically evaluable results. The comparison of
homologous strands of DNA is a powerful tool to investigate and quantify
diversity in organisms beforehand showing no discriminating characters. Albeit,
the method can have many drawbacks and needs careful investigation of the
study system, that can cost a lot of time and money. Reviews strongly suggest
validation of species defined by barcoding upon other lines of evidence. The
phylogenetic species concept comes within the limit of this topic. It assumes any
independent genetic lineage as species, subsequently asking what is to be
considered as “independent”. This species concept is somehow difficult to
handle as it allows a very subjective view on species. Consequently, purely
genetically defined species are unconvincing when not supported by other
independent lines of evidence such as morphology or geographical traits. While
the field of molecular systematic answers important and sophisticated questions
on diversity, speciation and evolution, a mere genetic species definition needs
further support to be applicable and accepted in everyday life.
There are many other possibilities to delineate species, as behaviour or
chromosomes. But all need to be validated upon other traits to proof their
402
reliability for species identification. Nevertheless all these methods are an

important basis to delineate diversity and subsequently investigate for the
possibility of cryptic species. In view of a modern approach the definition of
species should not be based upon a single character but should be carefully
checked for concordance with other characters and species concepts to increase
the reliability of the species description. This is especially true as new methods,
as barcoding, do not represent the Holy Grail to systematic and taxonomy, but,
in the light of accumulating experience in genetics, require most careful
validation upon other lines of evidence.
403
Spiders in agroecosystems of Kerala, India

Pothalil A. Sebastian1, Mundackatharappel J. Mathew2,
S. Murugesan3 & John Joseph1
1
Sacred Heart College, Kochi, India, drpothalil@rediffmail.com
2
IT@School Project, Kochi, India
3
Institute of Forest Genetics and Tree Breeding, Coimbatore, India
Introduction:
Spiders are an integral part of global biodiversity. They play many
important roles in ecosystems as predators and sources of food for other
creatures. Being insectivores, spiders are of economic value to human beings
because of their ability to suppress pest abundance in agroecosystems. Faced
with the need to reduce pesticide usage on crops and optimize natural biological
control, full investigation of the means by which spiders influence pest
abundance is long overdue. While arachnologists and others working in
agroecosystems have been encouraged by results of recent studies suggesting
that spiders can impact pest populations and reduce crop damage, most would
agree that agricultural arachnology is still in its infancy compared with the
breadth and depth of entomological research on Integrated Pest Management
(IPM) and biological control (Uetz et al. 1999). There is increasing evidence that
polyphagic predators, to which spiders belong, play an important role in the
regulation of the number of insects. To form a basis for research into the role of
spiders to determine the economic importance of them in different
agroecosystems, a survey of the araneid population along with the study of their
biology and ecology is necessary. Information of the spider species present, their
occurrence throughout the growing season and their abundance are important
and inevitable components of biological control and pest management in any
agroecosystem. Against this backdrop, a pioneering study was undertaken to
study the spider fauna in various agroecosystems in the state of Kerala in India.
Apart from documenting spiders, the study also envisaged to examine the
qualitative and quantitative aspects of their diversity, ecology, guild structure,
seasonality, etc in the various ecosystems.

The study area consisted of Ernakulam, Idukki, Trichur and Palghat
districts in central Kerala, India. Altitude in these districts ranges from seaboard
to 2695 m above MSL. The crops selected for the study were coconut (Cocos
nucifera L.) and vegetables including bitter gourd (Momordica charantia L.),
snake gourd (Trichosanthes cucumerina L.), ivy gourd (Coccinia grandis (L.)
Voigt, cowpea (Vigna unguiculata L. Walp.) and cabbage (Brassica oleracea L.
var. capitata). Sampling sites selected were Adimali (10o5'N, 77o44'E),
Murikkasseri (9o45'N, 77o6'E) and Kanthalloor (10o5'N, 77o4'E) of Idukki
district; Kothamangalam (9o58'N, 76o34'E), Piravom (9o58'N, 76o34'E),
404
Manjapra (9o58'N, 76o16'E) and Parakkadavu (11o15'N, 75o49'E) of Ernakulam

district; Vellanikkara (13o31'N, 76o13'E), Thekkumbagham (10o31'N, 76o13'E)
and Chuvannamannu (10o31'N, 76o13'E) of Trichur district; and Alathur
(10o39'N, 76o31'E) and Pathanapuram (10o39'N, 76o55'E) of Palghat district.
In vegetable ecosystems, samplings were done using quadrate method.
Random transect method using the technique adopted by Aiken and Coyle
(2000) was used for spider sampling in coconut plantations. The collected
spiders were identified with the help of literature (Tikader 1987, Barrion &
Litsinger 1995, Dippenaar-Shoeman & Jocque 1997, Deeleman-Reinhold 2000)
using stereoscopic microscopes (Leica MS5, Olympus SZ112).

In coconut, a total of 2120 individuals belonging to 55 species, 38 genera
and 14 families were sampled. The taxonomically dominant family was
Salticidae while the numerically dominant family was Lycosidae. P.
pseudoannulata was the most abundant species. Ground runners constituted the
dominant guild. Among the sites, Parakkadavu recorded the highest diversity
indices and species richness. Analysis of seasonality revealed the highest species
occurrence in the post-monsoon months (October to January), followed by pre-
monsoon months (February to May) and the least occurrence in the monsoon
period (June to August).
In bitter gourd, a total of 3504 individuals belonging to 66 species, 41
genera and 14 families were sampled. Araneidae was the taxonomically
dominant family. The numerically dominant family was Lycosidae. The
numerically dominant species was P. sumatrana. Ground runners constituted the
abundant spider guild in bitter gourd ecosystem. Murikasseri in Idukki district
recorded the highest diversity indices and species richness. Population growth
showed a steady increase as the crop advanced and reached a peak at early
flowering and early fruiting seasons, followed by a decline during the late
fruiting season.
In snake gourd, surveys yielded 1276 individuals belonging to 41 species,
29 genera and 11 families. Araneidae was the taxonomically dominant family.
The numerically dominant family was Lycosidae. The numerically dominant
species was P. sumatrana. Ground runners were the abundant spider guild in
snake gourd ecosystem. Manjapra in Ernakulam district recorded the highest
diversity indices. Population growth showed a steady increase as the crop
advanced and reached a peak at early flowering and early fruiting seasons,
followed by a decline towards the end of the vegetative growth of the crop.
In ivy gourd, 472 individuals belonging to 33 species, 23 genera and 10
families were sampled. Family Araneidae was the taxonomically dominant
family. The numerically dominant family was Lycosidae. The numerically
dominant species was P. pseudoannulata. Ground runners formed the abundant
spider guild in this ecosystem. Alathur in Palghat district recorded the highest
diversity indices and species richness. Population growth showed a sharp
increase as the crop advanced in growth and reached the peak at early flowering
and early fruiting seasons.
405
In cowpea, sampling yielded a total of 862 individuals belonging to 33

species, 23 genera and 8 families. Family Araneidae was the taxonomically
dominant family. The numerically dominant family was Lycosidae. The
numerically dominant species was P. pseudoannulata. Ground runners
constituted the abundant spider guild in cowpea ecosystem. Kothamangalam in
Ernakulam district recorded the highest diversity indices and species richness.
The population growth curve revealed a gradual increase as the crop advanced in
growth and reached the peak at the mid to late fruiting seasons. This peak was
then followed by slight decline towards the end of the vegetative growth of the
crop.
In cabbage, a total of 266 individuals belonging to 21 species, 15 genera
and 6 families were sampled. Family Araneidae was the taxonomically dominant
family. The numerically dominant family was Lycosidae. The numerically
dominant species was P. sumatrana. Ground runners formed the abundant spider
guild in the cabbage ecosystem. Population growth showed steady increase as
the crop grew and reached the peak at head formation stage. The population then
showed a slight decline towards the harvest period.
The results substantiate the fact that population densities and species
abundance of spider communities in agricultural fields can be as high as in
natural ecosystems. However, further investigations are warranted to study their
interaction with the environment, breeding behaviour, prey preference, predatory
potential and sensitivity to chemical and botanical pesticides in order to fully
utilize them as successful biological control agents in these ecosystems. Studies
worldwide indicate that spiders play a role in the suppression of insect pests and
that generalist predators such as spiders can limit exponential increases in pest
populations and need to form a part of integrated pest management strategies in
agro-ecosystems. However, despite their ubiquity and high densities, spiders
have not received the due recognition in this enterprise. The main reason for this
can be attributed to limited number of studies along these lines. Hence it is of
utmost importance to undertake elaborate studies on the ecology and biology of
agrobiont spiders, as well as the logistics of their conservation.
References
Aiken M. & Coyle F.A. 2000. Habitat distribution, life history and behavior of
Tetragnatha spider species in the Great Smoky Mountains National Park.
Barrion A.T. & Litsinger J.A. 1995. Riceland spiders of South and South-East
Asia. CAB International, UK and IRRI, Philippines, 700 pp.
Deeleman-Reinhold C.L. 2000. Forest Spiders of South East Asia: With a
Revision of the Sac and Ground Spiders (Araneae: Clubionidae,
Corinnidae, Liocranidae, Gnaphosidae, Prodidomidae, and
Trochanterriidae. Brill Academic Publishers, 591 pp.
Dippenaar-Schoeman A.S. & Jocqué R. 1997. African Spiders. An Identification
Manual. Plant Protection Institute Handbook No. 9. ARC, Plant Protection
Research Institute, Pretoria, 392 pp.
406
Tikader B.K. 1987. Handbook of Indian Spiders. Zoological Survey of India,

Calcutta, 251 pp.
Uetz G.W., Halaj J. & Cady A.B. 1999. Guild structure of spiders in major
crops. Journal of Arachnology, 27: 270-280.
407
The latest on the oldest: recent advances

in spider palaeontology
Paul A. Selden
The Paleontological Institute, Department of Geology, University of Kansas, Lawrence,
KS, USA, selden@ku.edu
Over the last few years, there have been many changes to our perception of
the evolutionary history of spiders, as evidenced by fossils. In particular, the
Mesozoic era, formerly almost barren of fossil spider records, is now much better
known. In this review of fossil spiders, I first discuss new methods for studying
spider fossils, including imaging techniques such as synchrotron x-ray CT
scanning.
Then recent finds of spider fossils, primarily from the Mesozoic era, are
discussed, with particular emphasis on how our perception of spider diversity
through geological time has changed as a result of these finds (see figure). Some
of the fossils in rock matrix preservation can be quite remarkably preserved, and
since the oldest amber with inclusions is early Cretaceous, rock matrix
preservation fossils extend our knowledge of fossil spider diversity much further
back in time. Finally, the reinterpretation of Attercopus, formerly the oldest known
spider, from the Devonian period, is discussed.
408
Comparison of male palps of orb-weavers between

Araneus sturmi and A. triguttatus, with notes
on identification characters 9
Anna Šestáková & Miroslav Krumpál
asestakova@gmail.com, krumpal@fns.uniba.sk
Archer (1951) brought into science a lot of synonyms, because his

description of males was based on median apophysis. Despite the Levi´s note
(1971) on insufficient value of median apophysis in identification due to its
function as a supporting sclerite during copulation, most of determination keys
still use it to distinguish males of A. sturmi (Hahn, 1831) and A. triguttatus
(Fabricius, 1793). During revision of Araneus species we noticed that the same
specimen could has different median apophysis on left and right palp, broken
terminal spines and other deformations. Locket & Millidge (1953) compared two
different “x” – marked sclerites, which put emphasis on general appearance of
palp (they are believed to be subterminal apophysis of A. sturmi and embolus of
A. triguttatus). After 50 years Almquist (2005) compared these species being
based on a shape of embolus. This character, however, is explicit and
comprehensible enough only with additional information on morphology and
position of other sclerites, because a small embolus of A. sturmi is difficult to
see. Our study follows up Locket & Millidge´s work. It is based on general
appearance of palpal organ. In the examined collections many specimens of
A. sturmi were misidentified as A. triguttatus. Comparing male palps of both
species we have found the more reliable characters. Palp of A. sturmi has wider,
fingered, elongated terminal apophysis, saddle-shaped conductor and relative big
cochleariform distended subterminal apophysis. These three sclerites completely
cover small short embolus. On the other hand palp of A. triguttatus has clearly
visible long pointed embolus, because it isn’t covered with subterminal
apophysis, which is very small and weakly sclerotized. Terminal apophysis has
smaller but more markedly separated fingered elongation. Another less
interesting character lies in comparison between button-shaped paracymbium of
A. sturmi and paracymbium without enlarging A. triguttatus.
9
The project is partly supported by grant VEGA 1/0176/09.
409
How many linyphiids (Araneae: Linyphiidae) are there

in tropical Africa?
Rimma R. Seyfulina
Department of Invertebrate Zoology, Faculty of Biology, Lomonosov Moscow State

University, Moscow, Russia, r-seyfulina@yandex.ru
Introduction
Linyphiidae are considered to be typical spiders for the northern
temperate zone, where they constitute the largest portion of the spider species
richness (e.g. Miller & Hormiga 2004). It is not surprising that most studies of
Linyphiidae have been focused on the Holarctic, leaving the fauna of other
regions rather understudied. Thus, the species richness of Linyphiidae in tropical
Africa was underestimated for a long time. Berland (1955) believed they are
very rare in Africa and absent from the western part of the continent because the
linyphiid fauna of the region was poorly known at that time. During the ensuing
years, the number of reported species quintupled, mostly with novel
descriptions. With more than 400 species described to date, this family is
outpaced for its diversity only by the Salticidae and Lycosidae from the
Afrotropical region. For comparison, more than 1350 linyphiid species are
reported from the Western Palaearctic (Europe and Northern Africa) bordering
with the Afrotropic and including, among others, cold temperate areas. This
number is probably close to the real species richness in the region, as at least
European linyphiid fauna is fairly well known and new species are rarely
described. So, the Linyphiidae are three times less diverse in tropical Africa than
in the adjacent zoogeographical region roughly equal to it in size. Is such a
pronounced difference due mostly to the shortage of taxonomic studies or does it
indeed reflect true biogeographical situation? In the attempt to answer this
interesting question, we estimated the total number of linyphiid spices in the
Afrotropical region using the unknown/known species ratio in samples collected
from several sites across tropical Africa by forest canopy fogging.

Samples were collected in 12 sites of the montane forest : Congo DR
(1993, 1 locality), Rwanda (1993, 3 localities), Kenya (1999, 2001-2003, 5
localities), Uganda (1997, 3 localities) and one site with lowland rainforest:
Ghana (2005, 1 locality). All sampling sites are located in tropical rainforest. In
some of them the identity of individually fogged trees was noted. In total, 15
species of trees were processed. Spiders were sampled with the pyrethrum
knockdown method following the protocol outlined by Stork (1987).
Spiders were identified during one year collaboration with the Royal
Museum for Central Africa, Belgium (under the financial support of the Belgian
Federal Science Policy Office).
410
The species number of Afrotropical Linyphiidae was estimated as follows:

N=D*(100/(100-P)), where N is the projected number of species, D – the
number of currently described species, P – the percentage of morpho-species.
This formula was used by Hodkinson (1992) for estimation of the total insect
species number.
Results
In this study, about 3350 specimens belonging to 84 species were
examined. The percentage of unknown species varies from 25% to 100%
between the localities. The total percentage of new species is 76, the weighted
mean is 69 per site. The Hodkinson’s formula produces the estimate of 1335
species. Taking into account that the arthropod fauna in canopy is twice as rich
in species as in the forest floor (Erwin, 1982) we reduced the estimated number
to 1000 species. Another calculation with using the data obtained by pitfall traps
in one of the studied localities (although not very representative, only fifty
specimens of 5 species were collected) yielded the same number.
Conclusions
The minimal estimated number of linyphiid spider species in tropical
Africa is 1000. The linyphiid species richness is similar between tropical Africa
and the Western Palaearctic or, more generally, does not differ drastically
between the temperate zone and the tropics. Probably the rain forests of tropical
Africa harbour as many linyphiids as boreal forests, a suggestion being in
agreement with the hygrophilous ecology of the Linyphiidae. However, in
temperate regions their fraction is much higher due to a sharp decrease in the
diversity of other spider groups.
References
Berland L. 1955. Les Arachnides De L’Afrique Noire Française. I.F.A.N. Dakar.
Erwin T.L. 1982. Tropical forests: their richness in Coleoptera and other
arthropod species. The Coleopterists Bulletin, 36(1): 74-75.
Hodkinson I.D. 1992. Global insect diversity revisited. Journal of Tropical
Ecology, 8: 505–508.
Miller J. & Hormiga G. 2004. Clade stability and the addition of data: A case
study from erigonine spiders (Araneae: Linyphiidae, Erigoninae).
Cladistics, 20: 385-442.
Stork N.E. 1987. Arthropod faunal similarity of Bornean rain forest trees.
Ecological Entomology, 12: 219-226.
411
Grassatores phylogeny based on ten molecular markers:

the evolution of a hyperdiverse arachnid infraorder
(Arachnida: Opiliones: Laniatores)
Prashant Sharma & Gonzalo Giribet
Department of Organismic and Evolutionary Biology, Harvard University, Cambridge,

MA, USA, psharma@fas.harvard.edu, ggiribet@oeb.harvard.edu
Of the four suborders of Opiliones, the largely tropical Laniatores

encompasses almost two-thirds (over 4000 species) of described opilionid
diversity, but has received far less than commensurate phylogenetic study.
Laniatores is presently divided into two tenuous infraorders, Insidiatores and
Grassatores, the latter harbouring most of the order’s striking examples of
morphological and behavioural diversity. Approximately half of all known
Opiliones species (and 22 of the 45 families) are within Grassatores. Previous
assessments of Grassatores relationships have recovered few stable clades and
fewer supported nodes. In the present study, the phylogeny of Grassatores is
reassessed with increased taxonomic sampling and ten molecular loci, under
dynamic homology using POY v. 4.1.2. Parsimony analyses strongly support the
monophyly of Grassatores and some constituent superfamilies (e.g.,
Gonyleptoidea). Other superfamilies, such as Samooidea, are polyphyletic and
require revision. One group of genera does not cluster with any described
families. The morphology of this clade and the potential for a new family of
Laniatores is discussed. The evolution of key morphological traits is discussed in
the context of the phylogeny.
412
Upstream colonization of Australasia by Neotropical

arachnids: phylogenetic relationships of the Zalmoxidae
(Arachnida: Opiliones: Laniatores)
Prashant Sharma & Gonzalo Giribet
Department of Organismic and Evolutionary Biology, Harvard University, Cambridge,

MA, USA, psharma@fas.harvard.edu, ggiribet@oeb.harvard.edu
The species richness and endemism of the Southwest Pacific are

traditionally held to result from dispersal from Australasian continental
landmasses (e.g., Australia, New Guinea, Southeast Asia), in concert with
prolonged isolation. In the present study, the phylogeny of the circum-Pacific
arachnid family Zalmoxidae was investigated using a multilocus dataset (six
molecular loci, totaling ca. 5800 bp), under dynamic homology using POY v.
4.1.2. Parsimony analyses support the monophyly of Zalmoxidae and a
Neotropical origin of this clade. Neotropical lineages within Zalmoxidae form a
paraphyletic grade at the base of a derived, monophyletic Indo-Pacific clade.
This topology and the relative phylogenetic placement of New Guinean and
Southeast Asian lineages suggest upstream colonization of Australasia by a
Neotropical radiation. This highly uncommon biogeographical signal is
contrasted with traditional views of Southwest Pacific biogeography.
413
Effect of adjacent crops and agronomic practices

on the active density and diversity of spiders
in wheat ecosystems
Sher Muhammad Sherawat1, Abida Butt & Hafiz M. Tahir2
1
District Officer Agriculture, Extension Malik Anwar Road, Sheikupura, Punjab,
Pakistan, sherawat25@hotmail.com
2
Department of Zoology University of the Punjab, Lahore, Pakistan,
hafiztahirpk1@yahoo.com
The study was aimed at to describe the effects of five wheat habitats
(which differed in adjacent crops and agronomic practices) on the activity
density, richness, evenness, diversity, guild structure of spiders and pest
populations. The study was conducted in district, Sheikhupura Punjab, Pakistan
during 2005-6 and 2006-7. A total of 23097 specimens of spiders belonging to
47 species, 31 genera and 12 families were collected from five different habitats.
Overall diversity and evenness of spiders did not differ among different habitats;
however activity density and richness of spiders was significantly different. All
sampled habitats had similar family and species composition. Significant
positive correlation was observed between active density of agrobiont spiders
and prey populations.
414
Diversity of spiders from the family Araneidae inhabiting

dry deciduous forest of Melghat Tiger Reserve (India)
Milind Shirbhate1, Amit Vairale B.2, Mahesh Chikhale3,

Deepa Kadu2 & Ujjwala Deshmukh4
1
Shankarlal Khandelwal College, Akola, India, m_shirbhate@yahoo.co.in
2
Department of Zoology, SGB Amravati University, Amravati, India
3
Indian Society of Arachnology, Amravati, India
4
Department of Zoology, Government Vidarbha Institute of Science & Humanities,
Amravati, India
Introduction
Melghat Tiger Reserve (21°15'N - 21°45'N, 76°57’E - 77°30'E, 312 m -
1178 m a.s.l.) is situated in the Satpura hill ranges of Central India (Melghat
forests, Amravati district, Vidarbha region of Maharashtra) and is bordered with
Madhya Pradesh in the North and East. It is a typical representative of Central
Indian Highland forming a part of the biogeographic zone “6 E-Deccan
Peninsula”. The area constitutes forests which are part of world’s fifth biologically
richest heritage country and the Reserve forms an important corridor between
forest areas of Madhya Pradesh and Maharashtra ensuring contiguity of forests in
Satpuras. It beholds one of the viable populations the Royal Bengal Tiger, out of 5
surviving tiger subspecies, including all the Tiger range countries. Out of 237 -
240 Tigers in Maharashtra in 2002, 75 - 80 (about 30%) were reported from
Melghat Tiger Reserve.
Observations
A survey of araneids was carried out in Melghat Tiger Reserve during 2007-
2009. We have reported 87 species from 11 genera. The maximum species
diversity was noted from August to January, 2007-2009. Out of 87 species 25 are
new. Among the collected species 28 are males and 59 are females.
Table 1. Araneid genera and species recorded from Melghat Tiger Reserve.
Genus Number of species
1. Araneus 11
2. Argiope 4
3. Chorizopes 3
4. Cyclosa 16
5. Cyrtarachne 2
6. Cyrtophora 7
7. Gasteracantha 2
8. Larinia 6
9. Neoscona 28
10. Poltys 2
11. Zygiella 6
415
Species
1. Araneus bilunifera Pocock Female
2. Araneus cucurbitinus Clerck Female
3. Araneus mitifica (Simon) Female
4. Araneus mitifica (Simon) Male
5. Araneus pachganiensis Tikader and Bal Female
6. Araneus pahalgaonensis Tikader and Bal Female
7. Araneus pahalgaonensis Tikader and Bal Male
8. Araneus sp. nov. Female
11. Araneus sp. nov. Male
12. Argiope aemula (Walckenaer) Male
13. Argione aemula (Walckenaer) Male
14. Argiope sp. nov. Female
15. Argiope sp. nov. Male
16. Chorizopes anjanes Tikader. Male
17. Chorizopes calciope (Simon) Female
18. Chorizopes khanjanes Tikader. Female
19. Cyclosa bifida (Doleschall) Female
20. Cyclosa bifida (Doleschall) Male
21. Cyclosa confraga (Thorell) Female
22. Cyclosa fissicauda Simon Female
23. Cyclosa hexatuberculata Female
24. Cyclosa insulana (Costa) Male
25. Cyclosa moonduensis Tikader Female
26. Cyclosa moonduensis Male
27. Cyclosa mulmeinensis (Thorell) Female
28. Cyclosa neilensis Tikader. Female
29. Cyclosa simoni Female
30. Cyclosa sp. nov. Female
33. Cyclosa sp. nov. Male
34. Cyclosa spirifera Simon. Female
35. Cyrtarachne bengalensis Tikader Female
36. Cyrtarachne sp. nov. Female
37. Cyrtophora bidenta Tikader Female
38. Cyrtophora cicatrosa (Stoliczka) Female
39. Cyrtophora citricola (Forskal) Female
40. Cyrtophora moluccensis (Doleschall) Female
41. Cyrtophora sp. nov. Female
42. Cyrtophora sp. nov. Female
43. Cyrtophora sp. nov. Male
44. Gasteracantha sp. nov. Female
416
45. Gasteracantha sp. nov. Female

