Manual of Tropical Bry Ology

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Manual of Tropical Bryology
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AN INTERNATIONAL JOURNAL ON THE BIOLOGY OF TROPICAL BRYOPHYTES
MANAGING EDITOR: J.-P. FRAHM

SCIENTIFIC EDITOR: W.R. BUCK
EDITORIAL BOARD:
Y. LEÓN-VARGAS (Universidad de los Andes, Mérida), B.J. O´SHEA (London), B.C. TAN (National
University of Singapore).
No. 23 www.bryologie.uni-bonn.de 2003
Manual of
Tropical Bryology
by
Jan-Peter Frahm
with contributions by
BRIAN O´SHEA, TAMAS POCS, TIMO KOPONEN, SINIKKA PIIPPO, JOHANNES ENROTH,
PENGCHENG RAO & YIN-MING FANG
ISSN 0935 - 5626

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Manual of tropical bryology 1
TROPICAL BRYOLOGY 23 (2003)

2 Frahm

Manual of
Tropical Bryology
by
Jan-Peter Frahm
with contributions by
TAMAS POCS, BRIAN O´SHEA, TIMO KOPONEN, SINIKKA PIIPPO, JOHANNES
ENROTH, PENGCHENG RAO & YIN-MING FANG

4 Frahm

Preface
It is still a fact that most bryologists per area are found in the temperate regions of the northern
hemisphere, who have spent up to 200 years (as in Europe) in the exploration of their bryofloras
with the result that these countries have not only floras for identification of the comparably low
number of species but some countries have already detailed grid maps of the distribution of all
species. On the other hand, there are vast regions in the tropics which are very insufficiently explored.
So far, the knowledge of bryophytes in these regions was predominantly provided by scientists from
North America, Europe or Japan. Still much work is done by scientists and - during the past decades
also increasingly - even by advanced amateurs from these countries. Regretably, these activities are
often misunderstood by local biologists in the tropics and especially by the authorities of these states
as scientific exploitation, and recently collecting of material for genetic studies as plundering of
genetic ressources, which is nonsense but makes research difficult or even impossible in such countries.
In this regard, the question raises why there are so few bryologists in tropical countries and even no
bryologists in many countries? Usually, the lack of ressources such as laboratories, money, libraries,
herbaria etc. is presented as arguments, which does not match the point, since many bryologists in
industrial countries suffer from similar restrictions and sometimes have worth working consitions
than colleagues in tropical countries, but make nevertheless valuable contributions to tropical bryology.
Even amateurs have contributed a lot to tropical bryology in the past. The fewest bryologists work in
such famous places as Missouri or New York Botanical Garden. Many of them are from eastern,
former communistic countries and never gave up to promote tropical bryology under these conditions.
In my opinion, the most crucial point is that students in tropical countries get not in contact with
bryophytes, and mainly because of the lack of literature. Nobody can expect that students pick up a
subject for their thesis if there is no literature available. This manual is therefore devoted to these
students. Possibilities to gain a bryological training in industrial countries and paid by these countries
were used only by few students, although available. And if these students do not come to us, we have
to go to them (which is even cheaper). This is the reason for the increased number of courses on
tropical bryology in the past.
Another group of potential users of this manual are bryologists, especially amateur bryologists from
industrial countries. The bryological exploration in these countries has made such progress during
the last 200 years that these countries are comparably good explored. Recent activities concern the
distinction of “small species” or mapping. Some countries e.g. Great Britain are even “ready mapped”.
Therefore the bryological activities are directed towards tropical countries. Checklists can even be
compiled without leaving the desk. In the 19. century, collections in tropical countries were usually
performed during scientific expeditions financed and operated by gouvernments of colonial states,
rarely by individuals, adventurers, who gained their lives by selling their collections. Later, scientific
institutions organized trips to the tropics to enhance their collections. Today, relatively cheap air
faires allow amateurs from industrial countries to spent their holidays in the tropics and to make new
discoveries, which they cannot make at home. By this way, some amateurs have made most valuable
contributions to the floristic knowledge of tropical countries, and by this way the members of the
British Bryological Society Tropical Bryology Working Group are more numerous than all bryologists
in Africa or SE-Asia. The acitivities within the society have led to the preparation of checklists,
detection of numerous new records and even new species in Malawi and Uganda. Local botanists in
the tropics should appreciate this and not interprete it as new scientific imperialism. It brings us
small steps forward towards an understanding of tropical bryology. Regretably, bryological acticities
of “tourists” were more numerous in some tropical countries than those of native full time bryologists.
Paradoxically, activities of bryologists from non-tropical countries in the tropics were and are generally

6 Frahm
still much larger than those of local bryologists. The activities of amateurs contradict also the argument
that bryological research in the tropics is not possible without large libraries, herbaria, lab space, or
personel. Even lacking budgets are no arguments: the German Research Foundation paid hardly
more for projects in the tropics than flights (which local botanists do not need) and a car rental
(which local botanists do not need because of an own car). And the professors stay in tents and cook
on kerosine stoves, something which many local botanists from tropical countries would not do.
Therefore this small manual may also stimulate bryologists from extratropical countries to focus
their interest in tropical bryology.
A short previous version of this manual was originally written by the author as script for the „curso
taller sobre briofitos tropicales“ held at the Jardin Botanico de Mérida, Venzuela, Feb. 24. - March 7.
1997 together with Tamas Pócs. It covered some topics of the course in a textfile, which was distributed
on disk, but was not complete. The printout as well as the disk version did not include illustrations
(which were presented during the course as slides). Furthermore, this script was focussed on the
neotropics.
Aim of this course and this script was to provide some knowledge of tropical bryology. It required,
however, a basic knowledge of bryology, for which textbooks shall be consulted. Therefore general
topics such as life cycle, anatomy, morphology, and systematics are omitted here.
More courses on Tropical Bryology have been helt for European postgraduate students in 1993,
1998 and 2000 in the Division of Systematic Biology, University of Helsinki, organized by Timo
Koponen and Johannes Enroth, made possible by funds of the European Community. Due to the
lack of alternatives, this Mérida-script was used as compendium for the courses in 1998 and 2000.
For that purpose, Timo Koponen and Johannes Enroth added illustrations and did some editorial
work. It was, however, still a fragment. Some efforts were later undertaken to get some colleagues
interested to complete this course script to a short manual, but in vain. Therefore I started to complete
this script myself with the consciousness that a joined work of several specialists would have been
better. I know that some colleagues argue that such a booklet should have better not written since it
is not complete or contains errors. The latter is true, but in my opinion the lack of such a small
compendium is even worth. But only those who do nothing make also no mistakes except for the
mistake to do nothing. It is therefore hoped that this small volume of „TROPICAL BRYOLOGY“
will raise interest in this much neglected group of plants, increase bryological activities in the tropics
and stimulate to include bryophytes in studies of biodiversity and ecology of tropical forests.
This manual has some limitations: because of copyright problems, the illustrations are mostly taken
from own publications or from the journal Tropical Bryology. Redrawing existing illustrations would
habe been too time consuming. Next, the explanations are mostly based on tropical rain forest bryology.
Other habitats are covered only to an lesser extend. Reason is simply that bryological research in the
tropics is focussed on rain forests, which harbours most of the tropical bryophytes. This will not say
that other bryological aspects in the tropics are not as interesting. And finally, the explanations are
not complete in some respects, it is not a complete textbook but still an extended course script, but
anyway, this manual will now be available not only for course participants but a wider range of
interested bryologists especially in the tropics, and - as mentioned before - better than nothing.
This volume is, like all volumes of „TROPICAL BRYOLOGY“ available as faksimile edition on
CD-ROM as pdf-file, which allows to print all or parts of it or to copy and paste parts of the text e.g.
references into other applications.
I wish to thank Brian O´Shea and Tamas Pócs, who contributed parts to this volume, as well as Timo
Koponen, Sinikka Piippo, Johannes Enroth, Pengcheng Rao and Yin-Ming Fang, who had completed
earlier versions of the course script.
Bonn, April 2003 Jan-Peter Frahm

CONTENTS
1. Introduction.....................................................................................................................................9
2. Diversity of bryophytes in the tropics............................................................................................13
2.1 Global diversity.....................................................................................................................13
2.2 Regional diversity..................................................................................................................14
2.2.1 Neotropics......................................................................................................................14
2.2.2 Tropical Africa................................................................................................................16
2.2.3 Tropical SE-Asia............................................................................................................18
2.3 Local diversity ......................................................................................................................18
3. Origin and age of tropical bryophytes...........................................................................................23
3.1 Evidence from fossils.............................................................................................................23
3.2 Evidence from plate tectonics................................................................................................26
4. Morphological adaptations to tropical rain forest environments.....................................................29
4.1 Life and growth forms............................................................................................................29
4.2 Water conducting and water storing structures......................................................................34
4.3 Life strategies.........................................................................................................................36
5. Ecology of tropical bryophytes.....................................................................................................39
5.1 Habitats...................................................................................................................................39
5.1.1 Epiphytes.......................................................................................................................39
5.1.2. Epiphylls......................................................................................................................44
5.1.3. Rotten wood.................................................................................................................47
5.1.4 Tree ferns......................................................................................................................48
5.1.5 Unusual substrates.........................................................................................................48
5.2 Altitudinal zonation................................................................................................................49
5.2.1 Determination of altitudinal belts with bryophytes.........................................................49
5.2.2 Results..............................................................................................................................50
5.3 Phytomass and water storing capacity...................................................................................53
5.4 Human impact........................................................................................................................56
6. Ecophysiology...............................................................................................................................59
6.1 Temperature, light and humidity............................................................................................59
6.2 Nutrients.................................................................................................................................64
6.3 Various topics.........................................................................................................................65
7. Bryosociology...............................................................................................................................69
8. Phytogeography.............................................................................................................................73
8.1 General tropical bryogeography.............................................................................................74
8.1.1 Endemism........................................................................................................................74
8.1.2 Relics...............................................................................................................................74
8.1.3 Disjunctions.....................................................................................................................75
8.1.4 Human influence.............................................................................................................79
8.2 Regional bryogeography.......................................................................................................79
8.2.1 Central and South America.........................................................................................79
8.2.2 Tropical Africa............................................................................................................85
8.2.3 Tropical Asia...............................................................................................................85
9. Bioindication...................................................................................................................................89
10. Conservation................................................................................................................................91

8 Frahm
Appendix...........................................................................................................................................95
I. Collecting bryophytes...............................................................................................................95
II. Herbarium management.........................................................................................................103
III. Identification.........................................................................................................................107
IV. Techniques of taxonomic revisions......................................................................................119
V. Vegetation analysis (Phytosociological method after
Braun-Blanquet).........................................................................................................................127
VI. Bryophyte checklists of tropical countries...........................................................................131
VII. Principles of preparing checklists........................................................................................139
VIII. Methodology of relevée studies.........................................................................................143
IX. The use of computers in the field.........................................................................................149
X. Photographing in the field......................................................................................................151
XI. A guide to collecting bryophytes in the tropics....................................................................155
Bibliography....................................................................................................................................175
Research needs and priorities...........................................................................................................195

1. INTRODUCTION
Bryophytes belong to the oldest land plants. They and on heavy metal rich soil, in canopies and
existed already in the Palaeozoic 300 mio years deserts, in dark caves and on exposed rocks. This
ago in forms which were hardly different from is rendered possible by various morphological,
the extant species. They remained relatively anatomical and physiological adaptations. Thus
unchanged with relatively low evolution rates bryophytes are not at all „primitive“ plants.
(and are thus often called a „conservative“ plant Bryophytes are a much neglected but nevertheless
group), but could successfully establish an important group of plants:
themselves in an always varying environment 1. They are the second largest group of green
from Devonian swamps to Permian forests, land plants (with ca. 15000 species).
Mesozoic deserts and as epiphytes in Tertiary 2. They play an important role in many
rainforests. They are not eaten by snails or ecosystems such as tundras, bogs and tropical
insects, and are resistant against fungi and rainforests (the extension of bogs in the boreal
bacteria. zone is much larger than that of the tropical rain
Whereas the species numbers of lycopods and forest, although this zone appears relatively small
horsetails have decreased over the past geological in certain map projections). In these ecosystems,
periods, bryophytes seem not to have been bryophytes play an important role in
decreasing but have perhaps even increased their - water storage
diversity by occupying new ecological niches, - nutrient uptake from rain
such as epiphytic existence in forests. - ecological interactions (habitat for animals).
Bryophytes are so called „lower plants“ which, 3. Because of their sensivity to water loss,
however, does not mean that they have primitive bryophytes are good indicators of
morphological structures. The presence of - microclimate
stomata, conducting tissues and a cuticula in - altitudinal zonation of rain forests.
many species (structures which are, interestingly, 4. This makes bryophytes useful also in
no more functional), shows that bryophytes had biodiversity research.
the ability to develop cormophytic structures, but 5. Bryophytes absorb water and nutrients
instead they chose a poicilohydric existence as exclusively or mostly by their surface and do not
an alternative to a cormophytic one. filter water and nutrients through soil and roots
Bryophytes grow in an astonishingly broad as flowering plants. They are therefore good
variety of substrates and habitats. They occur in indicators of
the snow vegetation with a 9 months‘ snow cover, - pollution of air and water
form masses in the tundra and in the boreal - heavy metal contamination
forests, cover tree trunks in tropical rain forests, - radioactivity.
grow 50 m deep in lakes and at 5000 m altitude 6. Due to direct reaction to climatic factors and
on mountain peaks, in extremely acid peat bogs, short life cycles and spore dispersal, bryophytes

10 Frahm
are also good and very fast indicators of climatic in the knowledge of geographic ranges, which
changes. reflect evolution and vegetation history.
- Study of phytochemical compounds of
Bryophytes of tropical rain forests have several bryophytes will give knowledge of chemical
advantages as compared with flowering plants: compounds with anti-microbial and anti-tumor
- They are much less numerous than flowering effects. Natural compounds will replace artificial
plants. There are about 250.000 species of substances e.g. as biocides.
flowering plants in the world as compared with - Knowledge of tropical species will enhance our
10-15.000 species of bryophytes. There are no knowledge of the systematics, since the majority
more than 5000 species of bryophytes in the of species is occurs in the tropics.
neotropics, presumably even less. In Colombia - Study of the anatomy of tropical species has
there are about 1000 species of mosses as brought extremely interesting results, e.g. the
compared with 45-55.000 species of flowering presence of vessel-like canals in the stems of
plants. Thus a wide knowledge of a regional Hypopterygium.
mossflora can be obtained relatively soon, but - Knowledge of the ecology of bryophytes,
never for flowering plants. especially of the epiphytic rainforest bryophytes,
- Bryophytes have much larger geographic ranges will enhance our knowledge of the importance
than flowering plants. A major part of the species of bryophytes in the ecosystem.
in different parts of the neotropics are identical. Bryological activities in the industrialized
The neotropic element accounts for about 40% countries are going rapidly down due to a
of the species. Many Andine species range from shortage of positions. The activities at
southern Mexico to northern Argentina. Thus universities are today focused on molecular
bryologists can work in different parts of the biology and, consequently, classical bryological
tropics with the same results. Nevertheless there positions are no more held and taxonomic or
are also endemic bryophyte species indicating phytogeographic studies can not even survive in
local phytogeographical characteristics. Also botanical museums. Therefore bryology must be
comparisons of species diversity of distant areas established in tropical countries „in situ“. The par-
is possible by the small ranges. tial aim of this volume is therefore to develop in
- The density of bryophytes is much higher than the students an interest in bryology, to stimulate
in flowering plants. Studying a hectare results in studies and to facilitate bryology in tropical
representative data for the whole altitudinal zone. countries.
- Sterile plants can usually be identified whereas
flowering plants need mostly to be in flower for
identification. General literature
Nevertheless, the knowledge of tropical
bryophytes is still very poor. There are only few Campbell, D.G. 1989. The Importance of
bryologists in the tropics. In Latin America, at Floristic Inventory in the Tropics. Pp.
present only about 25 in Mexico, Costa Rica, Pa- 6-30 in: Campbell, D,G. & Hamilton,
nama, Puerto Rico, Colombia, Venezuela, Brazil H.D. Floristic Inventory of Tropical
and Argentina. The situation in tropical Africa is Countries. New York Boztanical Gar-
much worth with few bryologists only in Nige- den.
ria, Kenya, Uganda and Malawi. Gradstein, S.R. 1992. The vanishing tropical
However, studying tropical bryophytes is very rain forest as an environment for
important because: bryophytes and lichens. Pp. 234-258 in:
- Floristic surveys will probably not result in the J.W. Bates & A.M. Farmer, Bryophytes
detection of many undescribed species as in other and Lichens in a Changing Environ-
groups of organisms (e.g. insects), but will result ment. Oxford.
in the knowledge of biodiversity (e.g. „hot spots“, Gradstein, S.R. & Pócs, T. 1989. Bryophytes.
centers of biodiversity); will give valuable Pp. 311-325 in: Lieth, H. & Werger,
information for nature conservation; will result M.J.A. (eds.), Tropical Rain Forest

Ecosystems. Ecosystems of the World Textbooks

Vol 14B. Amsterdam.
Pócs, T. 1982. Tropical forest bryophytes. Pp. Delgadillo M., C., Cárdenas S., A. (1990).
59-104 in: A.J.E. Smith (ed.), Bryophyte Manual de Briófitas. 134 pp., Mexico-
Ecology. London. City.
Prance, G.T. 1989. American tropical forests. This highly recommended book for Latin
Pp. 99-132 in: Lieth, H. & Werger, American students provides a basic knowledge
M.J.A. (eds.), Tropical Rain Forest of bryology (life cycle, morphology, physiology,
Ecosystems. Ecosystems of the World ecology, phytogeography, cytology, evolution
Vol. 14B. Amsterdam. and commercial use. It includes in addition some
Richards, P.W. 1984.The ecology of tropical proposals for practicals and a key and description
forest bryophytes. Pp. 1233-1270 in: of common bryophyte genera in Mexico.
R.M. Schuster (ed.), New Manual of Frahm, J.-P. 2001. Biologie der Moose.
Bryology, Part 2. Nichinan. Heidelberg (Spektrum). (in German,
Spanish translation in preparation).
Guide to bryofloras in the world: Shaw, A.J., Goffinet, B. 2000. Bryophyte
Biology. Cambridge (Cambridge
Greene, S.W. & Harrington, A.J. 1989. The University Press).
Conspectus of Bryological Taxonomic Schofield, W.B. 1985. Introduction to Bryology.
Literature. Part 2. Guide to national and New York.
regional literature. Bryophytorum
Bibliotheca 37.
Journal
TROPICAL BRYOLOGY
ed. J.-P. Frahm, University of Bonn. Published
irregularly at a price of $10 per 100 pages. CD
version half price Subscribers from tropical
countries get a reduction of 50%.
Societies
Sociedad Latinoamericana de Briología.

Publishes „Briolatina“ irregularly. Contact
address: Naris Salazar A., Smithsonian Tropical
Research Institute, Apartado 2072, Balboa, Pa-
nama.
British Bryological Society, Tropical Bryology
Working Group with activities in Malawi and
Uganda, see www.rbge.org.uk/bbs/bbs.htm.
Funding
Sastre, I., Tan, B.C. 1995. Directory of grants

and scholarships for bryologists. Bryol.
Times 83/84: 1-4.

12 Frahm

2. DIVERSITY OF BRYOPHYTE SPECIES IN THE TROPICS
2.1. Global diversity
The actual number of bryophyte species is given precipitation and humidity, but again in increase
in classical botanical textbooks as 25.000 (15.00 in the mountains. The same counts for the tropics.
mosses, 10.000 hepatics), a number probably Therefore the ratio hepatics : mosses can be
going back on a count of the species included in successfully and easily used to characterize dif-
the volumes of Engler-Prantl´s „Natürliche ferent climatic conditions of relevées in different
Pflanzenfamilien“ at the beginning of the 20th altitudes or distance to the ocean. The bryoflora
century. About 57.000 species of mosses have in the tropics may consists of 40% hepatics in
been described in total (Crosby et al. 1992).Their some regions but this percentage may increase
names are listed in the „Index Muscorum (Wijk to 90% in hyperhumid regions such as the Chocó
et al. 1959-69), which includes all names region in Colombia, which is one of the places in
published until 1963. The additions from 1963- the world with the highest precipitation.
1989 are included in the „Index of Mosses“
(Crosby et al. 1992), those from 1990-92 in a Two thirds of all bryophyte species or about 8000
supplement (Crosby & Magill 1994). Walther species occur in the tropics. This is the reason
(1983) has counted all legitimate moss species that tropical bryology needs to be focussed. In
in the „Index Muscorum“ and Supplements to contrast, there are about 1600 species of
1977 and came up with 16.455 species. On the bryophytes in Europe, which are covered in 32
other hand, critical revisions of genera result in floras (1:50). In contrast, the 8000 bryophyte
a considerable „loss“ of species. According to species in the tropics are covered by 16 floras (1:
Touw (1974), the percentage of recognized taxa 6000). With regard to the humid tropics, the
in revisions and monographs varies between 20 percentage of liverworts and hornworts is much
and 40% of the original number. Of the taxa higher than in other regions of the world.
described since 1930, 73% have been reduced to
synonymy. Examples of reduction rates in Crosby, M.R., Magill, R.E., Bauer, C.R. 1992.
taxonomic revisions are indicated in the Index of Mosses. Monographs in
taxonomy-chapter. Crosby et al. (1992) estimate Systematic Botany from the Missouri
that there are 10.000 species of mosses. There Botanical Garden vol. 42.
are no actual pproximations of numbers of Crosby, M.R., Magill, R.E 1994. Index of
hepatics in the world, which can be estimated with Mosses 1990-1992. Monographs in
about 4000. It has to be kept in mind that mosses Systematic Botany from the Missouri
and hepatics (with hornworts) have very diffe- Botanical Garden vol. 50.
rent diversities. Mosses are much more drought Gradstein, S.R. 1995. Bryophyte diversity of the
tolerant that hepatics (with a few exceptions of tropical rainforest. Archs. Sci. Geneve
Marchantiidae). Therefore hepatics can be used 48: 91-96.
as indicators for humid climates. Even in North Greene, S.W., Harrington, A.J. 1988. The
America and Europe, we have a decrease of Conspectus of Bryological Taxonomic
hepatics from the oceanic west coasts to the Literature. Part 1. Index to monographhs
interior of the continents with decreasing and regional reviews.. Bryphytorum
Bibliotheca 35.

14 Frahm
Touw, A. 1974. Notes on taxonomic and floristic - isolation (by islands, mountains, fragmentation
research on exotic mosses. J. Hattori of vegetation belts).
Bot. Lab. 36: 123-128.
Walther, K. 1983. Bryophytina, Laubmoose in
A. Engler (ed.), Syllabus der Pflanzen- 2.2.1 Neotropics
familien 13. Aufl.. Berlin, 108 pp.
Wijk, R. van der, Margadant, W.D., Flor- The numbers of bryophyte species recorded for
schütz, P. 1959-69. Index Muscorum. a country give an impression of the floristic
5 vols. Utrecht. richness, the comparison with other countries also
an impression of the state of exploration and the
need for floristic activities.
2.2. Regional diversity As an example, the species numbers for various
neotropical countries are:
Regional diversity depends on
- the climate of the area Mosses:
- the geological age and history of the area, Mexico: 946 (Sharp et al. 1994)
especial plate tectonics Colombia: 937 (Churchill & Linares 1995)
- the availability of species El Salvador: 233 (Menzel 1991)
- high diversity of habitats. Guyanas: 238 (Florschütz-de Waard 1990)
Size is not an important factor, since small regions Bolivia: 1222 (Hermann 1976)
can have much more species than large regions. Ecuador: 783 (Steere 1948)
Regions with high diversity (so called „hot Costa Rica: 542 (Bowers 1974)
spots“) are characteristed by a high diversity of Venezuela: 626 (Pursell 1973), 1010 (Moreno
habitats, a long geological history without 1992)
dramatic changes (arid periods, glaciation Brazil: 3690 (Yano 1981)
periods), a favourate climate and a rich source of Hepatics:
native species or immigrants and factors Guyanas: 375 (Gradstein & Hekking 1989)
stimulating the evolution (raise of mountains, Bolivia: 415 (Gradstein et al. 2003)
isolation on islands).
Diversity is expressed by species numbers per The present state of knowledge of the distribution
area. On a global scale, a worldwide used unit is of mosses in the neotropics is summarized in the
10.000 km2, however, diversity is also counted database LATMOSS (Delgadillo et al. 1995).
on smaller units down to a hectare or a tree. This catalogue indicated 4103 species and
Diversity in a place is composed varieties in the neotropics. Due to taxonomic
- from the original stock of species revisions, the number decreased to 3869 species
- by immigrants (e.g. through the chain of the and varieties (Delgadillo 2000), an effect, which
Andes, by mountain hopping in tropical Africa will continue for the next time since many genera
or island hopping in tropical SE-Asia have not yet been revised. The exact number of
- by secondary evolution. hepatics is not known. 44% of the taxa of mosses
Thus the diversity depends also on evolution rate. are endemic. The highest rates of endemism are
Evolution is influenced by found in:
- Possibility to occupy different ecological niches
(free habitats, habitat diversity) Brazil 46%
- genetic variability and different speed of Bolivia 26%
evolution in different systematic groups Paraguay 26%
- climatological changes (glaciations, changes Ecuador 15%
during continental drift) Venezuela 13%
- geological changes (uplift, submersion, Peru 10%
volcanism, break up or collision of continents) Colombia 9%
Mexico 9%

Countries of low endemism rates are e.g. Guate- of bryophytes is characteristic for cool-tmperate
mala, Belize, Honduras, Jamaica, the Guianas etc. climates and that the colonization of the tropics
The highest species numbers per area are found in the Tertiary started from the cool temperate
in Costa Rica (8.6 species per 10.000 km2) Gondwana flora (and to a smaller account from
This database allows comparison of the the northern hemisphere). For the Neotropics, the
mossfloras of certain countries and checking the Andes provided a suitable pathway for
similarities. Of the 946 species of mosses in immigration of taxa which subsequent speciation.
Mexico, and the 937 species in Colombia, 371 Therefore the lowland taxa are relatively young
are identical. 241 of those 371 occur also in the adaptations to the physiological harsh conditions
West Indies. Thus it can be concluded that one of tropical lowlands. The Lejeuneaceae adapted
third of the mossfloras of Mexico and Colombia to this kind of new habitat with explosive
is identical and that most of the species migrated radiation.
from S to N.
The similarities of the bryofloras of the Bowers, F.D. 1974. The mosses reported from
neotropical counries were calculated by Costa Rica. Bryologist 77: 150-171.
Delgadillo (2000) by means of a cluster analysis, Churchill, S. 1991. The floristic composition
which nicely indicates the floristic subunits and elevational distribution of Colom-
(Central America, Carribbean, Guianas, andine bian mosses. Bryologist 94: 157-167.
countries, Brazil.) and the level, on which the Churchill, S.P. & Linares, E.L. 1995. Prodro-
floras are related (at the present state of mus Bryologiae Novo-Granatensis. 2
knowledge). Interestingly, Belize floristically vols. Bogotá.
belongs to the Carribbean, Honduras and Nica- Churchill, S.P., Griffin, III, D., Lewis, M.,
ragua are separated from the rest of Central 1995. Moss diversity of the tropical
American countries and Paraguay has a separate Andes. Pp. 335-346 in: Churchill, S.P.,
position between Chile/Argentina/Uruguay on Balslev, E., Forero, E., Luteyn, J.L.
the one and Brazil on the other hand. (eds.) Biodiversity and Conservation of
The diversity if species varies much within a Neotropical Montane Forests. New
country in different altitudes. (see chapter on York Botanical Garden.
altitudinal zonation). Delgadillo M., C. 2000. Distribución geográfi-
Diversity is also different in hepatics and mosses. ca y diversidad de los musgos neotropi-
According to Gradstein (1995), the tropical cales. Bol. Soc. Bot. México 65: 63-70.
Andes have a hepatic flora of estimated 800 - Florschütz-de Waard, J. 1990. A catalogue of
900 species. Most species are, however, in the the bryophytes of the Guianas. II. Mu-
upper montane belt above 2000 m, and the lower sci. Trop. Bryol. 3: 89-104.
montane and subalpine forests have less species, Gradstein, S. R. 1982. Bryological exploration
although generally the species number of of tropical America: Summarizing re-
bryophytes raises with the elevation. There are marks. , In Geissler, P. and Greene, S.
also comparably many hepatic species in the W. (eds.). Bryophyte Taxonomy, Nova
lowland forests. The different diversity is Hedwigia 71: 537-538
determined by climatic, edaphic, physiological Gradstein, S.R. 1991. A view at the liverwort
and phytogeographical factors. flora of tropical America. , Bulletin of
Diversity differs not only within hepatics and the British Bryological Society (57): 13-
mosses but also within different genera and 15
families. Therefore spectra of families or genera Gradstein, S.R. 1995. Diversity of hepaticae and
are useful to determine differences between dif- Anthocerotae in Montane Forests of the
ferent regions or altitudes. In the Andes, monta- Tropical Andes. Pp. 321-334 in: Chur-
ne forests have 2.5 much more hepatic families chill, S.P., Balslev, E., Forero, E., Lu-
than lowland forests (Gradstein 1995). About teyn, J.L. (eds.) Biodiversity and Con-
70% of all the species in lowland forests belong servation of Neotropical Montane Fo-
to the Lejeuneaceae. Reason is, that the majority rests. New York Botanical Garden.