46. Larinia chloris (Audouin) Female
47. Larinia chloris (Audouin) Male
48. Larinia phtisica (L. Koch) Male
49. Larinia phtisica (L. Koch) Female
50. Larinia sp. nov. Female
51. Larinia sp. nov. Male
52. Neoscona achine (Simon) Female
53. Neoscona achine (Simon) Male
54. Neoscona bengalensis Tikader and Bal Female
55. Neoscona bengalensis Tikader and Bal Male
56. Neoscona chrysanthusi Tikader and Bal Female
57. Neoscona excelsus (Simon) Female
58. Neoscona laglaizei (Simon) Female
59. Neoscona lugubris (Walckenaer) Female
60. Neoscona molemensis Tikader and Bal Female
61. Neoscona mukerjei Tikader Female
62. Neoscona mukerjei Tikader Male
63. Neoscona nautica (L. Koch) Female
64. Neoscona nautica (L. Koch) Male
65. Neoscona odites (Simon) Female
66. Neoscona odites (Simon) Male
67. Neoscona pavida (Simon) Female
68. Neoscona rumpfi (Thorell) Female
69. Neoscona rumpfi (Thorell) Male
70. Neoscona shillongensis Tikader and Bal Male
71. Neoscona sinhagadensis (Tikader) Female
72. Neoscona sinhagadensis (Tikader) Male
73. Neoscona sp. nov. Female
74. Neoscona sp. nov. Female
75. Neoscona sp. nov. Male
76. Neoscona sp. nov. Male
77. Neoscona shillongensis Tikader and Bal Female
78. Neoscona theis (Walckenaer) Female
79. Neoscona theis (Walchenaer) Male
80. Poltys nagpurensis Female
81. Poltys sp. nov Female
82. Zygiella indica Tikader and Bal Female
83. Zygeilla indica Tikader and Bal Male
84. Zygeilla melanocrania (Thorell) Female
85. Zygeilla melanocrania (Thorell) Male
86. Zygiella sp. nov. Female
87. Zygiella sp. nov. Female
417
Application of bioinformatic tools in the analysis

of spider venom
Paweł Siuda
Adam Mickiewicz University, Poznan, Poland, kainael.ps@gmail.com
Bioinformatics is a discipline dealing with the use of Information

Technology and computer science to solve a wide variety of biological
problems. Using computers makes much easier the analysis of the large
quantitative data sets, which requires complicated mathematical calculations,
and inserting them in open-access databases too. Bioinformatics possesses many
research areas, for example the analysis of the genes expression, the proteins
expression and the genomes annotations. Other major research areas are the
protein structure prediction and the computational evolutionary biology.
Spider venoms are mixtures of different compounds, among other
peptides and proteins. Based on the amino-acid sequences, with the use of
bioinformatics tools such as SwissPDBViewer or PyMOL, we can create tertiary
structure models. This procedure will allow to detect the protein active center
and to learn about its biological functions.
Multiple sequence alignments of proteins, using for example Clustal,
allow us to create a phylogenetic tree, which will show the inferred evolutionary
relationships among various species of spiders. It can be useful for spiders
systematic.
To sum up, bioinformatics allows gathering more accurate knowledge
about spiders, mainly from the biochemical point of view.
418
Harvestmen (Arachnida: Opiliones) from Talysh, Azerbaijan

Natalya Yu. Snegovaya
Institute of Zoology, Baku, Azerbaijan, snegovaya@yahoo.com
The Talysh is a part of Azerbaijan territory (38°24'N - 39°22'N, 47°58'E -

48°52'E), with an area of 5370 km2. On the North it is bordered by the Mugan
steppe, on the East by the Caspian Sea and on the South and West by Iran.
Before the current research, only 12 species of Opiliones were known
from there (Morin 1937; Roewer 1917, 1923, 1950; Staręga 1966, 1978;
Martens 2006): Acropsolilio talischensis Morin, 1937, Opilio coxipunctus
(Sorensen, 1912), O. ejuncidus Thorell, 1876, O. lepidus L. Koch, 1878, O.
consputus (Simon, 1895), O. pallens Kulczyński, 1901, Zacheus bispinifrons
Roewer, 1911, Platybessobius caucasicus Šilhavý, 1966, Paranemastoma filipes
(Roewer, 1919), Phalangium punctipes (L. Koch, 1878), Mediostoma variabile
Martens, 2006, M. nigrum Martens, 2006. Most of Morin's determinations were
incorrect, however it is impossible to check all his identifications, because the
materials were lost during the World War II.
As a result of current research, two species listed above have been
confirmed (P. caucasicus, P. filipes, Ph. punctipes, M. variabile), two species
recorded as new for the Azerbaijan (Zachaeus birulai Redikorzev, 1936,
Dicranolasma ponticum Gruber, 1998 (earlier incorrectly determined as D.
giljarovi), two species have been reported as new for this territory (Opilio
parietinus (De Geer, 1778), O. lederi Roewer, 1911) and six species
(Phalangium armatum Snegovaya, 2005, Ph. staregai Snegovaya, 2005, Ph.
zuvandicum Snegovaya, 2005, Rilaena azerbaijanica Snegovaya, 2007, R.
talyshica (Snegovaya, 2007) comb. nov., Homolophus azerbaijanicus
Snegovaya, Staręga, 2008) described as new for science (Snegovaya 1999, 2004,
2005, 2007, Snegovaya & Staręga 2008).
419
Body size distributions of epigeic spider communities along

an environmental gradient
Marzena Stańska1, Izabela Hajdamowicz1 & Ulrich Werner2

1
Department of Zoology, Institute of Biology, University of Podlasie, Siedlce, Poland,
stanska@ap.siedlce.pl, hajdamo@ap.siedlce.pl
2
Department of Animal Ecology, Institute of Ecology and Environmental Protection,
Nicolaus Copernicus University of Toruń, Poland, ulrichw@umk.pl
Body size distributions of spiders in natural habitats are still not well
understood. In the present study we particularly aim at analyzing changes of
body sizes along environmental gradients (warm - dry to moist - cool) that might
indicate changes in prey size and availability. We studied spider communities at
the Bug river valley in the eastern Poland. The Bug valley represents well
preserved European rivers. We trapped spiders within four differed habitats: a
riparian forest, fresh meadow, psammophilous grassland and mesoxerophilous
grassland.
Material was collected from April to November 2007 with pitfall traps,
and preserved in 75% alcohol. From each of the 12 analyzed spider assemblages,
we assessed total length and cephalothorax width for both sexes of each species.
We used Ellenberg indicator values (modified after Zarzycki 2002) for habitat
classification.
Preliminary results show the decrease in body size from warm and dry to
cool and moist habitats.
420
Female cryptic choice and a cost of simultaneous polyandry

Jeffrey A. Stoltz & Maydianne C.B. Andrade
Department of Biological Sciences, University of Toronto Scarborough, Canada
Introduction
Conflict between the sexes can lead to traits that bias the outcome of
reproduction in favour of one sex while imposing costs on the other. There are
numerous costs associated with polyandry for females (Chapman et al. 1995),
however, these costs appear to be offset in some taxa by benefits that elevate the
fitness of females that mate multiply compared to those that do not. Genetic
diversity may be a particularly important indirect benefit of polyandry if
offspring disperse to unpredictable habitats (Garant et al. 2005).
If variation in the paternity of offspring affects females’ fitness, then there
can be selection on females to shape paternity (Jennions & Petrie 2000)
independent of mating decisions, leading to post-copulatory sexual selection
(cryptic choice, Eberhard 1996). The two independent sperm storage organs of
females in many spider species allows laboratory manipulations that segregate
sperm of competitors in different spermathecae. This can isolate effects of
cryptic choice on variation in sperm use, since physical separation removes the
possibility of direct sperm competition. This provides a rare opportunity to test
for evidence of biases in sperm use by females (Eberhard 2005).
We staged matings between females and pairs of males of the Australian
redback spider (Latrodectus hasselti Thorell), with re-mating intervals
mimicking those expected in nature. There are two natural windows of
opportunity for mating by female redbacks. First, during and shortly after a
female’s first copulation, active sex pheromones on the web may lead to the
attraction of rival males and possible re-mating (Stoltz et al. 2007). Webs are
largely dismantled by males during courtship however, and the silk females use
to rebuild them no longer contains pheromones (Stoltz et al. 2007). Second, later
in the reproductive season, females re-advertise their receptivity by resuming
pheromone production, providing a second window of opportunity for mating
(Peramalpadas et al. 2008). It is not known how the interval between copulations
affects paternity or potential costs and benefits of polyandry, but such effects
could determine female and male fitness in nature.

Each female was mated to a total of two males, each of which was allowed
a single copulation. In a simultaneous mating treatment, the rival was allowed
to mate within 24 hours of the first copulation, whereas in a delayed mating
treatment rivals mated approximately 60 days after the first copulation.
Paternity was assessed using a standard sterile male technique (Boorman &
Parker 1976).
421
Twenty females were mated in the simultaneous treatment. Seventy-six

females were placed in the delayed treatment as we anticipated that mortality
would reduce our sample size prior to the second mating. After mortality, our
data set included twenty-two delayed females in which the treatment was
completed. We collected egg sacs as they were made throughout the female’s
lifetime, and noted female post-mating longevity. Egg sacs were opened after 15
days, the eggs counted and classified as developed (spiderlings visible, fathered
by the normal male) or undeveloped (fathered by the irradiated male, see Snow
& Andrade 2005).
Results
Paternity. There was a significant effect of timing of insemination
(F1,37=9.53, p<0.001) on paternity. In the simultaneous treatment, average
paternity was mixed in most cases (mean P 2=49%, 95% CI: 37-61%) with P2
(paternity of second male) values between 20-80% in 79% (15/19) of the trials.
In the delayed treatment, after the second copulation the second male fathered
most offspring (mean P2=71%, 95% CI: 62-82%) with P2 values over 80% in
45% (10/22) of our trials. Thus, in the delayed treatment, although the first male
fathers all offspring produced in the first 2 months (prior to the second mating),
the 71% paternity of second males after this time reduced the overall first male
paternity to 47%., which is not significantly different from the 50% paternity
seen in the simultaneous treatment (t22=-0.37, p=0.71).
Longevity. To see whether there was a longevity cost of polyandry
compared to monandry, we compared survivorship of females in each treatment
in the 60 days following the first mating trial (prior to the second mating in the
delayed treatment). Delayed females (which had only mated once) had
significantly higher survivorship than simultaneous females (Wald=4.82, d.f.=1,
n=75, p=0.03). To see whether the cost of polyandry depended on the mating
interval, we also compared longevity of only those females that survived through
the first 60 days of the experiment (in this test, delayed females had also mated
a second time). Females with a delayed second mating lived significantly longer
(145+10 days) than those that mated with two males in quick succession
(simultaneous longevity: 104±15 days; ANCOVA F1,32=10.53, p=0.003).
Discussion
Our results show that, in redback spiders (1) females bias paternity to
favour genetic diversity of offspring, and the mechanism is selective sperm use
of sperm stored in different spermathecae, (2) polyandry decreases longevity of
females and (3) longevity costs of polyandry depend on the mating interval. Our
paternity data show that the overall effect of sperm use is that females roughly
equalize average paternity of the two males across the total number of offspring
produced in both simultaneous and delayed trials. This entails a shift from sperm
mixing to last-male sperm precedence when re-mating is delayed. This suggests
cryptic female choice for novel males. Moreover, our results show the timing of
female multiple mating has serious implications for female longevity and fitness.
Females that had simultaneous inseminations by two males had significantly
422
reduced longevity compared to females that had a two-month delay between

rival male inseminations. Thus, while genetic diversity may benefit females,
they should attempt to delay secondary matings. Strong support for the genetic
diversity hypothesis is provided by our results in the delayed re-mating treatment
where females produced a number of offspring with one male before mating
with a new male. Here, balanced paternity from both males would require an
increase in sperm use favouring the novel male. The shift to 71% paternity of the
second male at this point functionally equalized paternity across the two males.
Thus, across the female’s lifetime production of offspring paternity is equal for
both males.
Polyandry was clearly costly, but longevity costs depended on the timing
of a rival males’ copulation. Females that mated simultaneously with two rival
males had a significant reduction in lifespan relative to that of females that
mated with each after a two month delay. This may be caused by cumulative
effects of deleterious substances passed in the ejaculate. In redbacks copulation
typically lasts approximately 20 minutes but all of the sperm is transferred
within the first 5 minutes (Snow & Andrade 2005) suggesting ample time for the
transfer of accessory ejaculatory substances (Eberhard 1996). Females’ ability to
neutralize harmful chemicals contained within the ejaculate may depend on
dose. We propose females may be able to negate effects after a single
copulation, but not after two copulations in rapid succession. Interactions
between the dosage of seminal products and female fitness have only recently
been investigated directly (Radhakrishnan & Taylor 2007).
Here, we provide evidence that females exercise control over the outcome
of paternity by selectively using the sperm of novel males. However polyandry
can have costs in the form of reduced longevity when multiple mating occurs in
rapid succession. In comparison a delay in re-mating may allow time to recover
from the harmful effects of substances transferred during copulation. Such
considerations have important implications for female mating strategies dictating
the timing of mate attraction that optimizes fitness.
References
Boorman E. & Parker G.A. 1976. Sperm (ejaculate) competition in Drosophila
melanogaster, and the reproductive value of females to males in relation to
female age and mating status. Ecological Entomology, 1: 145-155.
Chapman T., Liddle L.F., Kalb J.M., Wolfner M.F. & Partridge L. 1995. Cost of
mating in Drosophila melanogaster females is mediated by male
accessory gland products. Nature, 373: 241-244.
Eberhard W.G. 1996. Female control: sexual selection by cryptic female choice.
Princeton, New Jersey: Princeton University Press.
Eberhard W.G. 2004. Why study spider sex: special traits of spiders facilitate
studies of sperm competition and cryptic female choice. Journal of
Garant D., Dodson J.J. & Bernatchez L. 2005. Offspring genetic diversity
increases fitness of female Atlantic salmon (Salmo salar). Behavioural
Ecology and Sociobiology, 57: 240-244.
423
Jennions M.D. & Petrie, M. 2000. Why do females mate multiply? A review of
the genetic benefits. Biological Reviews, 75: 21-64.
Perampaladas K., Stoltz J.A. & Andrade M.C.B. 2008. Mated redback spider
females re-advertise receptivity months after mating. Ethology, 114: 589-
598.
Radhakrishnan P. & Taylor P.W. 2007. Seminal fluids mediate sexual inhibition
and short copula duration in mated female Queensland fruit flies. Journal
of Insect Physiology, 7: 741-745.
Snow L.S.E. & Andrade M.C.B. 2005. Multiple sperm storage organs facilitate
female control of paternity. Proceedings of the Royal Society of London
Biological Sciences, 272: 1139-1144.
Stoltz J.A., McNeil J.N. & Andrade M.C.B. 2007. Males assess chemical signals
to discriminate just-mated females from virgins in redback spiders.
Animal Behaviour, 74: 1669-1674.
424
Diversity patterns of spiders along an elevational gradient

in Nelliyampathy hill ranges of the Western Ghats,
Kerala, India
Ambalaparambil Vasu Sudhikumar1, Frederik Hendrickx1,

Luc Lens1 & Pothalil Antony Sebastian2
1
Terrestrial Ecology Unit, Department of Biology, Ghent University, Belgium,
Ambalaparambil.Sudhikumar@UGent.be
2
Division of Arachnology, Department of Zoology, Sacred Heart College, Thevara,
Cochin, Kerala, India, drpothalil@rediffmail.com
Introduction
Systematics provides an essential foundation for understanding,
conserving, and using biodiversity. Yet for many groups of organisms we lack
even basic information as the identity and numbers of species found in the
Western Ghats, one of the biodiversity hotspots of the world. Spiders are an
especially diverse and ecologically important group whose ecological
dominance has been a subject of intense study. Spiders generally have humidity
and temperature preferences that limit them to areas within the range of their
physiological tolerances, which in turn makes them ideal candidates for land
conservation studies. Hence there is an urgent need to provide taxonomic
resources for groups from tropical ecosystems in view of current global
biodiversity crisis. A more complete inventory of spiders is essential to advance
understanding of their ecology, evolution, and behaviour, and to take full
advantage of their demonstrated value in conservation priority setting,
biomonitoring and biological control.
During the course of this study, a quantitative survey of spiders in the
Nelliyampathy hill ranges of the Western Ghats was attempted along an
elevation gradient, with an aim to investigate the patterns of spider species
richness along the elevation gradient.

The Western Ghats, running parallel to India's western coast, about 30 to
50 kilometres inland between the latitudes 8ºN to 20ºN and longitudes 73ºE to
77ºE, is one of the world's ten "Hottest biodiversity hotspots" and at least 325
globally threatened species occur here. Characteristic biota of the Western Ghats
has attracted the attention of researchers for more than a century. Due to its
fascinating plate tectonic history, the Western Ghats is considered as an
“exchange spot” of biotic elements between Africa and Southeast Asia.
Nelliampathy hill ranges (1033'45"-1032'34"N, 7638'27"-7646'39"E) are a
part of Peechi-Vazhani wild life sanctuary of the Western Ghats, located at an
altitude of 467 m to 1572 m above sea level, spreading over 82 km2. The
principal reason for selecting this as the study site is that this protected biota
425
represents different forest types viz., tropical evergreen forest, moist deciduous
forests, shola forest and thorny scrub forest as their primary vegetation type.
Spiders were collected at a weekly interval for one month from 5th
December 2009 to 10th January 2010 at three principal localities along an
elevation gradient. The inventories were conducted at the following sites and
habitats:
1. Elevation zone 515-575 m a.s.l. (1033'36.8"N, 7643'02.7"E, evergreen forest),
2. Elevation zone 900-960 m a.s.l. (1032'01.2"N, 7640'51.8"E, evergreen forest),
3. Elevation zone 1325-1375 m a.s.l. (1032'28.4"N, 7644'15.1"E, moist deciduous
forest).
The random transect method was used for spider sampling. This technique
involves a combination of four collection methods - ground hand collection,
aerial hand collection, beating and sweeping. Time was used as a measure of
sampling effort to make the methods comparable. One sample unit equalled
continuous one hour during which all spiders encountered were collected. The
collected specimens were preserved in 75% alcohol in separate flat bottomed
tubes with labels containing information regarding the collection.
Complementarity and overlap of the spider assemblages at different
elevations were assessed using distinctness and beta-diversity indices.
Complementarity was calculated using the Marczewski-Steinhaus (M-S)
distance index: CMS=(a+b – 2j)/(a+b – j) where j=number of species found at
both elevations, a=number of species at elevation A, and b=number of species at
elevation B. CMS was chosen because of its simple and statistically valid
approach to comparing two biota, ranging from a value of 0 where there is less
distinctness between the communities, to a value of 1 when there is high
distinctness between the communities. Beta-diversity (species overlap between
elevations) was calculated by Sørensen's similarity index. β=2c/s1+s2 where,
s1=the total number of species recorded in the first community, s2=the total
number of species recorded in the second community, and c=the number of
species common to both communities. The Sørensen’s index is a very simple
measure of beta diversity, ranging from a value of 0 where there is no species
overlap between two communities, to a value of 1 when exactly the same species
are found in both communities. The number of species unique to an elevation
and the number of species shared between elevations were also compared.
Results
Taxonomic survey of the spider fauna in three different locations in the
Nelliyampathy ranges of the Western Ghats resulted in the documentation of a
total of 515 individuals of spiders belonging to 210 species, 153 genera and 37
families. The most dominant family was Araneidae with 32 species and second
dominant family was Salticidae with 27 species. Thomisidae (20), Lycosidae
(14), Theridiidae (14), Sparassidae (13) and Tetragnathidae (10) were the other
dominant families. Cyclosa mulmeinensis of family Araneidae was the
numerically dominant species (11 individuals) and the second dominant species
was Leucauge celebesiana (10 individuals) of family Tetragnathidae.
426
Among the 210 species collected, 123 species were collected from an
altitude of 515-575 m a.s.l. A total of 101 species were collected from 900-960
m a.s.l. and a total of 51 species were collected from 1325-1375 m a.s.l. Out of
the 123 species collected from 515-575 m a.s.l., 38 species were shared with
900-960 m a.s.l. and 2 species were shared with 1325-1375 m a.s.l. Out of the
101 species collected from 900-960 m MSL altitude, 6 species were collected
both from 515-575 m a.s.l. and 1325-1375 m a.s.l. and another 17 species were
shared with 1325-1375 m a.s.l. Among the 51 species collected from the topmost
elevation, 23 species shared with mid elevation and 8 species with lowest
elevation were studied. This study revealed that elevation had measurable effect
on species richness, with the number of species at three elevations being
different. The number of species unique to lowest elevation was 77 while that of
mid elevation and high elevation was 40 and 20 respectively.
Measures of distinctness (CMS) and species overlap (β) between elevations
were significantly different especially between low elevation and high elevation
samples. CMS value was 0.795, 0.821 and 0.951 respectively between elevation
1-2, 2-3 and 1-3. Value of β was 0.339, 0.302 and 0.091 respectively between
elevation 1-2, 2-3 and 1-3. This analysis revealed that the greatest species
distinctness (CMS) and the lowest species overlap (beta) was between low
elevation and high elevation compared to low elevation – mid elevation and mid
elevation – high elevation.
Discussion
This study revealed the qualitative and quantitative richness of the spider
fauna in these ranges. The 37 spider families recorded from this region represent
62% of the total families reported from India. The number of species reported is
higher than the number recorded from any other regions surveyed in India. The
species richness is very high when compared to some other regions such as
Sikkim - 55 species and Andaman and Nicobar Islands - 65 species. The present
investigation is comparable to the study conducted at Purna Wildlife Sanctuary,
Gujarat where recorded 116 species and the study conducted at Parambikulam
Wildlife sanctuary of the Western Ghats where reported 147 species, which is
the highest number of species reported from an area in India. From these results,
it can be summarized that the spider fauna of Western Ghats of Kerala is rich
and diverse when compared to any other region in India. Because of the complex
interaction of various climatic factors like high rainfall and humidity, with
diverse topographical features, this region possesses many smaller but diverse
environmental niches that can support a diverse spider fauna.
Analysis of the diversity pattern revealed that diversity varied between
elevation gradient, indicating unique species compositions at each elevation.
Among the three elevations studied, the lower elevation recorded the highest
value of diversity. In this elevation, tropical evergreen forests form the chief
vegetation type, which is more complex in nature compared to other vegetation
types. The higher elevation recorded lower diversity and species richness.
Compared to low elevation and mid elevation, this area is occupied by moist
deciduous forest, which supports less spider diversity than evergreen forest. This
427
supports the correlation that exists between spider species diversity and habitat
types. A fundamental characteristic of mountain ecosystems is the drastic change
in vegetation as well as in climate from the base to the summit. Elevation
gradients create varied climates, along with resultant soil differentiation that
ultimately promotes diversification of both flora and fauna. The elevation
patterns of species richness are a consequence of many interacting factors, such
as plant productivity, competition, geographical area, historical or evolutional
development, regional species dynamics, regional species pool, environmental
variables, and human activity.
Despite being one of the most diverse groups of organisms, spiders have
largely been ignored by the conservation community and taxonomists alike.
Many threats to spider diversity have been documented, including habitat loss
and degradation due to deforestation, agriculture, grazing and urbanisation. The
major obstacle for spider conservation is an absence of public support, arguably
due to fear and ignorance. Conservation of spiders thus necessitates a greater
understanding by the general public, scientists, land use managers and
conservationists about the importance of bringing these fascinating creatures
onto the conservation radar screen.
428
Spider fauna from proposed Mahendri Wild Life Sanctuary
Kondulkar Sunil1, Dinesh Joshi2, Dinesh Vankhede3

& Atul Bodkhe4
1
M.F.M., Warud, India, sunil_kondulkar@rediffmail.com,
2
SGB Amravati University, Amravati, India,
3
Indian Society of Arachnology, Amravati, India,
4
JDPS Mahavidyalaya, Daryapur (Distt. Amravati, India
Introduction
Mahendri is a dry deciduous forest in Satpuda (India). It represents mixed
flora with dominance of tall trees and streams. It is a tiger land and has a good
herbivore population. Grazing is the threat for the existing flora and fauna. It lies
on 21°29'33.08"N and 78°20'05.80"E. The spiders are recorded from different
selected habitats which include riparian habitat, grasslands, dry deciduous forest,
mixed forest with tall trees and shrubs. Survey was also carried out for ground
spiders and spiders from slow flowing shallow streams, spiders from decaying
barks of trees, debris and crevices of rocks.
Results
The study included 238 species from 24 families and 77 genera (Tab. 1).
The spider diversity was in the order of Lycosidae, Thomisidae, Araneidae,
Gnaphosidae, Oxyopidae.
Within the rich spider diversity we have noted some morphological
variation in both sexes of Nephila pilipes occurring in small isolated habitats of
Mahendri.
In males (India: Maharashtra State, Amravati District, Mahendri village,
dry deciduous to riparian) three colour morphs were observed, showing
cephalothorax yellowish red to reddish brown; abdomen faint yellowish with
single longitudinal black line - to yellow with three longitudinal black colour
stripes.
In females the metatarsus of first and fourth legs showed colour variations.
Most of them (32) had black legs, 6 specimens (same population) had distinctive
yellow band on the anterior one-third portion of metatarsus of first legs and two
yellow bands on the fourth legs. The tarsus of fourth leg also showed yellow
band at its anterior tip. Some of the females (11) showed a bunch of bushy hairs
on the metatarsus. From the same habitat we have also collected females (5)
with yellow band only on the anterior tip of tarsus and not on metatarsus. These
colour morphs seem to be discontinuous from riparian to dry deciduous
ecosystems, suggesting distinctive phenotypical variation in Nephila pilipes
living in the same habitat.
429
Table 1. Spider families, genera and species recorded from Proposed Mahendri Wild
Life Sanctuary (2006-09).
Family Genus Species

1 Araneidae 10 34
2 Clubionidae 2 7
3 Corinnidae 1 3
4 Dictynidae 1 1
5 Eresidae 1 3
6 Gnaphosidae 12 25
7 Hersiliidae 1 2
8 Lycosidae 5 40
9 Miturgidae 2 5
10 Nephilidae 1 2
11 Oecobiidae 1 2
12 Oonopidae 1
13 Oxyopidae 2 23
14 Palpimanidae 1 1
15 Philodromidae 3 6
16 Pholcidae 3 5
17 Pisauridae 3 3
18 Salticidae 8 18
19 Scytodidae 1 3
20 Sparassidae 1 1
21 Tetragnathidae 3 6
22 Theridiidae 2 9
23 Thomisidae 11 35
24 Uloboridae 1 3
Total 77 238
430
The occurrence of agrobiont spiders indicates overgrazing
Csaba Szinetár1 & Ferenc Samu2

1
University of West Hungary, Savaria University Center, Zoological Department,
Szombathely, Hungary, szcsaba@bdf.hu
2
Plant Protection Institute, Hungarian Academy of Sciences, Budapest, Hungary,
samu@julia-nki.hu
In a 3-year project we have studied the effect of different grazing regimes

on the cursorial spider fauna of a dry pasture in a hilly region of Hungary. The
secondary grassland area, which had been originally forested, has been used for
pastoral farming for hundreds of years. In this system grazing is the primary
factor which keeps up the grassland. Without grazing the area would be
reforested and several protected grassland species would loose their habitat. To
study the short-term effect of different levels of grazing, we established three
treatment levels: 1) enclosed area by fencing, no grazing occurred; 2) sparsely
grazed area; 3) massively overgrazed area. In all three areas spiders were
collected by pitfall trapping and suction sampling during an early summer and
an autumn period in each year. Even during these 3 years we could observe a
quick succession in the enclosed area with litter accumulation, increase in
vegetation height and the occurrence of forest specialist spider species. The least
changes occurred in the sparsely grazed area. In the overgrazed area, in contrast,
the spider assemblage showed a significant change. Agrobiont and disturbance-
tolerant species turned up, which indicate a decrease in the conservation value of
the assemblage. These changes could be detected in the pitfall as well as in the
suction apparatus samples. Our results show that overgrazing will not only
change the physiognomy of the habitat, but its arthropod assemblages, too, and
may in the longer term lead to irreversible changes, such as desertification.
However, the lack of grazing lead to a rapid succession  another type of habitat
alteration. Therefore, finding the right balance in grazing intensity is vital if
pastoral landscapes with their exceedingly high biodiversity are to be preserved.
431
A preliminary global phylogeny of the giant goblin spiders

(Araneae: Orsolobidae)
Tamas Szuts1,*, Anthea Carmichael1, Alma Saucedo1,3

& Charles Griswold1,2,3
1
Francisco, CA, USA
2
Berkeley, CA, USA
3
Department of Biology, San Francisco State University, San Francisco, CA, USA
*
Corresponding author: tszuts@calacademy.org.
The dysderoid spider family Orsolobidae presents a strikingly

disjunct distribution at the southern end of the world. Orsolobids are
diverse in New Zealand, common in Chile, widespread in the moister
parts of Australia, and scattered among afromontane forests in southern
Africa. Extensive recent fieldwork suggests that they do not occur in
Madagascar. A global phylogeny for this family may allow us to test
alternative historical biogeographic scenarios in the southern hemisphere
including vicariance generated though the effects of continental drift on
the former Gondwanaland and austral transoceanic dispersal. We present
a multigene phylogeny for multiple orsolobid genera from Australia,
Chile, New Zealand, and South Africa, representatives of the other
dysderoid families Dysderidae, Oonopidae and Segestriidae, and
haplogyne outgroup taxa including representatives of the Tetrablemmidae
and Caponiidae.
432
The variation of some morphological structures within

Diaea s. lato (Araneae: Thomisidae) of Australia
Paweł Szymkowiak
Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, A.

Mickiewicz University, Poznań, Poland, pawel.szymkowiak@amu.edu.pl
The genus Diaea encountered in Australia presently includes 31 species.

Many species that were described in the past were classified according to such
features as external similarity, colouring and habitat, being compared against the
type species - Diaea dorsata (Fabricius, 1777) encountered in Palaearctic.
According to contemporary studies, the species within the genus are very
diversified as far as their morphological features are concerned. The most
striking differences can be observed with reference to body conformation, ocular
structures and reproductive structures. Within the discussed genus there can be
found females with a nearly spherical opisthosoma, e.g. Diaea dimidiata (L.
Koch, 1867), Diaea velata L. Koch, 1876, oval or pear-shaped opisthosoma, e.g.
Diaea blanda L. Koch, 1875, Diaea cruentata (L. Koch, 1874), Diaea
ergandros Evans, 1995, Diaea inornata (L. Koch, 1876), Diaea
kangarooblaszaki Szymkowiak, 2008, Diaea megagyna Evans, 1995, Diaea
pilula (L. Koch, 1867), Diaea prasina L. Koch, 1876, Diaea punctata L. Koch,
1875, Diaea pulleinei Rainbow, 1915, Diaea rosea L. Koch, 1875, Diaea
socialis Main, 1988, or visibly elongated opistosoma, e.g. Diaea caecutiens L.
Koch, 1876, Diaea circumlita L. Koch, 1876, Diaea rubropunctata Rainbow,
1920, Diaea tenuis L. Koch, 1875. Moreover, only in the latter group of species
the length of the prosoma slightly exceeds its width (length/width ratio > 1).
Some species are equipped with small cheliceral teeth: D. dimidiata, D.
velata, Diaea jucunda Thorell, 1881 (may be a synonym of D. adusta). There
are two types of ocular arrangements within Diaea: either narrow or wide in
proportion to the width of the prosoma. A prevailing number of species are
characterized by the trapezium of ocular area (MOA) shaped in such a way that
its width exceeds its length. In other species, as is the case in D. rubropunctata,
the proportions are just the opposite. There are also a few species with lateral
eyes arranged very wide, sitting on big tubercles stretching out of the outlines of
the prosoma: Diaea tristania (Rainbow, 1900), Diaea xanthogaster (L. Koch,
1875)). The construction of reproductive organs is the best indicator of
relationships within the genus. Among the species which belong to Diaea s. lato
there can be differentiated several dozen types of these organs. The largest
number of species have organs of two types: a) simple and pale reproductive
structures, which can be found in Diaea evanida (L. Koch, 1867), Diaea
haematodactyla L. Koch, 1875, Diaea multopunctata L. Koch, 1874, D. prasina,
433
D. punctata, Diaea punctipes L. Koch, 1875); b) complicated, distinctive

reproductive structures with a broad central hood in females and a long, massive
VTA in males, which can be found in (D. cruentata, D. ergandros, D. inornata,
D. kangarooblaszaki, D. megagyna, D. pilula, D. socialis, Xysticus periscelis
Simon, 1908 and Xysticus walesianus Karsch, 1878. The latter two species ought
to be transferred to Diaea as they evidently belong to this genus.
Many other species within the genus in question turn out to be synonyms
of others, and there are also some species which ought to be transferred to other
genera in the future, as was the case with Diaea cimicina (Thorell, 1881) and
Diaea adusta (L. Koch, 1867) (=Diaea tumefacta L. Koch, 1874=Diaea varians
Kulczyński, 1911), which were transferred to Mastira.
434
Diaea inornata L. Koch, 1876 - solving the taxonomy riddle

Paweł Szymkowiak & Agnieszka Dymek
Department of Animal Taxonomy and Ecology, Invertebrate Institute, Faculty of Biology, A.

Mickiewicz University, Poznań, Poland, pawel.szymkowiak@amu.edu.pl
Spiders of the genus Diaea belong to the family Thomisidae which

includes 2055 species within 171 genera worldwide (Platnick 2009). There are
125 species of thomisids in Australia. Diaea spiders inhabit flowers and leaves
of herbaceous plants. Their bodies are small (below 1 cm), the abdomen is oval
or round-shaped, sometimes oblate and sometimes pointed at the end. Body
colour is usually white, green or yellow while the abdomen remains whitish,
green, orange, pink, often with red, brown or orange spots or stripes.
Diaea inornata was described as Xysticus inornatus by Ludwig Koch in
1876 on the basis of one female individual encountered in Sydney. In 1966
Dondale transferred Xisticus inornatus to Diaea (new comb. Diaea inornata)
and while studying new material from ACT (Australian Capital Territory) he
completed the description of an unknown male of Diaea inornata.
Our revision has revealed that the specimens studied by Dondale (1966)
and redescribed by him under the name of D. inornata belong to a different,
probably unknown species of Diaea. Detailed studies of the holotype of X.
inornatus and other material from the Australian Museum (Sydney, Australia),
Canadian National Collection of Insects (Ottawa, Canada), Queensland Museum
(Brisbane, Australia) and the Zoological Museum (Hamburg, Germany) revealed
that D. inornata is a senior synonym of Diaea megagyna Evans, 1995.
The recent studies are the first step in establishing the taxonomic position
of several species of “pilula group”.
Diaea inornata has several characters which allow to distinguish it from other
similar species. It has a big body (especially in adult females), a high, dark
cephalothorax and a wide clypeus. The species has a sandy-coloured abdomen,
lighter than prosoma, with no pattern but with numerous grey, tiny spots
encircled by white rings and with a dark broad stripe situated posteriorly on the
ventral side of opistosoma. Females have bean shaped spermatchecae with short
reproductive ducts which are wide in the proximal part. Pedipalps are small,
with a slightly projecting cymbium. The embolus starts in a lower position and
leaves the tegulum in 170º; it circles the tegulum 1,5 times. Tibial apophysis
(VTA) is small and simple with a long seta. RTA is characterised by a
noticeably pointed tip and protruding transparent short fold in the lower position.
435
Can competing orb web spiders co-exist

in the same habitat?
Hafiz Muhammad Tahir1, Abida Butt2 & Muhammad Bilal2
1
2
Department of Zoology University of the Punjab, Lahore, Pakistan
Tetragnatha javana, Leucauge decorata, Neoscona theisi and Argeope

pradhani are common orb web spiders of rice ecosystems of central Punjab,
Pakistan. In spite of apparent competition among these spiders for resources they
co-exist in the same habitat. Present study was designed to investigate how
studied species segregate the available resources to minimize competition and
make their coexistence possible. Results of our study showed that orb web
spiders consumed same prey orders but in different proportions. Four orb web
spiders also differed in their habitat and in the prey size selection. Discriminant
function analysis also clearly separated the four species in three-dimensional
space. It is concluded that utilization of microhabitat and prey resources
differently reduces the competition among these spiders and make their co-
existence possible. Assemblage of predator species in the same habitat enhances
their biocontrol potential.
436
Relationship of web characteristics and body measures

of Leucauge decorate (Araneae: Tetragnathidae)
Hafiz Muhammad Tahir1, Abida Butt2 & Imtiaz Alam3
1
2
Department of Zoology University of the Punjab Lahore, Pakistan
3
Department of Zoology University of the Punjab Lahore, Pakistan
The study was conducted to investigate the relationships between web

characteristics and body measures of Leucauge decorata (Araneae:
Tetragnathidae). For this purpose a rice field (800 m2) in village Kirka located in
Lahore district was selected. The data of spider’s webs and prey was collected in
September 2007 and 2008. Most of the L. decorata constructed inclined webs at
the height that ranged from 40 cm to 123 cm above ground. Spiders recorded
from the higher webs (81-120 cm) were larger and heavier than the spiders
recorded from the lower webs (40-80 cm). Time required to complete a web was
1.00±0.21 hour. The average diameter of the web was 25±6.8 cm. Principal
component analysis (PCA) did not separate the collected spiders on the basis of
capture area (web character) and carapace width (body measure). Most of the
prey items recorded from the webs belonged to Diptera, Homoptera and
Lepidoptera. Prey types and their number collected from the webs of different
heights varied significantly. Capture area of L. decorata web showed a positive
correlation with carapace width and body weight. Web characteristics (i.e.,
number of radii, number of spirals and mesh height) were not correlated with
any of the body measures (i.e., carapace width, total length and wet weight). For
prey capture L. decorata construct webs of different sizes and at different
heights but always maintain the basic web architecture (i.e., number of radii,
number of spirals and mesh height).
437
Do Ricinulei possess a chemical defence mechanism?

Giovanni Talarico
Max-Planck-Institut für Chemische Ökologie, Evolutionäre Neuroethologie, Jena,

Germany, g.talarico@gmx.net
Little is known about the behaviour and morphology of the enigmatic

tropical hooded tick-spiders (Ricinulei, Arachnida). According to a single report
from the available literature, ricinuleids shall be capable of spraying a clear
liquid out of their anal opening over a distance of several centimetres.
Investigations of the ricinuleid alimentary system by means of light microscopy,
electron microscopy and X-ray micro computer tomography revealed that the
posterior part of the ricinuleid midgut tube shows peculiar modifications, which
may explain the behaviour mentioned above. The posterior midgut tube can be
bladder-like dilated and its epithelium has glandular character in defined areas.
The products of this rectal gland are stored in large compartments, which are
formed by apical extensions of the epithelial cells. It is supposed that the
glandular products are released into a central passage and then, aided by
contractions of a well developed muscularis, are pressed through the cuticle-
lined hindgut towards the anal opening. However, the rectal gland and its
products may have a physiological and/or defensive function.
438
DNA barcoding in spiders: exuviae for species

identification and matching male-female of dimorphic
species
Trina V.Z.Y. Tan1,3, David Court2, Laura-Marie Y.L. Yap1,
Sujatha Narayanan1, Youguang Yi1, Rudolf Meier1 & Daiqin Li1,4
1
2
The Raffles Museum for Biodiversity Research, Department of Biological Sciences,
National University of Singapore, Singapore
3
trinatan@nus.edu.sg
4
dbslidq@nus.edu.sg
DNA barcoding as a tool for species identification works by analyzing

sequence similarity in a 648-bp region of the mitochondrial gene, cytochrome c
oxidase 1 (COI), which also serves as a DNA barcode. In addition to its main
function in species identification, DNA barcoding can also facilitate species
discovery, and is said to be a valuable tool in a time of the taxonomic crisis.
DNA barcoding has shown its usefulness in highlighting cryptic species,
associating morphological disparate life stages of a species, matching male-
female dimorphic species and identifying materials derived from endangered
species. For many species, identifying species using DNA sequences may also
be faster than waiting for experts and it requires less expertise compared to using
traditional taxonomic techniques. In traditional taxonomy, the identification of a
species requires the use of morphological keys. These are tedious, effective only
across homologous semaphorants and sex, subjective and can be difficult to use
de to complex genitalia, sexual polymorphisms and phenetic plasticity.
However, the debate on the accuracy of DNA barcoding is still ongoing and
studies on various animal taxa yielded conflicting results. This study used the
species-rich arthropod group of spiders, serving as a useful framework to test the
validity of DNA barcoding and address five main issues. Firstly, we tested the
variability of the COI gene for DNA barcoding. The intraspecific and
interspecific genetic distances were checked to determine if a clear separation
existed between the two. Secondly, we examined the effectiveness of DNA
barcoding in spider species identification, using 711 species/2235 sequences
from Genbank and 257 new sequences, which were generated from 71 species of
spiders collected in SE Asia. Besides the inclusion of a wider sampling of
spiders, this study attempted to test DNA barcoding with morphologically
characterized 782 species. Through this study, we intended to fill in the
knowledge gap for South-east Asian spiders as well. Thirdly, we evaluated the
practicality of implementing DNA taxonomy. Fourthly, we assessed the
439
feasibility of using juvenile spider exuviae (i.e., cuticular moults) for species
identification in DNA barcoding. Finally, we generated preliminary hypotheses
for a crab spider species (Thomisidae) using techniques involving DNA
sequences. We found that distinguishing spider species was not as unambiguous
or straightforward as has been proposed. About 8% of all species shared
identical barcodes and the variability between species overlapped with the
differences within species. However these did not translate to an overall low
identification success. Considering only those species with multiple sequences,
92% of the sequences were correctly identified using “best match”. We also
demonstrated that spider exuviae (moults) can be used to identify juvenile
spiders. The logistical challenge of extracting DNA from exuviae and ensuring
no degradation or contamination can be met. Of nine exuviae from eight species
representing three families (Araneidae, Nephilidae and Salticidae), eight exuviae
were correctly identified. The DNA barcoding of exuviae is shown to be a non-
lethal, non-invasive method for juvenile spider identification and is especially
useful when the species is rare and endangered or adults are needed for
behavioural research. DNA barcoding as a tool to match male-female of
dimorphic species were tested using a pair of sexually dimorphic crab spiders
whose species is currently unknown. DNA barcodes together with somatic
morphological characters and ecological data provided an initial hypothesis
about the crab spider species pair. All together, this study shows that DNA
barcoding is of limited use when sampling is weak, but it can nevertheless be
useful as long as one is aware of the strengths and weaknesses. DNA barcoding
is useful because many routine identifications can be done by non-experts.
440
Attention please! Tuft decorations and barrier webs

of the spiny spider Thelacantha brevispina
may function to warn off avian predators
Huei-Jen Tseng
Department of Life Sciences, National Chung-Hsing University, Taiwan
Most studies on spider web decorations have focused on the cruciate

decorations of Argiope spiders. Relevant studies on silk tuft web decorations
built by spiny spiders are few. The E Asian spiny spiders Thelacantha
brevispina place silk tuft decorations on a 3D barrier web. In this study, we
investigated how barrier webs and silk tuft decorations are involved in prey
capture and predator defence of T. brevispina. The field experiments were
conducted in a tropical island in southern Taiwan. The presence of barrier webs
and the colour signal of silk tuft decorations were manipulated by painting tuft
decorations so they are indistinguishable to the background, and removing the
barrier web with burning incense. Therefore there are two treatments: (1)
decorations painted, (2) no barrier, (3) control. The interaction between
Thelacantha brevispina and its prey and predators were recorded by video
cameras. Interception of prey was higher in webs without barriers than the
control webs, but interception of prey in painted decorations was similar to
control webs. Silk tuft decorations on barrier webs, therefore, come at a foraging
cost. For all treatment groups wasps approached but never attacked the spiny
spiders. However, bird predation occurred on spiders in webs with painted
decorations. These results suggest that wasps are not the major predator of spiny
spiders and silk tuft decorations and barrier webs of T. brevipina function to
warn off avian predators.
Keywords: barrier web; Thelacantha brevispina; silk tuft decoration.
441
Factors shaping the lure signal design of

a nocturnal orb web spider
I-Min Tso* & Cheng-Hui Lai
Department of Life Science, Tunghai University, Taiwan