16 Frahm
Gradstein, S.R. & Hekking, W.H.A. 1989. A Sastre-De Jesus, I., and E. Santiago-Valentín.
catalogue of the bryophytes of the Guia- 1996. Bryology in Puerto Rico.
nas. I. Hepaticae and Anthocerotae. J. Knowledge prior to and after the
Hattori Bot. Lab. 66: 197-230. scientific survey of Porto Rico and the
Gradstein, S.R., van Reenen, G.B.A. & Grif- Virgin Islands. Annals of the New York
fin, D.G. 1989. Species richness and ori- Academy of Science 115:111-122.
gin of the bryophyte flora of the Colom- Sharp, A.J., Crum, H.A., Eckel, P.M. (eds.)
bian Andes. Acta Bot. Neerl. 38: 439- 1994. The moss flora of Mexico. 2 vols.
448. New York Botanical Garden.
Gradstein, S.R., van Reenen, G.B.A. & Grif- Steere, W.C. 1948. Contribution to the bryogeo-
fin, D.G. 1989. Origen de la flora de graphy of Ecuador I. A review of the
briofitas en el transecto Parque los Ne- species of Musci previously reported.
vados (Cordillera Central, Colombia). Bryologist 51: 65-167.
Stud. Trop. Andean Ecosyst. 4: 377- Yano, O. 1981. A checklist of Brazilian mosses.
384. J. Hattori Bot. Lab. 50: 279-456.
Griffin, D,G. & Gradstein, S.R. 1982. Bryolo-
gical exploration of the Andes: current 2.2.2 Tropical Africa
status. Beih. Nova Hedwigia 71: 513- (contributed by T. Pócs)
518.
Gradstein, S.R., Meneses, R.I., Arbe, B.A. Tropical Africa varies much in the level of the
2003. Catalogie of the Hepaticae and bryological exploration of its countries. The
Anthocerotae of Bolivia. J. Hattori Bot. known number of species is summarized from
Lab. 93: 1-67. the checklists of Hepaticae (Grolle 1995,
Hermann, F.J. 1976. Recopilacion de los mus- Wigginton & Grolle 1996) and for Musci (O´Shea
gos de Bolivia. Bryologist 79: 123-171. 1995). These lists reflect more the level of
Menzel, M 1992. Preliminary checklist of the exploration of the different countries than their
mosses of Peru. J. Hattori Bot. Lab. 71: real species richness. Especially, there is a great
175-254. discrepancy between the given species number
Moreno, E. J. 1992. Revisión histórica de la and the size and habitat diversity of the concerned
briología en Venezuela. Tropical country (see maps and graphs in Pócs 1982,
Bryology 6:139-146. O´Shea 1997). Therefore, the country or island
Moreno, E.J. 1992. Aproximación al is marked in bold which is underexplored based
conocimiento de las briofitas de on this discrepancy. The number of species is
Venezuela. Tropical Bryology 6: 147- marked in bold, where the number of species is
156. relatively high due to a reasonable high level of
Motito, A. M., E. Potrony, and M. D. Reyes. exploration.
1992. Estado actual y perspectivas Hep. Musci
futuras del estudio de los musgos Angola 58 125
cubanos. Tropical Bryology 6:157-160. Annobon 34 18
Pursell, R.A. 1973. Un censo de los musgos de Ascensión 21 14
Venezuela. Bryologist 76: 473-500. Benin 3 8
Reyes M., D. 1992. Acerca de la flora briológica Bioko 58 122
de Cuba. Tropical Bryology 6: 203 ff. Botswana 17 22
Salazar Allen, N., S. R. Gradstein, and S.P. Burkina Faso 0 13
Churchill. 1996. Bryophytes as non- Burundi 88 69
woody biodiversity indicators: a guide Cameroun 223 361
to the bryophytes of tropical America. Cabinda 0 2
A report. Anales del Instituto de Central Afr. Rep. 52 287
Biologia de la Universidad Nacional Chad 13 10
Autonoma de Mexico 67:59-65. Comores 122 191

Congo (Brazzav.) 49 76 unusual - that one of them was properly

Congo (Zaire) 291 579 investigated and the other component neglected.
Cap Verde 30 113 For example the Hepaticae of Ghana and Sierra
Côte d´Ivoire 77 178 Leone were much better explored by E.W. Jones
Equatorial Guinea 0 2 and A.J. harrington than the Musci by others.
Eritrea 15 75 Pócs 81992) has furher suggestions concerning
Ethiopia 119 285 the exploration of Hepaticae of the different
Gabon 51 250 countries.
Gambia 1 1 South Africa is exceptional, where modern local
Ghana 137 63 floras elaborate the great part of Musci (Magill
Guinea-Bissau 5 1 1981, 1987, Magill & van Rooy 1998) and the
Guinea 53 196 thallose liverworts (many works of Perold) and
Kenya 208 464 not a too old flora (Arnell 1963) deals with all
Liberia 3 55 hepaticae.
Lesotho 22 163 For tropical Africa, quite a number of revisions
Madagascar 354 751 are needed. The level of exploration is quite
Mali 2 15 uneven comparing Hepaticae (thanks to the
Malawi 122 223 activities of E.W. Jones and C. Vanden Berghen)
Mozambique 53 77 and Musci. Pócs & O´Shea compiled a list of
Namibia 31 56 basic taxonomic literature to identify tropical
Niger 0 6 African bryophytes, which contain all revisions
Nigeria 133 141 and monographs completed until that time,
Principe 24 14 similarly to Greene & Harrington (1988), but
Réunion 227 376 trying to be more specific to Africa.
Rodrigues 16 35 According to our present knowledge the number
Rwanda 223 293 of Hepaticae in tropical and South Africa is 894
South Africa 293 species. 713 occur on the continent and 436 on
Cape 389 the Indian Ocean Islands, and 255 are shared
Natal 340 between the two. For comparison: about 1250
Orange F.St. 137 species of hepatics occur in the neotropics. We
Transvaal 299 do not even know the approximate numbers for
Saô Tomé 84 74 tropical Asia.
Senegal 4 20 As can be seen from the above list, there is much
Seychelles 75 100 more to do for the taxonomy of Musci. At present,
Sierra Leone 144 99 there are 3048 taxa recorded for the Subsaharan
Socotra 12 25 Africa (O´Shea 1997), which will be reduced to
Somalia 4 19 estimated 1300 by revisions. O´Shea (1997)
St. Helena 21 30 deals in detail with the perspectives of African
Sudan 10 31 moss research.
Swaziland 21 89
Tanzania 389 780
Togo 33 83 Arnell, S. 1963. Hepaticae of South Africa.
Uganda 153 376 Stockholm.
Zambia 59 141 Greene, S.W., Harrington, A.J. 1989. The
Zimbabwe 135 265 Conspectus of Bryological Taxonomic
Literature. Part 1. Index to monographs
From the above list it is easy to see where inten- and regional reviews. Bryophytorum
sive research is still needed (the bold faced Bibliotheca 37: 1-321.
countries). The above data also suggest - where Grolle, R. 1995. The Hepaticae and
the ration between Hepaticae and Musci is Anthocerotae of the east African islands.

18 Frahm
An annotated catalogue. Bryophytorum 2.2.3 Tropical SE-Asia

Bibliotheca 48: 1-178.
Magill, R.E. 1981. Bryophyta. Part 1. Mosses. The bryoflora od SE-Asia is very diverse. The
Fasc. I. Sphagnaceae - Grimmiaceae. In Indian subcontinent harbours a great deal of
O.A. Leistner (ed.): Flora of Southern laurasian taxa and therefore has with almost 1600
Africa. Pretoria. species the most taxa (cf. tab. 2.1). Also in
Magill, R.E. 1987. Bryophyta. Part 1. Mosses. Indochina, part of the 1000 species are possibly
Fasc. II. Gigaspermaceae - attributed from the holarctic.The other parts such
Bartramiaceae. In O.A. Leistner (ed.): as Borneo or the Philippines have around 600
Flora of Southern Africa. Pretoria. species, which is less than comparable regions
Magill, R.E., van Rooy, J. 1998. Bryophyta. Part in the neotropics (e.g. in the Andes). The lowest
1. Mosses. Fasc. III. Erpodiaceae - species numbers are found in the densely
Hookeriaceae. In O.A. Leistner (ed.): populated lowland countries such as Bangladesh.
Flora of Southern Africa. Pretoria.
O´Shea, B. 1995. Checklist of the mosses of Sub-
saharan Africa. Tropical Bryology 10: Abeywickrama, B.A. 1960. The genera of
91-198. mosses of Ceylon. Ceylon Journal of
O´Shea, B.J. 1997a. The mosses of su-Saharan Science 3: 41-122.
Africa 1. A review of taxonomic Abeywickrama, B.A. & Jansen, A.B. 1978. A
progress. J. Bryol. 19: 509-513. check list of the mosses of Sri lanka.
O´Shea, B.J. 1997b. The mosses of su-Saharan UNESCO: Man and the Bosphere Nati-
Africa 2. Endemism and biodiversity. onal Committee for Sri lanka 2: 1-25.
Tropical Bryology 13: 509-513. Banu-Fattah, K. 2001. A comprehensive
Pócs, T. 1982. Hepaticological exploration in checklist of the bryophytes of
Subsharan Africa. Beih. Nova Hedwigia Bagladesh. Bangladesh J. Plant
71: 495-500. Taxonomy 8: 7-18.
Pócs, T. 1990. The exploration of the East Brühl, P. 1931. A census of Indian mosses.
African bryoflora. Tropical Bryology 2: Records of the Botanical Survey of India
177-192. 13: 1-135.
Pócs, T. & O´Shea, B.J. 1991. Quick reference Grolle, R., Piippo, S. 1984. An annotated
list of basic literature to identify tropical catalogue of western Melanesian
African bryophytes. Tropical Bryology bryophytes I. Hepaticae and
4: 69-84. Anthocerotae. Acta Bot. Fennica 125:
Wigginton, M.J., Grolle, R. 1996. Catalogue of 1-86.
the Hepaticae and Anthocerotae of Sub- Higuchi, M., Nishimura, N. 2003. Mosses of
Saharan Africa. Bryophytorum Bibl. 50: Pakistan. J. Hattori Bot. Lab. 93: 259-
1-267. 272.
Hoe, W.J. 1974. Annotated checklist of
The following files are available from the Hawaiian mosses. Lyonia 1: 1-45.
internet: http://britishbryologicalsociety.org.uk. Iwatsuki, Z. & Tn, B.C. 1979. Checklist of
Philippine mosses. Kalikasan 8: 179-
Checklist of Malawi bryophytes 210.
Koponen, T., Norris, D. H. & Piippo, S. 1992:
Checklist of mosses of sub-Saharan Africa Bryophyte flora of Western
Melanesia.A status report. -
African mosses - diversity & endemism Bryobrothera 1: 157-160.
Long, D.G. 1994. Mosses of Bhutan II. A
Checklist of liverworts of sub-Saharan Africa checklist of the mosses of Bhutan. J.
Bryol. 18: 339-364.
Meijer, W. 1953. The study of hepatics in the

Tab. 2.1: Number of mosses in SE-Asia
No. of taxa No. of endemics % endemics Ref.
Bangladesh 183 3 1.6 O´Shea 2003a

Pakistan 339 43 13 Higuchi & Nishimura
2003
Sri Lanka 561 11 O´Shea 2003b
India 1594 288 18 O´Shea 2003b
Indochina 1008 483 33 Tan & Iwatsuki 1996
Philippines 625 Tan & Iwatsuki 1991
Lesser Sunda Isl. 367 Touw 1992
Borneo 607 Touw 1978
Peninsular Malaya 475 20 4 Mohamed & Tan 1988
Malaysian Tropics. Bryologist 56(2): of Bangladesh, with a revised checklist.

95-98 J. Hattori Bot. Lab. 93: 259-272.
Menzel, M. 1988. An annotated catalogue of the O´Shea, B.J. 2003b. Bryogeographical
hepaticae and Anthocerotae of Borneo. relationships of the mosses of Sri Lanka.
J. Hattori Bot. Lab. 65: 145-206. J. Hattori Bot. Lab. 93: 293-304.
Miller, H. A., Whittier, H. O., Del Rosario, R. Tan, B.C. & Iwatsuki, Z. 1991. A new annotated
M., Smith, D. R. Bryological Philippine moss checklist. Harvard
bibliography of the tropical Pacific Papers in Botany 3: 1-64.
islands, especially Polynesia and Tan, B. C. & Iwatsuki, Z. 1996. A checklist of
Micronesia. Pacific Sci. Inform. Center, of Indochinese mosses. J. Hattori Bot.
Bernice P. Bishop Mus., Honolulu. 51 Lab. 74: 325-405.
Pp Tan, B. C. & Iwatsuki, Z. 1996: Hot spots of
Miller, H.A. & H.O. Whittier. 1990. Bryophyte mosses in East Asia. - Anales del
floras of tropical Pacific Islands. Instituto de Biologia de la Universidad
Tropical Bryology 2: 167-175. Nacional Autonoma de Mexico 67: 159-
Mohamed, H., Tan, B.C. 1988. A checklist of 167.
mosses of Peninsular Malaya and Touw, A. 1978. The mosses reported from
Singapore. Bryologist 91: 24-44. Borneo. J. Hattori Bot. Lab. 44: 147-
Norris, D.H. 1990: Bryophytes in perennially 176.
moist forests of Papua New Guinea: Touw, A. 1992. A survey of the mosses of the
Ecological orientation and predictions Lesser Sunda Islands (Nusa Tenggara),
of disturbance effects. - Bot. J. Linn. Indonesia. J. Hattori Bot. Lab. 71: 289-
Soc. 104: 281-291. 366.
Piippo, S., Koponen, T., Norris, D.H. 1987. Whittier, H.O. 1976. Mosses of the Society
Endemism in the bryophyte flora in New Isslands. 410 pp. Gainesville.
Guinea. Symposia Biol. Hungarica 35:
361-372.
Piippo, S. & Koponen, T. 1997: On the 2.3 Local diversity
phytogeographic biodiversity of We-
stern Melanesian mosses. J. Hattori Another important aspect is a determination of
Bot. Lab. 82: 191-201. diversity on a small geographical scale. This can
O´Shea, B.J. 2003a. An overview of the mosses be done on different levels:

20 Frahm
1. in different types of forests (primary - Allen, N. S., Gradstein, S. R. & Churchill, S.

secondary forest, young secondary - mature P. , 1996. Bryophytes as non-woody
secondary forest, lowland forest, montane forest), biodiversity indicators: a guide to the
interior of forest, edge of forest, periodically bryophytes of Tropical America. A
flooded alluvial forest, episodically flooded report , Anales Inst. Biol. Univ. Nac.
forest. Auton. Mexico, Ser. Bot. 67(1): 59-65.
2. on different types of habitats (trunk epiphytes, Churchill, S. P. 1996. Andean moss diversity
canopy epiphytes, soil, disturbed soil, rocks, and conservation: state of knowledge
leaves. and prequisites for the future. Anales
3. systematic diversity: percentage of families in del Instituto de Biologia de la
different regions, vegetation types or habitats. Universidad Nacional Autonoma de
Percentage of endemism. Mexico 67: 69-176.
4. succession studies in different old regrowths. Churchill, S. P., D. Griffin III, and M. Lewis.
The results of such studies give answers to 1995. Moss diversity of the tropical
questions such as: Andes. In S.P. Churchill, H. Balslev,
- which are the most species rich vegetation E. Forero & J.L. Luteyn (eds.).
types? Biodiversity and conservation of
- Which vegetation types need therefore the most Neotropical montane forests, pp. 335-
regard with respect to nature conservation? 346. New York Botanical Graden.
- Which species are lost by destruction of Bronx, New York.
vegetation types e.g. primary forests? Delgadillo, C., 1994. Tropical Bryophytes:
- Which species are indicators of primary forests Biology, diversity and conversation. ,
and need therefore to be placed in red lists? The Bryological Times 80: 6.
- Which species can survive in secondary Equihua, C. & S. R. Gradstein, 1995 .
habitats? Bryofloristic comparison between an
- Which species occur only in secondary habitats old field and a rain forest: preliminary
and are indicators of these habitats? results unpaginated tip-in. In C.
- Which species are indicators of disturbed, man- Delgadillo M (ed.), International
made habitats? (Serrano 1996). Bryological Conference: Tropical
- Which species can be used for determination Bryophytes: Biology, Diversity and
of altitudinal belts? Conservation. August 7-12, 1995.
As expressed above, bryophytes are easier tools Mexico City. Scientific Program,
for answering such questions than flowering Abstracts, Field Trips, Tourist Tips. ,
plants because they are less numerous, better Instituto de Biologia, UNAM, Mexico
indicator species because of their physiology of City.
water and nutrient uptake (e.g. for climatic Glime, J. M., P. S. Hudy, and S. Hattori. 1990.
conditions) and easier to be identified. Diversity and altitudinal niche width
The low number of bryophyte species worldwide characteristics for 35 taxa of the Papua
does not result in a low local species diversity. New Guinea Frullania flora with
In a lowland forest in French Guiana, Montfort consideration of sibbling pairs. Tropical
& Ek (1990) found 154 species of bryophytes Bryology 2:103-116.
(66 mosses, 88 hepatics) on 28 trees representing Gradstein, S.R. , 1991. A view at the liverwort
22 species. Four to five trees yielded about 75% flora of tropical America. , Bulletin of
of the total number of bryophyte species. In a the British Bryological Society (57): 13-
drier type of lowland forest, Cornelissen & ter 15
Steege (1989) found 79 bryophyte species (26 Gradstein, S. R. 1995. Bryophyte diversity of
mosses, 53 hepatics). the tropical rainforest. Arch. Sci. 48:91-
Wolf (1993) found between 55 and 140 species 95.
of bryophytes in a set of each four trees in Gradstein, S. R. 1995. Diversity of Hepaticae
elevations between 1500 and 3500 m. and Anthocerotae in montane forests of

the tropical Andes. In Churchill, S. P., Griffin, D. III., Gradstein, S. R., 1982.
H. Balslev, E. Forero, and J. L. Luteyn Bryological exploration of the tropical
(eds.). Biodiversity and conservation Andes: Current status. , In Geissler, P.
of Neotropical montane forests, pp. 321- and Greene, S. W. (eds.). Bryophyte
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Bronx, New York. 518
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neotropical páramos. Monographs of Bryophyte diversity in Belém, Para and
Systematic Botany (Missouri Botanical its potential as pollution indicator for
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Gradstein, S. R., 1982. Bryological exploration Emilio Goeldi, serie Bot. 11:199-225.
of tropical America: Summarizing Müller, U., Frahm, J.-P. 1998. Diversität epi-
remarks. , In Geissler, P. and Greene, S. phytischer Moose eines westandinen
W. (eds.). Bryophyte Taxonomy, Nova Bergregenwaldes in Ecuador. Tropical
Hedwigia 71: 537-538 Bryology 15:.29-44.
Gradstein, S. R., 1995. Diversity of Hepaticae O´Shea, B. , 1995 , Bryophyte biodiversity and
and Anthocerotae in montane forests of endemism in sub-Saharan Africa, pp.
the tropical Andes, pp. 321-334. In S. 38-39. In C. Delgadillo M. (ed.),
P. Churchill, H. Balslev, E. Forero & J. International Bryological Conference:
L. Luteyn (eds.), Biodiversity and Tropical Bryophytes: Biology,
Conservation of Neotropical Forests. , Diversity and Conservation. August 7-
The New York Botanical Garden, 12, 1995. Mexico City. Scientific
Bronx, NY. Program, Abstracts, Field Trips, Tourist
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Allen , 1995 , Bryophytes as non-woody Mexico City.
biodiversity indicators: A guide to the O’Shea, B. J. 1997. The mosses of sub-Saharan
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26. In C. Delgadillo M. (ed.). Tropical Bryology 13:75-86.
International Bryological Conference: Pinheiro De Costa, D. 1999. Epiphytic
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12. 1995. Mexico City. Scientific southeastern Brazil. The Bryologist
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Griffin III. 1989. Species richness and mass of epiphytic bryophytes and li-
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22 Frahm

3. ORIGIN AND AGE OF TROPICAL BRYOPHYTES
It should be stressed that mosses have undergone functional. Thus the sporophyte of mosses is
reduction during their evolution. The oldest principally not unsimilar to sporophytes of ferns
fossils were always costate and probably (although it is not independent). This allows
acrocarpous. Ecostate and pleurocarpous taxa speculations about the origin of mosses and
have apparently developed only later, as an hornworts from primitive tracheophytes but
adaptation to ecological niches in the understorey clearly not from algae.
of forests in the Tertiary. Insofar, tropical Similar cormophytic structures are found in the
bryophytes are predominantly the youngest stems of Polytrichaceae with a highly developed
branch of evolution. Recent results of molecular conducting system. With an internal hadrom and
studies show that some (of not many) are derived an external leptom, it resembles the conducting
from gondwanalandic anchestors. system of fossil Psilophytales. Even the leaves
We know nothing about the phylogenetic origin possess highly organized connducting tissues.
of bryophytes, thus their anchestors remain However, the leaf traces have no connection with
unknown. In the past, bryophytes were derived the central stand. This can also only be explained
from algae and pteridophytes from bryophytes, as reduction. Therefore bryophytes must be
because of „Haeckel´s law“ (the ontogeny repeats regarded as reduced tracheophytes.
the phylogeny). It was argued that bryophytes
share an algal stage (the protonema) in their
ontogeny and pteridophytes an bryophyte stage 3.1 Evidence from fossils
(the prothallium). Accordingly, the organisation
of mosses was and is commonly still regarded as The fossil history of bryophytes is not well known
thallophytic in many textbooks of botany. The because only relatively few fossil bryophytes
oldest known bryophyte fossil is Pallavicinites have been found. During the 19th century, fossil
devonicus from the Devonian of North America. bryophytes were only known from Quaternary
It resembles the extant thalloid liverwort genus and Tertiary and it was said that bryophytes
Pallavicinia (similar to the tropical genus cannot be preserved as fossils since they lack
Symphyogyna). Therefore bryophytes have lignine.
evolved even earlier and we have no fossil In 1959 , Savicz-Ljubitskaja & Abramov knew
evidence for the origin. It can, however, be 33 species of bryophytes from the Palaeozoic and
concluded from the drastic anatomical and Mesozoic, and Jovet-Ast (1967) raised the
morphological differences between liverworts, number to 68. The most recent survey of
hornworts and mosses that these groups may not bryophytes of the Mesozoic and Palaeozoic was
be monophyletic. It can be argued that mosses compiled by Oostendorp (1987).
as well as hornworts have developed from pri- The Devonian bryoflora consisted of thalloid
mitive tracheophytes by reduction, since they liverworts (Metzgeriales) and probably also a
show several cormophytic characters such as hornwort. First mosses were found in
stomata in the sporophyte, a cuticula and also Carboniferous deposits. The richest source of
conducting tissues, which are no more functional fossils from the Palaeozoic is known from the
and probably remnants of tracheophytic Permian of Russia and Antarctica. These mosses
ancestors. The stomata are no more needed but resemble extant taxa of the order Bryales.
originated surely in plants in which they were Permian deposits have also revealed the first leafy

24 Frahm
liverwort and the first moss with a differentiation graminicola, S. incompletus var. incompletus, and
of chlorocysts and hyalocysts, as in extant Thuidium erectum. All these genera and species
Sphagna, although with a different habit. are still existing on the island of Hispaniola (the
Mesozoic fossils are rich in Marchantiales, e.g. origin of Dominican amber). These are all genera
plants similar to Marchantia and Riccia, showing and species which are widely distributed in the
adaptations to dry climates. neotropics. It shows that the main stock of mosses
The fossil bryophytes of the Tertiary from Europe in the neotropics was already present in the
and North America, and also all fossils from the Tertiary and that these species are at least 25-45
Quaternary, are almost all extant species (Gams mio years old. Also the fossil bryophytes in Baltic
1932, Dickson 1973, Miller 1984). and Saxon amber are of interest for tropical
The best fossil from the Tertiary are preserved in bryology, since the climate in Oligocene and
amber. Tertiary amber is known from Europe, Miocene was subtropical in Scandinavia.
the island of Hispaniola (Dominican Republic) Therefore Baltic amber contains many
and Mexico. The European amber was produced subtropical elements. Again, the fossil hepatics
by pine trees in pine-oak forests in present were identified as extinct species of extant genera
Scandinavia in the Eocene. The resin flew over but with phytogeographical relationships to the
epiphytic bryophytes or single bryophytes were subtropics, especially of E-Asia. The fossil
blown in the resin by wind. The forest was mosses consist in part of extinct species, in part
drowned by raising sea level, the resin was of extant species. The moss flora of the amber
washed out and transported to the region of the forest was composed by species of Barbella,
former Eastern Prussia, today Russia, south of Hypnum, Campylopus, Campylopodiella,
the Baltic countries, and is called Baltic amber. Fabronia, Haplocladium and others. The most
From there is was partly transported to Saxony common epiphyte was Hypnodontopsis confertus
in Germany (called Saxon amber) and widely (Frahm 2001) described in the 19. century from
dispersed by glaciation during Quaternary. amber as Muscites confertus but in 1928 from an
Amber is the greatest source of fossil bryophytes extant collection as Hypnodontopsis mexicana
since the plants are perfectly preserved. Even oil from Mexico. The present rarity (the species was
bodies are preserved in hepatics (Grolle & Brau- found additionally only in Uganda) indicates that
ne 1988). The hepatics in amber were studied the formerly common species is getting extinct
by Grolle (1983). He listed 18 species from Baltic at present. Many species are still found in pine
and Saxonian amber and 12 species from oak forests; they have kept their habitat over
Dominican and Mexican amber. millions of years.
The earliest bryophyte fossils from the neotropics In conclusion, tropical genera were already
are found in Dominican and Mexican amber from existing in Tertiary and even many species at least
the Tertiary with an age of 25-45 mio years. The amongst the mosses were present 25-45 mio years
fossil hepatics were studied by Grolle (1983, ago, giving an estimate of the age of species. This
1984a, 1984b, 1985, 1987, 1988, 1990, does not mean that all bryophyte have that age.
1993).Grolle described all fossil hepatics as There are certainly younger species (e.g.
extinct species in extant genera. Therefore mountain endemisms) and probably also older
speciation of hepatics must have happend during (see chapter on molecular evidence).
the last 25-45 mio years. Gradstein (1993)
reported 10 hepatics from Dominican amber, of
which 3 could be recognized as extant species.
The mosses in Dominican amber were studied Dickson, J.H. 1973. Bryophytes of the
by Frahm (1993, 1994, 1996). Altogether 11 Pleistocene. Cambridge.
species of mosses could be identified: Frahm, J.-P. 1993. Mosses in Dominican am-
Adelothecium bogotense, Clastobryum sp., ber. Journal of the Hattori Botanical
Homalia sp., Hypnum sp., Mittenothamnium sp., Laboratory 74: 249-259.
Neckera sp., Octoblepharum cf. pulvinatum, Frahm, J.-P. 1994. Moose - lebende Fossilien.
Pilotrichella sp., Syrrhopodon africanus ssp. Biologie in unserer Zeit 24: 120-124.

Fig. 3.1: Syrrhopodon incompletus in Domini- Fig. 3.2: Pilotrichella sp. in Dominican amber.
can amber.
Fig. 3.3: ranges of moss species found in Dominican amber.

26 Frahm
Frahm, J.-P. 1996. New records of fossil mos- (ed.), Traite de Paléobotanique 2:17-
ses from Dominican amber. Cryptog. 186. Paris.
Bryol. Lichènol. 17: 231-236. Miller, N.D. 1984. Tertiary and Quaternary fos-
Frahm, J.-P. 1996. New records of fossil mos- sils. In: R.M. Schuster, New Manual of
ses from Dominican amber. Cryptoga- Bryology: 1194-1232. Nichinan.
mie, Bryologie Lichénologie 17(3): Oostendorp, C. 1987. The bryophytes of the
231-236. Palaeozoic and Mesozoic. Bryophyt.
Frahm, J.-P. 2001. Hypnodontopsis confertus Biblioth. 34: 1-112, XLIX pls.
comb. nov. from Baltic amber. Tropi- Savicz-Ljubitskaja, L.I. & Abramov, I.I. 1959.
cal Bryology 20: 79-82. The geological annals of Bryophyta.
Frahm, J.-P. 2001. New records of mosses from Revue Bryol. Lichénol. 28:330-342.
Dominican amber. Tropical Bryology
20: 39-42.
Gams, H. 1932. Quaternary distribution. In: F. 3.2 Evidence from plate tectonics
Verdoorn (ed.) Manual of Bryology, pp.
297-322. The Hague. Since there are only few fossils known, especially
Gradstein, S.R. 1993. New fossil hepaticae from the Mesozoic and Palaeozoic, and the
preserved in amber of the Dominican structures preserved in the fossils rarely allow a
republic. Nova Hedwigia 57: 353-374. detailed examination, the age of bryophytes can
Grolle, R. & Braune, W. 1988. Bazzania oleo- be concluded from their geographic ranges.
sa - ein Lebermoos mit erhaltenen Öl-
körpern in Dominikanischem Bernstein. How old are tropical bryophytes ?
Nova Hedwigia Beih. 90: 101-108. We have evidence from Gondwanalandic ranges
Grolle, R. 1983. Leucolejeunea antiqua n.sp., that certain bryophyte species were in existence
das erste Lebermoos aus Dominika- already 135 mio years ago. There are species,
nischem Bernstein. Stuttgarter Beitr. which are not only disjunct between SE-Brazil
Naturk. Ser. B, 96: 1-9. and SE-Africa but the same species occur also in
Grolle, R. 1984a. Bryopteris and Cyclolejeunea Sri Lanka and southern India (fig. 3.6). This type
fossil. J. hattori Bot. Lab. 56: 271-280. of disjunction cannot be explained by long
Grolle, R. 1984b. Cyrtolejeunea suzannensis distance dispersal. Also certain species in
spec. nov., ein weiteres fossiles Leber- common in tropical Africa and in the neotropics,
moos in dominikanischem Bernstein. which lack effective means of long distance
Cryptogamie Bryol. Lichénol. 5: 27-32. dispersal or which are not able for long distance
Grolle, R. 1985. Lejeunea palaeomexicana n.sp. dispersal because of lacking UV and frost
Stuttg. Beitr. Naturk. Ser. B. 108: 1-7. tolerance, are probably of a comparable age. Rain
Grolle, R. 1987. Radula steerei sp.nov., a further forests, as we know them today, originated in the
hepatic in Dominican amber. Mem. beginning of the Tertiary; they are thus less than
N.Y. Bot. Gard. 45: 259-263. 70 Mio years old. Thus we can assume that
Grolle, R. 1988. Bryophyte fossils in amber. tropical lowland species originated about at this
Bryol. Times 47: 4-5. time. The tropical montane bryoflora is derived
Grolle, R. 1990. Leucolejeunea antiqua (ein from invasions from the holarctic and, mainly,
Lebermoos aus Dominikanischem Bern- from the subantarctic. Many species (e.g.
stein) erstmals mit Gynözium, Perianth Lepyrodon tomentosus) which are widely
und Andrözium. Nova hedwigia 90: distributed through New Zealand, Tasmania and
101-108. Chile go up to the Andes. During this migration,
Grolle, R. 1993. Bryopteris bispinosa spec. nov. speciation has taken place as adaptation to new
(Lejeiuneaceae), ein weiteres habitats. Such genera (e.g. Chorisodontium) are
Lebermoos aus dominikanischem subantarctic in origin but have secondary centers
Bernstein. J. Hattori Bot.Lab. 74: 71-76. of evolution in the tropical mountains.
Jovet-Ast, S. 1967. Bryophyta. In: E. Boureau Campylopus is represented in the subantarctic

Fig. 3.4: Distribution of a. Campylopus introflexus, b. C. pilifer, c. C. aureus and C. julaceus (hat-
ched). C. introflexus can be regarded as ancestor, C. pilifer and C. aureus migrated into the tropics,
C. pilifer presumably at a time when the Gondwana continent was intact. C. julaceus lived along the
S-coast of the Gondwana continent and its range was split.
Fig. 3.5: Distribution of Campylopus pyriformis (a), C. fragilis (b), C. zollingerianus (c) and C
crispifolius (d). C. pyriformis can be regarded as anchestral species, from which the others develo-
ped.

28 Frahm
Fig. 3.6: Range of Campylopus controversus as example of a gondwanalandic range. The disjunct
occurrence in eastern south America, SE-Africa, the East African Islands and Sri Lanka is explained
as former continous range.
with 14 species, but has 40 species in the northern into SE-Asia from India and have partly
Andes, which may be derived from subantarctic undergone explosive speciation. Another
ancestors. Thus the youngest species in evolution pathway was detected very recently (Vitt 1990):
are those from young mountain systems with an SE-Asia was also colonized by bryophytes from
age of 10 (New Guinea) or only 3 (the Andes) Australia.
million years. Similar young species are found as endemics on
Most of the bryophyte species are found on young volcanic islands, although many of them
tropical mountains: species numbers as well as have turned out, or will turn out, in world wide
phytomass increases with elevation. This is due revisions to be more widespread., e.g. reductions
to the physiology, which resembles that of of endemic species of Campylopus in Hawaii
temperate bryophytes. (Frahm 1991) or in New Caledonia (Frahm
Pathways of migration from the subantarctic to 1990).
the tropics were: Bryophytes have a special role as indicators of
- the Andes, which provide a continuous continental drift since they are much older than
migration route. This is the reason why the flowering plants. The disjunctions of flowering
neotropics has the highest species numbers and plants concern genera, those of bryophytes
diversity. species!
- the mountains of tropical Africa, which could Frahm, J.-P. 1990. A short survey of the
only be reached by “mountain hopping”. Campylopus flora of New Caledonia.
- “Noahs arc”; species which lived on the Indian Cryptog. Bryol. Lichénol. 11: 369-375
continent, when it was part of the Gondwana Frahm, J.-P. 1991. A survey of the Campylopus
continent, and drifted on the Indian plate flora of Hawaii. Bryologist 94: 60-66.
northwards. There are disjunctions between Vitt, D.H. 1990. Desmotheca (Orthotrichaceae):
Madagascar and southern India /Sri Lanka, which Gondwanan fragmentation and the
show that at least part of the species survived the origin of a Southeast Asian genus.Trop.
long trek. However, some of the species migrated Bryol. 3: 79-88.

4. MORPHOLOGICAL ADAPTATIONS
The morphology of plants is to be understood as life form is the composition of individuals (e.g.
a reaction upon environmental factors and in tufts or mats). Mägdefrau‘s life form system,
therefore it can be used to characterize different however, includes also types of individuals (e.g.
ecological conditions. Analysis of the tails, dendroids) together with types of colonies
morphology is especially helpful in comparisons (mats, cushion etc.). Mägdefrau also
of different habitats or in studies of different distinguished annuals, short turfs, tall turfs,
relevés along a gradient, e.g. an altitudinal cushions, mats, wefts, pendants, tails, fans, and
transect. Even if the species are not exactly dendroids. It must be noted that annual is today
known, an analysis of morphological adaptations not regarded as a life form but a life strategy
provides useful insights in the altitudinal zonation (annual vs. perennial).
of rain forests. The adaptations concern In practice, it is hardly possible to see any exact
morphology and anatomy. distinction between short turf and tall turf (which
are now combined) and the distinction between
fans and tails seems to be superfluous. Therefore
4.1 Life and growth forms the system is modified by almost every author.
The system used in various studies during the
Giesenhagen (1910) was the first to define BRYOTROP projects is the following (fig. 4.1):
„Moostypen“ in rain forests in SE-Asia, which
were types of growth forms. crusts
Both the morphological features of individual wefts
bryophyte plants as well as the appearence of mats
aggregated individuals have been used, for which fans
the terms life form and growth form have been cushions
used. However, there is even still much confusion turfs
in terminology. Meusel (1935) was the first to pendants
introduce a system of growth forms of mosses dendroids
based on branching patterns. He distinguished
between orthotropous and plagiotropous mosses It is clear that life forms are adaptations to special
with subforms. Gimingham and Birse (1957) ecological niches and reflect habitats. They are
elaborated a system of growth forms for especially related to moisture conditions
temperate mosses and Birse (1957) tested it (Tobiessen et al. 1977, Pócs 1982, Thiers 1988,
experimentally. They described the general Proctor 1990).
appearence of colonies. For Mägdefrau (1969, Life forms have successfully been used to
1982), growth form is a morphological feature recognize altitudinal belts in rain forests of SE-
of a single plant (e.g. plagiotrop, orthotrop) and Asia (Frey et al. 1990), Central Africa (Frey et

30 Frahm
3 5
5 7
Fig. 4.1: Life forms of mosses: 1. turf, 2. cushion, 3. dendroid, 4. pendant, 5. tail, 6. mat, 7. fan.

Fig. 4.2: Distribution of pendants along a transect on Mt. Kinabalu, Borneo (from Frey et al.
1990)
al. 1995) and the Andes of Colombia (Groot et Frey, W., Gossow, R. & Kürschner, H. 1990.
al. 1993). Along altitudinal transects, crusts (of Verteilungsmuster von Lebensformen,
hepatics such as Lejeuneaceae) are highly wasserleitenden und wasser-
characteristic for lowland forests, wefts and fans speichernden Strukturen in
are typical for montane forests, whereas wefts epiphytischen Moosgesellschaften am
and mats are found at higher altitudes. Subalpi- Mt. Kinabalu (Nord Borneo). Nova
ne regions are rich in cushions, turfs and mats Hedwigia 51: 87-119
(fig. 4.2). Frey, W., Kürschner, H., Seifert, U.H. 1995.
In general, mats, wefts and cushions are Life strategies of epiphytic bryophytes
particularly effective in storing water and are of tropical lowland and montane forests,
characteristic for habitats with occasional ericaceous woodlands and the
desiccation. Pendants are highly characteristic of Dendrodenecio subpáramo of the
cloud belts, since this life form can effectively eastern Congo Basin... , Trop. Bryol. 11:
„comb“ humidity from mist. Dendroids are into- 129-149.
lerant of longer periods of desiccation, since the Giesenhagen, K. 1910. Die Moostypen der Re-
internal conducting system is not effective genwälder. Ann. Jard. Bot. Buitenzorg,
enough to transport water to the upper parts of 3, 1, 711-790,
the plant. Tails and fans seem to be an adaptation Gimingham, C.H., Birse, E.M. 1957.
for a better gas exchange and to avoid to be wetted Ecological studies on groth form in
along tree trunks, which reduces gas exchange. bryophytes. I.Correlations between
growth form and habitat. J. Ecol. 45:
Birse, E.M. 1957. Ecological studies on growth 533-545.
form in bryophytes. II. Experimental Groot, A.T., van Reenen, G.B.A., Wolf, J.H.D.
studies in growth forms of mosses. J. 1993. Life forms of tropical bryophytes
Ecol. 45: 721-733. along an altitudinal gradient in the

32 Frahm


34 Frahm
northern Andes. Mscr. Univ. Amster- 1. Presence of a central strand

dam. Central strands are present in many acrocarpous
Mägdefrau, K. 1969. Die Lebensformen der mosses. They consist of a „hadrom“ consisting
Laubmoose. Vegetatio 16: 285-297. of narrow, elongated cells with oblique end walls.
Meusel, H. 1935. Wuchsformen und Wuchs- This hadrom can conduct water but not
typen der europäischen Laubmoose. sufficiently. If external water conduction is
Nova Acta Leopoldina N.F. 3 No. 12. interrupted (e.g. by shaving the rhizoids and
Mägdefrau, K. 1982. Life forms of bryophytes. leaves from the stem) the moss dries up.
In: A.J.E. Smith (ed.), Bryophyte However, the central strand contributes to water
Ecology, pp. 45-58. London. conduction a certain amount. Mosses with a
Pócs, T. , 1982. Tropical forest bryophytes , In: central strand are growing on humid substrates
A.J.E. Smith (ed.), Bryophyte Ecology, but are exposed to frequent desiccation.
pp.59-104. London
Proctor, M.C.F. 1990. The physiological basis 2. Presence of external water conduction
of bryophyte production. Bot. J. Linn. External conduction of water is made possible
Soc. 104: 61-77. by a tomentum of rhizoids along the stem and by
Thiers, B.M. 1988. Morphological adaptations concave leaves in which the water is conducted
of the Jungermanniales (Hepaticae) to by capillarity. A tomentum is characteristic for
the tropical rain forest habitat. J. Hat- mosses growing on wet, exposed substrates (e.g.
tori Bot. Lab. 64: 5-14. Philonotis on dripping cliffs). Concave leaves are
Tobiessen, P.L., Mott., K.A., Slack, N.G. 1977. typical of pleurocarpous mosses with a prostrate
A comparative study of photosynthesis, growth.
respiration and water relations in four
species of epiphytic mosses in relation 3. Papillae on the leaf surface
to their vertical distribution. Bryoph. Papillae enable the plant to soak water rapidly
Biblioth. 13: 253-277. over the leaf surface. This can easily be tested by
putting two dry plants, one with a papillose leaf
lamina and one with a smooth leaf lamina, in
water. The papillose leaf will be wetted
4.2 Water conducting and water immediately. In addition, water can also be stored
storing structures between the papillae.
The use of life forms as indicators of environ- 4. Water sacks

mental factors, especially the humidity factor, is Water sacks are especially characteristic of
based on morphological characters. In the same certain groups of hepatics. In Scapaniaceae,
way also anatomical structures are adapted to the Radulaceae and Lejeuneaceae, the lower part of
water factor, since the period in which the plants a leaf is folded upwards and forms a „gab“ of
are turgescent is also the period for different size in which water can be stored. The
photosynthesis. The water conducting structures highest differentiation of this mechanism is found
enable the plant to take up water rapidly (within in the genus Frullania, in which this lower part
seconds) and to start photosynthesis of the leaf is separated and forms a closed bottle-
immediately. or cup-like structure.
Anatomical adaptations related to water 5. Alar cells

conducting and water storing are: Alar cells are cells at the basal angles of the leaves
of certain mosses. They are found in acrocarpous
mosses (e.g. Dicranaceae) as well as, and more
Figs. 4.3-4.4 on p. 30-31: Distribution of life forms along a transect on Mt. Kinabalu, Borneo
(from Frey et al. 1990)

Fig. 4.5: Water conducting and storing structures of bryophytes. 1-2. Alar cells (Sematophyllum
brachytheciiforme), 3-5. Concave leaves 3. Marchesinia excavata, 4. Evansiolejeunea roccatii, 5.
Pilotrichella profusicaulis. 6-9. Ciliate leaves. 6-8. Herbertus doggeltianus, 9. Leptoscyphus infus-
catus. 10. Hyalocysts (Campylopus nivalis). From Kürschner & Seifert 1995..

36 Frahm
frequently, in pleurocarpous mosses (e.g. zung der epiphytischen Moosvegetation

Hypnaceae, Amblystegiaceae Brachytheciaceae, in Regenwäldern NO-Perus. Beih. Nova
and especially Sematophyllaceae). They can be Hedwigia 88: 115-141.
firm-walled, incrassate and usually reddish Frey, W., Gossow, R. & Kürschner, H. 1990.
coloured, but more often thin-walled and inflated. Verteilungsmuster von Lebensformen,
They function in absorbing water into the leaf, wasserleitenden und wasser-
which is externally conducted along the stem by speichernden Strukturen in
rhizoids or concave leaves. epiphytischen Moosgesellschaften am
Mt. Kinabalu (Nord Borneo). Nova
6. Cilia Hedwigia 51: 87-119
Bryophyte leaves are organs for water uptake (in Kürschner, H. & Seifert, U.H. 1995. Wissen-
addition to photosyntheis, of course). In the schaftliche Ergebnisse der BRYOTROP
hepatics, the leaves can be incised one to several Expedition nach Zaire und Rwanda 6.
times and form acute lobes. The apices can be Lebensformen und Adaptationen zur
condensation points for humidity. If the leaves Wasserleitung und Wasserspeicherung
are split partially at the margins (e.g. Trichocolea) in epiphytischen Moosgesellschaften im
or totally (Kurzia, Blepharostoma) to form sin- östlichen Kongobecken. Trop. Bryol.
gle cell-rows (cilia), these structures enlarge the 11: 87-118
surface of the leaves drastically and facilitate
absorption of water.
4.3 Life strategy
7. Hyalocysts
Hyalocysts are large, empty, dead cells in the Under the term life strategy the different life
leaves of some moss families such as histories of bryophytes are understood. The term
Sphagnaceae, Dicranaceae (Paraleucobryum, life strategy makes not much sense since a
Campylopus) and Calymperaceae, rarely also in strategy means a adaptation to changing
some genera of Pottiaceae (Tortula, Tortella). conditions. This is not the case in genetically fixed
They can store large amounts of water, and are life cycles. During (1979) distinguishes between:
therefore interpreted as water storing structures,
although many of them grow characteristically Fugitives
in damp habitats, where storage is not necessary. Colonists
Water storing and conducting structures were Annual shuttle species
successfully used for an altitudinal zonation of Short lived shuttle species
bryophytes in rain forests in Peru (Frahm 1987), Perennial shuttle species
Borneo (Frey et al. 1990) and Central Africa Perennial stayers
(Kürschner & Seifert 1995).
In Zaire and Rwanda, bryophyte species with Life strategies are adaptations of the life history
cilia or hyalocysts occurred only at higher to different habitats. They concern primarily the
elevations (high montane, subalpine and alpine length of the life cycle (ephemeral, annual to
belt). Species with a central stand and a rhizoid perennial) but also different types of
tomentum were mainly found above 2000 m, reproduction(sexual, asexual), spore size and
whereas papillose and mamillose leaf surfaces growth form.
were found at all elevations .
During, H. J. 1998. De diasporenvoorraad in
Frahm, J.-P. 1987. Struktur und Zusammenset- de bodem van een savanne in
Fig. 4.6: Distribution of water conducting structures along a transect on Mt. Kinabalu, Borneo
(from Frey et al. 1990).


38 Frahm
Zimbabwe. Bauxbaumiella 45: 2-13.

During, H. J., and C. Moyo. 1999. The diaspore
bank of bryophytes in a Zimbabwean
savanna. Haussknechtia Beiheft
(Ricleft Grolle-Festschrift) 9:111-114.
Frey, W., and H. Kürschner. 1991. Lebensstra-
tegien epiphytischer Bryophyten im tro-
pischen Tieflandsund Bergregenwald
am Mt. Kinabalu (Sabah, Northern Bor-
neo). Nova Hedwigia 53:307-330.
Kürschner, H., W. Frey, and G. Parolly. 1999.
Patterns and adaptive trends of life
forms, life strategies and
ecomorphological structures in tropical
epiphytic bryophytes: A pantropical
synopsis. Nova Hedwigia 69:73-99.
Olarinmoye, S. O. 1986. Aspects of survival
strategies in three common mosses in
Ibadan, Nigeria. Cryptogamie,
Bryologie, et Lichénologie 7:213-218.
Van Leerdam, A., R. J. Zagt, and E. J. Vene-
klass. 1990. The distribution of epiphy-
te growth-forms in the canopy of a Co-
lombian cloud forest. Vegetatio 87:59-
71.
Zander, R. H., and H. J. During. 1999.
Neophoenix (Pottiaceae), a new African
moss species found through soil dispore
bank analysis. Taxon 48:657-662.

5. ECOLOGY OF TROPICAL BRYOPHYTES
5.1 Habitats poicilohydric bryophytes are almost all the time

photosynthetic active. The longer the light period,
The bryophyte habitats in the tropics are much in which bryophytes are wet, the higher is the
different from those in temperate, boreal or arctic net-photosynthesis (right temperature and light
regions. Predominant vegetation type is the forest. conditions presupposed) and the higher is the
Within the forest, the soil is not much covered production of phytomass. Furthermore, humidity
by bryophytes due to the large amount amount provided by rain supports nutrients,and the more
of litter and its fast decomposition. This is rain the more nutrients get the bryophytes. The
especially true for lowland forests, where only effect, that bryophyte phytomass increases with
rarely small quantities of Fissidens are found on elevation by higher humidity and increasing
bare soil and a few bryophytes on termite nests precipitation can also be observed in mountains
and ant gardens. The amount of bryophytes on of temperate regions, where the mass of epiphytic
soil is slightly increasing in montane forests with bryophytes can reach the amount of rain forests
increasing elevation, especially along roadside in relatively dry regions or not too high elevations
banks (there are no bryophytes along roadside (Frahm 2002).
bank in the lowlands!) and forests floors covered Epiphytic bryophytes are very sensitive to habitat
by bryophytes are only found in the subalpine. factors such as bark structure, bark pH,
Furthermore, there are usually no rocky habitats microclimate. The fact that they are exclusively
at lower altitudes in the humid tropics due to the supplied with water and nutrients from the
deep lateritic soils and fast chemical atmosphere makes them to excellent
decomposition. This confines the variety of bioindicators for air quality, which has been often
habitats in rain forests areas mainly to trees and used in Europe but so far only to a small extend
their leaves. in tropical countries (cf. chapter 8).
The tree habitat for epiphytes is very different
5.1.1 Epiphytes concerning its microhabitat conditions. Previous
The high to immense quantity of epiphytic authors such as Richards (1984, 1991)
bryophytes is a special character of (temperate differentiated between shade and sun epiphytes.
and tropical) rain forests. It is a direct result of Therefore the tree is usually divided into several
the high humidity. High humidity means that the zones (trunk, canopy with inner and outer

40 Frahm
branches and twigs). The most widely used a a determination of the cover of epiphytic
classification is that by Johannson (1974), see bryoohytes revealed that a high percentage of
fig. 5.1. trees had no bryophyte cover. These trees were
The bryophyte flora (as well as the flora and fauna trees with smooth bark and covered with
in general) of the canopy has formerly much been crustaceous lichens. Flaky bark is hardly
neglected. Canopy studies were usually confined colonized by bryophytes. Important is also the
to fallen trees. Since 25 years, alpine climbing water storing capacity of the bark. Soft bark stores
techniques, canopy walks, cranes (figs. 5.3, 5.4) more water and this water is released over a
and airships make canopy studies possible. longer time, resulting in more luxuriant epiphyte
Especially climbing techniques and crane studies grows. Another factor is the pH of the bark. This
have enhanced the knowledge of the bryoflora factor is not as important in the tropics. because
of the canopy. there are apparently no trees with basic reaction
The epiphytic bryophyte species of shady in contrast to temperate regions, were some trees
habitats are more vulnerable to forest destruction have barks with an pH of around 7. Measuremts
and disappear soon when the forest is opened. in different parts of the tropics (Frahm 1987,
They are good indicators for primary forests. 1990, 1994) revealed that the pH of bark of
Canopy bryophytes are, however, more tolerant tropical trees ranges between 4 and 6. As
to desiccation. They are also found epiphytic in phytosociological studies have shown, the
open habitats, e.g. savannahs and plantations. composition of species depends on differences
Therefore primary forests are more species rich in the bark pH. This explains that we have diffe-
than secondary forests. rent bryophyte communities on tree trunks in the
same study area on different trees.
The epiphyte diversity differs depending on: All these factors also concern host trees in the
1. the altitude of the study site extra tropics.
2. the host tree,
3. the height within a tree, 3. There is an uneven distribution of epiphytes
4. the number of trees studied. within a tree. The results of such studies,
however, differ. Richards (1984) studied 28 cut
1. Lowlands were regarded as less rich in species. down trees in a lowland rain forest in Guyana.
A comparison of an andine forest in Venezuela He found that the understorey is richer in species
and a lowland forest at the Upper Orinoco and the canopy has less species due to harsh
revealed distinctly less species (León-Vargas conditions of high ligt intensity, strong
2001). desiccation and high temperatures. In contrast, a
study of also 28 trees in French Guiana using
2. The bark structure and bark chemistry of the climbing techniques revealed an increase of
host tree has important influence on the number mosses, liverworts and lichens above 5 meters
and composition of epiphytic bryophytes. The (Montfort & Ek 1990). Therefore Gradstein
bark can be smooth, cracked or flaky to various called it a myth that lowland forests are poor in
degrees. Smooth bark does not much accumulate bryophyte species since their species richness is
humidity or humus and has usually lower located in the canopy (where the bryophytes have
numbers of species. Furthermore, this type of more light). A similar effect could be observed
bark is often colonized by crustaceous, even in a montane forest in Ecuador (Müller & Frahm
endophytic lichens. These lichens have an 1998), where only 14 of a total of 67 bryophyte
allelopathic effect on bryophytes and flowering species were found on the trunks of 10 trees. Also
plants and inhibit their colonization. In in a montane forest in Venezuela, the canopy had
experiments, aquaceous extracts of these lichens higher species numbers than the trunks (León-
inhibited spore germination of bryophytes and Vargas 2001). Especially the number of mosses
well as seed germination of Bromeliaceae (Frahm in the canopy is higher, because they are more
et al. 2000). This effect has been detected in a drought tolerant.
tropical lowlaqnd rain forest in Venezuela, where

Fig. 5.1: Epiphyte zones after Johannson Fig. 5.2: Species per tree number curve (from
(1974). Montfort & Ek 1990)
4. Also the number of trees studied is important. Leguminosae na mata pluvial tropical
A „minimum tree curve“ (fig. 5.2.) can give an (Dois Irmaos, Recife-PE).[English
estimate. In this case. a minimum of 5 trees per abstract] , Anais de VII Reuniao
study plot is required for a representative Nordestina de Botanica.
inventory. Cornelissen, J. H. C., and H. ter Steege. 1989.
Distribution and ecology of epiphytic
Akinsoji, A. 1991. Studies on the epiphytic flora lichens and bryophytes in dry evergreen
of a tropical rain forest in southwestern forest of Guyana. Journal of Tropical
Nigeria. II. The bark microflora. Ecology 5:131-150.
Vegetatio 92:181-185. Cornelissen, J. H. C., and H. ter Steege. 1986.
Augier, J. 1974. Bryophytes corticoles a l’etage Vertical distribution of epiphytes in a
submontagnard dans l’ouest rainforest of Guiana. Flora Guianas
camerounais. Annls. Fac. Sci. Univ. Newsletter 3:17-19.
Fed. Cameroun 17: 67-93. Cornelissen, J.H.C. & Gradstein, S.R. 1990.
Barbosa, D. C. de A., A. de J. R. da Silva, A. On the occurrence of bryophytes and
M. da S. Correia, M. das G. V. macrolichens in different lowland rain
Marinho & M. R. Fonseca 1985 . forest types at Mabura Hill, Guyana.
Levantamento preliminar de briofitas Tropical Bryology 3: 29-36.
corticicolas em Moraceae e Dilg, C., Frahm, J.-P. 1997. Untersuchungen zur

42 Frahm
Fig. 5.3: Crane above the rain forest canopy of a lowland rain forest, Surumoni-project, Upper
Orinoco.
Fig. 5.4: Bryological studies in the gondola of the crane in the canopy, Surumoni-project.