*
Corresponding author: spider@thu.edu.tw
Some predators use deceptive visual signals to lure prey and therefore the
design of visual signal is very important. Results of previous studies on diurnal
spiders show that the bright body colouration of some orb weaving spiders are
attractive to insects. However, the bright colour signals of spiders were also
found to be attractive to predators. Therefore, the current body colouration
pattern of numerous diurnal orb-weaving spiders might reflect a trade-off
between pressures of attracting prey and avoiding predators. Many orb-weaving
spiders hunt exclusively nocturnally and previous studies showed that some also
use visual signal to lure prey. Currently, the factors determining the visual lure
signal design of nocturnal spiders are still not clear? In this study, we evaluated
factors influencing nocturnal visual signal design by studying a nocturnal orb
weaving spider, Necoscona punctigera. We first measured reflectance spectra of
various kinds of collared cardboard papers and used them to make dummies. The
chromatic property, signal intensity and signal form of dummies were
manipulated to evaluate these treatments’ effects on nocturnal insect
attractiveness. The results of field experiments showed that webs containing
dummies with standard signal form attracted significantly more prey than webs
without dummies, indicating that chromatic signal alone is sufficient in
attracting nocturnal prey. When the signal arrangement pattern or intensity of
dummies was changed, their prey attraction rate did not significantly from that
of standard dummies. However, the prey attraction rate of dummies with signal
chromatic properties changed was significantly lower than that of standard
dummies. These results show that current body colouration pattern of N.
punctigera is a very effective visual lure. Why this nocturnal orb spiders’ luring
signal is so attractive and what important attributes of prey’s resources does this
signal mimic awaits further study.
442
A circular overlap resulting from a cascade of numerical

changes of chromosomes in the harvestman
Gagrellula ferruginea (Scelerosomatidae: Gagrellinae)
Nobuo Tsurusaki & Minako Kawaguchi
Laboratory of Biology, Faculty of Regional Sciences, Tottori University, Japan,

ntsuru@rstu.jp
When two populations representing both ends of a series of populations

along an extensive cline somehow meet in a restricted area without any
indication of interbreeding, we call the phenomenon “circular overlap” and the
series of interbreeding populations “ring species”. Examples of such ring species
include Greenish Warbler complex Phylloscopus trochiloides in Continental
Asia, Herring Gull Larus argentatus in the North Atlantic, and salamander
Ensatina in western United States (Mayr 1969, Ridley 1996, Coyne & Orr 2004,
Barton et al. 2007). These cases of ring species show that geographic variation
within a species can be extensive enough to produce a new species. However,
circular overlap has not been so extensively reported in invertebrates. Moreover,
ring species that involves chromosomal changes has not been known at least in
animals. We will report a possible case of circular overlap involving a cascade
of numerical change in chromosomes in a Japanese harvestman Gagrellula
ferruginea (Loman, 1902) (Opiliones: Sclerosomatidae: Gagrellinae).
Gagrellula ferruginea is a common species of harvestmen widely
distributed in the main three islands (Honshu, Shikoku, and Kyushu) of Japan.
This species is univoltine and overwinters as eggs and adults can be abundantly
found from July to October on herbs, shrubs, or on tree trunks. The species
shows enormous geographic differentiation in both external morphology and
chromosomes, and a total of 11 major geographic races have been recognized on
the basis of colouration and markings of the body and the diploid chromosome
number varies enormously from 10 to 22. Geographic variation of external
morphology (colouration) and that of chromosomes in this species do not
coincide one another in general. For example, the number of chromosomes
varies from 2n=14 to 20 only within the distributional range of the Kinki race
defined by colouration that occupies northern Kinki District of Honshu.
We surveyed chromosomes of the species for a total of 18 populations in
northern Shikoku (Ehime and Kagawa prefectures), where colouration is rather
uniform. The results obtained from the survey were as follows:
1) The number of chromosomes varies from a mountain mass to next in NW
Shikoku (Ehime Prefecture) where the species usually occurs only in the upper
parts of mountains higher than ca. 700 m: Is. Nakajima 2n=14; Mt. Takanawa
and Mt. Fukumi 2n=12/13/14; Mt. Ishizuchi and Mt. Kamegamori 2n=18,
Tengu Highland 2n=14, Ônogahara Highland 2n=16, etc.);
2) The diploid number of chromosomes increases from 2n=16 to 20 in NW
Shikoku (Kagawa Prefecture), through intermediate populations polymorphic
443
for the number, although populations in the easternmost part of the prefecture
including Is. Shôdo invariably show 2n=16;
3) Populations in the western part of Kagawa Prefecture are polymorphic
(2n=12/13/14) or monomorphic (2n=12).
Interestingly, distributional range of the 2n=12 populations in western part
of Kagawa Prefecture overlaps that of the 2n=20 populations which represent
westernmost end of the series of successive changes in chromosome number
from 2n=16 in easternmost part of Kagawa Prefecture, at Mt. Ryûô, without any
indication of hybridization (Eight of 10 males from a site near the summit of the
mountain showed 2n=12, while the other two were of 2n=20). This means that
the 2n=12 population is reproductively fully isolated from the 2n=20 population.
However, both the forms intergrade one another through populations with
intermediate numbers (2n=14, 16, 18), making narrow zones of contact in the
areas where two neighbouring populations with different chromosome numbers
abut in San-yô side of Chugoku District (Okayama and Hyogo Prefectures),
Honshu. Thus, the overlap of the distributional ranges of 2n=12 and 2n=20
populations with a series of intermediate populations connecting both ends can
be safely considered a case of circular overlap that arose from successive
increase (or decrease) of chromosome numbers.
References
Barton N.H., Briggs D.E.G., Eisen J.A., Goldstein D.B. & Patel N.H. 2007.
Evolution. Cold Spring Harbor Laboratory Press, Cold Spring Harbor,
New York, 833 pp.
Coyne J.A. & Orr H.A. 2004. Speciation. Sinauer Associates, Sunderland, MA,
545 pp.
Mayr E. 1963. Animal Species and Evolution. Belknap Press of Harvard
University Press, Cambridge, Mass., 797 pp.
Ridley M. 1996. Evolution. 2nd ed. Blackwell Science, Inc., Cambridge, Mass.,
719 pp.
444
Recent expansion of distributional range of

Phalangium opilio, a presumably introduced harvestman
in Hokkaido, Japan, with notes on the chromosomes
and male dimorphism
Nobuo Tsurusaki & Koji Takenaka
Laboratory of Biology, Faculty of Regional Sciences, Tottori University, Japan,

ntsuru@rstu.jp
Phalangium opilio (Phalangiidae) is a common species of harvestman

occurring in temperate to cool-temperate zones of Eurasia, especially in Europe,
and North America. This species favours open habitats such as meadows,
roadside, or gardens around human habitation (Spoek 1963, Clingenpeel &
Edgar 1966). Probably due to this anthropophilous habit of the species, this
species has also been recorded from New Zealand as introduced species (Forster
1962, Gruber & Hunt 1973, Vink et al. 2004). Furthermore, there is even a doubt
whether North American populations of the species are of native (Gruber &
Hunt 1973, Martens 1978).
Occurrence of this species in Japan was first confirmed in 1980 on the
basis of a few specimens (1 male, 1 female, and two juveniles) found from a site
along roadside of expressway (Dô-ô Expressway) under construction in the
Experiment Forest Station of Hokkaido University at Tomakomai, Hokkaido,
where timothy grass, red clovers, Kentucky blue grass, and etc., were planted for
the protection of the roadside slope (Suzuki & Tsurusaki 1981). However, no
additional records of the species had been reported after the first finding, except
for a juvenile found in the Engaru Station of Japan Railroad in the NW
Hokkaido in 1985. Phalangium opilio is known also from Siberia, Far East
(Gritzenko 1979), Sakhalin (Tsurusaki unpubl.), Moneron Island (Crawford &
Marusik 2006), and southern Kurils (Tsurusaki unpubl.). However, known
ranges of the species in these areas, which are adjacent to Hokkaido, also seem
to be limited to urban areas along Trans-Siberian Railway and islands suitable
for residence, indicating that populations of the species in these areas as well as
those in Hokkaido may have been established with the help of human agency.
In summer of 2003, one of the authors, Tsurusaki, found numerous
specimens of this species from following two sites in Hokkaido: the campground
of the Taisetsu Lakeside Park in central Hokkaido and the Wattsu Service Area
of the Dô-ô Expressway in Kita-Hiroshima City. To confirm whether this
species became more widespread than in the 1980s, we surveyed 40 sites in and
around Sapporo and Kita-Hiroshima City along expressway or urban parks for
the occurrence of the species in 2006. As a result, we newly found P. opilio from
12 sites in Kita-Hiroshima City, Sapporo City, and Iwamizawa City, including
two sites from which no specimens of the species were found in the former
collectings in 1980s by Tsurusaki.
445
It has been known that Phalangium opilio shows geographic variation in

the number of chromosomes. Diploid number of chromosomes of the species is
24 in France (Juberthie 1956) or 24/26 in Mainz, Germany, whereas the number
is 32 in Petersburg, Russia (Sokolow 1930) and Idaho in the United States of
America (Tsurusaki and Cokendolpher 1990; Tsurusaki 2007). Examination of
chromosomes of this species from two populations (Lake Taisetsu and Wattsu
Service Area) in Hokkaido showed invariably 2n=24. The result indicates that
these Japanese populations of the species have originated from Europe.
This species is well known for its conspicuous sexual dimorphism in size
and shape of chelicerae and palpi (Spoek 1963, Martens 1978, Willmart et al
2006). Distal segment of male chelicera is well developed and usually bears
hornlike protuberance dorsally at its basal end and male palpi is elongated in
response to development of the chelicerae. It has also been reported that size and
shape of male chelicerae are extremely variable and those of some males are
rather normal in shape (Spoek 1963, Martens 1978). Measurements of 10
characters for a total of 112 specimens (102 males and 10 females) collected
from a single population in Sapporo revealed that males of this species are
clearly dimorphic in size and shape of chelicerae and palpi. To confirm whether
this male dimorphism in the species relates to alternative mating strategies of
males such as territorial males versus sneakers, we conducted field observation
and some experiments in laboratory. However, unfortunately, no conclusion was
available for the subject due to scarcity of data.
References
Clingenpeel L.W. & Edgar A.L. 1966. Certain ecological aspects of Phalangium
opilio (Arthropoda: Opiliones). Papers of the Michigan Academy of
Science, Arts and Letters, 51: 119-126.
Crawford R.L. & Marusik Y.M. 2006. Harvestmen (Arachnida: Phalangida or
Opiliones) of Moneron Island. In: Flora and fauna of Moneron Islands of
Moneron Island (Materials of International Sakhalin Island Project).
Vladivostok: Dalnauka, pp. 196-201.
Forster R.R. 1962. A key to the New Zealand harvestmen Part 1. Tuatara, 10:
129-137.
Gritzenko N.I. 1979. Materials on the Opiliones fauna from Primorye region. In:
Ler P.A. (ed.), Terrestrial Arthropoda of the Far East. Academia Nauk
SSSR, Vladivostok, pp. 124-132.
Gruber J. & Hunt G.S. 1973. Nelima doriae (Canestrini), a south European
harvestman in Australia, and New Zealand (Arachnida, Opiliones,
Phalangiidae). Records of the Australian Museum, 28: 383-392.
Juberthie C. 1956. Nombres chromosomiques chez les Sironidae, Trogulidae,
Ischyropsalidae, Phalangiidae (Opiliones). Les Comptes rendus de
l'Académie des sciences, 242: 2860-2862
Martens J. 1978. Weberknechte, Opiliones. Die Tierwelt Deutschlands. 64 Teil.,
Gustav Fischer, Jena, 464 pp.
446
Sokolow I. 1930. Untersuchungen über die Spermatogenese bei den Arachniden.

IV. Über die Spermatogenese von Phalangiden (Opiliones). Zeitschrift
für Zellforschung und Mikroskopische Anatomie, 10: 164-194, pl. 6-8.
Spoek G.L. 1963. The opilionida (Arachnida), of the Netherlands. Zoologische
Verhandelingen (Leiden), 63: 1-70.
Tsurusaki N. 2007. Chapter 6. Cytogenetics. In: Pinto da Rocha R. et al. (eds.),
The Harvestmen: The Biology of Opiliones. Harvard University Press,
Cambridge, Massachusetts, 597 pp.
Tsurusaki N. & Cokendolpher J.C. (1990) Chromosomes of sixteen species of
harvestmen (Arachnida, Opiliones, Caddidae and Phalangiidae). Journal
of Arachnology, 18: 151-166.
Suzuki S. & Tsurusaki N. 1983. Opilionid fauna of Hokkaido and its adjacent
areas. Journal of the Faculty of Scence, Hokkaido University, Zool., 23:
195-243.
Vink C.J., Teulon D.A.J., McLachlan A.R.G. & Stufkens M.A.W. 2004. Spiders
(Araneae) and harvestmen (Opiliones) in arable crops and grasses in
Canterbury, New Zealand. New Zealand Journal of Zoology, 31: 149-159.
Willemart R.H., Farine J-O., Peretti A.V. & Gnaspini P. 2006. Behavioral roles of
the sexually dimorphic structures in the male harvestman, Phalangium opilio
(Opiliones, Phalangiidae). Canadian Journal of Zoology, 84: 1763-1774.
447
Higher level phylogenetic study of micronetine spiders

(Araneae: Linyphiidae: Micronetinae)
Lihong Tu1,2 & Gustavo Hormiga2
1
College of Life Sciences, Capital Normal University, Beijing, P. R. China,
tulh@mail.cnu.edu.cn
2
Micronesian Hull, 1920 is one of the largest and most diverse linyphiid
subfamilies, and their members have some of the most morphologically complex
genitalia. Micronetinae includes several highly heterogeneous and polyphyletic
genera (e.g., Lepthyphantes Menge 1866). The results of a recently published
phylogenetic analysis of Linyphiidae, based on morphological and nucleotide
sequence data, do not support the monophyly of Micronetinae. In the present
study, we have expanded our earlier micronetine taxonomic sampling to include
representatives of 35 genera (with each genus represented by its type species
and, in most cases, one or more additional representatives) to further test the
monophyly of Micronetinae and of many of its genera. We also study the
internal relationships as well as the placement of micronetines within
Linyphiidae. Outgroup taxa were represented by 25 species from other linyphiid
subfamilies, and Pimoidae (the sister group of Linyphiidae), as well as
representative species of Tetragnathidae and Theridiidae. More than 260
morphological characters were scored for our study taxa for phylogenetic
reconstruction. In addition to those characters used in recent analyses of
linyphiid phylogenetics, 50 new characters from epigynal morphology were
included in this study. Furthermore, we discussed the evolution of micronetine
genitalia based on the resulting phylogeny.
448
Distribution of epigeic spiders (Arachnida: Araneae)

in the forest mosaic
Ivan H. Tuf, Ondřej Machač, Jana Mišurcová & Marea Grinvald
Department of Ecology and Environmental Sciences, Palacký University, Olomouc,

Czech Republic, ivan.tuf@upol.cz
Introduction
The type of the forest growth includes many factors influencing the species
diversity and the number of soil invertebrates. Suitable interferences with the natural
forest, resulting in higher habitat diversity, have positive influence on diversity of
the soil invertebrates. The edge effect is one of the most important factors (Jokimäki
et al. 1998, Magura et al. 2002). On the other hand, a lot of forests have been split
into small isolated forest areas and clearings due to over cutting, which can lead to
small species diversity and even to the local extinctions (Siitonen & Martikainen
1994, Niemelä 1997). Clearings are also the invincible barrier for the most of the
forest soil invertebrates (Esseen et al. 1997).
This study is focused on the distribution of epigeic spiders (Arachnida:
Araneae) in the different age floodplain forests. Spiders are used as the ecological
model frequently because they play an important role as predators, the knowledge
on their biology is relatively high and the sampling methods are well developed.

The research has been done in Litovelske Pomoravi Protected Landscape
Area from March 2004 to March 2006 in the floodplain forest (association Querco-
Ulmetum), near the city of Olomouc, the Czech Republic. Spiders were caught using
the pitfall traps (plastic pots with metal cover) with 4% solution of formaldehyde.
Two lines of 17 traps crossed four types of forest environment: (1) 87-years-old
growth (Querco-Ulmetum); (2) 10-years-old Quercus monoculture; (3) 3-years-old
afforested clear-cut; (4) 127- years-old floodplain forest. Traps were controlled each
14 days and close surroundings of traps (2 meters diameter around each trap) were
characterised according to the coverage of the plant layers.
Data were analysed by Canoco for Windows 4.5 (ter Braak & Šmilauer 1998)
for the following environmental factors: age of the growth which responds to the
position of the trap in the forest mosaic; presence of shrubs, trees, herbs, litter-
coverage and litter thickness which was semi quantified at scale of 25%.
Results
Of almost 3,000 specimens of ground-dwelling spiders, 39 species were
identified. The most dominant species were lycosid Pardosa lugubris (40%),
amaurobiid Coelotes terrestris (16%), liocranid Agroeca brunnea and lycosid
Trochosa terricola (both 9%).
The amount of spiders caught in traps during the study period was the highest
in the ecotone zone between afforested 2 years old site and 10 years old oaks
449
monoculture, as well as in the ecotone between 10 years old oaks monoculture and
87 years old floodplain forest. Nevertheless, diversity indices in these positions were
among the lowest. The highest diversity was found in traps placed in the ecotone
zone between 2 years old site and 127 years old floodplain forest. The communities
sampled by traps inside 10 years old oaks monoculture had higher diversity than in
the other sites.
The whole CCA model explained 48.3% of species variability and was
significant after Monte Carlo test. All of the environmental factors were significant
for the prediction of spider catches, but the presence of shrubs and the litter coverage
can be considered as the most significant ones. Abundance of 14 species of spiders
had a significant response to the coverage of shrubs and 3 species of spiders to the
litter-coverage. Species Coelotes terrestris, Agroeca brunnea, Ozyptila pratensis
and Cicurina cicur were more abundant in the areas distant to shrubs, on the
contrary of other 11 species (e.g. Diplostyla concolor, Pardosa pullata, Pirata
hygrophilus and Zelotes subterraneus). Pardosa lugubris and Diplostyla concolor
preferred ground densely covered by leaf litter.
Conclusions
According to the results, we can conclude that the structure of the spider
communities, as well as abundance of spider species, are highly dependent on the
amount of leaf litter and on the presence of shrubs. Also, ecotone zone has a positive
effect on overall number of spiders caught during the research and as such presents
precious environment. It appears that spiders are really sensitive to the landscape
structure and therefore the structure of spider communities can be highly influenced
by the forest management.
Acknowledgement
We would like thanks to the Ministry of Environment of the Czech Republic
(No. SP/2D3/155/08) and Czech National Research Programme II (No. 2B 06101)
for support this work.
References
Esseen P.A., Ehnström E., Ericson L. & Sjöberg K. 1997. Boreal forests. Ecological
Bulletin, 46: 16-47.
Jokimäki J., Huhta E., Itämies J. & Rahko P. 1998. Distribution of arthropods in
relation to forest patch size, edge, and stand characteristics. Canadian Journal
of Forest Research, 28: 1068-1072.
Magura T., Tóthmérész B. & Bordán Z. 2002. Carabids in an oak-hornbeam forest:
testing the edge effect hypothesis. Acta Biologica Debrecina, 24: 55-72.
Niemelä J. 1997. Invertebrates and boreal forest management. Conservation
Biology, 11: 601-610.
Siitonen J. & Martikainen P. 1994. Occurrence of rare and threatened insects living
on decaying Populus tremula: a comparison between Finnish and Russian
Karelia. Scandinavian Journal of Forest Research, 9: 185-191.
450
Natural mating rates and potential for genetic benefits

in a polyandrous spider with high risks of inbreeding
Cristina Tuni & Trine Bilde
Department of Biological Sciences, Ecology and Genetics, Aarhus University, Denmark,

cristina.tuni@biology.au.dk
Direct benefits to females or indirect genetic benefits derived via enhanced

offspring quality are considered selective forces underlying the evolution of
polyandry. Experimental studies show that females of the spider Stegodyphus
lineatus suffer direct costs of polyandry through increased risk of reproductive
failure, production of fewer offspring, or loss of the brood to infanticidal males
coercing a second and smaller replacement clutch. If females are nevertheless
polyandrous, they may evolve counter-adaptations to reduce the cost of mating,
i.e. cryptic choice for good or compatible genes. We investigated the degree of
polyandry in two natural populations of S. lineatus using microsatellite markers.
Results from genetic parentage analysis reveal presence of multiple sires in up to
50% of the clutches. Given the high costs of re-mating in this system polyandry
is likely to be driven by sexual conflict over mating rate. Our data reveal
evidence for paternity bias and hence potential for cryptic processes which may
confer genetic benefits. However, relatedness estimates suggest co-ancestry
among neighbouring spiders, and together with low genetic variability among
sires and relatively low re-mating frequency, the potential for acquiring
outbreeding benefits or other types of genetic benefit are low and unlikely to
compensate high direct costs of polyandry.
451
The secret lives of dwarf spiders
Gabriele Uhl
University of Greifswald, Zoological Institute and Museum, Department of General

Zoology and Systematics, Greifswald, Gabriele.uhl@uni-greifswald.de
Ever since Darwin we have been increasingly aware of the important role
that sexual selection plays in shaping animal behaviour, physiology and
morphology. Eye catching extravagant traits such as the antlers of deer or the
train of male peacocks have attracted much attention. However, we are
surrounded by a wealth of tiny species that display equally extravagant traits.
Dwarf spiders (also called money spiders; Erigoninae: Linyphiidae) are minute
and most of them are strongly sexually dimorphic. Only males possess cephalic
regions that are modified into deep grooves, pits, tube-like depressions, bulges,
lobes, or even turrets. We investigated the anatomy of head structures and found
that they are always connected with extensive glandular tissue, whose secretions
are stored in large reservoirs. Behavioural observations showed that females
contact the male head structures during mating, release saliva onto the male head
to imbibe the fluid shortly after. These findings strongly suggest that these
modifications and their secretions evolved in the context of sexual selection.
Studies are presented that investigate this phenomenon on ultrastructural,
biochemical, behavioural and phylogenetic levels.
452
Spiders are special: fear and disgust evoked

by pictures of arthropods
Gabriele Uhl1, Antje Gerdes2 & Georg Alpers2

1
University of Greifswald, Zoological Institute and Museum, Department of General
Zoology and Systematics, gabriele.uhl@uni-greifswald.de
2
Department of Psychology, University of Würzburg, Germany
Because all spiders are predators and most subdue their prey by applying
poison, it has been suggested that fear of spiders is an evolutionary adaptation.
However, it has not been sufficiently examined, whether other arthropods
similarly elicit fear or disgust. We thus compared ratings derived from the visual
perception of spiders, beetles, hymenoptera (bees and wasps) and lepidoptera
(butterflies and moths). Our aim was to test whether all arthropods rate similarly
or whether only a subset elicits comparable responses. We predicted that based
on an evolutionary adaptation to potential harmfulness spiders and hymenoptera
should rate similarly.
75 students viewed pictures and rated them individually. We presented 15
gray scale pictures of each arthropod group in random order. Participants rated
anxiety, disgust, and how dangerous they thought the animal was and
categorized each animal into one of the four animal groups. In addition, we
assessed a screening for fear of spiders. The picture ratings demonstrate that
spiders elicit significantly more fear and disgust than any other arthropod and
they are rated as more dangerous. The extent of fear of spiders is highly
correlated with the ratings of spiders but not with the ratings of other arthropods.
We conclude that harmfulness itself cannot explain why spiders are feared
so often.
References
Gerdes A.B.M., Alpers G.W. & Pauli P. 2009. Spiders are special: fear and
disgust evoked by pictures of arthropods. Evolution and Human Behavior,
30: 66-73.
453
Community structure and composition of spiders (Araneae)

in Western Himalayas, India
Virendra Prasad Uniyal* & Shazia Quasin
Wildlife Institute of India, Chandrabani, Dehradun, India

*
Corresponding author: uniyalvp@wii.gov.in
Spiders are amongst the highly species-rich group of invertebrates and, as

general predators, are important regulators of the ecosystem.
The study has been proposed to describe the spider community structure
and composition in Western Himalayas, Nanda Devi Biosphere Reserve
(NDBR) and to estimate the species diversity in different sites of altitudinal and
vegetation gradient. It also emphasized the need for conservation of spider
diversity by characterizing species diversity and highlighting rare and endemic
species in NDBR. This systematic approach will help to pave way for better
understanding of the Himalayan spider biodiversity, leading to improvised long
term ecological monitoring of the environment. Stratified random sampling was
employed in select sites to collect the spider samples from June - July 2009.
Spider specimens were collected by five semi quantitative techniques viz., pitfall
trapping, ground hand collection, aerial hand collection, sweep netting, aerial
hand collection and litter sampling.
A total of 1303 individuals (142 species/morphospecies, 63 genera, 25
families) were reported, 28.9% and 71.1% of them identified to species or genus
level, respectively. Species richness within families varied: 22 sp. (15.3% of the
total species) for Araneidae; 17 sp. (11.8%) for Thomisidae; 16 sp. (11.2%) for
Salticidae; 10 sp. (13.8%) for Linyphiidae; 10 sp. (13.8%) for Lycosidae; 8 sp.
(5.6%) for Gnaphosidae. Other families were rare and were represented by 60
species (42.1%) altogether.
Spiders were divided into three functional groups: the plant wanderers
(Thomisidae, Salticidae, Oxyopidae, Philodromidae and Sparassidae), the web
builders (Araneidae, Theridiidae, Linyphiidae, Uloboridae and Pholicidae), and
the ground wanderers (Gnaphosidae, Lycosidae, Selenopidae and Agelenidae).
Overall the number of web building spiders was greater than that of the ground
and plant wanderers. The web builders comprised of 57.9% of the total 1303
individuals, followed by plant wanderers (22.4%) and ground wanderers
(19.6%). The 25 families recorded in the area represent 41.7% of the total
families in India. This result suggests that the Himalayan region has a rich
diversity of spiders. Furthermore, the high percentage of web builders was
attributed with the rich floral diversity. Araneidae was found to be most diverse
in terms of species richness and diversity. Further investigation of spider fauna
may provide interesting results on new, endemic, rare and range restricted
species of these areas.
454
The distribution of Horus Chamberlin

(Arachnida: Pseudoscorpiones: Olpiidae)
Jacques van Heerden
University of Venda, Thohoyandou, Limpopo, South Africa,

Jacques.vanheerden@univen.ac.za
Horus Chamberlin is represented by nine species in Southern Africa and

one from the Côte d’Ivoire. The distribution of the identified specimens from
Southern Africa were plotted on a map of the region and revealed a rather
disjunct distribution of seven of the nine Southern African species10.
Horus species are usually found in rock crevices (such as thin cracks in
mudstone) – in only two cases have they been reported from under the loose
bark of trees. Field observations led to the conclusion that the species do not
employ phoresy (at least not as a rule) and therefore their dispersal rate is slow.
This is incompatible with the reported distribution of the species.
Preliminary DNA analysis conducted on samples from twenty sites in the
southern half of South Africa showed that species are geographically confined;
in general, the farther away the localities are from one another, the more distant
their relationship.
It is evident that the current definitions of the species of Horus need to be
revised because the taxonomic characters originally nominated by Chamberlin
(1931) are of limited value. This may also apply to other pseudoscorpion
species.
10
The two most abudant species are H. granulatus (Ellingsen) and H. obscurus
(Tullgren), with the former more common in the drier, western part of the region and H.
obscurus widely distributed in the eastern section; however, the two species also overlap
in the centre. The other seven Southern African species are less common: H. brevipes
Beier and H. modestus Chamberlin have been recorded from one site only, H. asper
Beier from two localities, and H. gracilis Beier from three localities. H. montanus Beier
and H. transvaalensis Beier have been recorded from six sites each and H. zonatus Beier
from seven.
455
Mating behaviour in spiders. Do we know everything?