Flora der epiphytischen Moose der N.M. 2001. Epiphytic bryophytes of

südlichen Drakensberge (Südafrika). Monteverde, Costa Rica. Tropical
Tropical Bryology 13: 35-46. Bryology 20: 63-70.
Frahm, J.-P. 1987. Ökologische Studien über Montfoort, D., and R. van Ek. 1990. Vertical
die epiphytische Moosvegetation in distribution and ecology of epiphytic
Regenwäldern NO-Perus. Nova bryophytes in a lowland rain forest of
Hedwigia, Beih. 88:143-158. French Guiana. MS.c. Thesis,
Frahm, J.-P. 1987. Struktur und Zusammen- University of Utrecht, Utrecht.
setzung der epiphytischen Moosvege- Müller, U., Frahm, J.-P. 1998. Diversität epi-
tation in Regenwäldern NO-Perus. phytischer Moose eines westandinen
Beih. Nova Hedwigia 88: 115-141. Bergregenwaldes in Ecuador. Tropical
Frahm, J.-P. 1990. The ecology of epiphytic Bryology 15:.29-44.
bryophytes on Mt. Kinabalu, Sabah Pócs, T., and A. Szabo. 1993. The epiphytic
(Malaysia). Nova Hedwigia 51:121- vegetation the endemic giant groundsel
132. (Senecio barbatipes) of Mt. Elgon,
Frahm, J.-P. 1994. Scientific Results of the Kenya. Opera Botanica 121:189-194.
BRYOTROP Expedition to Zaire and Osada,T., and T. Amakawa. 1956. An
Rwanda 1. The ecology of epiphytic observation on the ecological
bryophytes on Mt. Kahuzi (Zaire). distribution of pendulous bryophytes in
Tropical Bryology 9: 137-152. the Tsushima Islands. Journal of the
Frahm, J.-P. 2002. Untersuchungen zur Höhen- Hattori Botanical Laboratory 17:52-54.
gliederung der Moose und der Wasser- Rebelo, C. F., Y. Struftaldi-De Vuono, and M.
speicherung von epiphytischen Moosen Domingos. 1995. Estudio ecológico
entlang eines Transektes durch den de comunidades de briófitas epifitas da
Westhang der Vogesen. Bull. Soc. Hist. Reserva Biológica de Paranapiacaba,
Nat. Colmar 64:67-78. SP, en trechos de floresta sujeitos à
Frahm, J.-P., Kürschner, H. 1990. What drives influência da polução aérea. Revista
mosses onto trees? German Res. 2/90: Brasileira de Botanica 18:1-15.
6-9. Rhoades, F. M. 1995. Nonvascular epiphytes in
Frahm, J.-P., Kürschner, H. 1989. Was Moo- forest canopies: Worldwide distribution,
se auf die Bäume treibt. Untersuchun- abundance, and ecological roles. In
gen im Regenwald. Forschung, Mitt. der Lowman, M. D., and N. M. Nadkarni
DFG 4/89: 18-22. (eds.). Forest canopies, pp. 353-408.
Frahm, J.-P., Specht, A., Reifenrath, K., León- Academic Press, New York.
Vargas, Y. 2000. Allelopathic effects Richards, P. W. 1984. The ecology of tropical
of crustaceous lichens on epiphytic forest bryophytes. In Schuster, R. M.
bryophytes and vascular plants. Nova (ed.). New manual of bryology vol. 2,
Hedwigia 70: 245-254. pp 1233-1270. Hattori Botanical
Johannson, D. 1974. Ecology of vascular Laboratory, Miyazaki, Japan.
epiphytes in West African rain forest. Richards, P. W. 1991. A bryologist in British
Acta Phytogeogr. Suecica 59: 1-136. Guiana and the West Indies. Journal of
León-Vargas, Y. 2001. Diversity of epiphytic Bryology 16:437-441.
bryophytes in a montane cloud forest Russell, K. W., and H. A. Miller. 1977. The
in the Venezuelan Andes. PhD Diss. ecology of an elfin forest in Puerto Rico,
Univ. Bonn. 17. Epiphytic mossy vegetation of Pico
Matelson, T. J., N. M. Nadkarni, and J. T. del Oeste. Journal of the Arnold
Longino. 1993. Longevity of fallen Arboretum 58:1-24.
epiphytes in a neotropical montane Sillett, S. C. 1991. Canopy bryophyte
forest. Ecology 74:265-269. communities of a lower montane wet
Mervin, M.C., Gradstein, S.R., Nadkarni, forest in Costa Rica: Quantitative

44 Frahm
sampling in intact forest and isolated family Lejeuneaceae, which counts for 95% of
trees. M.Sc. thesis, University of all epiphylls, which have especially adapted to
Florida, Gainsville. this kind of habitat. The morphological
Sipman, H. J. M. 1997. Observations on the adaptations include:
foliicolous lichen and bryophyte flora - short life cycle
in the canopy of a semideciduous - reduction of the gametophyte (neoteny)
tropical forest , Abstracta Botanica - diaspore production after a short time
21(1): 153-161 - frequent production of gemmae
Wolf, J. H. D. 1993. Ecology of epiphytes and - adaptation for fixation on the leaf by mucilage
epiphytes communities in montane rain secreted by plants or gemmae, develoment of
forests, Colombia. Ph.D. dissertation, rhizoid plates.
University of Amsterdam, The Nether- Diversity studies (e.g. Lücking 1995, 1997)
lands. include the analysis of diversity on single leaves,
Wolf, J. H. D. 1994. Factors controlling the single localities and larger geographical areas.
distribution of vascular and nonvascu- Ecology studies include the determination of the
lar epiphytes in the northern Andes. influence of altitude, leaf preferences,
Vegetatio 112:15-28. composition of species, micro climate, and hu-
Wolf, J. H. D. 1995. Nonvascular epiphyte di- man influence. Few species are confined to the
versity patterns in the canopy of an lowlands; most species occur from the lowlands
Upper Montane Rain Forest, Central to 1600, from the lowlands to 3000 m or betwen
Cordillera, Colombia. Selbyana 16: 500 and 1600, 500 and 3000 or 1600 and 3000
185-195. m (Eggers 2001). The increase with the elevation
is caused by increasing facultative epiphyllous
species. A decrease of diversity is caused by dry
climate and human influence, resulting in covers
5.1.2 Epiphylls of small, closely appressed species, whereas in
strongly humid conditions, the epiphyllous
The study of epiphylls has gained much attention species can even change to upright growth. Three
since epiphylls are said to be a special microsites can be differentiated: shady
characteristic of tropical rainforests, although understorey, light gaps and the canopy. The most
they are more a characteristic of all evergreen species rich sites are light gaps with sufficient
rainforests and occur also in temperate rain forests humidity, especially sites along streams. Another
(e.g. the Valdivian rain forest, where is probably topic of interest are interactions with other
the origin of this life syndrome). Epiphyllous foliicolous organisms and the host leaf. Epiphylls
growth requires high rainfall and the presence of absorb light intensity. Especially in shady habitats
suitable leaves. Primarily, the leaves function for with less than 2% relative light intensity and in
bryophytes simply as substrate which is shown leaves with a high cover of epiphylls, this effect
by the fact that bryophytes can also grow on can harm the host plant. On the other hand,
lizzards, turtles, beetles or coke bottles. Even epiphyllous bryophytes with their antifeedant
filmy ferns can be colonized as well as other effects might deter leaf herbivores including leaf
bryophytes. Only some bryophytes are substrate cutter ants. An important interaction is that with
specific, many epiphyllous species can be found cyanobacteria, which live in close contact with
on other substrates as well. The degree of bryophytes which provide a humid micro-
epiphyllism increase with humidity and the high- environment for them. The nitrogen fixed by the
er the humidity the more corticolous bryophytes cyanobacteria can be taken up by the epiphyllous
shift from bark to leaves. bryophytes as well as the host leaf (Bentley &
The habitat is shared with lichens, fungi, green Carpenter 1980, 1984, Bentley 1987, 1989,
algae and cyanobacteria. It forms a special Carpenter 1992). Competition, e.g. with
environment called „phyllosphere“. epiphyllous lichens, has not yet been studied
Conspicuously, most epiphylls are hepatics of the much. It may not be so important, since both seem

to have avoidance strategies. Hepaticae of Kowloon Peninsula (Hong

Kong). Tropical Bryology 4: 17-22.
But, P.P.P.H., Wu, P-C., Wang, M.-Z. 2000.
Aguirre, C. J. & E. Castillo , 1993 , [Abstract] Epiphyllous liverworts on rosette leaves
Tropical and sub-Andean epiphyllous of Ardisia species (Myrsiniaceae) in
bryoflora in El Macizo de Tatama, China. Tropical Bryology 19: 27-30.
Colombia. , AAU Reports 31: 9, Carpenter, E. J. 1992. Nitrogen fixation in the
Allorge, V., and P. Allorge. 1939. Sur la epiphyllae and root nodules of trees in
répartition et l’écologie des hépatiques the lowland tropical rainforest of Costa
épiphylles aux Açores. Boletim Rica. Acta Oecologica 13:153-160.
Sociedade Broteriana, Coimbra 13:211- Coley, P. D., and T. A. Kursar. 1996. Causes
231. and consequences of epiphyll
Baudoin, R. 1985. Analyse de la distribution colonization. In Mulkey, S. S., R. L.
des bryophytes épiphylles sur la Chazdon, and A. P. Smith (eds.).
soufrière de Guadeloupe. Compte Tropical forest plant ecophysiology, pp.
Rendu des Séances de la Société de 337-362. Chapman and Hall, London.
Biogéographie 61:47-57. Coley, P.D., T.A. Kursar & J.-L. Machado.
Bentley, B. L. 1987. Nitrogen fixation by 1993. Colonization of tropical rain
epiphylls in a tropical rainforest. Annals forest leaves by epiphylls: effects of site
of the Missouri Botanical Garden and host plant leaf time. Ecology 74:
74:234-241. 619-623.
Bentley, B. L. 1989. Nitrogen fixation by Daniels, J. D. 1998. Establishment and succes-
epiphylls in a tropical rainforst: The sion of epiphyllic bryophyte assembla-
ecological conversion of leaves to roots. ges on the fronds of the Neotropical
, American Journal of Botany 7(6 understory palm Geonoma seleri (Co-
Suppl.): 4 sta Rica). Ph.D. dissertation. Universi-
Bentley, B. L., and E. J. Carpenter. 1980. ty of North Dakota. 114 pp.
Effects of desiccation and rehydration Eggers, J. 2001. Epiphyllous Lejeuneaceae in
on nitrogen fixation by epiphylls in a Costa Rica. Contributions to the
tropical rainforest. Microbial Ecology altitudinal distribution of selected
6:109-113. species. Tropical Bryology 20: 109-116.
Bentley, B. L., and E. J. Carpenter. 1984. Direct Eze, J. M. O., and G. K. Berrie. 1977. Further
transfer of newly-fixed nitrogen from investigations into the physiological
free living epiphyllous micro-organisms relationship between an epiphyllous
to their host plant. Oecologia 63:52-56. liverwort and its host leaves. Annals of
Bentley, B.L. & Carpenter, E.J. , 1980 , Effects Botany 41:351-358.
of desiccation and dehydration on Freiberg, E. 1994. Stickstoffixierung in der
nitrogen fixation by epiphylls in a Phyllosphäre tropischer
tropical rainforest , Microbial Ecology, Regenwaldpflanzen in Costa Rica.
6, , 109-113, Ph.D. Dissertation, Faculty of Natural
Berrie, G. K., and J. M. O. Eze. 1975. The Science, University of Ulm, Germany.
relationship between an epiphyllous Freiberg, E. 1998. Microclimatic parameters
liverwort and host leaves Annals of influencing nitrogen fixation in the
Botany 39:955-963. phyllosphere in a Costa Rican
Busse, W. 1905. Über das Auftreten premontane rain forest. Oecologia
epiphyllischer Kryptogamen im 117:9-18.
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Berichte Deutsche Botanische epiphyllae. Thesis. Dept. of Botany,
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University of Georgia, Athens, Georgia. Bryology 8:275-289.
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Mitteilungen des Bundes für Forst und flaccida Lindenb. & Gottsche to leaf
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114.

48 Frahm
5.1.3 Rotten wood were found on Cyathea, two on Dicksonia trunks.

Tree ferns are very ancient plants; they existed
There is little information on bryophytes from already in the Mesozoic and provided a constant
rotten wood, which is one of the few habitata for microhabitat for more than 100 mio years.
bryophytes in lowland forests. This habitats is Therefore the relation between the bryophyte
usually occupied by Sematophjyllaceae, vegetation on tree fern trunks in SE-Brazil and
Hookeriaceae and Leucobryaceae. In montane in New Zealand was studied. The associations
forests, the bryophyte flora on rotten wood is in both parts were not just composed by identical
richer since the trunks are not so fast decayed. In species but by vicariant species. For instance,
Tanzania, 102 species of bryophytes were Hymenodon aeruginosus in New Zealand is
identified from rotten woods (Mattila & Koponen replaced in Brazil by H. pilifer, Pyrrhobryum
1999). The bryophyte vegetation varies bifarium by P. spiniforme, Hypopterygium
depending on the stage of decomposition. An didictyon by H. tamarisci. This shows that in dif-
easy method to determine this stage is the ferent parts of the world species of the same genus
determination of the depth of penetration of a compose associations on comparable habitats,
knife (e.g. 1: knife does not penetrate, 2. due to their common habitat preferences.
penetrates one centimeter, 3, penetrates several
centimeters, 4. penetrates to the handle).
. Ahmed, J., Frahm, J.-P. 2002. Moos-
Germano, S. R., and K. C. Pôrto. 1997. gesellschaften auf Baumfarnstämmen in
Ecological analysis of epixylic Südostbrasilien. Tropical Bryology 22:
bryophytes in relation to the 135-178.
decomposition of substrate
(Municipality of Timbaúba-
Pernambuco, Brazil). Cryptogamie,
Bryologie, et Lichénologie 18:143-150. 5.1.5. Unusual substrates
Mattila, P., and T. Koponen. 1999. Diversity
of the bryophyte flora and vegetation The higher the humidity, the lower is the
on rotten wood in rain and montane influence of the substrate. If there is sufficient
forests in northeastern Tanzania. atmospheric suplly with water and nutrients,
Tropical Bryology 16:39-164. bryophytes (and especially also lichens) can grow
Sastre-De Jesús, I. 1992. Estudios preliminares on any substrate, including shells of water turtles,
sobre comunidades de briofitas en old shoes, traffic signs ot old cars. Therefore it is
troncos en descomposición en el bosque not surprising that bryophytes from tropical
subtropical lluvioso de Puerto Rico. forests can also grow epizoic on beetles and
Tropical Bryology 6: 181-192. lizards.
Another unusual substrate has been reported from
Peru (Frahm 1985) and was also observed in
5.1.4. Tree Ferns Venezuela: bryophytes growing on ant gardens
in the canopy (fig. 5.7).
Tree ferns are an ideal substrate for bryophytes,
causing a rich vegetation. They have, however, Frahm, J.-P. 1985. A Bryophyte in an Ant
gained no special interest in the past. The first Garden. Bryol. Times 34:1.
and only study was performed in SE-Brazil on Gradstein, S. R., and C. Equihua. 1995. An
Cyathea and Dicksonia trunks. A total of 142 epizoic bryophyte and algae growing on
species was found on these trunks. Most species the lizard Corythophanes cristatus in a
grew only by chance in this habitat, only 20 of Mexican rain forest. Biotropica 27:265-
the species was found in more than 10% of the 268.
tree fern trunks studied. Five bryophyte Gradstein, S. R., D. H. Vitt, and R. S.
associations could be described, of which three Anderson. 1984. The epizoic

Fig. 5.7: The moss (Brachymenium columbicum) growing in an ant garden (NE-Peru)
occurrence of Daltonia angustifolia ve than vascular plants and have the advantage,
(Musci) in Papua New Guinea. that they are much less numerous. Whereas it
Cryptogamie, Bryologie et makes enormous difficulties to determine the
Lichénologie 5:47-50. vascular flora along an altitudinal transect, the
Gressit, J.L., Samuelson, G.A. & Vitt, D.H. determination of bryophytes is much easier. This
1968. Moss growing on living Papuan could be proved all around the world, in Peru,
moss-forest weevils. Nature (London) Colombia, Central Africa and East Asia.
217: 765-767. There are several different attempts to determine
the altitudinal zonation of rain forests:
- by floristic analysis
For that purpose, the species composition is
5.2. Altitudinal zonation determined along a transect. It is sufficient to
focus the analysis on representative (undisturbed)
5.2.1 Determination of altitudinal belts areas. Intervalls of 200 m altitude are sufficient.
As a representative area, one hectare has proved
As figured out in chapter 4, bryophytes are useful, since such an area is most widely used
adapted to different ecological conditions by and allows comparisons, however, also 25 x 25
morphological and anatomical structures. These m can be sufficient depending on the structure
adaptations vary according to the environment. of the forest. All species are noted, collected and
Since the environmental conditions vary with the identified and arranged in a table. It can be seen
altitude, we found differently adapted bryophytes from such a table that certain species cluster
in different altitudes. Therefore bryophytes are a together. These species are grouped together and
useful tool for elaboration of altitudinal allow a vertical classification. The composition
zonations. Bryophytes react much more sensiti- of species or the restriction of certain genera to

50 Frahm
vertain altitudes is so characteristic in tropical - by determination of life form spectra

forests, that a bryologist with some experience Autecological adaptations reflect changing
can often determine the altitude as exact as 200 environmental conditions, too, as explained in
m without altimeter by bryophytes only, since chapter 4. By this way, useful altitudinal
many species and genera are indicator species zonations can be derived even without any
for certain altitudinal belts. The absolute elevation determination of species.
is only modified by the humidity: the higher the
humidity the more are the altitudinal belts - by estimation of phytomass
depressed. In hyperhygric conditions, the species Similarly to the estimation of cover, the
characteristic of higher altitudes go far more determination of the phytomass of (preferably
lower down than in drier regions.. epiphytic bryophytes) give valuable result for
constructing altitudinal zonations (see also
- by species numbers chapter 5.3).
Beside the species composition, also the number
of species per area is significant for different In total, all methods described here give identical
altitudes and shows a characteristic curve along results.
a transect. Therefore it is not necessary to identify
all species but to determine the number per area
(fig. 5.8). Species numbers in the frest 5.2.2 Results
understorey are comparably low in the lowland
forest, increase a bit in the submontane forest, There are numerous classifications and
show another increase in the lower montane terminologies characterizing the altitudinal belts
forest and a highly significant increase in the in the wet tropics. They were differrently derived
upper montane forest and a decrease at and above from the aspect of the forest (height, presence of
the forest line. palms, tree ferns etc.), which was commonly used
by geographers, or by various groups of animals
- by determination of floristic discontinuities includsing butterflies and birds. The belts are
A variation of the both previous method is the differently named with local names (tierra
following: all species have lower and upper ends caliente, tierra templada, tierra fria or selva
of their ranges along a transect. Therefore it is neotropical inferior, bosque subandino or bosque
determined from the table showing the andino), as rain forest, montane forest, cloud
occurrences of the single species in the relevées forest, mist forest, or with numbers. The
along a transect, how many species have their terminology, however, should be worldwide
uppermost viz. lowermost occurrence in every comparable and thus use the same system. A
altitude. The results are plotted in a graph (fig. classification into lowland, submontane, lower
5.9). The altitude, in which a maximum number montane, upper montane and subalpine forest is
of species with lower- or uppermost occurrences therefore recommended. For indication of the
is found, indicates a shift in the species tropical conditions, the prefix tropical should be
composition, and thus a change in the zonation. added.
The altitudinal zonation derived from bryophytes
- by estimation of cover is primarily a climatic one: climate factors
Also the determination of the cover of bryophytes determine the species number, species
either on soil and rocks or on trees gives an clear composition, adaptation and phytomass. It is
altitudinal zonation (cf. fig. 5.10). Reason is that therefore not a classification for bryophytes but
the amount of phytomass is determined by for the ecological conditions of the forest.
physiological factors, which change along a In equatorial latitudes, the altitudinal belts are as
transect (see chapter 5.3) and phytomass is an follows:
expression of these changing ecological
conditions. - 3-400 m: tropical lowland forest
- 1100-1300 m: submontane forest

- 1800 m: lower montane forest

- 2800 m: upper montane forest
- forest line: subalpine forest
The limits of the altitudinal belts vary moderately.

They depend on:
- the elevation of the mountain. Low mountains

have the altitudinal zonation compressed and a
distinct peak effect at top, causing subalpine
conditions even at lower altitudes. Higher
mountains have the altitudinal belts extended. Tis
is explained by higher temperatures, caused by
Fig. 5.8: Characteristic species curve along a
the „mass effect“ of the mountain.
transect in Colombia between 950 and 3500 m
(after Van Reenen & Gradstein 1983).
- isolated mountains differ from mountain ranges
- the exposition. As everywhere in the world, N-
exposed slopes have lower limits than S-exposed (Canary Islands). Ms. Univ. Newcastle
slopes. This is a strong indication that the On Tyne.
zonation is also caused by the temperature. Churchill, S. P. 1991. The floristic composition
and elevation distribution of Colombian
- the humidity. Regions with higher precipitation mosses. The Bryologist 94:157-167.
have depressed altitudinal belts. The reason is Enroth, J. 1990. Altitudinal Zonation of
amongst other factors that the temperatures are Bryophytes on the Huon Peninsula,
lowered. Another reason is, that the high Papua New Guinea. Tropical Bryology
humidity, which is found e.g. in the high monta- 2: 61-90.
ne belt, and which is dependend for certain Frahm, J.-P. 1990. The altitudinal zonation of
bryophytes, is realized already at lower altitudes. bryophytes on Mt. Kinabalu. Nova
Hedwigia 51:133-149.
- the latitude. The forest line is declining from Frahm, J.-P. 1994. Scientific Results of the
the equator towards the poles, until it reaches sea BRYOTROP Expedition to Zaire and
level in the arctic/antarctic. By this way, the Rwanda 2. The altitudinal zonation of
altitudinal belts are not compressed but they and the bryophytes on Mt. Kahuzi, Zaire.
at sea level depending on the latitude following Tropical Bryology 9: 153-168.
the model by Troll (fig. 5.7). This means that for Frahm, J.-P., and S. R. Gradstein. 1991. An
instance in SE-Brazil, we have no lowland forest altitudinal zonation of tropical rain fo-
but the forest starts with an submontane forest at rests using bryophytes. Journal of Bio-
sea level. This forest in SE-Brazil has the same geography 18:669-678.
„mossiness“ and comparable phytomass, cover Glime, J.M., P.S.Hudy & S. Hattori
and species numbers as a tropical forest in 800 1990.Diversity and altitudinal niche
m in equatorial latitude. In Patagonia, the subal- width characteristics for 35 taxa of the
pine forest ands at sea level. In fact, the Papua New Guinea Frullania flora with
bryological characteristics of the forest in consideration of sibbling pairs. Tropical
Patagonia and in Ecuador at 3300 m are very Bryology 2: 103-116.
similar, and even many species are found in both Gradstein, S. R., and J.-P. Frahm. 1987. Die
forests. floristische Höhengliederung der Moose
entlang des BRYOTROP-Transektes in
Bines, T.J. 1964. Mosses in relation to zonation NO-Peru. Beih. Nova Hedwigia 88:105-
on Hierro, Expedition report, 1964 113.
expedition to Gomera and Hierro Harrington, A.J. 1976. Vegetational zonation

52 Frahm
Fig. 5.9: Determination of the floristic continuities along a transect from the lowland rain forest to
the forest line in NE-Peru (after Gradstein & Frahm 1987). The peaks indicate elevations, in which
most species have either their upper- or lowermost occurrence thus indicating a floristic change.
Fig. 5.10: Bryophyte cover values along a transect in Colombia (after Van Reenen & Gradstein
1983). I: lowland forest, II. submontane rain forest, III: lower montane rain forest, IV: upper monta-
ne rain forest, V: paramo. SR group: terrestrial and saxicolous sapecies, TL-group: epiphytes.

of the bryophytes of West Africa. Bull.

Brit. Bryol. Soc. 27: 00-00
Kürschner, H. 1995. Wissenschaftliche Ergeb-
nisse der BRYOTROP-Expedition nach
Zaire und Rwanda. 5. Höhengliederung
epiphytischer Moose im östlichen Kon-
gobecken und den angrenzenden Ge-
birgsstöcken (Parc National de Kahuzi-
Biega/Zaire, Forêt de Nyungwe/Rwan-
da). Tropical Bryology 11: 77-86.
Russell, S. 1981. Humidity gradients and
bryophyte zonation in the Afro-montane
forests of the eastern Cape, South
Africa, XIII Int. Bot. Congress
Abstracts: 291. Sydney
Russel, S. 1982. Humidity gradients and
bryophyte zonation in the Afro-montane
forests of the Eastern Cape, South Fig. 5.7: Scheme of the altitudinal zonation
Africa. J. Hattori Bot. Lab. 52: 299-302. between 40° S and N (after Troll).
Pócs, T. 1994. The altitudinal distribution of
Kilimanjaro bryophytes. Pp. 797-812 in:
Seyani, J.H. & Chikuni, A.G. (eds.)
Proceedings of the XIIth Plenary Colombia. Journal of the Hattori
Meeting of AEFTAT, Zomba, Malawi, Botanical Laboratory 56:79-84.
1991. Vitt, D. H. 1991. Distribution patterns, adaptive
Seifriz, W. 1924. The altitudinal distribution of strategies, and morphological changes
lichens and mosses on Mt. Gedeh, Java. of mosses along elevational and
Journal of Ecology 12:307-313. latitudinal gradients on South Pacific
Van Reenen, G. B. A. 1987. Altitudinal Islands. In P.L. Nimis, P. L., and T. J.
bryophyte zonation in the Andes of Crovello (eds.). Quantitaive approaches
Colombia: A preliminary report. to phytogeography, pp. 205-231.
Symposia Biologica Hungarica 35:631- Kluwer Academic Publishers,
637. Dordrecht, The Netherlands.
Van Reenen, G. B. A., and S. R. Gradstein.
1984. Analisis de la vegetacion de
briofitas en el transecto Buritca - La 5.3 Phytomass and water storage
Cumbre (Sierra Nevada de Santa Marta, capacity
Colombia). Studies on Tropical Andean (with collaboration by T. Pócs)
Ecosystems 2:189-202.
Van Reenen, G. B. A., and S. R. Gradstein. An important ecological rôle of bryophytes in
1983. Studies on Colombian tropical rain forest areas is the water storage
Cryptogams XX. A transect analysis of function. By this way, bryophytes
the bryophyte vegetation along an - regulate the level of water courses
altitudinal gradient in the Sierra Nevada - prevent soil erosion
de Santa Marta, Colombia. Acta - warrants water suplly for a much larger area
Botanica Neerlandica 32:163-175. for
Van Reenen, G. B. A., and S. R. Gradstein. -- drinking water
1984. An investigation of bryophyte -- irrigation
distribution an ecology along an -- energy.
altitudinal gradient in the Andes of

54 Frahm
Beside, deforestation has the effects of in Colombia and Frahm (1994) in Zaire. A survey
- seasonality of watercourses (desiccation and (at the former state of knowledge) of bryophyte
inundation) phytomass in tropical ecosystems was given by
- drying and warming up the local climate Frahm (1990).
topsoil erosion (up to 4 cm per year) In a less mossy forest at Mt. Kilimanjaro, Pócs
- torrential rains causing landslides. (unpubl.) determined a water storage capacity of
Therefore catchment forests are required in mon- 4500 l/ha in an intact forest but only 2700 l/ha
tane areas. To determine the rôle of bryophytes, after logging.
studies of the water storing capacity of epiphytic Phytomass increases with elevation (see chapter
bryophytes have been undertaken in several parts on altitudinal zonation). A considerable amount
of the tropics. The results are difficult to compare, of epiphytic phytomass is reached above 2000
since some authors determined the phytomass of m, which stresses the importance of high monta-
single trees, others per square meter, and ne forests. Below 1000 m, the epiphytic bryomass
accordingly estimations of phytomass per hectare is with 10-12 g/m2 quite low in all tropical regions
vary. On horizontal branches, humus studied (Frahm 1987, 1990, 1994). Maxima are
sccumulated below the bryophytes may be realized at the forest lines. There, the phytomass
included in the calculation or removed before can be 1000 times as high as in the lowland forest.
measurement. The same concerns the Reason are high temperatures and low light
determination of the water storage capacity. Part intensities at lower altitudes (see chapter
of the water is stored by the plants, another part ecophysiology). Local microclimatic conditions
is stored between the plant (interception water). (cloud belts, „elfin forests“ at mountain ridges)
The storage capacity of bryophytes is usually can have a strong influence on these values.
three times of the dry weight. A higher percentage The highest phytomass is found on branches,
of Leucobryaceae and Calymperaceae with especially horizontal branches, and not on the
specialized water storing adaptations can raise trunks of trees. Accordingly, the water storage
the value to 5 times. Interception water is difficult capacity of branch epiphytes is higher (Björk &
to determine since it easily runs off when the Mareby 1992). Naturally, the larger the tree the
bryophytes are balanced. In any case, these higher the water storage capacity, which
studies give an idea of the relations. emphazises the value of old growth forests.
Jacobsen (1978) was the first who made Although the values for water storage capacity
phytomass measurements of epiphytic are impressive, the following points have to be
bryophytes in Transvaal. He found 34 g/m2 and considered (Frahm 1994):
34 kg/ha. Pócs (1976, 1980) was the first who - in rain forest climates, the bryophytes are rarely
determined phytomass and water storage really dry and cannot take up the full amount of
capacity of epiphytes in the Uluguru Mtns., water. During rain fall, the bryophytes are soaked
Tanzania. He found 2130 kg phytomass/ha at very fast with water and the additional rain water
1415 m and 10.000 kg/ha phytomass at 2100 m cannot be stored. To eliminate this effect from
elevation. The water storage capacity was 15.000 the calculations of water storage, the interception
viz. 50.000 l/ha. Of the 10.000 kg total water can be neglected.
phytomass, 8.000 kg were found in the canopy. - the percentage of the total rain fall stored by
In the Nguru Mtns, Tanzania, Björk & Mareby bryophytes is very low. In an area with 4000 mm
(1992) found 13.000 kg/ha with 53.000 l water of annual rain fall, we have a precipitation of 40
storage. Wolf (1993) determined 2000 kg million liters per hectare and year. Assumed it is
phytomass/ha with 10.000 l water storage in an raining every day, we have 109.589 l/ha rainfall
Andean forest in Colombia at 1500 m elevation per day. If the water storing capacity of
and 20-23.000 kg/ha with 80-1000.000 l at 3500 bryophytes is 10.000 l/ha, only 10% of the rain
m. fall is stored by bryophytes, presumed that the
Additional calculations of phytomass and water bryophytes were totally dry before rainfall (what
storage capacity was made Frahm (1987) in Peru, they not are). At lower altitudes with accordingly
Frahm (1990) in Borneo, Hofstede et al. (1993) lower rates of phytomass, the percentage of rain

stored by bryophytes gets much lower to rates of Björk, L., Mareby. J. 1992. Epiphytic mass and
1-2% of the precipitation and is at least even water storage capacity of epiphyte
neglible in lowland forests. vegetation in a mossy montane forest,
The effects of phytomass and water storage Nguru Mountains, Tanzania. Swedish
capacity are not confined to tropical forests but Univ., Internal Rural Devel. Centre
similar in other types of humid forests such as Working Paper 213, Uppsala.
temperate rain forests or montane forests in Clark, K. L., N. M. Nadkarni, and H. L. Gholz.
temperate regions. They are simply related with 1998. Growth, net production, litter
the humidity, which is increasing with elevation decomposition, and net nitrogen
in almost all mountains. accumulation by epiphytic bryophytes
Most of the retained water presumably in a tropical montane forest. Biotropica
evaporates, contributing to a high air humidity, 30:12-23.
condensation of clouds and again to rainfall. Frahm, J.-P. 1990. Bryophyte phytomass in
Bryophytes have therefore importance for a tropical ecosystems. Botanical Journal
balance of the climate and avoid drying and of the Linnean Society 104:23-33.
warming of a landscape. Frangi, J. L., and A. E. Lugo. 1992. Biomass
Beside of the phytomass of epiphytic bryophytes, and nutrient accumulation in ten-year
also the terrestrial bryophytes play an old bryophyte communities inside a
important rôle for water storage. This especially flood plain in the Luquillo Experimental
concerns subalpine forests, where the ground is Forest, Puerto Rico. Biotropica 24:106-
densely covered with bryophytes. So far, no 112.
studies of the water storage capacity of Hofstede, R. G. M., J. H. D. Wolf, and D.
bryophytes in have been performed in the tropics. Benzing. 1993. Epiphytic biomass and
It can be assumed that these bryophytes nutrient status of a Colombian upper
contribute at least the same amount if not more montane rainforest. Selbyana 14:37-45.
to the water storage. The ecological function of Jacobsen, N. H. G. 1978. An investigation into
the bryophytes on the ground is that they store the ecology and productivity of
water to a certain amount and evaporate it to the epiphytic mosses. Journal of South
atmosphere. If the bryophytes are sturated with African Botany 44:297-312.
water, additional water does not run off causing Klinge, H. & Herrera, R. 1983. Phytomass
erosion but feeds the ground water. structure of natural plant communities
A much neglected effect of epiphytic bryophytes on spodosols in southern Venezuela: the
is the uptake of nutrients. Rain water is the tall Amazon caatinga forest. Vegetatio
source of nutrients for all epiphytic and most 53: 65-84.
epigaeic bryophytes. If rain water would run off, Munshi, J. D. 1974. Seasonal changes in
also the nutrients would run off. If it is taken up standing crop and annual net production
by bryophytes, the nutrients are incorporated to of two moss communities at Varanasi.
built up organic phytomass. If this phytomass is Tropical Ecology 15:28-38.
later decomposed, the nutrients are incorporated Pentecost, A. 1998. Some observations on the
in the ecosystem. This effect concerns especially biomass and distribution of cryptogamic
epiphytic „moss balls“, which can grow to epiphytes in the upper montane forest
enormous seize and weight, and brak down from of Rwenzori Mountains, Uganda.
canopy branches when they got too heavy. Global Ecologya nd Biogeography
According to Veneklas (1982), the humus Letters 4:273-284.
produced by rotten epiphytes is in average 10% Pócs, T. 1976. The role of epiphytic vegetation
of the epiphytic phytomass (it varies between 2 in the water balance and humus
and 65%). This humus can be 2500 kg/ha (Pócs production of the rainforests of the
1976, 1980). Uluguru Mountains, East Africa.
Bossiera 24:499-503.

56 Frahm
Pócs, T. 1976. Bioclimatic studies in the Uluguru amplitude for light and humidity. Even selective
Mountains (Tanzania, East Africa). cutting of tress results in changing humidity
Acta Bot. Acad. Scient. Hungaricae 22: conditions and changing bryophyte composition.
163-183. Florschütz-de Waard & Bekker (1987) observed
Pócs, T. 1980. The epiphytic biomass and its in the Guianas that the epiphytic species in the
effect on the water balance of two plantations were the same as in scrubby
rainforest types in the Uluguru vegetation around. These species were again the
Mountains (Tanzania, East Africa). same as those from the rain forest canopy. A small
Acta Botanica Academy Scientificae tree near the ground in the open is ecologically
Hungarica 26:143-167. comparable to the canopy in 25 m elevation and
Pócs, T. 1987. Changes in the biomass and bothh are colonized by sun epiphytes. Hyvönen
productivity of bryophytes in East et al. (1987) stated that the species confined to
African rainforests. XIV Int. Bot. primary rain forests in papua New Guinea are
Congress Abstracts: 264. shade epiphytes
Thompson, S. L., D. J. Yates, and D. M. 2. Species of secondary and primary forests have
Bergstrom. 1994. Water holding different reproduction strategies. The species of
capacity of epiphytic bryophytes in primary forests have facilitiesto propagate
subtropical forests. Australian vegetatively and can survice in sterile conditions.
Bryology Newsletter 31:6. In terms of life strategies, they are stayers. After
Van Dunne, H. J. F., and M. Kappelle. 1998. a clear cut, the regrowth has to be colonized from
Biomass diversity relations of epiphy- larger distance. In terms of life stategies, these
tic bryophytes on small Quercus are colonists. The recolonizsation of the
copeynsis stems in a Costa Rican mon- secondary forest has to start from zero with dif-
tane cloud forest. Revue Biologique ferent species.
Tropical 46:217-224. Therefore secondary forests have a very diffe-
Veneklaas, E. J., R. J. Zagt, A. van Leerdam, rent epiphytic bryophyte flora. Pócs (1982)
R. van Ek, A. J. Brockhoven, and M. observed that 90% of the species of native forests
van Genderen. 1990. Hydrological did not occur in plantations. Nevertheless,
properties of the epiphyte mass of a secondary forests may have a higher diversity
montane tropical forest, Colombia. than primary forests, at least with regard to trunk
Vegetatio 89:183-192. epiphytes. Epiphytic bryophytes can be very ab-
Wolf, J. H.D. 1993. Diversity patterns and bio- undant in coffee, citrus, mango, rubber or cocoa
mass of epiphytic bryophytes and li- plantations to such an extend that they must be
chens along an altitudinal gradient in the removed. Therefore species numbers alone does
northern Andes. Annals Missouri Bot. not tell too much, the indicator value of the
Garden 80: 928-960. species must be considered. The quantity of
species is nit sufficient but the quality.
5.4 Human impact Similarly, foliicolous bryophytes can be sensiti-
ve for disturbance. However, except for human
Bryophytes are a very valuable tool for measuring made disturbances, there are also natural
disturbance of habitats. This concerns especially disturbance effects such as the effects of El Nino
the recognition of primary forests and its on the cryptogamic vegetation of Galapagos, the
differentiation from secondary forests for two effects of hurricanes in the West indies, those of
reasons: volcanic eruptions including the effects of sour
1. Secondary forests have a different structure ashes and plants and the forest floor, climatic
and micro climate. The more open structure fluctuations, especially wetter and drier periods.
favours sun epiphytes and reduces shade
epiphytes. Species of secondary forests are more
drought resistant and have a broader ecological
Equihua, C. & S. R. Gradstein 1995 .

Tab. 5.1: Phytomass of epiphytic bryophytes in kg/m2 and kg/ha in different regions of the
tropics.
Bryofloristic comparison between an

old field and a rain forest: preliminary
results. In C. Delgadillo M (ed.),
International Bryological Conference:
Tropical Bryophytes: Biology,
Diversity and Conservation. August 7-
12, 1995. Mexico City. Scientific
Program, Abstracts, Field Trips, Tourist
Tips. Instituto de Biologia, UNAM,
Mexico City.
Hyvönen J., Koponen T. & Norris D. H. 1987.
Human influence on the mossflora of
tropical rainforest in Papua New Gui-
nea , Symposia Biologica Hungarica 35:
621-629
Norris, D. H. 1990. Bryophytes in perennially
moist forests of Papua New Guinea:
ecological orientation and predictions of
disturbance effects. Botanical Journal of
the Linnean Society 104:281-291.
Romero, C. 1999. Reduced-impact logging
effects on commercial nonvascular
pendant epiphyte biomass in a tropical
montane forest in Costa Rica. Forest
Ecology and Management 118:117-
125.
Serrano, Y. 1996. The mosses of disturbed areas
in the municipality of Bayamon, Puerto
Rico. The Bryologist 99:81-84.
Sillett, S. C., S. R. Gradstein, and D. Griffin
III. 1995. Bryophyte diversity of Ficus
tree crowns from cloud forest and
Fig. 5.8: Phytomass of epiphytic bryophytes per pasture in Costa Rica. Bryologist
hectare along a transect on Mt. Kinabalu, Bor- 98:251-260.
neo. The low values around 2700 m are caused
by a forest type on ultrabasic soil. The higher
values at 20 m are caused by the nearby coast.

58 Frahm

6 ECOPHYSIOLOGY
The ecophysiology of tropical bryophytes is still phytomass of bryophytes on the one hand and
a much neglected field, because only few the light, temperature, and humidity. However,
physiologists have studied bryophytes in the it could not be explained why bryophytes did not
tropics and the classical bryologists usually did occur in the lowlands e.g. along road cuttings
no physiological work. Furthermore, the working where they as much light or more as at higher
conditions for physiologists in the tropics are not altitudes. The key to solve this problem was a
the best. An understanding of the ecological combination of factors as suggested already by
behaviour of bryophytes can, however, only be Richards (1984). Richards regarded a
reached if their physiology is known. combination of temperature and light intensity
as important, when he wrote: „Studies of
photosynthesis, respiration and net assimilation
6.1. Temperature, light and humidity rates in lowland and montane forest bryophytes
might be of great interest in this connection“.
A conspicuous effect of the bryophyte vegetation Therefore first laboratory experiments were
in the tropics is the increase of „mossiness“ with performed (Frahm 1987, 1990) to test this
the altitude. Tis effect has fascinated even hypothesis. Tropical monaten bryophytes were
botanists in the 19. century. Several attempts kept in a clima chamber and treated under
have been made to explain this effect.Increasing lowland conditions (high temperature of 30°, low
precipitation or humidity with the altitude was light intensity of 300 Lux) and montane
argued, although the precipitation in the lowlands conditions (10° and 1500Lux). The result was,
can already be quite high, decreasing that bryophytes did not gain net photosynthesis
temperatures were also introduced as argument, under lowland conditions because of high
more light intensity at higher elevations, respiration. The respiration during night time was
desiccation in the lowlands, but all arguments higher than the phososynthesis over day, resulting
could not explain this effect satisfactorily because in a negative net-photosynthesis. Similar results
they were gaps in the argumentation or were obtained by Lösch et al. (1994) with mate-
exceptions. During the BRYOTROP expedition rial collected in Zaire. Zotz et al. (1997) and Zotz
in Peru, attempts were made to clarify this (1999) could support this explanation with field
problem with ecological measurements. There experiments performed in Panama in various
were in fact good correlations between the altitudes betwen 30 and 1200 , also for folious

60 Frahm
lichens, which have a similar physiology as turgescent. Dark phases in wet state must be
compared with bryophytes. deducted from the balance because they cause
The photosynthesis of tropical montane respiration. And this is a problem with species in
bryophyte species does not differ from those from the understorey, which live under conditions of
temperate regions. The physiology nicely low light intensity over day but strong respiration
supports that these species are phylogenetically loss over night. It has so far not yet studied how
derived from temperate species. the bryophytes in the understorey of lowland
It is still an open question how the (fewer) forest can reach net photosynthesis. One possible
bryophytes in the tropical lowlands can survice explanation could be that they reach highjer rates
under these conditions, which montane species of photosynthesis from light spots shining
cannot tolerate. One hypothesis is that the low through the canopy and wandering with the sun
ligt intensity is compensated by higher ober the ground of the forest. This could explain
carbondioxide, which originates from rotten plant why bryophytes have such a scattered, uneven
material close to the ground. This would explain distribution at the bottom of the forest.
the effect that sometimes epiphylls in lowland
forests have a distinct upper limit in about 1 m. Changes of factors with the elevation.
Another possibility would be that they have dif- Within a transect from the lowland to the subal-
ferent phytochrome systems to gain more pine forest, the important factors of light,
photosynthesis under low light conditions. temperature and humidity are changing, which
Species in the outer part of the canopy gain more is the reason for the different life form spektra,
light and thus have a higher photosynthetic rate. the different growth forms, and various
They are, however, exposed to severe drying anatomical adaptations of species as well as the
especially at noon. increase of phtomass of bryophytes.
The temperature decreases constantly by 0.6°C
Changes of factors within the forest. per 100 m elevation (fig. 6.4). The mean annual
Within the forest, there is the well known distinct temperature can easily be determined in the
increase of light from the ground to the canopy tropics by measuring by the soil temperature in
(fig. 6.2). There is only 1% of the light intensity 30 cm depth.
at the bottom of lowland forests, which is The light intensity within the forest is increasing
responsible for the low number of species. With with the elevation due to lower forests with
The light intensity in the canopy is almost that of smaller leaves (fig. 6.5). This causes an increase
open places. On the other hand, the humidity is of bryophytes in the ground layer from almost
declining, being high at the bottom of the forest zero to almost 100% in the subalpine forest.
for all over the day and night and reaching even It is a myth which is propagated in many ecology
extreme ow values in the canopy at noon (fig. textbooks that the tropical rain forest has
6.3). Thus the bryophytes at the bottom have to consistent high humidity. Measurements in many
suffer from low light bu have high humidity, parts of the tropics revealed that there is a
those in the canopy have to suffer from strong decrease of the humidity at noon, which can reach
desiccation and high temperatures but have more 60% rH or less (fig. 6.6). In fact, we have an
than sufficient light. Both requires morphological incrasing air humidity with the elevation because
and anatomical adaptations. The critical value of auf the decreasing temperatures because the water
the humidity for bryophytes is 80% rH. This is contents of the air declines with the temperature.
based on observations in the field, according to Thus the air at high temperatures in the lowlands
which bryophytes dry up below 80% but stay takes up more humidity than the air at lower
turgescent above 80%. Reason is, that bryophytes temperatures in the mountains, resulting in a
can take up humidity from the air by their high decrease of the relative humidity. If the air cools
osmotic values. As poikilohydric plants, down continuously with the elevation, the
bryophytes have metabolism only in wet state, saturation point will be reached causing mist or
so long as they are turgescent. They have clouds. This is the reason that cloud forests are
photosynthesis only in light phases when found only at higher altitudes.

Fig. 6.1: Photosynthesis of Plagiomnium rhynchophorum under various temperature conditions at

1500 Lux. The montane forest conditions (5°, 15°) give a sufficient net phyotosynthesis, the lowland
condition (25°) gives photosynthesis over day but no net-photosynthesis. 35° gives no net photosyn-
thesis during day.
Fig. 6.2: Profile of light intensity in an Amazon Fig. 6.3: Profile of humidity in an Amazon low-
lowland forest (Surumoni, upper Orinoco). From land forest (Surumoni, upper Orinoco). From
León-Vargas 2001. León-Vargas 2001.
Instructions for performing measurements of tor for resistance to water stress in some
ecological factors such as temperature, humidity Nigerian species of corticolous bryoph-
and light are given in the appendix. ytes. Tropical Ecology 26:80-84.
Biebel, R. 1967. Temperaturresistenz tropischer
Urwaldmoose. Flora 157:25-30.
Akande, A. O. 1984. Anhydrobiosis in cortico- Biebl, R. 1964. Austrocknungsresistenz tropi-
lous bryophytes. Tropical Ecology scher Urwaldmoose auf Puerto Rico.
25:255-259. Protoplasma 59:277-297.
Akande, A. O. 1985. Osmotic potential - a fac- Frahm, J.-P. 1987. Which factors control the

62 Frahm
Fig. 6.4: Mean annual temperatures (derived from soil temperature) along an transect in Eastern
Congo/Uganda. The decrease is almost constant.
Fig. 6.5: Minima and maxima of relativ light intensity in percent (as compared to the situation
outside the forest) at the bottom of rain forests in Eastern Cong/Uganda. The light increases
slightly from the montane to the high montane forest and reaches maxima in the subalpine forest.

Fig. 6.6: Daily curves of temperature and air humidity in different elevations along a transect at the
E-slope of the Andes in NE-Peru. At 300 m, the temperature raises at noon to 30° but the humidity
drops down to 45%. With increasing elevation, the temperature decreases causing a higher relative
humidity. The overlapping parts of the curves show not saturated air humidity (only short time at
noon).

64 Frahm
growth of epiphytic bryophytes in tro- Phyllogonium fulgens] , Selbyana 13:

pical rainforests? Symposia Biologica 148-149.
Hungarica 35:639-648. Coxson, D. S., D. D. McIntyre & H. J. Vogel ,
Frahm, J.-P. 1990. The effect of light and tem- 1992. Pulse release of sugars and
perature on the growth of the bryophy- polyols from canopy bryophytes in
tes of tropical rain forests. Nova Hed- tropical montane rain forest
wigia 51:151-164. (Guadeloupe, French West Indies). ,
Johnson, A., and P. Kokila. 1970. The Biotropica 24: 121-133.
resistance to desiccation of ten species Nadkarni, N. M. 1983. The effects of epiphytes
of tropical mosses. The Bryologist on nutrient cycles within temperate and
73:682-686. tropical rainforest tree canopies. Ph.D.
Lösch, R., Mülders, P., Fischer, E., Frahm, J.- dissertation, University of Washington,
P. 1994. Scientific Results of the Seattle.
BRYOTROP Expedition to Zaire and Nadkarni, N. M. 1984. The biomass and
Rwanda 3. Photosynthetic gas exchange nutrient capital of epiphytes in a
of bryophytes from tropical lowland and neotropical elfin forest. Biotropica
mountain forests of eastern Central Af- 16:249-256.
rica. Tropical Bryology 9: 169-186. Nadkarni, N. M. 1986. The effects of epiphytes
Zotz, G. 1999. Altitudinal changes in diversity on precipitation chemistry in a
and abundance of non-vascular neotropical cloud forest. Selbyana 9:44-
epiphytes in the tropics - an ecological 51.
explanation. Selbyana 20:256-260. Nadkarni, N. M., and T. J. Matelson. 1992.
Zotz, G., B. Büdel, A. Meyer, H. Zellner & O. Biomass and nutrient dynamics of
L. Lange , 1997 , Water relations and epiphyte litterfall in a neotropical
CO2 exchange of tropical bryophytes in montane forest, Costa Rica. Biotropica
a lower montane rain forest in Panama , 24:24-30.
Botanica Acta 110: 9-17. 4 fig. 2 tab. Veneklaas, E. 1990. Rainfall interception and
aboveground nutrient fluxes in
6.2 Nutrients Colombian montane tropical rain forest.
Diss. Univ. Utrecht.
Akande, A. O., S. O. Olarinmoye, and A. Veneklass, E.J. 1990. Nutrient fluxes in bulk
Egunyomi. 1985. Nutrient studies of precipitation and throughfall in two
some corticolous bryophytes in Nigeria. montane tropical rainforests, Colombia.
Cryptogamie, Bryologie, et J. Ecol. 78: 974-992.
Lichénologie 6:121-133.
Coxson, D. S. 1991. Nutrient release from
epiphytic bryophytes in tropical
montane rain forest (Guadeloupe).
Canadian Journal of Botany 69:2122-
2129.
Coxson, D. S. & Vogel, H. J. , 1989. Drought
exposure and nutrient cycling; The role
of canopy ephiphytes in a tropical cloud
mist forest. , American Journal of
Botany 76(6 Suppl.): 5
Coxson, D. S. et al , 1991 , The release of sugars
and polyols from epiphytic bryophytes
in tropical montane rain forests
(Guadeloupe). [Frullania atrata,

6.3 Various topics Octoblepharum albidium Hedw. in

western Nigeria II. Phenology and
The remaining literature to various other topics water relations. Nova Hedwigia
of the ecology of tropical bryophytes is compiled 31:377-390.
here: Equihua, C. & S. R. Gradstein , 1995 ,
Akiyama, H. , 1992. Rheophytes of bryophytes old field and a rain forest: preliminary
in wet tropics of southeastern Asia. , Nat. results unpaginated tip-in. In C.
Envir. Sc. Research 5: 43-55. Delgadillo M (ed.), International
Bien, A. 1982. Substrate specificity of leafy Bryological Conference: Tropical
liverworts (Hepaticae: Lejeuneaceae) in Bryophytes: Biology, Diversity and
a Costa Rican rainforest. M.Sc. Thesis, Conservation. August 7-12, 1995.
State University of New York, Mexico City. Scientific Program,
Stonybrook. Abstracts, Field Trips, Tourist Tips. ,
Breen, R. S. 1953. Tropical mosses on Instituto de Biologia, UNAM, Mexico
limestone. The Bryologist 56:1-7. City.
Bryant, E. H., B. Crandall-Stotler, and R. E. Florschütz-de Waard, J., and J. M. Bekker.
Stotler. 1973. A factor analysis of the 1987. A comparative study of the bry-
distribution of some Puerto Rican ophyte flora of different forest types in
liverworts. Canadian Journal of Botany West Suriname. Cryptogamie, Bryolo-
51:1545-1554. gie, et Lichénologie 8:31-45.
Delgadillo, M.C. 1976. Estudio botanico y Frahm, J.-P. 1996. Diversity, life strategies,
ecologico de la region del Río origin and distributions of tropical Inselberg
Uxpanapa, Veracruz. No. 3. Los bryophytes. Anales del Instituto de Biolo-
Musgos. Publ. Inst. Invest. Recurs. gía de la Universidad Nacional Autonoma
Bioticos, 1, 2, 19-28. de Mexico 67: 73-86.
Egunyomi, A. 1978. Autecology of Frahm, J.-P., Porembski, S. 1994. Moose von
Octoblepharum albidum Hedw. in Inselbergen aus Westafrika.. Tropical
western Nigeria I. Growth and Bryology 9: 59-68.
competition. Nova Hedwigia 29: 665- Frahm, J.-P., S. Porembski, R. Seine, and W.
674. Barthlott. 1996. Mosses from
Egunyomi, A. 1979. The viability of spores of inselbergs of the Ivory Coast and
some tropical moss species after Zimbabwe. Nova Hedwigia 62:177-
longtime storage and their survival 189.
chances in nature. Journal of the Hattori Frahm, J.-P., and H. Kürschner. 1992. Moose
Botanical Laboratory 45:167-171. tropischer Regenwälder. Spektrum der
Egunyomi, A. 1984. A survey of asexual Wissenschaft 10:58-67.
reproductive disaspores in the Nigerian Fulford, M., B. Crandall, and R. Stotler. 1971.
moss flora. Journal of the Hattori The ecology of an elfin forest in Puerto Rico.
Botanical Laboratory 56:115-121. Part 15. A study of the leafy hepatic flora of
Egunyomi, A., H. J. Harrington, and S.O. the Luquillo Mountains. Journal of the
Olarinmoye. 1980. Studies on Arnold Arboretum 52:435-458.
regeneration from the leaves of O. Fulford, M., Crandall, B. & Stotler, R. 1970.
albidium Hedw. (Musci). Cryptogamie The ecology of an elfin forest in Puerto
Bryologie, et Lichénologie 1:73-84. Rico 11. The leafy Hepaticae of Pico
Egunyomi, A. 1979. Bryophytes and the del Oueste. J. Arnold Arboretum 51: 56-
Nigerian vegetation types. Misc. Bryol. 69.
Lichenol.4: 67-68. Gates, F. C. 1915. A Sphagnum bog in the
Egunyomi, A. 1979. Autoecology of tropics. Journal of Ecology 3:24-30.
Germano, S. R., and K. C. Pôrto. 1997.