Peter J. van Helsdingen
European Invetebrate Survey – Netherlands, Leiden, Netherlands, helsdingen@nnm.nl
In spiders sperm induction is an essential phase in the reproductive

sequence. A literature survey shows that sperm induction remained unobserved
in many observations of spider mating behaviour. An overview of what is known
in different families is presented: the moment of the sperm induction, the
construction of the sperm web, the position of the male when the palps are
charged. A correlation with the presence or absence of epiandrous glands is
discussed.
456
On three new species of the genus Oxyopes Latreille from

central India (Arachnida: Araneae: Oxyopidae)
Ganesh Vankhede1 & Shivaji Deshmukh2

1
Department of Zoology Sant Gadge Baba Amravati University, Amravati – 444602,
India, vganeshan2001@rediffmail.com
2
Indian Society of Arachnology
India has been represented by 4 genera and 69 species from Oxyopidae

family. Three new species of the spiders from Oxyopidae family are described
from central India from meadows after resettlement of forest villages Kund,
Koha and Bori in Melghat. Development of grassland in the undisturbed forest
has made favourable conditions for predators like spiders.
Key words: new species, taxonomy, Oxyopes, India.
457
Wolbachia-induced sex ratio distortion in the solitary

spider Oedothorax gibbosus and its potential implications
on sexual selection
Bram Vanthournout1 & Frederik Hendrickx2
1
University of Ghent, Belgium, bram.vanthournout@ugent.be
2 Royal Belgian Institute of Natural Sciences, frederik.hendrickx@naturalsciences.be
According to the sex-allocation theory of Fisher, a 50:50 sex ratio is

considered to be an evolutionary stable strategy. In a population with uneven
numbers of males and females, the underrepresented sex will have a fitness
advantage at the time of mating, hence increasing in the population. This
ultimately leads to the evolution of a stable 50:50 sex ratio.
Despite this theory, many examples of distorted sex ratios can be found in
nature. This is also the case in the male dimorphic and solitary dwarf spider
Oedothorax gibbosus, for which populations are female biased. Specific
breeding designs show that the sex ratio distortion is primarily maternally
inherited and correlates with the presence of the endosymbiotic bacteria
Wolbachia. This first direct evidence of a female biasing effect of this bacterium
in spiders was further confirmed by antibiotic curing of females, which restored
the sex ratio to an equal proportion of males and females. Endosymbiotic
bacteria can potentially have profound effects on its host ecology and evolution
by altering sexual selection pressures. Therefore, the potential interaction of
maternally inherited bacteria on the effect of the morph specific reproductive
strategies are discussed within the framework of the apparently stable
dimorphism observed in this spider species.
458
Spider assemblages in urban habitats

from Rennes (Brittany, France)
Marion Varet1, Julien Pétillon2 & Françoise Burel1
1
ECOBIO, University of Rennes 1, Rennes, France, marion.varet@sfr.fr
2
Evolutionary Ecology Group, University of Antwerp, Antwerpen, Belgium
During the last decades, urban population has strongly increased, creating
intensive urban areas (Douglas 1992, Fenger 1999, Weber 2003). Urbanization
can be defined as the installation process of anthropogenic structures (e.g.
buildings, roads) to the detriment of natural or farming areas, to satisfy human
population requirements (Croci et al. 2008). At the same time, social demands
for nature and biodiversity within the city (at individual or local authority levels)
are increasing (Chiesura 2004, Clergeau 2007). These two statements lead to the
development of studies about « biodiversity and urbanization ». A large part of
studies on this topic are based upon the analysis of assemblages along urban-
rural gradients. These studies mainly focused on three biological models: birds,
ground beetles and plants. Studies on other models are more marginal, including
one on spiders (Shochat et al. 2004).
Within a PhD project, we focus on a particular, under-investigated habitat:
residential areas, located in the peri-urban zone. 50% of the urbanized area is
dedicated to green space (private garden, public green space.). Since few years, a
new type of residential form has been developed in France. They promote public
green spaces (where the use of pesticides is very low or null) and reduce private
green spaces. In this study, we will test if this new type of residential areas are
characterized by higher animal species richness (notably for spiders, but also for
ground beetles, butterflies and birds) compared to those of classical residential
types (mainly housing estate). We are also interested in studying biodiversity at
the edge between urban and rural habitats.
The experimental design encompass two types of urban forms, each
spatially replicated three times: new urban form and housing estate (old form).
Sites were selected for presenting the same age and similar surface. At each site,
between 40 and 45 sampling points were randomly set up in public hedges. For
each sample point, local (vegetation cover, nature of the litter, temperature,
hygrometry …) and landscape (composition and density of hedge network,
neighbourhood structure…) variables were estimated and integrated in a GIS.
We studied the effect of different local and landscape factors on spider
assemblages. Additionally, three urban-rural transects of 1.200 meters long, and
including seven sampling points (established in hedgerows), were sampled. Sites
were located in peri-urban cities around Rennes (Brittany, France). The trapping
has been conducted by pitfall traps, continuously between April 20 and June 17,
2009.
In total, 5,063 individuals belonging to 137 species were collected. 25
families were represented, of which Lycosidae was dominant (42% of
459
individuals), followed by Linyphiidae (21.3%), Thomisidae (7.4%), Dysderidae

(4.6%), Liocranidae (4.1%), Gnaphosidae (4.0%), Clubionidae (2.9) and Zodariidae
(2.5%), notably.
Three categories of local variables are distinguished: variables dependent
on choices of planners during the establishment of hedgerows, variables
dependent on choices of gardeners and variables independent of man. The
partition of variance (Fig. 1) revealed that the choices of planners and gardeners
are equally important but there is little interaction between these different
choices.
The regression displayed a decrease in the number of individuals with the
proximity of urbanized area (log spider abundance=1.2697+0.0004 * Distance of
edge; r²=0.38; p=0.002). This is mainly due to changes in abundance of
individuals belonging to the family of Lycosidae. It is notable that the number of
individuals belonging to species considered as forest-inhabiting did not change
along the urban-rural transect. We did not neither observe a decrease in species
richness but instead a replacement of species. Zelotes pedestris is as such rather
associated with rural habitats whereas Pardosa hortensis is more associated with
urban (residential) areas (CA analysis).
Fig. 1. Diagram representing the variance partitioning using partial canonical

correspondence analysis (CCA).
This study finally showed that, in city, spider species' richness is not
dependent on population density. Population density is estimated by the number
of houses (dwellings, residences) in each hectare (richness=72.6135+0.5335 *
housing/ha; r²=0.1; p=0.2). The neighbourhood design could be one factor.
Currently, we are investigating and testing the effect of hedgerow connectivity
on spider populations, in particular for forest-living species (e.g. Pardosa
saltans).
460
References
Chiesura A. 2004. The role of urban parks for the sustainable city. Landscape
and Urban Planning, 68: 129-138.
Clergeau P. 2007. Une écologie du paysage urbain. Apogée, Rennes, 142 pp.
Croci S., Buter A., Georges A., Aguejdad R. & Clergeau P. 2008. Small urban
woodlands as biodiversity conservation hot-spot: a multi-taxon approach.
Landscape Ecology, 23: 1171-1186.
Douglas I. 1992. The case for urban ecology. Urban Nature Magazine, 1: 15-17.
Fenger O. 1999. Urban air quality. Atmospheric Environment 33: 4877-4900
Shochat E. Stefanov W.L., Whitehouse M.E.A. & Faeth S.H. 2004.
Urbanization and spider diversity: influences of human modification of
habitat structure and productivity. Ecological Applications, 14: 268-280.
Weber C. 2003. Interaction model application for urban planning. Landscape
and Urban Planning 63: 49-60
Weller B. & Ganzhorn J.U. 2004. Carabid beetle community composition, body
size, and fluctuating asymmetry along an urban-rural gradient. Basic and
Applied Ecology, 5: 193-201.
461
The influence of leaf litter structure on spiders in an

Atlantic Forest remnant in north-eastern Brazil
Sheila Luzia de S. Varjão, Kátia R. Benati, Tércio S. Melo,

Marcelo Cesar L. Peres, Alessandra R. S. Andrade,
Marcos Vinicius A. Guimarães & Marcelo A. Dias
Catholic University of Salvador, Centro de Ecologia e Conservação Animal, Brazil,
varjaoecoa@yahoo.com.br
Introduction
Organic plant and animal matter in different degradation stages constitutes
the leaf litter structure, providing a range of microhabitats, influencing
temperature and light conditions (Barbosa & Faria 2006) and supporting variety
of animals (Vallejo et al. 1987).
The studies in temperate forest provided data on influence of leaf litter on
spider composition. Uetz (1976) showed also that spider abundance and
diversity was affected by the flooding regime. Stevenson and Dindal (1982)
found that the space inside the deposited leaves, the lower surface of twisted
leaves and the gaps between the leaves may serve as microhabitat for smaller
species.
Wagner et al. (2003) found that the leaf litter depth and stratification
influence families, guilds and body size by creating foraging conditions.
However, it is still unclear the way in which spiders respond to leaf litter
structural changes in tropical forests. Spiders may use the environment structure
as refugia from predators, foraging, breeding, shelter or web building (Uetz
1976, Vallejo et al. 1987).
Understanding the structural patterns of leaf litter composition may help in
understanding its influence on spider biology, composition and habitat use and
may contribute to management in natural biota.
The objective of this study was to verify the leaf litter structural influence
on the richness and abundance of spiders in an Atlantic Forest remnant in the
municipality of Salvador, Bahia, Brazil.

The study was conducted in 2008. The survey included 15 randomly
arranged spots, each 50 x 50 cm. They were sampled twice: in dry and wet
seasons. During surveys, we measured the litter depth and the litter samples
were placed into the Winkler´s extractor (48 hours) to get spiders. The leaf litter
was then sieved and passing material was classified as large leaves (> 5 cm) that
could be large flat or curved and small leaves (≤ 5): small flat and little curved.
The sticks and trunk pieces were quantified and classified in slightly straight and
slightly curved. To test the influence of the leaf litter structure on the abundance
462
of spiders, we applied the Multiple Regression test (Graphpad InStat © 3.0),

since all the structural covariates passed the normality test (Kolmogorov-
Smirnov).
Results
We collected 90 spider species, representing 11 families. The most
frequent were Theridiidae (n=33), Oonopidae (n=14), Salticidae (n=14) and
Scytodidae (n=14), making 83.33% of all species. The most abundant was
Coleosoma floridana Banks, 1900 (n=14).
The leaf litter structure influenced the spiders abundance during the first
sampling season (p <0.0001, r ² 1.0000). Small flat leaves, large flat leaves, large
curved leaves and litter depth were the covariates positively influencing the
spider abundance. Small curved leaves, slightly straight and slightly curved
influenced negatively. There also was leaf litter structural significant influence
during second sample season on spiders abundance (p <0. 0001; r² 1.000). The
covariates slightly straight and slightly curved influenced positively and small
flat leaves, small curved leaves, large flat leaves, large curved leaves and litter
depth influenced negatively.
Discussion
Our results agree with the literature data. Several studies in a variety of
ecosystems suggest that spiders are susceptible to various environmental
variables and structural complexity (Uetz 1976, Wagner et al. 2003, Stevenson
& Dindal 1982. The negative influence of the leaves in the second sample
season may be as a result of more intense rainfall (average 149.1 mm, Inmet
2008), generating a lower production and increasing leaf litter degradation
(Souto 2006).
The negative influence of the leaf litter depth was not expected. Uetz
(1976) showed that the growing leaf litter depth increases the abundance of
spiders. We found a relationship between the depth of the leaf litter and the
amount of leaves. In temperate forests Uetz (1976) found that during the rainy
season the availability of prey is more important than the complexity of the
habitat, which may be expressed here.
The study demonstrated the importance of analyzing the leaf litter
structure in analysing spider faunas and assemblages and provides the useful
data for microhabitat management.
References
Banks N. 1900. Some new North American spiders. Canadian Entomologist. 32:
96-102.
Barbosa J.H.C. & Faria S.M. 2006. Aporte de serrapilheira ao solo em estágios
sucessionais florestais na Reserva Biológica de Poço das Antas, Rio de
Janeiro, Brasil. Rodriguésia, 3: 461-476.
463
Souto P.C. 2008. Acumulação e decomposição da serapilheira e distribuição de

organismos edáficos em área de caatinga na Paraíba, Brasil. Centro de
Ciências Agrárias da UFP, 146 pp.
Stevenson B.G. & Dindal D.L. 1982. Effect of leaf shape on forest floor spiders:
Community organization and microhabitat selection of immature
Enoplognatha ovata (Clerck) (Theridiidae). The Journal of Arachnology,
10:165-178.
Uetz G.W. 1976. The Influence of variation in litter habitat on spider
communities. Revista Oecologia, 40: 29-42.
Vallejo L.R., Fonseca C.L. & Gonçalves D.R.P. 1987. Estudo comparativo da
mesofauna do solo entre áreas de Eucaliptus citriodora e mata secundaria
heterogênea. Revista Brasil Biologia, 47: 363-370.
Wagner J.D., Toft S. & Wise D.H. 2003. Spatial stratification in litter depth by
forest- floor spiders. Revista The Journal of Arachnology, 31: 28-39.
464
The subsocial spider Anelosimus cf. studiosus can adopt

juveniles recently emerged
Carmen Viera & Maria de Fatima da Rocha Dias
Lab. Entomología, Facultad de Ciencias, Universidad de la República, Montevideo,

Uruguay
Lab. Ecología del Comportamiento, Instituto de Investigaciones Biológicas Clemente
Estable, Montevideo, Uruguay, anelosimus@gmail.com, faldias@yahoo.com.br
Anelosimus cf. studiosus (Theridiidae) is a sub-social species from

Uruguay (36ºS). Spiders live in low branches of perennial trees, building nests
with dry leaves. When the individuals reach adulthood, they leave the communal
nest, but sometimes adult females can share a big nest, but maintaining their own
territory (Viera et al. 2007). There are two kinds of nests: uni-female and multi-
female nests. The uni-female one is composed by the mother and one or two
broods (30-60 individuals), and the multi-female ones are composed by many
females with their broods. In spite of living together, adult females show
territoriality, being intolerant among them, limiting the degree of sociality. In
multi-female nests the females are located in separate retreats and they do not
cooperate in prey capture and do not show cooperative maternal care, either. The
social species have originated from sub-social ancestors through prolongation of
tolerance and cooperation to adulthood. One of the limits to reach the sociality is
the intolerance among adult females forcing the dispersion. In spite of the
intolerance among adult females, the cannibalism is inhibited and the sub social
spiders show high tolerance and individual maternal investment. In this species,
the maternal effort is high, due to the cost of egg-sac building, they are bigger
than their mother’s abdomen size, and the permanent care is to avoid parasitism
affecting mother’s feeding during this period. When the time of development
inside of the egg-sacs is finished, each mother makes a big hole to allow the exit
of spiderlings. The process of opening the egg-sac is synchronized by fine
endogenous mechanism that was activated at the moment of the oviposition and
the alarm clock strikes approximately 21 days later (Viera et al. 2008). The
mother feeds the juveniles, capturing prey by wrapping, cutting in pieces, and
regurgitating. Usually mothers die when the juveniles reach the 4th instars and
are cannibalized by them. The female territorialism and asynchrony among
oviposition of different females makes the cooperative maternal care apparently
impossible. Although the adult females with egg-sacs show tolerance to the
foreign juveniles, the cannibalism is inhibited. This distinguishes this species
from other more social because the mother´s presence is absolutely necessary to
allow the exit of juveniles and to feed at the early instars of development.
The aggression among adult females increases when they become
mothers, defending actively their egg-sacs against other females. In spite of this
465
aggression, we have already observed adoption of egg-sacs in the laboratory

conditions. Even though the mothers do not recognize their own egg-sac, the
delicate mechanism of opening egg-sac make the cooperative care difficult.
In reference to the importance of the first meals by mother regurgitations
to the juveniles, we tested if the young ones could obtain the regurgitation
substances from other adult females, which would act like stepmothers. We used
two experimental groups: 20 adult virgin females and 20 adult females
copulated. We maintained both groups under the same laboratory conditions at
the average temperature of 25ºC and 70% humidity. The individuals were fed
with Drosophila spp. and small Tenebrio molitor larvae. Each adult female was
raised isolated, like in a uni-female nest, in small Petri dish. The data of the
copula and laying the egg-sacs were registered. We put a group of recently
emerged juveniles with virgin adult females and other group with female
mothers but not their own mothers. We used only juveniles because we had
previously found under experimental conditions that they were able to capture
small prey by themselves since the second instars (Ghione et al. 2004), but until
then, they needed the presence of the mother for supplementary feeding by
regurgitation.
We observed that all the juveniles showed a solicitation behaviour
consisting of many beats on the mothers’ mouth, very closely to the chelicerae
and being exposed to possible aggression. The group of virgin females showed
high tolerance behaviour toward spiderlings, they did not cannibalize, but were
unable to feed. In the second group with we observed a total adoption
mechanism. In two cases the real mother was dead and the substitute mother was
able to feed juveniles with parts of prey and regurgitation like a “real mother”.
They did not recognize their own brood and could replace the dead mother.
Other care behaviour observed was the repair of the communal web, cleaning of
prey’s rest, protection of egg-sacs and early juveniles against parasitism of Ficus
and bacteria. When the prey arrived to the communal web, the adult females
captured it and made forelegs display on the threads and the juveniles responded
and approached to be fed. The mothers stayed calm to avoid causing any injury
to the spiderlings. During the feeding process the mothers watched if all the
juveniles were fed, helping them carefully all the time. The mothers did not feed
themselves until the juveniles finished eating. The high mother´s investment was
prolonged and continuous, and our findings suggest that even though the real
mother was dead, other females sharing the same nest can substitute the mother
and take care of their relatives, making the entire colony survive. The substances
transferred to the juveniles by regurgitation could be very important, since this
procedure occurs among sub-adults individuals and the males fed by their sisters
acquire bigger size and they would be more reproductive successful than the
small ones (Viera et al 2007). Further research about adoption must be done,
together with future investigations of the importance of the substances
transferred by regurgitations for the healthy development of the juveniles.
466
References
Ghione S., Viera C. & Costa F.G. 2004. Ability to capture prey in early instars
of the subsocial spider Anelosimus cf. studiosus (Henz, 1850) from
Uruguay (Araneae, Theridiidae). Bulletin of the British Arachnological
Society, 13(2): 60-62.
Viera C., Costa F.G., Ghione S. & Benamú-Pino M.A. 2007. Progeny,
development and phenology of the sub-social spider Anelosimus cf.
studiosus (Araneae, Theridiidae) from Uruguay. Studies on Neotropical
Fauna and Environment, 42(2): 145-153.
Viera C., Ghione S. & Costa F.G. 2006. Regurgitation among juveniles in the
subsocial spider Anelosimus cf. studiosus (Araneae, Theridiidae). Journal
of Arachnology, 34(1): 258-260.
Viera C., Ghione S. & Costa F.G. 2007. Mechanisms underlying egg-sac
opening in the subsocial spider Anelosimus cf. studiosus (Araneae,
Theridiidae. Ethology, Ecology & Evolution, 19(1): 61-67.
Viera C., Ghione S. & Costa F.G. 2007. Post-embryonic development of the
sub-social spider Anelosimus cf. studiosus (Araneae, Theridiidae). Bulletin
of the British Arachnological Society, 14(1): 30-32.
467
The invasive Australian redback spider,

Latrodectus hasseltii Thorell, 1870 (Araneae: Theridiidae):
current and potential distributions
Cor J. Vink1, Craig B. Phillips2, José G.B. Derraik3 & Phil J. Sirvid4
1
Biosecurity Group, AgResearch, Lincoln, New Zealand & Entomology Research
Museum, Ecology Department, Lincoln University, New Zealand,
cor.vink@agresearch.co.nz
2
Biosecurity Group, AgResearch, Lincoln, New Zealand
3
Disease and Vector Research Group, Institute for Natural Sciences, Massey University,
Auckland, New Zealand
4
Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand
Populations of the Australian redback spider, Latrodectus hasseltii Thorell