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Ecological analysis of epixylic Sphagnum in Bolivia. Tropical

bryophytes in relation to the Bryology 13:65-74.
decomposition of substrate Mohamed, M.A.H. 1987. The limestone moss
(Municipality of Timbaúba- flora of Malaya. Symposia Biologica
Pernambuco, Brazil). Cryptogamie, Hungarica 35: 649-663.
Bryologie, et Lichénologie 18:143-150. Moyá, M.T. 1992. Phenological observations
Giesenhagen, K. 1910. Die Moostypen de Re- and sex ratios in Marchantia chenopoda
genwälder. Annales Jardin Botan. de L. (Hepaticae: Marchantiaceae).
Buitenzorg (suppl. 3, pt.2):711-790. Tropical Bryology 6: 161-168.
Hoffmann, A. & Riverón, R. 1992. Biorelacio- Nadkarni, N. M., A. R. Cobb, and R. Solano.
nes entre los musgos y su acarofauna 2000. Interception and retention of
en México. Tropical Bryology 6: 105- macroscopic bryophyte fragments by
110. branch substrates in a tropical cloud
Kuc, M. 2000. Adaptations of lowland jungle forest: an experimental and
mosses to anthropogenic environments demographic approach. Oecologia
in Guayana. Tropical Bryology 18: 49- 122:60-65.
54. Newton, A. E. , 1995 , A comparison of the
Linares, E. L. 1986. Estudio taxonomico y species composition of bryophyte
ecologico de la brioflora en la franja alto populations on forest and pasture trees
andina de “El Tablazo”, Cundinamarca. at La Selva, Costa Rica, p. 37. In C.
Ph.D. Thesis, Universidad Nacional de Delgadillo M. (ed.), International
Colombia, Bogotá. 308 pp. Bryological Conference: Tropical
Lisboa, R. C. L. 1976. Estudos sobre a vegetaçâo Bryophytes: Biology, Diversity and
des campinas amazonicas V. Conservation. August 7-12, 1995.
Brioecologia de uma campina Mexico City. Scientific Program,
amazonica. Acta Amazonica 6:171-191. Abstracts, Field Trips, Tourist Tips. ,
Mägdefrau, K. 1983. The bryophyte vegetation Instituto de Biologia, UNAM, Mexico
of the forests and páramos of Venezuela City.
and Colombia. Nova Hedwigia 38:1-63. Norris, D. H. 1990. Bryophytes in perennially
Marinho, M.D.G.V. & D.C. de A. Barbosa , moist forests of Papua New Guinea:
1987 , [Abstract:] Levantamento de ecological orientation and predictions of
briófitas corticicolas em Parkia pendula disturbance effects. Botanical Journal of
Benth. na mata pluvial tropical (Jardim the Linnean Society 104:281-291.
Botanico do Curado, Recife-PE) , in: Odu, E. A. 1981. Reproductive phenology of
XXXVIII Congresso Nacional de some tropical African mosses.
Botanica, Resumos. P. 106. Cryptogamie, Bryologie, and
Universidade de Sao Paulo, Sao Paulo. Lichénologie 2:91-99.
Mattila, P., and T. Koponen. 1999. Diversity Odu, E.A. 1982. Phenology of West Tropical
of the bryophyte flora and vegetation African mosses. J. Hattori Bot. Lab.
on rotten wood in rain and montane 52:283-286.
forests in northeastern Tanzania. Pôrto, K.C. 1992. Bryoflores d’une forêt de
Tropical Bryology 16:39-164. plaine et d’une forêt d’altitude moyenne
McQueen, B. C. 1991. Niche breadth and dans l’état de Pernarnbusco (Brésil). 2.
overlap of four species of Sphagnum in Analyse écologique comparative des
southern Ecuador. The Bryologist forêt. Cryptogamie, Bryologie, et
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Richards, P.W. 1935. Ecological notes on the basin, Department of Loreto, Peru.
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Richards, P. W. 1954. Notes on the bryophyte flanc méridional du Massif Sud
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sobre comunidades de briofitas en ecología de musgos y hepáticas (datos
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Schuster, R. M. 1988. Ecology, reproductive Los Nevados (Introucion y datos
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Sillett, S. C. 1991. Canopy bryophyte strategies, and morphological changes
communities of a lower montane wet of mosses along elevational and
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Bryology 5: 73-78.

7. BRYOSOCIOLOGY
Like all other plants, also bryophytes are growing Bryophyte associations have also been described
together in distinct compositions of species, mainly from Europe, rarely also from Japan, and
which are called associations. These are only few attempts were made to describe
composed of species with similar habitat bryophyte associations in the tropics (Akande &
preferences. The species composition is very Olarinmoye 1982, Augier 1972, 1974, Giacomini
regular and typical. The name of an association & Cifferi 1950). In part, these „associations“ or
implies a combination of numerous species and „communities“ were not described in a
allows to characterize certain habitats, altitudes phytosociological way (Griffin 1974, Jovet-Ast
and climatic conditions. As species can be 1949, Miller 1954). Even for the German
grouped by similarity to genera, families and BRYOTROP project, first approaches to
orders, also associations can be grouped in a determine the epiphytic bryophyte vegetation in
hierarchial system. Peru included only descriptive and statistical
This field called plant sociology has been methods (Frahm 1987). (The tree trunks were
developed in Europe and applied mainly in wrapped with plastic, the patterns of bryophyte
Europe, later also in comparable regions (Japan). specis was copied with colour ink pens on the
In North America, different techniques in the way plastic, the cover was calculated and the affinities
of vegetation analysis and statistics is used between the species calculated by Chi-Square
instead. They have no standard methods for the tests). Later, the work of Kürschner (1990a,
field, for tbale work and for evaluation have no 1990b, 1995a, 1995b) in Borneo and Zaire
classification and therefore do not allow to revealed that distinct epiphytic bryophyte
compare plant communities from different associations can be distinguished also in the
regions. tropics, which are characteristic for different
For a long time it was supposed that the altitudes. Therefore Kürschner & Parolly (1998)
European plant sociology technique can not completed the analysis of tropical epiphyte
successfully be applied in the tropics. This may studies in Peru and could (Kürschner & Parolly
still concern certain types of rain forests with a 1999) show that worldwide in the tropics there is
high diversity of trees. First attempts in the a system of comparable epiphytic bryophyte
páramo vegetation of Colombia (Cleef 1981) and communities. This is based on the fact that species
later studies by Mrs and Mr Miehe (plant of the same genera, in some cases even the same
geographers from the university of Marburg, species, are growing in comparable elevations in
Germany) in tropical African mountains revealed the tropics, forming vicariant communities
that these methods can be used in the tropics quite (Kürschner & Parolly 1998, 1999). Although this
well with the result that vegetation units can be methods was either ignored or heavily criticized
typified and classified.

70 Frahm
by North American botanists, it has proved to be Giacomini, V. & Ciferri, R. 1950.

very succesful and useful. Meanwhile, it has also Un’associazione crittogamica a
sucessfully be used to determine bryophyte Polytrichadelphus e Cora (Coreto-
communities on tree fern trunks in SE-Brazil Polytrichadelphetum ciferrii) su rocce
(Ahmed 2002) and also be used very successfully della Foresta delle nebbie in
in temperate rain forests in New Zealand and Venezuela. Atti Ist. Vot. Univ. Pavia,
Chile. Ser. 5, 9: 211-217.
The importance of phytosociology is based on Gill, L. S., and H. I. Onyibe. 1986.
the fact that plant species cannot be seen isolated. Phytosociological studies of epiphytic
They grow - determined by ecological factors - flora of oil palm (Elais guineensis) in
together with other species and even animals (in Benin City, Nigeria. Feddes Repert.
biocoenoses). Therefore changes of ecological 97:691-695.
parameters (altitude, disturbance) affect not sin- Griffin III, D. 1974. Observations on bryophy-
gle species but associations. Furthermore, the te community of the Poas Volcano,
name of an association represents and symbolizes Costa Rica. Miscellanea Bryologica Li-
a complex of species with all its determining chenologica 6:174-175.
ecological factors and characterizes a certain Jovet-Ast, S. 1949. Les groupements des
habitat. muscinées épiphytes aux Antilles
To promote this method in the tropics, a short françaises. Revue Bryologique et
instruction of the field technique is given in the Lichenologique 18:682-686
appendix. Kürschner, H. 1990a. Die epiphytischen Moos-
gesellschaften am Mt. Kinabalu (Nord-
Borneo, Sabah, Malaysia). Nova Hed-
Ahmed, J., Frahm, J.-P. 2002. Moos- wigia 51:1-76.
gesellschaften auf Baumfarnstämmen in Kürschner, H. 1990b. Höhengliederung (Ordi-
Südostbrasilien. Tropical Bryology 22: nation) von epiphytischen Laub- und
135-178. Lebermoosen in Nord-Borneo (Mt.
Akande, A. O., S. Olarinmoye, and A. Kinabalu). Nova Hedwigia 51:77-86.
Egunyomi. 1982. Phytosociological Kürschner, H. 1995a. Epiphytic moss
studies on some corticolous bryophytes associations in the Eastern Congo basin
in Ibadan, Nigeria. Cryptogamie, and the adjoining highlands (National
Bryologie, et Lichénologie 3:235-248. Park Kahuzi-Biega/Zaire, Nyungwe
Augier, J. 1972. Groupments de bryophytes Forest/ Rwanda): Scientific results of
terricoles sur le campus universitaire de the BRYOTROP-expedition to Zaire
Yaounde. Annls. Fac. Sci. Univ. Fed. and Rwanda. No. 4. Nova Hedwigia
Cameroun 9: 73-85. 61:1-64.
Augier, J. 1974. Bryophytes corticoles a l’etage Kürschner, H. 1995b. Wissenschaftliche Ergeb-
submontagnard dans l’ouest nisse der BRYOTROP-Expedition nach
camerounais. Annls. Fac. Sci. Univ. Zaire und Rwanda. 5. Höhengliederung
Fed. Cameroun 17: 67-93 epiphytischer Moose im östlichen Kon-
Cleef, A.M. 1981. The vegetation of the páramos gobecken und den angrenzenden Ge-
of the Colombian Cordillera Oriental. birgsstöcken (Parc National de Kahu-
Dissertationes Botanicae 61. zi-Biega/Zaire, Forêt de Nyungwe/
Fiedler, P.L. 1984. Preliminary observations on Rwanda). Tropical Bryology 11:77-86.
the structure of a neotropical cryptogam Kürschner, H., and G. Parolly. 1998. Life
community. Brenesia 22: 85-93. strategies of epiphytic bryophytes in
Frahm, J.-P. 1987. Struktur und Zusammen- rain forests on the Andean slope in the
setzung der epiphytischen Moos- Amazonian lowland in Northern Peru.
vegetation in Regenwäldern NO-Perus. Nova Hedwigia 67:1-22.
Beih. Nova Hedwigia 88: 115-141. Kürschner, H., and G. Parolly. 1998. Shade

Fig. 7.1: Plant sociological table composed from different vegetation analyses in a montane rainfo-
rest in Venezuela (Monte Zerpa, Mérida). The data were elaborated by students during a course on
tropical bryology. The table shows that the epiphytic bryophyte vegetation in this forest can be
attributed to one association with several constant species as well as two subassociations reflecting
different host trees.

72 Frahm
epiphytic bryophyte communities on the tropical montane rainforest in the nort-

eastern slope of the Andes and in hern Andes I. Lower montane commu-
Amazonian lowlands of Peru. Nova nities. Phytocoenlogia 22:1-52.
Hedwigia 66:1-87.
Kürschner, H., and G. Parolly. 1998.
Syntaxonomy of trunk-epiphytic
bryophyte communities of tropical rain
forests: A first pantropical approach.
Phytocoenologia 28:357-425.
Kürschner, H., and G. Parolly. 1999.
Pantropical epiphytic rain forest
bryophyte communities – Coeno-
syntaxonomy and floristic-historical
implications. Phytocoenologia 29:1-52
Miller, H.A. 1954. Field observations on asso-
ciations of Hawaiian mosses. Bryolo-
gist 57: 167-172.
Petit, E., and F. Symons. 1974. Les bryophytes
des bois artificles de Cupresses et
d’Acacia au Burundi. Analyse
factorielle de la végétation
bryophytique. Bulletin Jardin Botanique
National e Belgique 44: 219-247.
Sastre-De Jesús, I. 1992. Estudios preliminares
sobre comunidades de briofitas en
troncos en descomposición en el bosque
subtropical lluvioso Puerto Rico.
Tropical Bryology 6:181-192.
Sillett, S. C. 1991. Canopy bryophyte
communities of a lower montane wet
forest in Costa Rica: Quantitative
sampling in intact forest and isolated
trees. M.Sc. thesis, University of
Florida, Gainsville.
Volk, O.H. 1984. Pflanzenvergesellschaftungen
mit Riccia-Arten in Südwestafrika (Na-
mibia). Vegetatio 55: 57-64.
Went, F.W. 1940. Soziologie der Epiphyten
eines tropischen Urwaldes. Annales
Jardin Botanica de Buitenz 50:1-98.
Wolf, J. H. D. 1989. Comunidades epífitas en
un transecto altitudinal en la Cordillera
Central, Colombia: datos inciales sobre
la cantidad de espescies de briófitas y
líquenes. In van der Hammen, T., S.
Diaz-Piedrahita, and. Alvarez (eds.).
Studies on Tropical Andean Ecosy-
stems, 3, pp. 455-459. J. Cramer, Va-
duz.
Wolf, J. H. D. 1993. Epiphyte communities of

8. BRYOGEOGRAPHY
Each phytogeographical analysis must be number as in the Andine countries over a much
unsufficient at present due to unsufficient larger area, and there are 1248 species in the
botanical exploration. It has to be considered whole North America north of Mexico.
that botanical exploration in the last century was Compared with a number of 45-55.000 species
almost confined to the surroundings of cities, later of flowering plants in Colombia, bryophytes
along roads, and they were usually not carried count only for about 4% of the species number
out systematically. At present it is confined to of flowering plants, which makes them more
the availablity of roads, however, even such a easily usable.
large area as the Chocó in western Colombia is
accessed by only two roads. There are so far no The basis for any diversity studies are checklists!
systematical explorations in the tropics such as It is also the basis for any bryological studies in
the grid mappings in Europe, not even on the the tropics to know what is known from a region
basis of the UTM grid (50x50 km). or country. Therefore checklists are not only
Another problem is the lack of monographs and useful but essential. A list of checklists from
revisions and the high numbers of taxonomic tropical countries is given in the appendix.
synonyms. The knowledge about the bryofloras Because checklists are lacking for many countries
of the tropics has been greatly increasing over (at least actual ones), instructions how to compile
the last decades. This can be exemplified by the a checklist are also given in the appendix.
species of mosses which were known from Peru
in different years: The oldest bryogeography textbook was written
1951: 345 by Herzog (1924). It included chapters on the
1975: 568 autecology (still valid), types of ranges and
1985: 834 descriptions of floristic realms , which are
1987: 903 outdated in many respects. This is result of the
1992: 889 (Menzel 1992) increased knowledge of tropical bryophytes.
The recent decrease of numbers of species is due Example: Herzog knew only about disjunctions
to the results of revisions and monographs. between Aouth America and Africa on the genus
In Colombia we know at present a little less than level; this is a result of old taxonomist (e.g. Carl
1000 species. We can estimate that a similar Müller), who described (for religious reasons)
number of mosses as in Colombia or Peru (900- thesame species from different parts of the tropics
1000) is present in Venezuela. under different names or bryologists having
For comparison, we have 965 species of mosses relatively uncritically described bryophytes from
in Canada, that means approximately the same all over the world (Brotherus, Dixon, Thériot,

74 Frahm
Cardot etc.). Worldwide (!) revisions in the past Jungermanniaceae 25%

decades revealed, however, an increasing number Plagiochilaceae 55%
of species identical in the Neotropics and Africa Schistochilaceae 74%
(and other parts of the world). Furthermore, Her- Frullaniaceae 61%
zog did not consider (as all contemporary Metzgeriaceae 28%
colleagues) aspects of the plate tectonics, Herbertaceae 0%
although Wegener´s continental drift theory was Cephaloziaceae 0%
published already 10 years earlier but was not Pallaviciniaceae 0%
accepted at that time. Marchantiaceae 0%
A modern, long but also lengthy and eloquent
overlook of the phytogeography of bryophytes In total, 168 out of 440 species of hepatics are
is found in Schuster (1983). It has, however, kept endemic in Western Melanesia (38.2%) - more
in mind that phytogeographical „explanations“ than 50% occur in the montane forest belt.
based on the shape of ranges are always
hypothetical. Only recently proofs of these Piipo, S. 1994. Phytogeography and habitat
hypotheses are possible by molecular studies, e.g. ecology of Western Melanesian
by determination of genetic distances between endemic Hepaticae. J. Hattori Bot. Lab.
disjunct populations. 75: 275-293.
Piippo, S., T. Koppen, and D. H. Norris. 1987.
Herzog, Th. 1924. Geographie der Moose. Fi- Endemism of the bryophyte flora in
scher, Jena 439 pp. New Guinea. Symposia Biologica
Schuster, R.M. 1983. Phytogeography of the Hungarica 35:361-372.
bryophyta. Pp. 463-626 in R.M. Schu-
ster (ed.) New Manual of Bryology,
Nichinan. 8.1.2 Relics
Relics are called species with former continuous

8.1. General tropical bryogeography range, which survived in isolated situations.
(with contributions by T. Pócs) Reasons for the isolation are usually climate
changes. As compared with flowering plants,
8.1.1 Endemism there are relatively many examples of relics
because of the fact that bryophytes are able to
The rate of endemism is much higher in the survive in small ecological niches, e.g. rock
tropics than in the extratropics but always much fissures.
lower than endism of vascular plants: There are several cases of relics:
1. Species with tropical affinities in the temperate
Vasc. Bryoph. zones.
Galapagos Islands 50% 10% Examples are Leptoscyphus cuneifolius,
Cuba 50% 12% Adelanthus decipiens, A. lindenbergianus,
Kilimanjaro 6% Teleranea nematodes, Lepidozia cupressina,
Usambara Mtns. 3% species which are found in the Neotropics, in the
Réunion 9% African mountains as well as in the highly oceanic
Mauritius 6% parts of Europe. Some species are found in the
neotropics, on the Makaronesian Islands as well
The rate of endemism varies in different as in western Europe such as Marchesinia
taxonomic groups, e.g. in Western Melanesia mackaii, Harpalejeunea molleri,
(Piippo 1994): Drepanoclejeunea hamatifolia, Colura
calyptrifolia, Jubula hutchinsiae, Plagiochila
Lepidoziaceae 27% bifaria amongst the hepatics (mainly
Lejeuneaceae) and Campylopus shawii,

Leptodontium flexifolium and others amongst the the species as in Patagonia are found through the
mosses. Some tropical montane species are Andes (Lepyrodon tomentosus), in other cases
confined to the hyperoceanic parts of Europe (e.g. this accompanied by speciation into new
the Lejeuneaceae), others occur also in varieties, subspecies (Monoclea gottschei ssp.
suboceanic regions (Campylopus fragilis, C. gottschei and ssp. elongata) or even species (e.g.
flexuosus), others also in dry regions Chorisodontium). The speciation of complexes
(Campylopus pilifer). Some species with main of tropical species within some genera can be
occurrence in the tropics are found in the southern explained from subantarctic anchestors.
Alps (e.g. Braunia alopecura), others
(Calymperes erosum, Trematodon longicollis) 8.1.3 Disjunctions
around fumaroles in Italy.
It is argued that these species had a wider range Disjunct ranges can include (parts of) continents,
in Tertiary, including Europe, and survived in islands or mountains. Reasons for disjunct
Europe the glaciations of the Quaternary along occurrences include continental drift, formation
the coast of the Atlantic Ocean. (Since the sea of mountains by uplifting or volcanism,
level was 170 m lower at that time, presumably formation of islands by plate tectonics and
in regions which are now inundated.) In situ volcanism. Disjunct ranges can be old (plate
survival in the southern Alps is unlikely because tectonics) or young (colonization of neovolcanic
the habitats of these species were glaciated, if it islands e.g. Galapagos islands) by long distance
is not assmed that they are „migration relics“ annd dispersal. It is difficult to decide whether a
survived their in other places. disjunct range is a result of long distance dispersal
or a relictic occurrence. Long distance dispersal
2. Temperate, boreal and arctic species in the is only possible in air streams at high altitude.
tropical mountains. Spores must therfore be resistent to frost and UV-
This concerns especially the New World, where radiation. It is the merit of van Zanten (1976,
a continuous mountain range from Alaska to 1978) (see also van Zanten & Gradstein 1987,
Tierra del Fuego allows migration of species in 1988, van Zanten & Pócs 1981) having tested
different altitudes from North to South and vice the ability of spores of species for long distance
versa. See also Gradstein & Vana (1987, 1994). transport in the lab and in vivo (by carrying spores
In Africa, some temperate and mediterranean attached to a plane). Long distance dispersal can
species are found on mountains, usually above be assumed for weedy species (e.g. along
3000 m. An explanation could be that these roadside banks) and for alpine or subalpine
species have moved wouth during Quaternary, species, who can easily be dispersed around the
when Europe was glaciated and all vegetation world, especially in the innertropics with the
belts were shifted southwards. During innertropical convergence (ITC, jetstream). Note:
interglacial times and after the last glaciation, the ITC goes from East to West. A spore released
these species stayed in the tropical mountains in the Andes cannot be transported across the
were they found appropriate ecological Atlantic Ocean but across the Pacific Ocean.
conditions in the according elevations. The mean Tramnsport is only possible from Africa to South
annual temperatures of the localities of these America.
species in the tropics resemble those in Europe Types of disjunctions are
(Frahm 1994). A. in the tropics:
- pantropical disjunctions
3. Austral and subantarctic species in the tropics - Neotropics - tropical Africa (see 7.1.3.1)
By the same way (migration through mountain - tropical Africa - tropical SE-Asia (see 7.1.3.2)
chain in South America, mountain hopping in - tropical Asia - Oceania - South America
Africa, island hopping in SE-Asia), subantarctic B. in the extratropics
species were able to invade the tropical - amphipacific
mountains. This is especially the fact for South - amphiatlantic
America with its ideal conditions. In some cases, - gondwanalandic

76 Frahm
bipolar disjunctions dans l‘hemisphère austral distribution de

Beside there are some worldwide scattered quelques hépatiques tropicales. ,
disjunctions, which are hard to explain. Mémoires de la Société de
Examples: Scopelophila cataractae with Biogéographie sér. 3, 4: 95-110.
occurrence in Japan, Java, Bolivia, Mexico, Van Zanten, B.O. 1976. Preliminary report on
scattered localities in N-America. Brothera leana the germination experiments designed
occurs in disjunct in North America and east Asia to estimate the survival chances of moss
but was found in malawi (Pócs 1993). spores during aerial transoeceanic long-
Fragmented ranges can be either relics of range dispersal in the Southern Hemi-
formerly continuous ranges and local extinction sphere, with particular reference to New
or occasional long distance dispersal. An Zealand,Journal of the Hattori Botani-
important and often not considered factor in cal Laboratory 44:455-482
phytogeography is chance. Chance can play a rôle Van Zanten, B. O. 1978. Experimental studies
in all irregularities. on trans-oceanic long-range dispersal of
moss spores in the Southern hemisphe-
Frahm, J.-P. 1994. Scientific Results of the re. Journal of the Hatttori. Botanical
BRYOTROP Expedition to Zaire and Laboratory 44:455-482.
Rwanda 1. The ecology of epiphytic Van Zanten, B. O., and S. R. Gradstein. 1987.
bryophytes on Mt. Kahuzi (Zaire). Tro- Feasibility and long-distance transport
pical Bryology 9: 137-152. in Colombian hepatics, preliminary
Gradstein, S. Rob & Vana, Jiri , 1987. On the report. Symposia Biologica Hungarica
occurrence of laurasian liverworts in the 35: 315-322.
Tropics , Memoirs of the New York Van Zanten, B.O. van & Gradstein, S.R., 1988.
Botanical Garden 45: 388-425 Experimental dispersal geography of
Gradstein, S. R. & J. Vana, 1994. A boreal neotropical liverworts. Beih. Nova Hed-
bryophyte community in a tropical wigia 90: 41-94
montane forest of Mexico. , Tropical Van Zanten, B. O., and T. Pócs. 1981.
Bryology 9: 31-34. Distribution and dispersal of
Pócs, T. 1982. Examples of the significance of bryophytes. Advancs in Brylogy 45:
historical factors in the composition of 479-562.
bryofloras. Beih. Nova Hedwigia
71:305-311
Pócs, T. 1993. Brothera leana (Sull.) C. Muell.,
a Laurasian species in tropical Africa. 8.1.3.1 Disjunctions Neotropics - tropical
Bull. Jard. Bot. Belg. 62: 221-224. Africa.
Schofield, W.B., Crum, H.A.,1972. Disjunctions
in Bryophytes,Ann. Missouri Botanical At present, 4103 species of mosses are known
garden 59:174-202 from the neotropics (Delgadillo et al. 1995) and
Schofield, W.B.,1974. Bipolar disjunctive 2939 species of mosses from tropical Africa (O’
mosses in the southern hemisphere, with Shea 1995). That means we have about 25 %
particular reference to New more species in the neotropics.
Zealand,Journal of the Hattori Botanical At present, 334 species of mosses are known to
Laboratory 38:13-32 occur in the neotropics and tropical Africa. This
Schuster, R.M.,1969. Problems of antipodal dis- is 8% of the neotropical mossflora (Delgadillo
tribution in lower land plants,Taxon 1993). The number will certainly increase in the
18:46-90 future with a better exploration of the tropics and
Sharp, A. J. 1938. Tropical bryophytes in the a better knowledge. For instance, during the 4
southern Appalachians. , Ann. Bryol. weeks of fieldstudies during the BRYOTROP
11: 141-144 project in Zaire and Rwanda, many species were
Tixier, P. , 1994 , Contribution à la biogéographie even found new to Africa, most of which were

Fig. 8.1: Range of Campylopus nivalis, example of an African-American alpine disjunction.
Fig. 8.2: Range of Campylopus savannarum, example of an African - American loewland disjunc-
tion.