1870, were first recorded in New Zealand in the early 1980s and in Osaka, Japan
in 1995. Reliable records suggest that naturalised populations of L. hasseltii in
New Zealand are present only in Central Otago and New Plymouth. In Central
Otago, L. hasseltii feeds on endangered invertebrates. Latrodectus hasseltii is
also a hazard to the New Zealand endemic L. katipo through interbreeding and
competitive displacement.
The computer programme CLIMEX was used to model the potential
global distribution of L. hasseltii based on current climate, and using ArcGIS
9.2, areas of suitable climate in New Zealand were overlaid with suitable
habitats to identify areas most suitable for L. hasseltii establishment. The shelter
that urban areas offer L. hasseltii were modelled in CLIMEX and incorporated
into ArcGIS to produce maps indicating cities and built up areas where the
species could establish. The presence of L. hasseltii in New Zealand and Japan,
and its possible spread to other parts of the world, is of human health
significance, and it may also impact on native biodiversity.
468
Using real-time remote diagnostics to examine

valuable specimens
Cor J. Vink1, John Marris2, John M. Kean3 & Trevor K. Crosby4
1
Biosecurity Group, AgResearch, Lincoln, New Zealand & Entomology Research
Museum, Ecology Department, Lincoln University, New Zealand,
cor.vink@agresearch.co.nz
2
Entomology Research Museum, Ecology Department, Lincoln University, New
Zealand
3
Biosecurity Group, AgResearch, Lincoln, New Zealand
4
New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand
Real-time remote diagnostic tools have the potential to change the way
arachnological specimens are examined and identified. We selected and trialled
four internet-based video conferencing software systems for their applicability to
real-time remote diagnostics. Trials were conducted using standardised tests for
image resolution and latency, and real diagnostic challenges were set using
spider specimens. The tests were conducted within and between research
organizations in New Zealand, and with an international collaborator in Western
Australia. The most important features of a good remote microscopy system
were identified as cost, ease of set-up and use, image quality, the ability to
capitalise on high-speed research networks, vocal communication capability, and
a remote pointer. The greatest impediment was obtaining access through
institutional firewalls. More generic solutions to this problem are required if the
full potential of remote communication technologies is to be realised.
Real-time remote diagnostic tools have particular potential for use in
taxonomic studies, to avoid time delays and possible damage to valuable
specimens during transportation. A study of the costs and benefits of remote
diagnostics for accessing spider and insect specimens in the New Zealand
Arthropod Collection with be discussed.
469
Sperm storage and sperm activation in the orb-weaving

spider Argiope bruennichi (Araneidae, Araneae)
Oliver Vöcking, Peter Michalik & Gabriele Uhl
Zoologisches Institut und Museum Greifswald, Germany, Oliver.Voecking@gmx.de,

Michalik@uni-greifswald.de, Gabriele.Uhl@uni-greifswald.de
In spiders, spermatozoa are transferred to the female in a coiled and

encysted state and stored in the spermathecae until oviposition. Little is known
of the processes involved in sperm activation in spiders (Brown 1985,
Berendonck & Greven 2005). This study focuses on sperm activation in the
genital system of the sexually cannibalistic Argiope bruennichi in which male
genital mutilation serves to plug the insemination duct after mating. Most males
are cannibalized during their first insertion and are thus able to monopolize one
spermatheca only. Post-mating female mate choice may thus occur by selective
activation of the sperm from rival males that are stored in different
spermathecae.
Virgin females were raised in captivity and double-mated under controlled
conditions. By amputating one pedipalp each, both males were restricted to one
insemination into opposite spermathecae. Post-mating, the spermathecae were
dissected out after different time periods to study morphological changes in the
stored spermatozoa by means of light microscopy and electron microscopy.
Shortly after copulation, the spermatozoa show the typical secretion sheath
known from spiders and other Araneidae. Stages before oviposition show coiled
spermatozoa without the secretion sheath or almost fully uncoiled spermatozoa,
however, in a few specimens we found spermatozoa with and without secretion
sheath, the latter mainly occurring close to the spermathecal wall. Hence, we
assume that the activation is triggered by a female signal released from the
glands that surround the spermathecae. After oviposition, spermatozoa
remaining in the spermatheca were always decapsulated which might suggest
that females are able to hold back sperm for a subsequent oviposition bout.
Our data suggest that sperm activation takes place in the female´s
spermathecae by means of an activation trigger produced by the female.
Whether females are able to selectively activate different sperm stores remains
to be investigated.
470
Taxonomic and ecological data on spiders from

Saranda, Albania (Arachnida: Araneae)
Blerina Vrenosi1 & Christo Deltshev2
1
Museum of Natural Sciences, Faculty of Natural Sciences, Tirana University, Albania
2
Institute of Zoology, Bulgarian Academy of Sciences, Bulgaria
Spiders play significant role as ecological bioindicators and are important

part of the food chain. Geographic position, sea influence, high biodiversity of
relieves, the elevation from the sea level and diversity of specific climate and
microclimates conditions influence the species richness.
In six different habitat types (Saranda Hills, Mirror Beach, Butrinti Hills,
Ksamil Beach, Blue Eye Hills and Kakome Beach), 27 species representing 21
genera and 8 families have been recorded. Most of the species have a low
ecological valence, but a few of them are more widespread and have higher
ecological valence. The calculation of the similarity coefficient between hilly
habitats of Saranda showed a similarity of araneofauna (K=20%) between city’s
hills and protected area’s hills (Blue Eye and Butrinti). Further and extended
studies will be undertaken at Saranda in the future.
471
Spider fauna in paddy fields in Thailand

Wipada Vungsilabutr & Wimolwan Chotiwong*
Entomology and Zoology Division, Department of Agriculture, BangKhen, Bangkok,
Thailand, * yuri_socool@hotmail.com
A survey of spiders in paddy fields was carried out from October 1994 to
September 1996 at Pathumthani, Suphanburi, Chachengchao, Nontaburi,
Nakhonsawan, Chiang-Mai, Ratchaburi, Songkhla and Nakonrachasima
provinces. The collected fauna included 14 families, 36 genera and 50 species,
21 species newly recorded for Thailand. The species were general predators,
feeding on green rice leafhoppers, rice brown plant hoppers, rice cutworms, stem
borers and stink bugs.
472
The origin and speciation of Solenysa spiders

on the Japan Archipelago
Fang Wang & Lihong Tu
College of Life Sciences, Capital Normal University, Beijing, P.R. China,

wycgbiochemical@yahoo.com.cn, tulh@ioz.ac.cn
According to the biological species concept (BSC), morphological

variations of genitalia are often used to identify species. However, how much
difference would lead to reproductive isolation and speciation is highly
debatable. The advent of relatively inexpensive and rapid DNA sequencing has
made it possible for biologist to detect and differentiate morphologically similar
species. The spider genus Solenysa, a group of highly specialized Linyphiidae,
has many exclusive features of both genitalic and somatic morphology and
endemic distribution. The four species occurring on the Japan Archipelago have
unique genitalia, which only differ in fine details. Furthermore, they share
similar habitat and distribute along the islands with little range overlap. In
present study we reconstructed a species level phylogeny of the Japanese species
based on the molecular sequences data (mitochondrial COI, 16S; nuclear 28S,
18S, histone H3) and morphological data. The most parsimonious trees and
Bayesian trees revealed four monophyletic clades that is congruent with the
previous interpretation based on morphological data. Furthermore, the
phylogenetic relationships suggest that the Japanese species were derived from
the species occurring on mainland China and dispersed from south to north on
the Japan Archipelago. In combination with historical geographical information,
we discussed the origin and speciation of the Solenysa species on the Japan
Archipelago and the historical caused that produced the current distribution
patterns.
473
A comparison of Argiope bruennichi and

Araneus quadratus population density
in different habitats and regions of Poland
Wioletta Wawer
Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw, Poland,

w.wawer@miiz.waw.pl
The spectacular expansion of Argiope bruennichi (Scopoli, 1772) gives an

opportunity to study the dispersal conditions and adaptability of species. The
wasp spider is of Mediterranean-Pontian origin. In the Holocene, it has been
gradually spreading to the north. Nowadays it is distributed over a considerable
part of the Palearctic.
In Poland A. bruennichi was first noted in the second half of the 19th
century. A conspicuous expansion in western Poland has been taking place since
the 1940s. Over the next decades an increasing number of wasp spider stations
were observed but only single sites were discovered in the 1960s and 1970s in
the South-East regions of the country. By the end of the 1990s ca. 100 sites were
known in many regions, except NE Poland, where the climate is harsh. Currently
the wasp spider is distributed over the whole territory of Poland and it has
become a very common species.
A. bruennichi spreads relatively fast and the character of its expansion is
not well recognised. The wasp spider controls the abundance of other spiders,
wins competition with other species or seizes free niches. An expansive species
is frequently identified as an invasive species. Some arachnologists (Kajak &
Łuczak 2003) have emphasized the substantial biocoenotic role of A. bruennichi
in terms of changes in number of other spiders (mainly web spiders) occupying
the same area.
The impact of the wasp spider on other spider species is not yet clear.
Particularly strong competitive interactions can, for instance, occur between
Argiope bruennichi and Araneus quadratus (Clerk, 1754). The four spot orb
weaver has a one-year life cycle, with young individuals appearing in May and
females forming sacks on September (Kajak 1965). The life cycle of A.
bruennichi is very similar. Both species occupy the same habitats and have
similar food preferences. I suppose that A. bruennichi limits the abundance of
orb weaver spiders. I would like to test this hypothesis through research on the
competition of the two species.
The purpose of my investigations is to compare the densities of different
web spiders in similar habitats but in different regions and geographical
conditions within the distribution range of web-building spiders. It is possible
that in the vicinity of the 'Polish cold pole', the abundance of the four spot orb
weaver is higher than the numbers of the more thermophilous wasp spider,
because of the harsh climate.
474
In my presentation I show the results of the first season of my

investigation of A. bruennichi and A. quadratus density in selected habitats and
regions.
The study was conducted in August 2009 in several areas characterised by
different climate conditions: Pojezierze Mazurskie (Mazury Lake District)
(53°41'N, 21°54'E), where the wasp spider has been observed since 2000, and
Kotlina Sandomierska (Sandomierz Valley) (50°10'N, 21°43'E), where the wasp
spider has been present since the 1970s. Six locations were selected in each
region (e.g. natural meadows, arable meadows and wastelands). The method of
direct counting in 25 m2-squares was used, with 36 squares selected in Kotlina
Sandomierska and 32 in Pojezierze Mazurskie.
The early results suggest a higher abundance of A. bruennichi than A.
quadratus. Within Kotlina Sandomierska A. bruennichi considerably
predominated in 32 squares. Within the Pojezierze Mazurskie in 5 squares A.
quadratus was slightly more numerous than A. bruennichi.
The abundance of A. bruennichi in north-eastern and south-eastern Poland
did not differ significantly (Student's t-distribution). However, A. quadratus
density in north-eastern Poland (Pojezierze Mazurskie) was significantly higher
than its abundance in Kotlina Sandomierska.
Little data concerning A. quadratus density are available from several
dozen years ago, since the wasp spider was not present in many regions of the
country at that time. Basing on information provided by Kajak (1960) about A.
quadratus abundance in the 1950s (Biebrza basin, north-eastern Poland) this
species dominated in many communities. Moreover, abundance fluctuations
were observed over the years, but the pattern of change during each season was
similar.
The available information indicates a tendency of spatial passing of A.
bruennichi and A. quadratus. Subsequent studies should verify the assumptions
concerning the wasp spider potential for dislodging the four spot orb weaver
from its natural habitats.
References
Kajak A. 1960. Changes in the abundance of spiders in several meadows.
Ekologia Polska, 9: 199-228.
Kajak A. 1965. An analysis of food relations between the spiders – Araneus
cornutus Clerk and Araneus quadratus Clerk – and their prey in meadows.
Ekologia Polska, 32: 717-764.
Kajak A. & Łuczak J. 2003. Spiders – meaning, quantity, structure, spatial
localization. In: Andrzejewski R. (ed), Kampinoski Park Narodowy, vol.
Wydawnictwo Kampinoski Park Narodowy, Izabelin, pp. 539-563.
475
The role of web placement in structuring

linyphiid spider communities and food webs
Kelton D. Welch, Eric G. Chapman & James D. Harwood
Department of Entomology, University of Kentucky, KY, USA, kdwe222@uky.edu
The Linyphiidae are a numerically dominant family of small, web-building

spiders whose ecology is of central importance to the function of arthropod
communities in agroecosystems. However, their importance within these
communities has not translated into widespread or systematic attention in the
ecological literature. Of particular interest is the diet of linyphiid spiders, which is
known to include economically important pest species, such as aphids and
leafhoppers. This suggests that these spiders can fill an augmentative role in
biological control programs. However, before such programs can be designed, an
understanding of the factors that impact consumption of pests by linyphiid spiders
must be investigated. The capacity of web-building spiders to contribute to the
biological control of insect pests can be mediated or facilitated by the availability
of alternative prey or the density of competing predators, both of which will be
impacted by the placement of spider webs. This makes the selection of
microhabitats for the placement of webs a particularly influential ecological
variable.
In order to character web-placement patterns, and to elucidate the effects of
web placement on trophic and community ecology of linyphiids, data on spider
webs was collected in two alfalfa fields in central Kentucky using a quadrate-
based sampling protocol. Spider species were shown to make differential usage of
available structures for web attachment points, and consistently selected distinct,
narrow ranges of heights for web placement. Tennesseellum formicum (Emerton)
most commonly inhabited web-sites at the base of plant stems at heights averaging
3-4 mm off the ground. Erigone autumnalis Emerton build webs on open ground
at heights 1-2 mm off the ground. A common tetragnathid spider, Glenognatha
foxi (McCook), placed its orb-webs approximately 10 mm off the ground, and
rarely utilized the ground as an attachment point.
Springtails (Hexapoda: Collembola) constitute a majority of the potential
prey for linyphiid and other web-building spiders. Although not typically
considered pests, springtails (and other alternative prey items) can serve to sustain
spiders in the absence of prey, thus allowing spiders to maintain populations ahead
of pest outbreaks and maximize their potential impact on these pests. Sticky trap
collections in the same two alfalfa fields revealed differential access to Collembola
between epigeal and low-foliar web-sites, indicating that web height may be a
means of trophic niche partitioning among web-building spiders. To draw
correlations between consumption of prey and web placement, we used PCR to
screen spider guts for traces of Collembola DNA. Significant levels of trophic
specialization were observed: 74% of T. formicum, 40% of E. autumnalis, and
14% of G. foxi, tested positive for Collembola. T. formicum, the most abundant
476
spider (n=370), showed greater consumption of Collembola at web heights

between 2 and 5 mm than at higher or lower heights, and at web-sites at the base
of plant stems than at epigeal or foliar web-sites, but the differences were not
significant.
These data provide valuable insights into the partitioning of trophic and
spatial niches among web-building spiders, and form a foundation of basic
ecological knowledge that will contribute to the development of conservation
biological control programs that incorporate these abundant spiders.
477
A scanning electron microscopic survey of the cuticle in mite

harvestmen (Arachnida, Opiliones, Cyphophthalmi) with
the description of novel sensory and glandular structures
Rodrigo H. Willemart1 & Gonzalo Giribet2

1
Escola de Artes Ciências e Humanidades, Universidade de São Paulo, São Paulo, SP,
Brazil, willemart@usp.br, corresponding author
2
Department of Organismic and Evolutionary Biology & Museum of Comparative
Zoology, Harvard University, Cambridge, MA, USA, ggiribet@oeb.harvard.edu
The cuticular surfaces of Cyphophthalmi (Opiliones) were studied in

detail with scanning electron microscopy, covering a wide range of their
taxonomic diversity. Previously unknown structures are described, including a
sexually dimorphic row of spines and glandular openings on leg I of Fangensis
cavernarum. Scanning electron micrographs of the prosomal paired hairs and the
subapical process are provided for the first time. Evidence for the multi-pored
nature of the shaft of solenidia as well as the hollowed nature and absence of
wall pores of sensilla chaetica are also shown for the first time using scanning
electron microscopy. The prosomal paired hairs may constitute a novel
synapomorphy for the suborder Cyphophthalmi, as they are absent in all studied
members of the other Opiliones suborders. Finally, the variation in shape of
some of the structures examined may be of great taxonomic value.
478
Harvest-ironman: heavy armature and not its defensive

secretions protects a harvestman (Opiliones)
against a spider (Araneae)
Rodrigo H. Willemart & Elene da Silva Souza
Escola de Artes Ciências e Humanidades, Universidade de São Paulo, São Paulo, SP,
Brazil, willemart@usp.br
The use of scent gland secretions against predators is a costly defense that has
convergently evolved in several taxa. In detailed analyses of arthropod behaviour,
chemicals have consistently been shown be the responsible for repelling the specific
predators. We studied the interaction between the harvestman Discocyrtus invalidus,
a heavy bodied animal which bear a pair of scent glands, and the spider
Enoploctenus cyclothorax, a generalist predator. In the first experiment, we left E.
cyclothorax with either D. invalidus or a cricket (control) for 5 days in a recipient
(n=16). The survival rate of harvestmen was 100%, and that of crickets was below
25%. In the second experiment, we observed details of the interaction by digitally
recording 32 spiders divided in 2 treatments (harvestmen and crickets, n=16). Most
spiders rejected the prey but in none of the videos could we notice the release of
scent gland secretions. In an attempt to explain why rejection occurred, we designed
a third experiment where we tested the hypothesis that harvestmen would release
little amounts of secretions, invisible to the human eye, and that these would be
responsible for the rejection by the spiders. We digitally recorded 52 spiders, divided
into 4 treatments of 13 spiders each (harvestmen with gland obstructed with glue,
harvestmen with glue on the dorsum, crickets with glue on the dorsum and crickets
with no glue). Harvestmen were significantly less preyed than crickets, the glue had
no effect and blocking the glands did not interfere in the results. A comparative
fluxogram between the behaviour of spiders against crickets and harvestmen
allowed us to clearly show how spiders behave differently when facing these two
prey. We also tested the effect of the harvestmen secretion per se, by applying
harvestmen secretions between the chelicerae of a spider, immediately after it
captured a cricket (control: distilled water). None of the spiders released their prey.
Since the role of chemicals in rejection were ruled out, we designed a fifth
experiment to test whether E. cyclothorax could pierce the integument of D.
invalidus and we found that only one out of ten spiders did it. Finally, we provide
SEM micrographs to show that only the articulations, mouth and tip of the legs are
not covered by a hard integument in several harvestmen species. Our combined
results have implications in the understanding of proximate mechanisms of prey-
predator interactions: this is the first experimental evidence that chemically defended
harvestmen do not use their scent gland secretions to repel a much larger predator
but rather rely on their heavily built body.
479
Archaeid and mecysmaucheniid spiders and their relatives

(Araneae: Archaeidae, Mecysmaucheniidae): phylogeny,
biogeography and evolution of the carapace morphology
Hannah M. Wood1,2,3, Charles E. Griswold1,2,

Rosemary G. Gillespie2 & Damian O. Elias2
1
Francisco, CA, USA
2
Berkeley, CA, USA
3
Corresponding author: hwood@berkeley.edu
The spider families Archaeidae and Mecysmaucheniidae are revised.

Molecular and morphological phylogenetic analyses are performed for
Malagasy, South African and Australian archaeid lineages, for Chilean and New
Zealand mecysmaucheniids, and for outgroup taxa representing 20 different
spider families. Fossil archaeids are examined using X-ray Computed
Tomography in order to understand phylogenetic placement of extinct lineages.
Biogeographic findings within different continents as well as between continents
are discussed. Evolution of the carapace shape is examined in relation to
predatory behaviours and biomechanical properties.
480
The antimicrobial properties of spider silk

Simon Wright
University of Nottingham, United Kingdom, plxsw5@nottingham.ac.uk
Spider silk is a remarkable material in nature, showing levels of strength,

torsionality, flexibility, lightness that are unmatched by man-made materials.
Another notable feature of spider silk is its longevity. Spider webs often remain
in the environment long after the spider has expired.
The longevity of spider silk indicates that the material has some properties
of resisting decomposing by microorganisms. Studies by Volrath et al have
investigated the compounds present in spider silk and have found that spider silk
contains molecules that are known to have antimicrobial properties.
Preliminary tests carried out on Tegenaria web silk have shown that when
isolated and placed in solution with E. coli, the ability of the E. coli to grow and
reproduce is significantly diminished.
Further tests to be carried out will include investigating if this
antimicrobial activity is effective across a wide range of bacteria and fungi and
also if a wide range of spiders are able to produce silk with these characteristics.
481
Taxonomic study of Myrmarachne (Araneae: Salticidae)

from Borneo
Takeshi Yamasaki
Graduate school of Science and Engineering, Kagoshima University, Japan,

howyadpincreep@yahoo.co.jp
Introduction
The members of the genus Myrmarachne (Salticidae) resemble ants
morphologically and behaviourally. Around 250 species are recorded all over the
world, of them around 80 from Southeast Asia and 10 from Borneo.
The purpose of this study is to describe new species and to redescribe the
known species from Borneo, and to review the taxonomic system of Myrmarachne
in Southeast Asia.
M. borneensis and M. shelfordii are described from Borneo in 1907.
However, it is difficult to identify these two species without comparison with type
specimens because there are no drawings in these descriptions. The redescriptions of
M. borneensis and M. shelfordii are needed to facilitate to identify these species.
Most Myrmarachne species have been described based on a few specimens
in previous studies. This has resulted in the misunderstanding of male/female
combination of a certain species because taxonomists have to guess the combination
based only on morphological data. Males and females of the same species have been
described separately as different species. It is expected that a number of specific
names in Southeast Asia may be junior synonyms. Commonly the males of
Myrmarachne become the adults earlier than the conspecific females, and wait for
them becoming adults near their nests. The direct observation in the field is
considered important in particular for the study of this group of spider.

Spiders were collected from various sites in Sabah and Lambir Hills
National Park in Sarawak, Borneo. Manual collecting including beating and
sweeping was conducted for specimens. When spiders were juveniles, spiders were
kept in empty bottles for make them adults. Collected adult spiders were preserved
in 75% ethanol.
Specimens were sorted and identified mainly based on the structures of
chelicerae and palpi in males, and epigyne in females.
Type specimens of M. borneensis and M. shelfordii were loaned from The
Museum of Comparative Zoology, Harvard University for redescription and
comparison with collected specimens.

Twenty-two species were collected from present study and 10 of them
were identified to species. M. borneensis and M. shelfordii were not included
among these 22 species. Only 4 species are mentioned as temporary results.
482
M. borneensis Peckham & Peckham, 1907: M. borneensis is a dark brown

(in ethanol) and robust species. Chelicerae are relatively short compared with the
length of carapace. This species has thick long hairs like spines under tibial
apophysis on their palpal tibia. This character is unique among Myrmarachne
species.
M. shelfordii Peckham & Peckham, 1907: M. shelfordii is a light brown (in
ethanol) species. Chelicerae are slender with numerous teeth. Tibial apophysis is
well developed.
M. hanoii Żabka, 1985 and M. topali Żabka, 1985: M. hanoii was
described based on only male and M. topali based on only female. I observed a
male of M. hanoii waited for a subadult female of M. topali, suggesting that M.
hanoii and M. topali are a single species.
483
A new serrular structure and its implications for the

phylogeny of the scytodids (Araneae: Scytodidae)
Laura-Marie Y.L. Yap1, David J. Court2 & Daiqin Li1*

1
2
Honorary Research Associate, Raffles Museum of Biodiversity Research, National
University of Singapore, Singapore, araneus@singnet.com.sg
*
dbslidq@nus.edu.sg
A new form of serrula in some large cave-dwelling scytodids (Araneae,

Scytodidae) from the south west of China is reported and figured. The
configuration of the serrula within the Araneae is reviewed and the new form is
compared with the serrulae of other members of the Scytodidae, with those of
other scytodoids, and with those of the much less closely related Mesothelae,
Mygalomorphae and non-haplogyne Araneomorphae. Instead of the commonly-
observed single-rowed serrula the new form is bi-cusped almost to the extent of
being double rowed. A cladistic analysis has been performed and we now
consider it most parsimonious to treat this bi-cusped trait as being a unique
apomorphic character which partially defines a clade within the Scytodidae.
Although the serrula is nearly double-rowed we suggest that it is unlikely to be
synonymous with the multi-rowed serrula of the Hypochilidae. It is speculated
that the bi-cusped serrula functions as an instrument which ruptures a hard but
brittle exoskeleton of an item of the spider’s prey.
484
Brotherly love or sibling rivalry? The presence of

older siblings influences body condition in the social spider
Delena cancerides (Sparassidae)
Eric C. Yip
Department of Entomology, Cornell University, Ithaca, NY, USA; Research School of

Biology, Australian National University, Canberra, ACT, Australia, ECY7@cornell.edu
Cooperative foraging is an important benefit of group living for a variety

of animals and for the social spiders in particular. The social huntsman spider,
Delena cancerides, unlike most other social spiders that build capture webs,
lives under tree bark. Laboratory and field data show that, while spiders share
prey while inside the retreat, the vast majority of prey is captured outside the
retreat when spiders forage solitarily. While prey sharing inside or outside the
retreat is a rare event, it might still be an important benefit, particularly to young
spiders that have a limited ability to capture large prey. To investigate the
possibility that prey sharing, though rare, might provide a measurable benefit to
spiders, I measured the condition of spiders at collection and related the
condition of spiders to colony demographics. I collected and measured 2821
spiders from 102 colonies, using the deviation from the expected weight based
on carapace width as an indication of “condition.” Young spiders (3rd-4th instar)
had significantly greater condition when in the presence of older siblings than
young spiders with siblings only of the same size. Older spiders (7th instar-
subadult) tended to do worse in the presence of younger siblings. The data
suggest that occasionally sharing prey with older siblings is an important benefit
for second or third cohort young. Conversely, other benefits of group living
besides prey sharing must provide sufficient benefits to first cohort offspring to
remain in the natal nest.
485
Molecular phylogeny indicates intercontinental dispersal

of euophryine spiders in evolutionary history
(Araneae: Salticidae)
Junxia Zhang & Wayne P. Maddison
Department of Zoology, University of British Columbia, Vancouver, BC, Canada,

jxzhang@interchange.ubc.ca, wmaddisn@interchange.ubc.ca
The Euophryinae has worldwide distribution and is the only major group
of jumping spiders (Salticidae) to have diversified into many genera in both the
Old and New World, which makes its historical biogeography particularly
interesting. As one of the largest subfamilies of jumping spiders, the
Euophryinae currently contains at least 95 genera and more than 800 species.
However, its phylogeny is poorly studied. To clarify the phylogeny of the
subfamily and understand its historical biogeography, we amplified and
sequenced five genes (mitochondrial: COI, 16SrDNA, NADH1; nuclear: 28S
rDNA, Actin) from species collected from the major distribution areas of this
subfamily. In total, 265 euophryine species (218 identified species of 69 genera
and 47 unidentified species of 12 potentially new genera) and 30 outgroups are
included in the phylogenetic analysis. The result strongly supports the
monophyly of euophryines and sheds some light on problems that have long
existed in their systematics, for instance, the taxonomic position of the South
American Euophrys species. The subfamily's divergence time is estimated by a
Bayesian approach using fossil calibration. The result shows the Euophryinae
likely originated in late Eocene or early Oligocene after the continents had
moved into their current positions. This finding is consistent with other analyses
that suggest much of salticid diversification occurred after the separation of the
continents of the Old World and New World, and indicates dispersal in
evolutionary history is the major mechanism to result in the current
intercontinental distribution pattern of this subfamily. This study also finds two
hotspots of euophryine species: Papua New Guinea and the Caribbean Islands.
Most jumping spiders known from these two areas are euophryines.
486
Harvestmen (Opiliones) thermo-preference reactions

under industrial pollution
Maxim P. Zolotarev & Ivan A. Kshnyasev
Institute of Plant and Animal Ecology, Ural Division of the Russian Academy of
Sciences, Ekaterinburg city, Russia, zmp@ipae.uran.ru, kia@ipae.uran.ru
Introduction
An ecosystem transformation under industrial pollution is one of classic
issues in contemporary ecology. At the same time much attention is focused on
functional or taxonomic/population changes in communities. The physiological
reactions are important basic components in population adaptation. The
preference reactions may be used as the indicator of ecological features of
species or populations. The long-term emissions of a copper-smelting factory
changes plant cover and temperature conditions and should affect thermo-
preference reactions of epigeic invertebrates. Harvestmen have a short range of
individual activity (approximately 200 m), cannot move over long distances by
air like insects or spiders so they are convenient research subject. Our goal in
this study is to test the assumption that harvestmen dwelling in close vicinity of
the copper-smelting factory have specific thermo-preference reactions.
Methods
The investigation was carried out in Pervouralsk district of Sverdlovsk
region (Middle Urals, Russia) near the Revda town during two summers (2007,
2008). The native biotopes in this region are fir forests with admixture of birch,
aspen and pine. Two sites located at the different distances from the
Sredneuralsk copper-smelting factory (SUMZ) were used: background zone (30
km) where pollution is at the average regional level and impact zone (2 km)
where the amplitude of daily temperature fluctuations in forest litter is higher
than on the unpolluted area. The most numerous three species of harvestmen has
been chosen for study: Nemastoma lugubre, Oligolophus tridens and Lacinius
ephippiatus. Harvestmen were caught by soil traps (2 days exposition). Live
individuals were delivered to a laboratory and placed inside experimental device
with temperature gradient from+8ºС to+30ºС. Six rounds of measurements
during day and night with 3 hours intervals were taken. The experiments have
been executed on August, 14th, 17th, 21st, 23rd, 29th and on September, 7th in 2007
(evaluation units (n) is individual): 25 L. ephippiatus, 89 N. lugubre and 5 О.
tridens from the background population and 5 О. tridens from the impact
population. Eleven rounds have been executed in 2008: on July, 14th, 17th, 22nd,
25th, 28th, on August, 10th, 13th, 17th, 21st, 26th and on September, 3rd (47 L.
ephippiatus, 70 N. lugubre and 57 О. tridens from the background and 5 О.
tridens from the impact population). For the temperature registration in litter top
horizon (depth 1 cm) during the period of laboratory research in 2007 we
installed the wireless sensors THERMOCHRON DS 1921 GF50. The statistical
487
analyses were carried out using Generalized Regression Models in Statistica

program (StatSoft, Inc. 2001). The mean daily temperature (meteorological
data), time within a day and a species (or pollution zone for two О. tridens
population) were used as continuous or categorical predictors for explanation of
variability in harvestmen preference temperature.
Results
Litter temperature
The temperature in forest litter where harvestmen spend many time at
their early stages has considerable influence on their activity. The similar
average temperature of the litter top horizon (registration unit (n) is sensor) in
the background zone (16.0±0.3ºC, sd=1.4, n=6) and in the impact zone
(16.2±0.5ºC, sd=2.4, n=4) was observed (F(1; 648)=0.2) but the variability of
daily (time*zone F(23; 648)=6.3) and seasonal (season*zone: F(4; 648)=20.8)
temperatures is higher in the impact zone.
Preference reactions
The daily average air temperature during the capture set in 2007 was less
by 1.3ºC than that in 2008 (F(1; 2422)=59.4). It is remarkable that in 2007 the
preference values of О. tridens (hereinafter М±se: 16.8±0.9) and N. lugubre
(17.8±0.3ºС) were a little lower (18.1±0.4, 18.2±0.2ºC in 2008 accordingly). On
the contrary L. ephippiatus in 2007 chose higher temperatures (20.1±0.5ºC) than
in 2008 (18.7±0.4ºC). The thermo-preference of all three species varied slightly
among two years (F(1;2309)=2.66). Significant interspecies differences in
thermo-preference were observed (F(2;2310)=16.1). L. ephippiatus is more
thermophilic species (19.2±0.3ºC). О. tridens and N. lugubre showed similar
reactions (18.0±0.4ºC and 18.0±0.2ºC accordingly). Contrary to expectations
thermo-preference of O. tridens inhabiting impact and background zone was
similar (18.4±0.7ºC, F(1; 532)=0.0002). The influence of average daily air
temperature on harvestmen thermo-preference is revealed (F(1; 2310)=8.58),
with decrease air temperature it decrease too.
Circadian dynamics in harvestmen preference temperature
All three species show similar circadian dynamics of preferred
temperatures (time: F(7; 2310)=4.9): the harvestmen chose high values at night
but low values at day, such feature is peculiar to crepuscular and nocturnal
invertebrates. There is only one coherent peak in all three species preference
temperature dynamics within 3 a.m. and 6 a.m. L. ephippiatus chose minimum
temperatures within the period since 12 a.m. till 3 p.m., background population
of N. lugubre and О. tridens – since 3 p.m. till 6 p.m. О. tridens inhabiting
impact territory displayed maximum width daily range thermo-preference
(5.1ºC). The thermo-preference circadian dynamics in both populations of О.
tridens are similar (zone: F(1; 532)=0.0002; time zone: F(7; 532)=1.28).
Harvestmen’s norm of reaction to temperature
Thermal preference reactions allow characterized organism as eury- or
stenothermic and we used a standard deviation as evaluation for it. N. lugubre
has a lowest standard deviation (2.0ºC), impact population of О. tridens (2.3ºC)
488
and L. ephippiatus (2.6ºC) characterized by intermediate value, background

population of О. tridens has a largest standard deviation (2.8ºC).
Discussion and conclusions

According to effects relative importance in explanation of thermo-
preference variability, mean daily air temperatures and its seasonal dynamics has
the greatest relevance (p=0.004), the circadian rhythm (p=0.014) and
interspecies differences (p=0.013) are significant too. Thus the influence of daily
average air temperature evidences the harvestmen acclimation to environmental
temperature dynamics and it is similar to reactions in insects. There is a point of
view that the temperature mode in early ontogenesis, average summer
temperature, physiological condition, age, gender etc affects on thermo-
preference. Because of the standard deviations of all three species are similar
(Hartley statistic Fmax=1.9), it is possible to say about narrow/wide species norm
of reaction only as a weak tendency. The distribution of the three species
somehow corresponds to its presence in polluted zones: N. lugubre has not been
catch in the impact area while L. ephippiatus and О. tridens has been found in
the odds 1:10. L. ephippiatus is larger than О. tridens and its distribution
possibly more depends on food supply and spatial restrictions (favourable places
for oviposition and young development). The relative narrow norm of reaction in
О. tridens impact population may be partially explained by its low abundance
and genetic diversity. Local microclimate differences in two habitats are not
enough for induced strong harvestmen physiological response. Nevertheless
some weak tendency exists, and it is possible that under more contrast
environmental conditions the difference will be stronger.
Acknowledgements
This work was supported by the program “Biological diversity”. We are
grateful to A.B. Kohan for his help in experiment execution and to K.V.
Maklakov for manuscript translation improvement.
489
Gravity and spider webs

Samuel Zschokke

Introduction
Most studies on animal displacements focus on horizontal movements,
which are equally easy and possible in all directions. In contrast, many web-
building spiders also move up- and downwards, because their webs have a
vertical expansion. Vertical displacement is not equally easy in both directions,
especially for larger organisms, because gravity causes climbing upwards to be
more exerting than walking downwards.
Vertical orb-webs usually have a form that deviates from a perfectly round
structure. Because this deviation is usually directed downwards – i.e. the part
below the hub is larger – it can be safely assumed that this asymmetry is
somehow linked to gravity. Similarly, almost all orb-webs are elongated, i.e.
they are higher than wide, and in almost all orb webs, spiders face downwards
when waiting for prey on the hub. In my presentation, I review the possible
reasons for the vertical asymmetry and elongation of orb-webs, and I review the
explanations why spiders face downwards. In addition, I propose two
hypotheses, how the influences of gravity may have been a driving force behind
the evolutionary transition from orb-webs to the derived three-dimensional webs
of theridiid and linyphiid spiders.
Functional explanations for orb-web asymmetry and elongation

Several reasons have been suggested for the vertical asymmetry and
elongation of orb-webs. Among the first explanations was one by a physicist,
who suggested that the stability of the web was improved, when the hub with the
relatively heavy spider is above the geometric centre of the web (Langer 1969).
Masters & Moffat (1983) were the first to suggest that the vertical
asymmetry of orb-webs is an adaptation to the different running speeds of
spiders in upward and downward direction, and that the distances from hub to
top and hub to bottom of the web are directly proportional to the spider's running
speeds in the two directions. This view was supported by the study of ap Rhisiart
& Vollrath (1994) and by the recent study of Zschokke & Nakata (2010). The
latter study additionally suggested that the different upward and downward
running speeds can also explain the almost universal downward orientation of
orb-web spiders on the hub and that this downward orientation further increases
the vertical asymmetry of orb-webs. In addition, it suggested that prey tumbling
also influences web asymmetry and that prey tumbling can explain why some
Cyclosa spiders face upward and build webs with larger upper parts.
490
The view that web asymmetry is an adaptation to prey capture was

challenged by Nentwig (1985) on the ground that Masters & Moffat had
assumed an equal distribution of prey in the web, which is not the case. Instead,
Nentwig found that more prey was caught in the upper part of the web. This
would, however, suggest that orb webs should generally have larger upper parts,
which is not the case.
Herberstein & Heiling (1999) found that web asymmetry increases with
spider weight (natural weight or experimentally added lead weights) and
suggested without further tests that the added weight interfered with spiral
placement in the upper web region, and that web asymmetry was a result of this
physical constraint during web construction. Coslovsky & Zschokke (2009)
could not repeat these results and found, in contrast, that web building in the
upper part of the web is more efficient than in the lower part.
Most orb-webs show a moderate web elongation, which is probably also
an adaptation to prey capture. There are examples of extremely elongated orb-
webs, some of which are an adaptation to capture moths (Eberhard 1975),
whereas others are adaptations to environments where only narrow webs can be
built (Kuntner et al. 2008).
I conclude that orb-web asymmetry is primarily an adaptation to prey
captures, and that it reflects the ability of the spider to run downwards faster than
upwards. It is likely that other factors (like web building) also play a role, but
they seem to be less important.
Gravity and evolution of araneoid webs

Orb-webs are the only spider webs where the two functions of intercepting
and retaining flying prey are spatially very close together. This approach
requires the entire web to be invisible, which in turn requires a regular
meshwork of very thin threads (Zschokke 2002). Such a meshwork cannot easily
be repaired without losing its properties, which, together with the fact that glue
droplets in ecribellate orb-webs become ineffective after some time, requires
regular re-building of the entire meshwork.
Building orb-webs requires some special adaptation of the spider, like the
ability to produce sticky silk and the ability to produce extremely strong silk. In
addition, since orb-webs can contain several thousand connections, orb-web
spiders require the ability to fasten two silk strands together in a very short time.
With theses extreme requirements, it is not surprising that orb-web spiders hold
the world record for the strongest natural material and, as far as I know, also
hold the world record for the fastest hardening glue: Araneus diadematus
requires less than 0.2 seconds to attach the sticky silk to a radius with glue from
its piryform gland.
Araneoid spiders date back to at least the early Cretaceous, and
phylogenetic analyses suggest that among them, orb-web spiders are ancestral to
linyphiid and theridiid spiders (Blackledge et al. 2009), suggesting that their
491
spatial webs have been derived from orb-webs. However, the transition of the
web structure during the evolution from orb web to spatial webs is unknown and
no hypothesis has been suggested how this transition may have occurred.
Most linyphiid and some theridiid spiders build horizontal sheet-webs
with knock-down threads above. While the sheet webs built by both groups are
very similar, they differ in the resting position of the spider. Linyphiid spiders
spend their time on the underside of their sheet, whereas theridiid spiders live in
a retreat above the sheet. Since spiders of both groups overwhelm their prey
from beneath, through the sheet, the resting place in theridiid sheet-webs
requires the spider to rush down and through the sheet when prey has been
caught.
I propose the hypothesis that linyphiid sheet webs were derived from
horizontal orb-webs, with additional knock down threads above. In a second
step, the strength of the horizontal web was increased, which made regular
rebuilding of the entire orb uneconomical, which in turn lead to a loss of the
regularity of the orb.
I further propose the hypothesis that the theridiid sheet-webs have evolved
from vertical orb-webs in which the area at the bottom of the web with high
sticky silk density became increasingly pronounced to catch all prey tumbling
down along the web. In a second step, the lower part of the web then became
wider and the upper part of the web lost its sticky silk and started to function as a
mere knockdown trap. In a third step, the dense web at the bottom also lost its
sticky silk.
References
ap Rhisiart A. & Vollrath F. 1994. Design features of the orb web of the spider,
Araneus diadematus. Behavioural Ecology, 5: 280-287.
Blackledge T.A., Scharff N., Coddington J.A., Szuts T., Wenzel J.W., Hayashi C.Y.
& Agnarsson I. 2009. Reconstructing web evolution and spider diversification
in the molecular era. Proceedings of the National Academy of Sciences USA,
106: 5229-5234.
Coslovsky M. & Zschokke S. 2009. Asymmetry in orb-webs: an adaptation to web
building costs? Journal of Insect Behaviour, 22: 29-38.
Eberhard W.G. 1975. The 'inverted ladder' orb web of Scoloderus sp. and the
intermediate orb of Eustala (?) sp. Araneae: Araneidae. Journal of Natural
History, 9: 93-106.
Herberstein M.E. & Heiling A.M. 1999. Asymmetry in spider orb webs: a result of
physical constraints? Animal Behaviour, 58: 1241-1246.
Kuntner M., Haddad C.R., Aljancic G. & Blejec A. 2008. Ecology and web
allometry of Clitaetra irenae, an arboricolous African orb-weaving spider
(Araneae, Araneoidea. Nephilidae). Journal of Arachnology, 36: 583-594.
Langer R.M. 1969. Elementary physics and spider webs. American Zoologist, 9: 81-
89.
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Masters W.M. & Moffat A.J.M. 1983. A functional explanation of top-bottom

asymmetry in vertical orbwebs. Animal Behaviour, 31: 1043-1046.
Nentwig W. 1985. Top-bottom asymmetry in vertical orbwebs: a functional
explanation and attendant complications. Oecologia, 67: 111-112.
Zschokke S. 2002. Form and function of the orb-web. In: Toft S. & Scharff N. (eds),
European Arachnology 2000. Aarhus University Press, Aarhus, pp. 99-106.
Zschokke S. & Nakata K. 2010. Spider orientation and hub position in orb webs.
Naturwissenschaften, 97: 43-52.
493
Spiral and web asymmetry in orb webs of

Araneus diadematus (Araneae: Araneidae)
Samuel Zschokke
Introduction
Most araneoid orb-web spiders build vertically asymmetric webs, in which
the capture area below the hub is larger than above the hub. Empirical and
theoretical studies suggest that this asymmetry is mainly an adaptation to the spider's
prey capture behaviour, and that it reflects the spider's ability to run downwards
faster than upwards.
The basic structure of an orb web consists of radial threads converging to a
central point, the hub. Around the hub, a sticky thread is placed in concentric spiral
loops, forming the capture area. Such a basic web structure implies a round capture
area with the hub in its geometric centre. However, as stated above, real orb webs
are vertically asymmetric, and there must therefore be modifications to this basic,
round web structure.
The modifications to the basic round web which spiders have been shown to
use to achieve vertically asymmetric webs are: 1) asymmetrically placed (eccentric)
sticky spiral loops and 2) additional sticky spiral threads below the hub. In addition,
spiders could build webs with a larger average mesh size below than above the hub.
The first aim of the present study is to describe these modifications in webs of
A. diadematus in detail and to assess their contributions to web asymmetry.
All orb-web spiders build their web in a specific order. After completing the
frame and the radii, they build the widely meshed, non-sticky auxiliary from the hub
outwards and then the sticky spiral from the periphery inwards. As most orb-web
spiders are virtually blind, the orientation during web building must follow non-
visual cues. These cues include gravity and the position of earlier laid threads in the
spider's reach. Consequently, when the orientation of the web relative to gravity is
altered, we can expect changes in web building. Similarly, we can expect changes in
web building, when previously laid threads are altered,
The second aim of the present study is to assess the influence of gravity on
the different web modifications and to describe the relationships among these
modifications during web building, i.e., how do these modifications influence each
other?