78 Frahm
known from the neotropics before. moose. In: Fittkau et al. (eds.), Biogeo-
Part of the species common in the neotropics and graphy and Ecology in South America
Africa can be dispersed by spores. This may pp. 562-582. W.Junk, The Hague
especially count for the species in the high mon- Ochyra, R., Kempa, R., Buck, W.R. 2000.
tane and alpine belt (fig. 8.1). However, since Plagiothecium lucidum (Hook. f. &
the main wind currents go from E to W in the Wils.) Paris in tropical Africa. Tropi-
tropics, it must be assumed that the species from cal Bryology 18: 147-152.
tropical Africa reached the Andes, which is most O’ Shea, 1995. Checklist of the mosses of sub-
unlikely, since the Andes can be regarded as saharan Africa. Trop. Bryol. 10: 91-
centers for evolution of tropical montane and 198.
tropical alpine species. In this case, a spore Reese, W. D. , 1985 , Tropical lowland mosses
dispersal from the Andes westwards („around the disjunct between Africa and the
world“) to Africa must be assumed. Americas, including Calyptothecium
Another part of the species identical in tropical planifrons (Ren. & Par.) Argent, new
Africa and the neotropics is probably a remnant to the Western Hemisphere. Acta
of the time (in Jurassic) when the continents were Amazonica, Suppl. 15(1-2): 115-121.
connected (fig. 8.2). This means they are at least
135 mio years old.
This concerns species: 8.1.3.2 Disjunctions tropical Africa - tropical
- with a disjunction between Brazil and E-Africa SE-Asia
- species of tropical lowlands, since tier spores
cannot easily get into higher air currents, and The relations between the tropical African and
especially SE-Asian bryofloras have been calculated by
- species which are sterile or produce spores not Pócs (1976, 1992). It shows that such calculations
capable of long distance dispersal. Examples are: depend on the intensity of fieldwork and
Archidium, e.g. the almost pantropical A. taxonomic studies. The first account (Pócs 1976)
ohioense with a spore diameter of 250 µm; reported 35 species of liverworts and 73 species
Campylopus carolinae, occurring in the Cerrado of mosses with this type of disjunction. The new
regions and Brazil, two localities in Rwanda and investigation (Pócs 1992) raised the number to
recently found in S Africa, which has lived almost 70 viz. 108. Beside, there are also vicariant taxa
buried in sand and has capsules hidden in the (genera and species) in both parts. Interestingly,
perichaetial leaves. the affinity of the bryoflora of the neotropics and
There are also vicariant taxa or subspecies in both tropical Africa is less close than between tropical
continents showing that the mossflora of tropical Africa and SE-Asia. There are 70 species of
Africa and the neotropics had a common origin hepatics in common between tropical Africa and
and separate evolution took place in each SE-Asia but only 52 between tropical Africa and
continent. tropical America. This result seems to support
the concept of the Palaeotropics also in
Delgadillo, C.M. 1993. The Neotropical - bryophytes. It has be kept in mind, however, that
African disjunction. The Bryologist all such calculations are preliminary due to the
96:604-615. unsufficient exploration of the tropics (and thus
Delgadillo, C., Bello, B. & Cárdenas, A. 1995. a good motivation for such studies). The reasons
LATMOSS. A catalogue of neotropical for this type of disjunction has not been studied
mosses. 191 pp. Missouri Bot. Garden. yet. Whereas South America and Africa share a
Frahm, J.-P. 1982. Grossdisjunktionen von common geological history, the situation in SE-
Arealen südamerikanischer Campy- Asia, which is split into many islands, is more
lopus-Arten. Lindbergia 8:36-45. complicated. Tropical wind systems can disper-
Grolle, R.,1969. Grossdisjunktionen in Arten- se bryoohytes only from SE-Asia to Africa and
arealen Lateinamerikanischer Leber- not vice versa.

Pócs, T. 1976. Correlations between the tropical Tan, B.C. 2002. The affinity of Moss Floras of
African and Asian bryofloras, I. Journal Japan, Taiwan and the Philippines
of the Hattori Botanical Laboratory revisited: Old problems, new insight and
41:95-106. more questions. Acta Phytotax. Geobot.
Pócs, T. , 1990. Correlation between tropical 53: 77-84.
African and Asian bryofloras. II , p 8
in: T. Koponen & J. Mänttäri (eds),
Congress of East Asiatic Bryology, 8.1.4 Human influence
Programme and Abstracts, University of
Helsinki, Helsinki. In contrast to Europe or North America, or parts
Pócs, T. 1992. Correlation between the tropical of the southern hemisphere such as New Zealand,
African and Asian bryofloras. II. where introductions of bryophytes are more often
Bryobrothera 1:35-47. recognized, an introduction of species in the
tropics is less often observed but probably not
less frequent. For example, Campylopodium
8.1.3.3 Disjunctions Tropics - Extratropics medium, a species widespread in E-, SE- and
austral Asia has several times been found on
The floristic contrast between the tropics and the Puerto Rico.
extratropics countries is much higher as between Beside, many distribution patterns cannot
tropical regions, but much less as in satisfactorily be explained. This conv´cerns for
phanerogams. For instance, 142 species of example the genus Cinclidotus, a genus of aquatic
mosses are in common between Europe and species endemic to Europe and the Near East, of
tropical Africa and 146 species are in common which C. aquaticus was undoubtedly found near
between Europe and tropical America (Frahm Valdivia, Chile, and C. fontinaloides in the
1995). Except for 10% cosmopolitan species, the Central African mountains. Thus we can offer
species common in Europe and the tropics are explanations for many types of distribution (of
found in the tropics in (upper) montane regions, which some may be erroneous) but not for all.
where they have the same growth conditions as
in Europe (e.g. the same mean annual
temperature). 8.2 Regional bryogeography
There are also few examples of tropical species
in temperate regions, for example 8.2.1 Central and South America
Heterophyllium affine from northern South and
Central America, which was found several times The austral element
between 1820 and 1860 in Central Europe The oldest elements in the bryoflora of South
(caused by spore dispersal by an volcanic America is the austral element, which is confined
eruption?). Hyophila involuta, a pantropical to southern Chile and southern Argentina. These
species from wet limestone, occurs in central regions share e.g. about 200 species of mosses
Europe along the shores of some lakes (which with the bryoflora of New Zealand (Blöcher &
were, however, glaciated until 13.000 years ago). Frahm 2002). This austral plant realm covers
The affinities between the bryophyte flora of the southern South America, southern Africa,
austral regions and tropical regions have not southern Australia, Tasmania and New Zealand.
yet been calculated, wlthough the austral region All these regions were still connected 100 mio
is considered as important phyolgenetic origin years ago and formed the southern coast of the
for the tropical bryoflora. Gondwana continent. Important is, that - even
after separartion of this continent, these regions
Frahm, J.-P. 1995. Correlations between the remained in perhumid climates, providing a
European, tropical African, and tropi- climatic consistency and also a floristic
cal American moss floras. Fragmenta consistency. Although many species may have
Floristica et Geobotanica 40:235-250.

80 Frahm
a genetic exchange by long distance dispersal confined to the Jungermanniales, showing that
through the westwind drift (roaring forties), there the Metzgeriales are the more ancient element.
are species which are not enabled to long distance There are over 38 genera of Jungermanniales
dispersal because of lacking tolerance agisnst endemic in the neotropics., often highly
freezing or UV-radiation. These species can be apomorphic or reduced (even confervoid or
interpreted as remnants of the gondwana flora. thalloid, or neotenic). They are typically
The austral element in South America has an stenotypic with only 1-3 species each and are
importance for the andine element in the way that found in the upper montane zone to the páramo
the austral species were able to migrate through (Schuster 1990). These endemic genera are
the Andes to Central America. derived from cool-Gondwanalandic suborders.
Part of the endemics are found in the Guyana
shield, a very ancient and partially isolated region.
The Neotropics Principally, the floristic elements of bryophytes
As shown by fossils in Dominican amber, the are more or less the same as those of flowering
basic stock of bryophyte species in the neotropics plants. There are, however, differences in the way
has probably originated in Tertiary as a result of that many bryophytes are more easily distributed
long lasting periods with tropical temperatures. and that extinctions by climatic events can be
Except for climate changes during the balanced by subsequent dispersal. Evaluations of
Pleistocene, to which the species could at least types of ranges have not yet been made on a broad
partially adapt by migrations, this climate base. Therefore ranges of species of Campylopus
persisted until today. Since the neotropic realm were used (fig. 8.3). This species rich genus can
is situated in between the holarctic and the aus- serve as example and case study. In general, we
tral plant realm, it has been invaded by elements can distinguish between the following floristic
from these both realms by a floristic pathway elements:
provided by the cordillera strechting from Alaska
to Tierra del Fuego. The floristic interchange in Neotropical element
Central America has been treated by Delgadillo These species are widespread from Mexico to
(1987a,b, 1988, 1992, 1995). The laurasian southern Brazil, including the Carribbean, part
element in the neotropics were treated by Grads- of them extending to SE- North America (Gulf
tein & Vana (1987, 1994). There are also coastal plain, especially Florida, southern
neotrpüocal bryophyte species found along the Alabama), where they might have been
gulf coastal plain of North America and Florida introduced by hurricanes. A part of them are not
(Pursell & Reese 1970) as well as in the southern found in the Amazon lowlands since they are
Appalachians and the Ozark Mtns. in Missouri submontane to high-montane. Species of this
(Sharp 1984), which may be interpreted as reöicts distribution are present in Dominican amber with
from the tertiary, which survived there the an age of about 20 mio years. Others are not found
glaciation periods. in the Andes, because they are lowland species.
Thanks to a compilation of species of mosses and This is the most floristic ancient element. Th
their distribution (Delgadillo 1992), their lowlamnd flora is found in rhe Guianas and the
phytogeography could be evaluated. In total, Amazon basin. More than 80% of the hepatic and
there are about 4050 species of mosses in the moss flora of the Guianas belongs to this element.
neotropics. Accordingly, the rate of endemism is with 2.5%
North and South America share about 675 species very low (Gradstein et al. 1990). Some of the
of mosses, which is 16.7 % of the neotropical species are also found in tropical Africa and it is
moss flora, either with a continuous range or a an open question whether they are relicts of a
disjunct distribution (Delgadillo 1995). The former closed range (and in this case of mesozoic
disjunct distribution pattern of 118 species may origin) or distributed by long distance dispersal.
have originated by tectonic or climatic changes A mesozoic origin is supported by the fact that
or by long distance dispersal. these are often species from dry habitats
Within the liverworts, endemism is almost (savannahs, cerrados).

A B
C D
Fig. 8.3.: Distribution patterns in the Neotropics based on species ranges of the genus Campylo-
pus. A. andine, B. SE-Brazilian, C. caribbean, D. neotropic.

82 Frahm
An important impact on the present distribution in Campylopus carolinae, which occurs also
may have had the climatic fluctuations during along the coastal plains of SE North America.
the Quaternary. During the cold periods, the This is remarkable insofar as there was never a
savannahs expanded and the rain forests areas landbridge between the regions. Some of the rain
were retreated to small isolated parts, in which forest species show disjunctions with E Africa
the rain forest flora and fauna survived. This or even India (e.g. Campylopus controversus).
effect was postulated by the German ornithologist They show relations to the austral flora and are
Haffer, who introduced the „refuge theory“ based remnants of a gondwana flora, when SE-Brazil
on the distribution of endemic birds and centers was situated beside E-Africa, Madagascar and
of diversity of birds. It was later confirmed by India along the south coast of the gondwana
palaeoclimatological data, applied by continent. There are still species which also occur
phanerogamists and even applied in tropical in E-Africa or Madagascar (e.g. Campylopus
Africa and Australia, but not yet really used in controversus, even in Sri Lanka), which must
bryogeography except an attempt made by by already have been present when the south Atlantic
Frahm (1990). These (about 8) refuge areas are had just been opened and there was a continuous
„hot spots“ for biodiversity of other organisms, shoreline from Brazil to Madagascar, about 135
because many species were not able to expand mio years ago. Other species are represented with
their ranges after the cool period with the rain vicariant subspecies in both regions (Campylopus
forest, which possibly also concerns bryophytes. trachyblepharon with ssp. trachyblepharon and
One of these „hot spots“ for phanerogamists is ssp. comatus, C. julaceus with ssp. julaceus and
the Chocó region at the pacific slope of the Andes ssp. arbogasti, interestingly all species from costal
of Columbia. The rain forests species shall here sand habitats), showing that there was at least a
be trapped between the ocean and the alpine by small separate evolution by isolation within this
the uplift of the Andes, causing a high rate of long time span. There are more examples,
endemism in plants, birds and butterflies. It is however, the trend to recognize every taxon on a
also the place with the highest precipitation in species level wipes out any chance to demonstrate
the neotropics (12-15 m/yr). Only few bryophytes the evolutionary relations by infraspefecific
were known from that region (31 species in the categories. This flora persistet in humid climatic
Dept. Chocó). A analysis based on floristic conditions since that time. It is today isolated in
studies in 10 hectar plots in various altitudes the north by arid cerrado regions, to the east by
financed by the National Geographic Society lowlands from the Andes and to the south by the
revealed that 60% of the mosses are widely arid steppes from Patagonia, which explains its
neotropical in distribution, 12% are pantropical rate of endemism (which has not yet been
to cosmopolitan, 6.6% are andine, 15% are meso- calculated for all bryophyte species from that
american, which come down from the coast of region).
the Caribbean and the darien gap (Frahm 1994).
Only three recently described (all epiphytic) Caribbean element
species of mosses are endemic to that TheWest Indies have strong continental affinities,
phytogeographical regions, in contrast to the high Originally it was thougt that the floristic affinities
rates of endemism of flowering plants. This are strongest with central America (Crosby 1969),
outlines that bryophytes are genetically more but a database founded analysis showed that the
conservatice and are not evolving as fast as the affinities are strongest with South America
flowering plants (which are almost 200 mio years (Delgadillo 1993). Crum & Steere (1958) and
younger). Steere (1984, 1985) stated that the moss flora of
the West Indies, especially of the Greater Antilles,
is more closely related to that of the sandstone
Species confined to SE-Brazil and granitic mountains of Venezuela, the Guianas
These are either dry-adapted species of the and eastern Brazil than to the floras og more
cerrado regions or species of the coastal calcareous mountains and high plateaus of
rainforests. A remarkable disjunction is found northern South America, Central America,

Mexico and the southern Appalachians of z´the from Costa Rica to Ecuador, others are even
United States (Delgadillo 1993). confined to parts of Colombia or Ecuador. The
Generally, the endemism is higher in the Greater distribution of species of higher elevations is
Antilles as in the Lesser Antilles. The rate of controlled by a humidity gradient, which allows
endemism is 12.2% in Cuba (Duarte 1982), which páramo species to occur only from Ecuador to
is comparably high for bryophytes. The flora of Costa Rica but Puna species in a range covering
the Greater Antillean islands (Cuba, Jamaica, Pu- Peru and Bolivia.
erto Rico, Hispaniola) is mainly determined by The Andes are a relatively young mountain chain
the geological history, by climatic changes with an estimated age between 10 mio and 3
duringthe pleistocene, by their altitude and the years, when the present altitude was reached. This
distance to the mainland. As in all island does not mean that we have young species in the
phytogeography, it has to considered that the Andes which recently evolved. These mountains
water level of the oceans have been 100-170 m provided an ideal pathway for austral species,
lower during the past pleistocene glaciation which were able to migrate northwards in
periods, resulting in (sometimes) landbridges or elevations with appropriate temperatures.
at least closer distances to the neighbouring Examples are species such as Polytrichadelphus
islands and continents. A survey of the magellanicus, Lepyrodon tomentosus or
bryogeography of the Greater Antillean islands Rhizogonium novae-hollandiae. Forest and
is given by Buck (1990). Low islands such as paramo species could migrate in warmer viz.
Jamaica or Puerto Rico were submerged during cooler periods, since the vegetation belts had dif-
their geological history and therefore have no ferent extensions in the different climatic periods
bryogeographically significant flora as well as of the Pleistocene. There were changes in the
low biodiversity. In contrast, the bryoflora of altitudinal forest limit between 3300-3600 m and
Hispaniola was not submerged and its flora could less than 2000 m during about 20 different colder
respond to climatic changes with according periods, when the paramos expanded and fused
variation of altitudinal ranges. All islands have to larger areas.
a stock of widespread neotropical species plus Some of the andean species are also found in the
endemic caribbean elements. Some species of the erra do Itatiaia in SE-Brazil, which they reached
latter element are also found along the coast of possibly by long distance dispersal rather than
the caribbean sea („circum-caribbean“ in Belize, migration in coller periods.
Venezuela Suriname), where they probably have Some of the andean alpine bryophyte species are
secondaryly dispersed. The high altutides of also found in the mountains of tropical Africa,
Hispaniola harbout a few temperate and boreal probably caused by long distance dispersal. It
elements (e.g. Aulacomnium palustre, Calliergon must, however, kept in mind,that the wind
trifarium), perhaps as relicts from cooler periods systems in the tropics go from E to W and that
in the Pleistocene, and also a considerable amount these species may have originated in Africa (or
of andean elements (e.g. Rhizogonium lindigii, cam to Africa from the Andes but across the
Racomitrium crispulm, Lepyrodon tomentosus), Pacific Ocean and SE-Asia).
which underlines the role of long distance
dispersal. Jamaica and Cuba are composed of
limestone rocks and therefore have a limited Blöcher, R., Frahm, J.-P. 2002. A comparison
biodiversity. The Leasser Antillean Islands are of the moss floras of Chile and New
of recent volcanic origin and thus colonized from Zealand. Tropical Bryology 22: 81-92.
their surroundings. Buck, W.R. 1990.Biogeography of the Greater
Antillean Mosses. Tropical Bryology 2:
Andine element 35-48.
About 40% of the mosses are confined to the Crosby, M.R. 1969. Distribution patterns of
Andes to a smaller or larger extent. Some are very West Indian mosses. Annals of the
widespread and reach from southern Mexico to Missouri Botanical Garden 56: 409-416.
northern Argentina, others have narrower ranges

84 Frahm
Crum, H., Steere, W.C. 1958. A contribution tropical mosses of Mexico. , Bryologist
to the bryology of Haiti. Amer. Midland 79(4): 511-513
Natur. 60: 1-51. Duarte Bello, P.P. 1982. Musgos cubanos: su
Delgadillo M., C. 1995. Neotropical moss floras: presencia mundial. Acta Bot. Cubana 9:
Species common to North and South 1-19.
America. Tropical Bryology 10: 1-6. Frahm, J.-P. 1982. Grossdisjunktionen von
Delgadillo M., C. 1993. The Antillean Arc and Arealen südamerikanischer und afrika-
the distribution of neotropical mosses. nischer Campylopus-Arten. Lindbergia
Tropical Bryology 7: 7-12. 8:45-52.
Delgadillo, C. M. 1979. Mosses and Frahm, J.-P. 1990. Campylopus, a modern and
phytogeography of the Liquidambar successful genus? Trop. Bryol. 2: 91-
forest of Mexico. The Bryologist 101.
82:432-449. Frahm, J.-P. 1990. The origin and distribution
Delgadillo, C. 1990. Advances in Mexican of Neotropical species of Campylopus.
Bryology. Tropical Bryology 2: 49-52. Tropical Bryology 3: 1-18.
Delgadillo M., C. 1992. El banco de datos de Frahm, J.-P. 1994. A contribution to the
los musgos neotropicales. Tropical bryoflora of the Chocó region,
Bryology 6: 61-64. Colombia I. Mosses. Tropical Bryology
Delgadillo M, C.. 1987a. The Meso-American 9: 89-110.
element in the moss flora of Mexico. Fulford, M. 1951. Distribution patterns of the
Lindbergia 12: 121-124. genera of leafy Hepaticeae of South
Delgadillo M., C. 1987b. Moss distribution and America. Evolution 5:243-264.
the phytogeographical significance of Gradstein, S. R. & J. Vana, 1994. A boreal
the neovolcanic belt of Mexico. J. of bryophyte community in a tropical
Biogeography 14: 69-78. montane forest of Mexico. , Tropical
Delgadillo M., C. 1988. Floristic corridors for Bryology 9: 31-34.
moss distribution across the Gradstein, S. Rob & Vana, Jiri , 1987 , On the
Neovolcanic belt of Mexico I. The occurrence of laurasian liverworts in the
Tuypan corridor. Journal of Bryology Tropics , Memoirs of the New York
15: 165-175. Botanical Garden 45: 388-425
Delgadillo, C. M. 1992. Moss interchange: Gradstein, S. R., D. Montfoort & J.H.C. Cor-
bryofloristic similarities between nelissen 1990. Species richness and
Mexico and Colombia and the phytogeography of the bryophyte flora
phytogeographical role of the Central of the Guianas, with special reference
American bridge. The Bryologist to the lowland forest. Tropical Bryolo-
95:261-265. gy 2: 117-126.
Delgadillo, C. M. 1994. Endemism in the Gradstein, S. R., G.B.A. Van Reenen & D.
Neotropical moss flora. Biotropica Griffin III. 1989. Species richness and
26:12-16. origin of the bryophyte flora of the
Delgadillo M, C. 1998. Los musgos, la Colombian Andes. Acta Botanica
diversidad y sus causas en el Neerlandica 38: 439-448.
Neotrópico. Monographs of Systematic Gradstein, S.R., Weber, W.A.,1982,
Botany (Missouri Botanical Garden) Bryogeography of the Galapagos
68:61-67. Islands. Journal of the Hattori Botanical
Delgadillo, C. M., and A. S. Cárdenas. 1989. Laboratory 52:127-152
Phytogeography of high-elevation Grolle, R. 1969. Grossdisjunktionen in Arten-
mosses from Chiapas, Mexico. The arealen Lateinamerikanischer Leber-
Bryologist 92:461-466. moose. In: Fittkau et al. (eds.), Biogeo-
Delgadillo, C. M., Reese, W. D. 1976. Notes on graphy and Ecology in South America
pp. 562-582. W.Junk, The Hague

Kuc, M. 2000. Altitudinal additamenta to the species from South America are also recorded
uppermost ranges of mosses in Ecuador. from Africa and vice versa. As for the neotropics,
Tropical Bryology 18: 39-48. an anlysis of distribution types has not yet made
Kuc, M. 2000. The distribution of Hemiragus on a broad scale. Therefore the ranges of the
aurea (Brid.) Ren. & Card. species of Campylopus are again used as
(Hookeriaceae, Musci) and related notes examples (fig. 8.4)
of interest. Tropical Bryology 18: 55-
64. Egunyomi, A. 1984. Affinities between the moss
Ochyra, R., Bednarek-Ochyra, H., Arts, T., flora of East and West Africa. Yushania
Lewis Smith, R.I. Occurrence of the 1: 1-6.
neotropical moss Dicranella hilariana O´Shea, B.J. 1997.The mosses of sub-Saharan-
(Mont.) Mitt. in the Antarctic. Tropical Africa 2. Endemism and biodiversity.
Bryology 18: 153-160. Tropical Bryology 13: 75-86.
Pócs, T. Biogeography of the Cuban bryophyte O´Shea, B.J., Ochyra, R. 2000. Families and
flora. Taxon 37: 615-621. genera of mosses no longer believed to
Pursell, R.A., Reese, W.D. 1970. occur in sub-Saharan Africa. Tropical
Phytogeographal affinities of the mosses Bryology 18 : 119-128.
of the Gulf Coastal Plain of the United Ochyra, R., Kempa, R., Buck, W.R. 2000.
States and Mexico. J. Hattori Bot. Lab. Plagiothecium lucidum (Hook. f. &
33: 115-152. Wils.) Paris in tropical Africa. Tropical
Robinson, H. 1986,. Notes on the Bryology 18: 147-152.
Bryogeography of Venezuela,The Pócs, T. 1975. Affinities between the bryoflora
Bryologist 89: 8-12. of East Africa and Madagascar.
Schuster, R.M. 1990. Origins of neotropical Boissiera 24:125-128.
Leafy Hepaticae. Tropical Bryology 2: Pocs, T. 1991. Geography and ecology of
239-264. Usambara´s bryophytes. Pp. 71-85 in
Sharp, A.J. 1984. Geographical relationships in Hedberg, I, & Persson, E. (eds.),
the bryoflora of Mexico. J. Hattori Bot. Research for Conservation of Tanzanian
Lab. 56: 15-17. Catchment Forests.
Steere, W.C. 1984. The continental affiliations Spence, J.R. & Pócs, T. 1989. Distribution
of the moss flora of Hispaniola. J. Hat- patterns in the Afroalpine moss flora of
tori Bot. Lab. 56: 19-20. East Africa. Pp. 291-307 in: W.C.
Steere, W.C. 1985. On the continental affiliations Mahaney (ed.), Quaternary and
of the moss flora of Hispaniola. Environmental Research on E African
Monographs in Systematic Botany from Mountains. Balkema, Rotterdam.
the Missouri Botanical Garden 11: 155-
173.
8.2.3 Tropical Asia
8.2.2 Tropical Africa
Plant realms are delimited by lines of the highest
Tropical Africa harbours at present 2788 species floristic contrast. To determine the borders
of mosses (O´Shea 1997, there were 2939 species between plant realms, the changes of species and
indicated two years before, O´Shea 1995; as the genera are calculated. This has been done for
the whole tropics, the number goes down due to tropical Asia by Tan (2002). Tan could show
taxonomic revisions). 77% of the taxa are that 57% of the Philippine osses are not found in
endemic to tropical Africa. As compared with Taiwan and 64% of Taiwan mosses not in the
flowering plants, a percentage of 23% of species Philippines. This strong barrier is called Merrill´s
occurring also outside tropical Africa is Line. Within Asia, the Wallace Line is famous in
remarkably. This percentage will increase as a zoology. It seems, however, to have not much
result of worldwide revisions and more and more significance for bryophytes.

86 Frahm
Fig. 8.4: Distribution types of African mosses exemplified by the genus Campylopus. A. panafrican
range, B. East African range (there are also species confined to the East African islands). C. Guinea
coast range, D. subantarctic range. The afroalpine range is not illustrated. This range includes the
summits of the east African mountains as well as Mt. Kameroon.