Second year juvenile and sub-adult A. diadematus were kept under laboratory
conditions in Plexiglas frames. In order to assess the effects of web orientation
(vertical vs. horizontal) during web building, some of these webs were laid
horizontally either during auxiliary spiral building or during sticky spiral building or
throughout spiral building. The webs were photographed and the attachment
positions of both spirals were measured. The coordinates of both spirals were then
converted into series of ovals of corresponding shape, of which I calculated the
494
eccentricity, i.e. their vertical displacement compared to the hub, using the formula
(upper-lower)/(upper+lower), yielding auxiliary spiral eccentricity and sticky spiral
eccentricity. In addition, the number of sticky spiral loops, the average mesh size of
the sticky spiral and the distance from the hub to the outermost turn of the sticky
spiral were assessed at the top and the bottom of the web. For these parameters,
asymmetry was calculated as above, yielding sticky spiral ratio, mesh size
asymmetry and web asymmetry respectively.
Results
Control webs (i.e. those built entirely in a vertical orientation) had on average
a negative web asymmetry, implying that the hub was placed above the geometric
centre of the capture area (as it is the case in most orb webs). Similarly, all sticky
spiral modifications (sticky spiral eccentricity, sticky spiral ratio and mesh size
asymmetry) were on average negative in the control webs, indicating that they all
contributed to the observed web asymmetry.
Web asymmetry and all modifications except mesh size asymmetry were less
negative in webs laid horizontally during auxiliary spiral building, suggesting that
they were in some way influenced by the orientation of the web during auxiliary
spiral building.
The orientation of the web during sticky spiral building had a significant
influence only on mesh size asymmetry. Interestingly, however, the (non significant)
influence of the orientation during sticky spiral building was opposite to that of the
orientation during auxiliary spiral building; a vertical orientation during sticky spiral
building tended to reduce web asymmetry and its modifications. This means, that
the webs with the strongest asymmetry were those, which were in a vertical position
during auxiliary spiral building and in a horizontal position during sticky spiral
building, and not those in vertical orientation during the entire web building.
A causal model suggested that web orientation during auxiliary spiral
building influenced auxiliary spiral eccentricity, which in turn influenced sticky
spiral eccentricity and web asymmetry. Web asymmetry influenced sticky spiral
ratio and, together with sticky spiral ratio and web orientation during sticky spiral
building influenced mesh size asymmetry.
Conclusions
1. All three modifications (sticky spiral eccentricity, sticky spiral ratio and mesh size
asymmetry contribute to the overall web asymmetry in A. diadematus.
2. The eccentricity of the auxiliary spiral has a strong influence on the eccentricity of
the sticky spiral, which supports the theory that the auxiliary spiral is used as a
guiding line during building of the sticky spiral in A. diadematus.
3. The spider uses different rules to build the two spirals. During auxiliary spiral
building, gravity has a great influence, whereas during sticky spiral building the
spider mainly relies on previously laid threads.
495
Salticidae (Arachnida: Araneae) of Australia – a synthesis
Marek Żabka
Department of Zoology, University of Podlasie, Siedlce, Poland, marekzabka@ap.siedlce.pl
The beginnings of the taxonomic studies on Salticidae of Australia go

back to the second half of ninetieth century. Since then almost 360 species
representing more than 70 genera have been described/recorded; large part of
them studied and/or described by the author over the last 25 years of field and
laboratory research.
Due to long-term post-Gondwanan isolation, unique geological, biotic and
climatic history, the great majority of species and most genera are endemics of
local origin. Some have spread towards New Guinea, adjacent islands and single
species – even to New Zealand. Several genera (e.g. Neon-complex) so far
thought to be of “northern origin” seem to have originated in Australia and show
impressive diversity. Others (e.g. Phintella, Langona) enlarge the list of (likely)
northern immigrants.
The author’s results on relationships and biogeography are based upon
morpho-geographical and biotic-climatic data. Recent molecular results
provided by Maddison and co-workers complete and verify our view of the
Australian salticid fauna.
The talk presents the summary of current knowledge on Australian salticid
taxonomy and biogeography.
496
Males prefer mid-aged females that are UV-induced

fluorescent brighter and greener
Jia Fen Seah1, Matthew L.M. Lim1 & Daiqin Li1,2

1
2
dbslidq@nus.edu.sg
Males are typically more elaborately ornamented than females; and they
are classically considered to be the competing, but not the choosing, sex,
because their investment in offspring is much lower. Females in some species
are highly ornamented, but adaptive significance of female ornamented is largely
unknown although recent studies suggest that selection acting on females might
be a widespread cause of female ornamentation. As a sexually selected trait,
female-specific ornamentation, like that of males, may be often condition
dependent, indicting female quality, on which males may make mate choice
decisions. Here, we investigate Cosmophasis umbratica, a jumping spider that
has sexual dimorphic UV-induced fluorescent ornamentation, in which females
have palps with UV-excited bright green fluorescence that is absent in males,
and provide evidence that female palp fluorescence is strongly age dependent in
this species. Mid-aged females are brighter and greener than younger and older
females. When given a choice between two similar-sized females with different
ages (i.e., young vs. mid-age and mid-age vs. old) which are correlated with palp
fluorescence, male spent more time courting mid-aged females with brighter and
greener fluorescent palps and less time courting young or old females that have
dull and yellow fluorescent palps. This provides strong experimental evidence
that C. umbratica males make use of the female’s age-dependent fluorescent
markings as a reliable signal in their choice of mate, supporting the hypothesis
that female ornamentation is sexually selected.
497
Index of Authors
Abbas, Jelodar Zadeh 309

Abdigoudarzi, Mohammad 168
Adamowicz, Sarah J. 84
Agnarsson, Ingi 51, 52, 83, 169, 240
Aisenberg, Anita 53, 55
Alam, Imtiaz 437
Alberti, Gerd 57, 250
Albo, Maria José 58, 59, 61
Almeida-Silva, Lina M. 62
Alpers, Georg 453
Álvarez-Padilla, Fernando 64, 65, 114
Andrade, Alessandra R.S. 290, 462
Andrade, Maydianne C.B. 421
Angyal, Dorottya 207
Araujo, Diego P. 66
Araujo, Douglas 124, 284
Arnedo, Miquel A. 67, 80, 114, 240, 243, 260, 320
Avilés, Leticia 69, 173
Azarkina, Galina 70
Baba, Yuki 71
Baehr, Barbara C. 73
Barth, Friedrich 74
Bartos, Maciej 75, 76
Bayer, Steffen 77
Benati, Katia R. 290, 462
Benavides, Ligia Rosario 79, 306
Bergler, Helmut 148
Bidegaray-Batista, Leticia 80, 260
Bilal, Muhammad 436
Bilde, Trine 59, 61, 263, 451
Binford, Greta J. 82, 281
Birch, Debra 163
Bird, Tharina 217
Blackledge, Todd A. 52, 83, 169, 329
Blagoev, Gergin A. 84, 112, 143
Blamires, Sean J. 85
Blick, Theo 86, 194
Bodkhe, Atul 87, 429
Bolzern, Angelo 89
Bonaldo, Alexandre B. 94, 251, 371
Boniecki, Paweł 315
Bonte, Dries 342
Botta-Dukát, Zoltán 390
Bowden, Joseph J. 91
Braddy, Simon J. 117
Bradley, Robert 331
498
Braga, Pilar L. Maia 172

Bragagnolo, Cibele 93
Brescovit, Antonio Domingos 62, 124, 259, 285, 312, 350
Brown, Georges 251
Buchar, Jan 115
Buddle, Christopher M. 91
Bumrungsri, Sara 352
Burel, Françoise 459
Butt, Abida 414, 436, 437
Canard, Alain 246, 354
Candiani, David F. 94
Cardoso, Pedro 320
Carmichael, Anthea 170, 432
Carvalho, Leonardo Sousa 94, 97, 124, 251, 285
Cera, Inese 100
Chapman, Eric G. 322, 476
Cheng, Ren-Chung 368
Chiarle, Alberto 104
Chikhale, Mahesh 415
Chishiki, Yuki 301
Chotiwong, Wimolwan 472
Christophoryová, Jana 108
Coddington, Jonathan 52, 83, 329
Conner, Ashley 388
Copperi, Sofia 135, 139
Cotterill, Melanie A. 110
Court, David 439, 484
Crosby, Trevor K. 469
Cruickshank, Rob H. 265
da Fonseca Ferreira, Rafael 172
da Rocha Dias, Maria de Fatima 465
da Silva Souza, Elene 479
Dankaninová, Lenka 157
DaSilva, Marcio B. 111
de Santana Varjão, Sheila Luzia 290, 462
de Souza Sá, Fernanda 172
Decae, Arthur 342
Decaёns, Thibaud 251
Deltshev, Christo 112, 471
Derraik, José G.B. 468
Deruytter, David 342
Deshmukh, Shivaji 457
Deshmukh, Ujjwala S. 113, 415
Dias, Marcelo A. 290, 462
Dimitrov, Dimitar 114, 189, 243
Dippenaar-Schoeman, Ansie S. 145
Dolejš, Petr 115
Dulíková, Lenka 227
499
Dunlop, Jason A. 117, 118, 300

Durrant, Bradley J. 119
Dymek, Agnieszka 435
Eberhard, William G. 120
Edwards, G.B. 121
Elgar, Mark A. 122
Elias, Damian O. 480
Esposito, Lauren A. 123
Fagundes de Mattos, Viviane 124, 285
Fain, Leighton 388
Faltýnek, Fric Z. 311
Farley, Roger D. 127
Fauler, Günter 148
Fedoriak, Mariia M. 130
Felska, Magdalena 156, 289
Fernández, Miguel-Angel 320
Fernández-Montraveta, Carmen 369
Ferretti, Nelson 135, 139
Fet, Victor 143
Fetykó, Kinga 191
Figueiredo, David F. 97
Foellmer, Matthias W. 144
Foord, Stefan H. 145
Forman, Martin 146
Föttinger, Petra 148, 362
Framenau, Volker W. 119, 151
Freire Jr., Geraldo B. 152
Freire, Ronald Bastos 391
Frick, Holger 155
Friederichs, Anja 118
Gabryś, Grzegorz 156
Gajdoš, Peter 157
Gale, George A. 352
Gao, Jie 159
García, Luis Fernando 160
Gardzińska, Joanna 162
Gavish-Regev, Efrat 164
Gawryszewski, Felipe M. 163
Gerdes, Antje 453
Gillespie, Rosemary G. 80, 480
Giribet, Gonzalo 57, 114, 165, 256, 306, 412, 413, 478
Gnelitsa, Valery A. 166
Gomes, Jerriane O. 97
Gonzaga, Marcelo O. 173
González, Alda 139
Gonzalez, Edmundo 167
González, Macarena 55
Gonzalez-Filho, Hector M.O. 259
500
Goudarzi, Hamid Reza 168, 298

Green, David I. 331
Gregorič, Matjaž 169, 233
Grinvald, Marea 449
Griswold, Charles 64, 65, 170, 432, 480
Guadanucci, José Paulo Leite 171, 172
Guevara, Jennifer 173
Guimarães, Marcos Vinicius A. 290, 462
Gutjahr, Melanie 57, 250
Haddad, Charles R. 146, 174, 221, 227, 230
Hajdamowicz, Izabela 176, 315, 420
Harms, Danilo 177, 178
Harvey, Mark S. 73, 119, 177, 178, 179, 359, 375
Harwood, James D. 322, 476
Haupt, Joachim 180
Hedin, Marshal 227
Hendrickx, Frederik 425, 458
Henriques, Sérgio S. 227
Herberstein, Marie E. 163
Hesselberg, Thomas 181
Hoefler, Chad D. 388
Höfer, Hubert 194, 361, 399
Holm, Christina 183
Honda, Yoshiko 184
Hore, Upamanyu 191
Horigane, Mari 184
Hormiga, Gustavo 79, 114, 164, 189, 243, 256, 257, 306, 448
Horváth, András 390
Horváth, Roland 193
Hou, Chueh 190
Hsieh, Samuel Yu-Lung 196, 197
Huber, Bernhard A. 199, 230
Huckstorf, Katarina 200
Hudson, Peter 151
Hula, Vladimír 201, 311
Indicatti, Rafael P. 251
Isaia, Marco 104, 202, 325
Ivanova, Natalia 143
Jäger, Peter 304, 305, 394
Jastrzębski, Piotr 204
Jocqué, Rudy 205
Joseph, John 282, 404
Joshi, Dinesh 429
Judson, Mark L.I. 206
Kadu, Deepa 415
Kancsal, Béla 207
Kanniparambil, Sunil Jose 208
Karaman, Ivo M. 212, 362
501
Katušić, Luka 215

Kawaguchi, Minako 443
Kayedi, Mohammad Hassan 310
Kean, John M. 469
Klann, Anja E. 217
Klußmann, Bastian J. 211
Kojima, Jun-ichi 359
Komnenov, Marjan 143
Kondulkar, Sunil 395, 429
Konrad, Jessika 218
Koponen, Seppo 219
Koraimann, Günther 148
Korenko, Stanislav 220
Kořínková, Tereza 221, 227, 230
Košulič, Ondřej 201
Kovblyuk, Mykola M. 224, 307
Kraeski, Marcia G. 285
Král, Jiří 67, 146, 221, 227, 230
Kralj-Fišer, Simona 233, 235
Kropf, Christian 155
Krumpál, Miroslav 237, 409
Krumpálová, Zuzana 108, 237
Kshnyasev, Ivan A. 487
Kuntner, Matjaž 169, 233, 235, 240
Kunz, Karin 241
Kupryjanowicz, Janusz 242
Labarque, Facundo M. 243
Laborda, Álvaro 55
Lacava, Mariángeles 160
Lai, Cheng-Hui 442
Laška, Vratislav 386
Lavelle, Patrick 251
Lazarov, Stoyan 112
Lehtinen, Pekka T. 244
Leis, Hans-Jörg 148
Lens, Luc 425
Leroy, Boris 246
Li, Daiqin 66, 233, 439, 484, 497
Lim, Matthew L.M. 497
Linsenmair, Karl Eduard 196, 197
Lipke, Elisabeth 57, 250
Llandres, Ana L. 163
Lokovšek, Tjaša 240
Lo-Man-Hung, Nancy F. 97, 251
Longhorn, Stuart J. 254, 255
Lopardo, Lara 114, 256, 257, 258
Lubin, Yael 295, 322
Lucas, Sylvia M. 259
502
Lukanc, Tjaša 235

Machač, Ondřej 449
Macías-Hernández, Nuria 260
Maddison, Wayne P. 261, 383, 486
Magura, Tibor 193
Maia, Marília G. 262
Majer, Marija 183, 263
Malumbres-Olarte, Jagoba 265
Marhabaie, Mohammad 268
Marichal, Raphaël 251
Marris, John 469
Marson, Michael 144
Martins, Marlucia 251
Marusik, Yuri M. 269, 272, 331
Masta, Susan 255, 277
Mathew, Elizabeth V. 278
Mathew, Mundackatharappel J. 282, 404
Maxwell, Elise N. 281
Mąkol, Joanna 156, 206, 289
McNeil, Andrew 331
Meier, Rudolf 439
Melo, Tércio S. 290, 462
Meriste, Mart 293
Merrell, Andrew V. 82
Mestre, Laia 295
Michalik, Peter 296, 297, 470
Mikula, Jan 386
Miller, Jeremy 268, 281
Mirakabadi, Abbas Zare 298
Mišurcová, Jana 449
Mitov, Plamen G. 299, 300
Miyashita, Tadashi 71, 301
Mohammadpur, Naser 298
Monod, Lionel 303
Moradmand, Majid 304, 305
Motta, Paulo C. 152, 335
Moya-Laraño, Jordi 144
Muelelwa, Mulalo I. 145
Müller, Carsten H.G. 385
Murienne, Jerome 306
Murugesan, S. 404
Musilová, Jana 227, 230
Muster, Christoph 194
Nadolny, Anton A. 307
Narayanan, Sujatha 439
Navidpour, Shahrokh 309, 310
Nayebzadeh, Hassan 310
Neckel-Oliveira, Selvino 97
503
Neidert, Dóra 390

Nentwig, Wolfgang 155
Niedobová, Jana 201, 311
Nogueira, André do Amaral 312
Olejniczak, Izabela 315
Oleszczuk, Maria 176, 316
Omelko, Mikhail M. 317
Ono, Hirotsugu 319
Opatová, Věra 67, 320
Opatovsky, Itai 322
Oromí, Pedro 260
Ortiz-Villatoro, David 339
Ott, Ricardo 119
Ovtcharenko, Vladimir I. 323
Ozimec, Roman 324
Panzera, Alejandra 339
Papler, Tadeja 235
Paschetta, Mauro 325
Pastuchová, Markéta 230
Paterson, Adrian M. 265
Patoleta, Barbara 204, 328
Paula, Felipe S. 259
Paulsch, Detlev 194
Pekár, Stano 220, 221, 329
Peng, Po 330
Penney, David 272, 331
Perdomo, Cintya 339
Pereira, Rommel B. 335
Peres, Marcelo Cesar L. 290, 462
Peretti, Alfredo V. 53
Pérez-Miles, Fernando 135, 139, 339
Pétillon, Julien 246, 342, 354, 459
Phillips, Craig B. 468
Piacentini, Luis N. 243
Pinto-da-Rocha, Ricardo 93
Piterkina, Tatyana V. 343
Planas, Enric 344, 369
Platen, Ralph 345
Polchaninova, Nina Yu. 346
Polotow, Daniele 350
Pompozzi, Gabriel 135, 139
Ponksee, Booppa 352
Pons, Joan 243
Postiglioni, Alicia 58
Postiglioni, Rodrigo 55
Praxedes, Catarina 251
Prendini, Lorenzo 123, 167, 303
Preziosi, Richard F. 331
504
Prószyński, Jerzy 353

Psota, Václav 201
Puzin, Charlène 354
Quasin, Shazia 357, 454
Rahmadi, Cahyo 359
Ramírez, Martín J. 243, 296
Raspotnig, Günther 148, 362, 396
Raub, Florian 361, 399
Raven, Robert J. 365
Rayor, Linda S. 367
Řezáč, Milan 67, 230
Rheims, Cristina Anne 376, 379
Ribera, Carles 344, 369
Ricetti, Janael 371
Richardson, Barry J. 373
Rittschof, Clare C. 297
Rix, Michael G. 375
Rosenstand, Marie 61
Rozwałka, Robert 204, 242
Ruiz, Gustavo R.S. 381, 383
Runge, Jens 385
Růžička, Vlastimil 386
Rypstra, Ann L. 388
Salomon, Maxence 389
Samu, Ferenc 390, 431
Santana de Sá, Rodrigo 391
Santos de Souza, Rita de Cássia 391
Saraceno, Tomas 394
Satish, Akarte 395
Saucedo, Alma 432
Schaider, Miriam 362, 396
Scharff, Nikolaj 52, 83, 164
Scheuermann, Ludger 399
Schönhofer, Axel L. 401
Schuster, Reinhart 250
Seah, Jia Fen 497
Sebastian, Pothalil Antony 278, 282, 404, 425
Selden, Paul A. 408
Sensenig, Andrew 52, 83
Šestáková, Anna 409
Seyfulina, Rimma R. 410
Sharma, Prashant 412, 413
Sherawat, Sher Muhammad 414
Shirbhate, Milind 395, 415
Silva, Paulo Roberto R. 94
Simó, Miguel 55
Sirvid, Phil J. 468
Siuda, Paweł 418
505
Smrž, Jaroslav 115

Snegovaya, Natalya Yu. 419
Sotthibhundhu, Sunthorn 352
Stam, Edward M. 145
Stańska, Marzena 176, 315, 420
Steineck, Dieter 394
Stoltz, Jeffrey A. 421
Sudhikumar, Ambalaparambil Vasu 278, 425
Suesdek, Lincoln 94
Svenning, Jens-Christian 263
Szinetár, Csaba 193, 207, 431
Szita, Éva 390
Szuts, Tamas 432
Szymkowiak, Paweł 433, 435
Tahir, Hafiz Muhammad 414, 436, 437
Takenaka, Koji 445
Talarico, Giovanni 438
Tan, Trina V.Z.Y. 439
Taylor, DeMar 184
Toft, Søren 59, 61
Tóthmérész, Béla 193
Trontelj, Peter 240
Tselouiko, Stéphanie 251
Tseng, Huei-Jen 441
Tso, I-Min 85, 442
Tsurusaki, Nobuo 443, 445
Tu, Lihong 159, 448, 473
Tuf, Ivan H. 237, 386, 449
Tuni, Cristina 61, 451
Turner, Steven 254
Ubick, Darrell 64, 65
Uhl, Gabriele 241, 258, 452, 453, 470
Uniyal, Virendra Prasad 191, 357, 454
Vairale, Amit B. 415
van Heerden, Jacques 455
van Helsdingen, Peter 456
Vankhede, Dinesh 429
Vankhede, Ganesh 457
Vanthournout, Bram 458
Varet, Marion 459
Vasconcellos, Alexandre 262
Vasconcellos-Neto, João 173
Velasquez, Elena 251
Velikonja, Urška 235
Venticinque, Eduardo Martins 312
Viera, Carmen 160, 465
Vink, Cor J. 265, 468, 469
Vítková, Magda 227, 230
506
Vöcking, Oliver 470

Voger, Alfried 453
Vollrath, Fritz 181
Vrenosi, Blerina 112, 471
Vungsilabutr, Wipada 472
Waldock, Julianne M. 119
Wang, Fang 473
Wawer, Wioletta 474
Weintraub, Phyllis G. 322
Welch, Kelton D. 476
Werner, Ulrich 420
Willemart, Rodrigo H. 478, 479
Winther, Gudrun 61
Wirkner, Christian S. 200, 211
Withers, Philip J. 331
Wolf, Marcos José 124
Wood, Hannah Marie 170, 480
Wright, Simon 481
Wulff, Christof 394
Yamasaki, Takeshi 482
Yap, Laura-Marie Y.L. 439, 484
Yi, Youguang 439
Yip, Eric C. 485
Ysnel, Frédéric 246, 354
Zakharov, Boris 323
Zemljič, Larisa 235
Zhang, Junxia 486
Zhang, Shichang , 233
Zhukovets, Evgeni M. 130
Ziegler, Ashley 388
Zobel-Thropp, Pamela A. 82
Zolfagharian, Housein 298
Zolotarev, Maxim P. 487
Zschokke, Samuel 394, 490, 494
Żabka, Marek 48, 50, 496
507

Book of Abstracts - 18th International Congress of Arachnology 2010

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University of Podlasie

Editor: Marek Żabka

Congress Honorary Committee

Congress Organizing Committee

Technical assistance: Mrs. Małgorzata Kozłowska, Mr. Łukasz Nicewicz

Congress logo: Joanna Gardzińska

To Colleagues and Friends – Arachnologists,

Organizacja Kongresu to efekt zbiorowego wysiłku i życzliwości wielu osób,

Władze Akademii Podlaskiej, Dziekani Wydziałów Przyrodniczego i Nauk

W bieżącym roku obchodzimy pięćdziesiątą rocznicę Pierwszego

Spotykamy się w Uczelni, w której przed 40 laty Prof. Jerzy Prószyński

Wielu gości Kongresu jest w Polsce po raz pierwszy, a większość nigdy

W imieniu Komitetu Organizacyjnego,

We are grateful to the Arachnologists from 6 continents and almost 50

The Congress is being held at the University of Podlasie, where 40 years

Many Congress participants and accompanying persons have not been in

Phylogeny and sociogeography of Anelosimus spiders

The diversity and distribution of species across habitats and landmasses

Behavioural and biomaterial coevolution in spider orb webs

Ingi Agnarsson1, Todd A. Blackledge2, Andrew Sensenig3,

Mechanical performance of biological structures, such as tendon, byssal

Costs of burrow-digging and sexual dimorphismin immune

Burrow digging on the sand has been reported as an energetically expensive

implant was removed and photographed under a dissecting microscope. Immune

Spatial distribution along the sand dunes and temperature

Anita Aisenberg1, Macarena González1, Álvaro Laborda2,

Allocosa brasiliensis and Allocosa alticeps are two sympatric and

according to the stage in A. brasiliensis (χ2=4.92, df=2, P=0.09). A. alticeps

Fine structure of dimorphic sperm in mite harvestmen

The spermatozoa of mite harvestmen have received special attention in

Male meiosis: sex chromosomes and heterochromatin

Nuptial gifts are heritable characters associated with reproduction, and

Nuptial gift implies a high cost for males of the nursery

In both vertebrates and invertebrates poor feeding or an unbalanced

Worthless donations as a male reproductive tactic

Maria José Albo1, Gudrun Winther2, Marie Rosenstand2,

Nuptial gifts such as captured prey, regurgitations, glandular fluids, part of

Cladistic analysis of the spider family Titanoecidae

¹ Universidade de São Paulo, Brazil, linamas@gmail.com

Family Titanoecidae was proposed by Lehtinen to include five genera:

Anuvinda remains monotypic. The female of Anuvinda escheri Reimoser is

Interfamilial phylogenetic relationships of goblin spiders,

Cybertaxonomy has made possible simultaneous collaboration among

Three new genera of goblin spiders from Madagascar

Fernando Álvarez-Padilla1, Darrell Ubick2 & Charles Griswold3

Currently goblin spiders, the spider family Oonopidae, are subject to

Shooting stars in an Asian tropical forest:

Animal colours are achieved by two processes: chemical (in case of

Does karyotype change drive speciation in the woodlouse

Intrinsic and extrinsic factors in social evolution and

Department of Zoology, University of British Columbia, Vancouver, Canada,

The geographical distribution of species with different social systems

Aelurillinae (Araneae: Salticidae) of the world

Siberian Zoological Museum, Novosibirsk, Russia, urmakuz@gmail.com

Aelurillinae is one of 20 subfamilies of Salticidae with ca 5000 described

Host-associated differentiation in the relative leg length

Material and methods

the walking speed of A. kumadai between the population on experimental

Results and discussion

Who are these Goblin Spiders?

Barbara C. Baehr1 & Mark Harvey2

In 2002 Burger, Nentwig and Kropf described a quite unusual Opopaea

Spider senses: how it all fits together

Life Sciences, Department of Neurobiology, University of Vienna, Austria,