Gradstein, S. R. 1991. Diversity and distributi- Tan, B., Engel, J.J. 1995. A preliminary study
on of Asian Lejeuneaceae subfamily of the affinities of Philippine, Bornean
Ptychanthoideae. Tropical Bryology and New Guinean Hepatics. Tropical
4:1-16. Bryology 11: 265-272.
Horikawa, Y. 1954. The occurrence of tropical Thiers, B.M. 1990. The floristic affinities of the
bryophytes in the Japanese Archipelago Lejeuneaceae of tropical Australia , p
and its phytogeographical significance. 13 in: T. Koponen & J. Mänttäri (eds),
VIII Congr. Inter. Bot. Rapp. et Comm. Congress of East Asiatic Bryology,
Sect. 16: 91-92 Programme and Abstracts, University of
Hu, R.1990. Distribution of bryophytes in China Helsinki, Helsinki.
Tropical Bryology 2: 133.
Iwatsuki, Z. 1990. Origin of the New Caledonian
bryophyte flora. Tropical Bryology 2:
139-148.
Jia Yu, Wu Pan-Cheng & Luo Jian-Xin , 1995.
The Mossflora of Mt. Jiuwan, Guangxi
and its significance in dividing the
boundary line between tropical and
subtropical regions in China. , Acta
Phytotaxonomica Sinica 33: 461-468.
1 fig.
Piippo, S. 1994. Phytogeography and habitat
ecology of Western Melasian endemic
Hepaticae. Journal of the Hattori
Botanical Laboratory 75:275-293.
Piippo, S. 1992. On the phytogeographical
affinities of temperate and tropical
asiatic and australasiatic hepatics. J.
Hattori Bot. Lab. 71: 1-35.
Piippo, S. 1994. On the bryogeography of
Western Melanesian Lejeuneaceae, with
comments on their epiphyllous
occurrence. Tropical Bryology 9:43-58.
Piippo, S., T. Koppen, and D. H. Norris. 1987.
Endemism of the bryophyte flora in
New Guinea. Symposia Biologica
Hungarica 35:361-372.
Redfearn, P. 1990. Tropical component in the
moss flora of China. Tropical Bryology
2: 201-222.
Schuster, R.M.,1972. Continental movements,
Wallace Line and Indomalayan-Au-
stralasian dispersal of land plants: some
eclectic concepts. Bot. Review 38:3-86.
Tan, B.C. 2002. The affinity of Moss Floras of
Japan, Taiwan and the Philippines
revisited: Old problems, new insight and
more questions. Acta Phytotax. Geobot.
53: 77-84.

88 Frahm

9. BIOINDICATION
Although bryophytes are frequently used in the Escocia Ariza, S. C. 1998. Estimados del
northern hemisphere, especially Europe, as deterioro de la flora briológica en
bioindicators for water quality, air quality, localidades industrializadas del area
nuclear contaminations, climate change, there has metropolitana de San Juan, Puerto Rico.
been most regretably hardly made any use of it M.Sc. thesis, University of Puerto Rico,
in the tropics. Especially in the tropics, where Mayaguez. 93pp.
environmental damages, air and water pollution Jayasekera, R., and M. Rossbach. 1996.
are locally still much more relevant than in Background levels of heavy metals in
industrial countries, the use of bryophytes as plants of different taxonomic groups
bioindicators is highly recommended. from a montane rainforest in Sri Lanka.
Base of this application is the fact that bryophytes Environmental Geochemistry and
as poicilohydric plants take up water and nutrients Health 18:55-62.
through their surface. They have no protection Lisboa, R. C. L., and A. L. I. Borges. 1995.
against toxic substances as flowering plants, Bryophyte diversity in Belém, Para and
which have cuticles and bark and take up water its potential as pollution indicator for
and nutrients buffered through the soil. Therefore urban areas. Boleton Museo Paraense
toxic substances are automatically taken up by Emilio Goeldi, serie Botanica 11:199-
bryophytes, either in lakes and rivers or from the 225.
atmosphere. Odum, H. T., G. A. Briscoe, and C. B. Briscoes.
The effects of air pollution on bryophytes has 1970. Fallout radioactivity of epiphytes.
only be studied by Duran et al. (1992) in Mexico- In Odum, H. T. (ed.). A tropical
City, Lisboa & Borges (1995) in Belém, Brazil, rainforest: A study of irradiation and
and Esciócia Ariza (1998) in Puerto Rico. Natu- ecology at El Verde, Puerto Rico.
ral heavy metal accumulation has been measured Atomic Energy Commission,
by Jayasekera & Rossbach (1996). Radioactive Washington D.C.
fallout has been determined in bryophytes by Steere, W. C. 1970. Bryophyte studies on the
Odum et al. (1970) and Steere (1970). irradiated and control sites in the
rainforest at El Verde, Puerto Rico. In
Odum, H. t. (ed.). A tropical rainforest:
Durán D., A., Cisneros C., A.E. & Vargas V., A study of irradiation and ecology at El
A. 1992. Evaluación briológica de los Verde, Puerto Rico. Atomic Energy
efectos de la contaminación atmosférica Commission, Washington D.C.
en la Ciudad de México. Tropical
Bryology 6: 71-82.

90 Frahm
Bioindication does not only concern the Rico. The Bryologist 99:81-84.
indication of human impact or pollution. Any Sillett, S. C., S. R. Gradstein, and D. Griffin
bryophyte, any organism is an indicator of its III. 1995. Bryophyte diversity of Ficus
habitat preferences and ranges, geographical tree crowns from cloud forest and
ranges and vertical ranges. The narrower the pasture in Costa Rica. Bryologist
ecological niche inhabited by a species, the better 98:251-260.
the indicator value. The indication covers
parameters of soil and climate:
- nitrogen or phosphorous rich habitats indicated
by Bryum argenteum, Funaria calvescens and
others.
- heavy metal rich habitats indicated by species
of Mielichhoferia, Scopelophila, Merceya.
- primary and secondary forests, disturbed and
non disturbed sites (Equihua & Gradstein 1995,
Hyvönen et al. 1987, Norris 1990, Romero 1999,
Serrano 1996, Sillet et al. 1995).
- altitudinal belts, cloud belts (cf. chapter 6).
- phytogeographical provinces (cf. chapter 8).
Equihua, C. & S. R. Gradstein 1995 .

old field and a rain forest: preliminary
results unpaginated tip-in. In C.
Delgadillo M (ed.), International
Bryological Conference: Tropical
Bryophytes: Biology, Diversity and
Conservation. August 7-12, 1995.
Mexico City. Scientific Program,
Abstracts, Field Trips, Tourist Tips. ,
Instituto de Biologia, UNAM, Mexico
City.
Hyvönen J., Koponen T. & Norris D. H. 1987.
tropical rainforest in Papua New Gui-
nea , Symposia Biologica Hungarica 35:
621-629
125.

10. CONSERVATION
The sensivity of many bryophytes to disturbance, Holstein in Germany by Eigner & Frahm in 1974.
the indication for narrow ecological niches and Meanwhile almost every state in Germany as
the rarity of many species designates bryophytes most countries of Europe have red lists.
as valuable tools for nature conservation. Basis Interestingly they are not used outside Europe,
for this use is the classification into into categories although they have proved extremely useful. In
such as Europe, the value of an region or habitat is
- extinct or vanished measured by the number of endangered species
- endangered occurring in this area. Therefore the presence of
- vulnerable, and rare bryophytes can be used as argument for
- rare, or protecting areas or for the value of protected
0 = extinct or vanished, 1 = threatened by areas.
extinction, 2 = strongly threatened, 3 = To my knowledge, there exists not yet red list
threatened, 4 = potentially threatened. for a tropical country, which should be a
This classification requires a good knowledge of stimulation to set up such lists. The purpose of
the frequency and distribution of bryophytes, Red Lists is, that not only the endangered species
which is not as good in tropical countries. shall be protected but its habitat. If areas are to
However, this should not be a reason not to make be considered for conservation, it is asked how
use of these properties. Bryologists have to care many red-List-sapecies are found in this area and
that not only mammals, birds, orchids or bby this way the value of an area is determined.
bromeliads are used for nature conservation. There more „mossy“ a habitat is, the more ur-
Principally, it has to be shown th authorities that gent is it to include bryophytes in nature
bryophytes not only exist but are valuable tools conservation activities. There is, however, a
for nature conservation. So the logical worldwide red list issued by the International
consequency of a checklist, which is a necessary Association of Bryologists (Tan et al. 1994),
base, is the estimation of the frequency of species which includes 91 species. Amongst these species
and the estimation of their threads. Even without are mostly tropical species, which are known only
a checklist (for a country or province), single from the type collection and which habitat is
cases of rare and endangered species can be endangered.
picked up and made public to nature conservation In Europe, bryophyte 29 species are listed in
authorities. annexe 2 of the Convention on the Conservation
Such lists of threatened bryophytes, so called „red of European Wildlife and Natural Habitats. The
lists“, were first elaborated in Europe. The first European countries have lists of protected
red list was compiled for the state of Schleswig- bryophyte species to a various amount, ranging

92 Frahm
from a few enfangered taxa (e.g. Sphagnum species on cacao trees in western Ecua-
species) to all endangered red list species. dor. Bryobrothera 5:81-86.
Red lists can be elaborated on different Hallingbäck, T., and B. Tan. 1996. Towards a
geographical levels: world, continent (e.g. global action plan for endangered
Europe), country and provinces. A species bryophytes. Anales del Instituto de
common in one province can be rare in another Biologia de la Universidad Nacional
and neds protection on this regional basis. Abso- Autonoma de Mexico 67:213-221.
lute rarity exists only on a wordlwide level. Hedenäs, L. , 1995. How do we select species
Therefore the most intensive efforts have to be for conservation?, pp. 28-29. In C.
undertaken to protect the worldwide endangered Delgadillo M. (ed.), International
species. Thus every tropical bryologists should Bryological Conference: Tropical
be orientated which species of the world red list Bryophytes: Biology, Diversity and
occurs in his country, which other species are Conservation. August 7-12, 1995.
similarly endangered (this world list is surely not Mexico City. Scientific Program,
complete but a begin) and which efforts can be Abstracts, Field Trips, Tourist Tips. ,
undertaken for the protection of these species. Instituto de Biologia, UNAM, Mexico
This would include not only protection of habitats City.
(„in situ“) but active steps for preservation of Koponen, T. 1992. Endangered bryophytes on
species, e.g. in culture („ex situ“). Similarly as a global scale. Biological Conservation
endangered animals are protected in zoological 59:255-258.
gardens and rare plants in botanical gardens, it Sastre-D. J., I & B. Tan 1993. Problems of
cannot be waited until certain bryophytes get bryophyte conservation in the tropics:
extinct. Often, nature conservation has got a neotropical and paleotropical cases. ,
burocratic act including filling lists of endangered American Journal of Botany 80 (6,
species, ranking the rarity and finally adding a Supp.): 6.
cross mark after the species name when it has Sastre-D. J., I. & B. Tan 1995. Problems of
got extinct. This is administration of extinction, bryophyte conservation in the tropics: a
which should be avoided. discussion, with case example from
Puerto Rico and the Philippines. ,
Delgadillo, M. C. , 1987. About endangered Caribbean Journal of Sciene 31: 200-
tropical areas. P. 5. , The Bryological 206.
Times 41: 5 Söderström, L. , 1995 , Islands-endemism and
Delgadillo M., C. , 1994. Tropical Bryophytes: threatened bryophytes, pp. 43-44. In C.
Biology, diversity and conversation. Delgadillo M. (ed.), International
The Bryological Times 80: 6. Bryological Conference: Tropical
Delgadillo, C. M. 1996. Moss conservation in Bryophytes: Biology, Diversity and
Mexico. Anales del Instituto de Biologia Conservation. August 7-12, 1995.
de la Universidad Nacional Autonoma Mexico City. Scientific Program,
de Mexico 67:177-181. Abstracts, Field Trips, Tourist Tips. ,
Gamundí, I. J., and C. M. Matteri. 1998. La Instituto de Biologia, UNAM, Mexico
problemática de la conservación en las City.
criptógamas avasculares. Monographs Streimann, H. 1994. Conservation Status of
of Systematic Botany (Missouri Bryophytes in Eastern Australia.
Botanical Garden) 68: 287-299. Tropical Bryology 9: 117-122.
Gradstein, S.R. 1992. Threatened bryophytes of Sastre-D. J., I & B. Tan , 1993 , Problems of
the neotropical rain forest: a status re- bryophyte conservation in the tropics:
port. Tropical Bryology 6: 83-94. neotropical and paleotropical cases. ,
Gradstein, S. R. 1999. On the rediscovery of American Journal of Botany 80 (6,
Spruceanthus theobromae (Le- Supp.): 6.
jeunaceae, Hepaticae), an endangered Sastre-De Jesus, I., and B. C. Tan. 1995.

Problems of bryophyte conservation in

the tropics: a discussion with case
examples from Puerto Rico and the
Phillipines. Caribbean Journal of
Science 32:200-206.
Tan, B., Geisler, P., Hallingbäck, T. 1994.
Towards a World Red List of
Bryophytes. Bryol. Times 77: 3-6.
Human impact and disturbance
Hyvönen J., Koponen T. & Norris D. H. , 1987.

tropical rainforest in Papua New Guinea
, Symposia Biologica Hungarica 35:
621-629
Hyvönen, J. , 1970. Human Influence on the
moss flora of tropical rainforest in Papua
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125.
Rico. The Bryologist 99:81-84.

94 Frahm

APPENDIX 1: COLLECTING TROPICAL BRYOPHYTES
Collecting in tropical countries officially requires countries getting problems with „inofficial“
usually permits and is usually due to severe collecting. And picking up small samples of
restrictions. The conditions vary from country to bryophytes for identification is usually not that
country. Frahm (1996) has tried to compile the what even national park officials understand as
available data, which are, however, not complete plant collecting. Often bryophytes are not
and have changed in many countries since. Local regarded as „real plants“
botanists are not as affected as foreign botanists, The same problems as with collecting permits
for which in some cases collections are made concern the export of specimens, which concerns
impossible. An official argument - also used by again foreign botanists and is not discussed here,
bryologists from tropical countries - is that they because this manual is primarily addressed to
are said to plunder the genetical ressources of tropical bryologists, which generally do not have
tropical countries. This is political propaganda these problems. It is important to know, that (at
and nonsense. Foreign botanists visiting tropical present) bryophytes are not on the CITES list and
countries have, of course, to consider some points may be legally introduced to the homeland of
which Mori & Holm-Nileson (1981) and the collector.
Delgadillo (1987) have outlined. On the other Bryophyte collecting is much easier as collecting
hand it has to kept in mind that usually and flowering plants. Bryophytes need not to be
predominantly foreign botanists have done and pressed, and they need relatively little care during
are currently doing the botanical exploration in fieldtrips. O´Shea (1989) has written a short guide
the tropics and most what is known about tropical for planning collecting trips and collecting
bryophytes has been elaborated by foreign bryophytes in the tropics especially for visitors
bryologists, else most of the tropical countries from non-tropical countries. Buck & Thiers
would be terra incognita. (1996) published guidelines for collecting
As everywhere in the world, collecting in natio- bryophytes in the tropics. Gradstein et al. (2001)
nal parks is not allowed without permits, and give also instructions for that purpose. Special
permits are often only very circumstantially to instructions for collecting corticolous bryophytes
apply for. Thanksfully, collecting of bryophytes as well as foliicolous bryophytes and lichens are
is much more inconspicuous as that of flowering given by Gradstein et al. (1995).
plants (especially cacti or orchids), and there is
no example known of a bryologists in tropical

96 Frahm
There are several instructions for planning and 1. Fieldstations

preparing fieldtrips in tropical countries (see
refs.). For visitors from non-tropical countries, The easiest way to stay in the field is to use
there are logistic problems concerned with the fieldstations. These fieldstations can be very
style of the fieldwork such as health care, sending differently equipped, from bamboo huts or roofs
equipment by air cargo, car rental, staying abroad in the jungle to small labs with microscopes and
in the field, nutrition, food, which are not covered dryers. The use of these stations requires,
here. Another problem is the requirement of however, always official permits, which can be
collecting permits in most countries, which have difficult to get. A local bryological counterpart
to be obtained in advance, and also eventual can be very helpful in this respect, if there is
export permits of specimens. In this case, contact somebody willing to act as such. There are often
to local bryologists will be helpful. no possibilities for accomodation in the field that
Recommemndations for botanists visitimng the camping is required instead, which bears still the
tropics in this concern are given by Mori & Hom- risk of getting robbed in parts of the tropics.
Nielsen (1981) and Delgadillo (1987). Years ago, an attempt has been made to compile
a list of fieldstations in the neotropics, which
Buck, W.R., Thiers, B.M. 1996. Guidelines for remained fragmentary and my be outdated today.
collecting bryophytes. Selectes
Guidlines for Ethnobotanical Research: Delgadillo M., C. 1989. A Guide to fieldstations
A field manual: 143-146. in the tropics I., Mexico. Tropical
Delgadillo-M., C. 1987. Additional Bryology 1: 1-4.
recommendations for bryologists Frahm, J.-P. 1992. A Guide to Fieldstations in
visiting the tropics. Bulletin of Bryology the Tropics III. Colombia. Tropical
XXIII. Taxon 36: 289-291. Bryology 5: 23-26.
Frahm, J.-P. 1996. Directory of Bryophyte Salazar Allen, N. 1989. A Guide to fieldstations
Collecting. Bryol. Times 89 Special in the tropics II. Panama. Tropical Bryo-
Issue. Includes regulations for 30 logy 1: 5-8.
countries.
Gradstein, S. R., P. Hietz, R. Lücking, A.
Lücking, H. J. M. Sipman, H. F. M.
Vester, J. Wolf, and E. Gardette.
1996. How to sample the epiphytic 2. Fieldwork
diversity of tropical rainforests.
Ecotropica 2:59-72. Collecting bryophytes in rain forests was so far
Mori, S. A., and L. B. Holm-Nielsen. 1981. and is usually still confined to collections of trunk
Recommendations for botanists visiting epiphytes below 2.5 m height or epiphylls, since
neotropical countries. Taxon 30:87-89. there are few bryophytes on soil in rain forests.
O’Shea, B. J. 1989. A guide to collecting The canopy has much been neglected. By this
bryophytes in the tropics. British way, only a small portion of the diversity of rain
Bryological Society Special Volume forests has been registered. This limitation has
No. 3: 1-30. British Bryological now been overcome by introducing alpine
Society, Cardiff. climbing techniques for canopy research (or
O’Shea, B. J. , 1992. Beginners guide to tropical locally by cranes, canopy walks or use of
bryology - The BBS Tropical Bryology balloons). Climbing has been tought in some of
Group trip to Mount Mulanje, Malawi. our courses, but it is resigned to give instructions
Bulletin of the British Bryological here, because self teaching of these climbing
Society (59): 12. techniques may be too risky.

Checklist for field equipment: 2. Collecting. The bryophytes are removed from
Fieldbook the substrate by hand or a knife. (These knifes
waterproof marker or pen with cord are often lost, there for use cheap ones or attach
altimeter and/or GPS it by a chain to your belt). Very convenient is the
pocket knife with cord use of scrapers, which can be bought in do-it-
paperbags youself-shops for very few money (buy half a
bagpack dozen, they disappear within the time, but 50 cost
1. Locality data. As for collecting of every as much as a swiss knife), especially to remove
biological specimen, note the locality data in a bryophytes from soil For better drying, the water
notebook with hard cover. This is done with a is removed by pressing the specimen with the
fine waterproof marker or a special pens hand, removing soil or litter, and flattening the
(„astronauts pens“, Fisher pen) with allow to specimen. Cushions are cut in slices. The
write even on wet paper. Pens are easily lost in specimen is put in a paper bag (small brown paper
the field, so have spare pens or attach the pen to bags used in shops in the Americas, paper lunch
a cord and wear it around the neck or fix it at bags elswhere). If no bags are avaliable, folded
your belt. Every collecting spot gets a number. double (!) layers of newspaper can be used. Every
The notes include specimen gets a number with a marker, usually
- the locality (state, county, parish, road from... and preferably the collectors consecutive number.
to... kilometer ..., geographical coordinates, today There are also systems in use with a combination
easily taken from a GPS). The data should allow of the date (02041523 = 23. specimen on April
somebody else to localize the place, e.g. a local 15th, 2002), country code or locality codes (e.g.
botanist to refind a species or a monographer to CR 31/17 = 17th specimen on locality 31 during
map it. For the latter purpose, geographical the Costa Rica trip). At least the locality number
coordinates are best, else indication of the next is necessary, specimen numbers can also be added
city found on a world atlas (e.g. 45 km NE of later. In addition, the microhabitat has to be noted
Bogotá). This is more valuable (for this purpose) on the paper bag, for which abbreviations may
than the indication of villages which nobody will be used (e.g. ct: corticolous, ru: rupestrial, ep:
find on a map. Military recording systems such epiphyllous, tc: terricolus).
as the UTM system not really been used by The use of plastic collecting bags should be
botanists, although they allow to determine every avaided, even for short time. The physiological
place on the world by special maps. National grid problem is that wet specimen are metabolic active
systems are useful for national grid mapping. and start to dissimlate oif they have no light and
- the habitat. no CO2. This causes severe damages. Even if it
- the elevation. Barometric altimeters can only is intended to cultivate the specimen later, it is
be trusted if they are corrected every day and even recommended that they are transported dry. This
during a day, the air pressure can change due to concerns also transport e.g. by air mail. It is part
changing wheather conditions that the readings of the bryophyte life strategy that the plants can
are not correct. The change of one millibar stand dry periods in anabiosis. Ypou may start a
airpressure results in a change of 7 m altitude culture easily with a dry specimen which is
reading! Today, wrist watches with built in soaked or sprayed with water after transport.
altimeters are even cheaper than separate 3. Storing. During the day, the specimens are
altimeters. Electronic altimeters are better shock- kept in a net or bagpack. If the specimens cannot
proof than mechanical altimeters (such as the be dried soon, they can be stored in nets (plastic
famous swiss Thommen). Satallite navigators nets) or cotton bags (flour bags, pillows,
can give the altitude if 4 satellites are available; blankets), but never in plastic bags (except for
they are only as exact as 100 m (as you can see protection against rainfall). Some authors have
from the altitude, which gos up and down even recommended to use plastic bags filled with silica
if you stand in a place). If the place is known, gel, but the amount of water in the specimen is
the altitude is best taken from a map (if available). so much that kilograms of silica gel must be used
- date. .

98 Frahm
4. Drying. Specimens can be dried in between in If the specimens are dry, they are sealed in plastic
the sun. Spreading the specimens on the soil bags. Litter bags are convenient for that purpose.
makes problems during windy weather. In this
case, the specimens can be hung up in their nets Croat, T. B. 1979. Use of a portable propane
for drying, or can be spread in tents, if these have gas oven for field drying plants. Taxon
mosquito nets and the air can pass through the 28:573-580.
tent. If the wheather does not allow or the days Frahm, J.-P., and S. R. Gradstein. 1986. An
are used for field studies, they can be dried in apparatus for drying bryophytes in the
the evening over kerosine or butan stoves. Croat field. Bryological Times 38:5.
(1979) gave instructions for a propane heater Greene, S. W. 1986. Keeping them dry.
mounted on a pick-up truck. The easiest Bryological Times 38:6.
arrangement is the place a metal bar (e.g. from a Howard, R. A., Howard, E. S. 1981 Labels for
plant press) over two aluminium boxes, stones, wet tropical research. Biotropica 13: 77-
walls made from stones (like a barbecue grill), 78.
put the wet specimens on the bar and place a
kerosine stove below. For this purpose, the
construction of light weight aluminium frames Special instructions for collecting epiphylls
have also be recommended (Frahm & Gradstein
1986). In the lab, similar frames can be used for Special instructions for collecting epiphyllous
drying specimens. They can be heated by electric bryophytes are given by Lücking & Lücking
radiators placed below or by electric bulbs (1996). Epiphyllous bryophytes are collected
mounted at the bottom of wooden boxes (bulb with adhaerent leaves and thus the same drying
dryer). Greene (1986) recommended the use of techniques as for leaves are used, at the best in
siliga gel for drying. In my experience, the small plant presses. They can be made from iron
amount of silica gel required to dry wet bars or wooden plates in the size of folded
specimens is enormous and raises weight and newspaper. Although special drying cartoon is
storage problems. In addition, the problem raises available, newspaper can be more easily obtained
how to get the silica dry. The use of silica gel is, and exchanged. A good trick ist to insert pieces
however, recommended for specimens for DNA of cardboard of the same size between the
extraction. In general, specimens for DNA newspaper sheets. by this way, the humidity
extraction need to dry up fast, that the protein soaked up from the leaves by the newspaper is
chains get not broken. If the specimen is kept soaked up the the cardboard and can evaporate
wet for a longer time during the trip, especially through the wholes in the cardboard to both sides.
in plastic bags, it starts to mould, resulting in a If epiphylls are collected as saide collection,
degeneration of proteins. The ability of older leaves or part of them may also be put in paper
herbarium specimens for later DNA extraction bags. From large leaves, small portions are cut.
depends on the way they have been dried. To covr the whole diversity , the collection of
Specimens which were properly dried can even numerous leaves is necessary. According to R.
be extracted after 20 years, those having not & A. Lücking, the collection of 9 leaves in a low
properly dried can no more be used. Therefore a diversity area in Costa Rica (with a total of 48
quick desiccation is required. The quickest is epiphyllous species) resulted in 50% of the
shock drying in silica gel. For that purpose, 50 species, but 90% of the species were only reached
or 100 ml plastich bottles are filled 3/4 with silica by the collection of 62 leaves. In a high diversity
gel. Next several single bryophyte plants are area, 360 leaves were necessary to get 90% of
picked up with tweezers, put upon the silica gel the species. Epiphylls are especially found in
and the bottle is closed. The silica gel the understory of the forests (often with a sharp
immediately extracts all humidity from the plants, upper delimitation in 1 m height) or in light gaps
which is a guarantee for a successful extraction. and along creeks. Both reflects the physiological
If no silica gel can be used, the specimens must problems of epiphylls in the tropical forest: light
carefully be dried else. intensity in the understory of such forests is

Fig. I.1: A portable drier made from an aluminium frame, heated with kerosine stoves (Frahm
Frahm & Gradstein 1986).
Fig. I.2: Drying specimens in the field over a kerosine stove (BRYOTROP Peru).

100 Frahm
(e.g. Pócs 1978, Winkler 1967) should be

consulted.
Lücking, R., Lücking, A. 1996. Foliicolous

bryophytes and lichens. In: Gradstein,
S. R., P. Hietz, R. Lücking, A. Lücking,
H. J. M. Sipman, H. F. M. Vester, J.
Wolf, and E. Gardette. How to sample
the epiphytic diversity of tropical
rainforests. Ecotropica 2:59-72.
Pócs, T, 1978. Epiphyllous communities and
their distribution in East Africa. Bryoph.
Bibl. 13: 681-713.
Winkler, S. 1967. Die epiphyllen Moose der
Nebelwälder von El Salvador, C.A. Rev.
Bryol. Lichénol. 35: 303-369.
3. Fieldwork
Hectareplot studies
Fig. I.3: Dryer from fig. I.1 in action, front re- Collecting should preferably not be at random.
moved to show the stoves (Irangi, Congo, for- In the past, collecting in thr tropics was usually
mer fieldstation of King Leopold II. done by travelling around, collecting here along
a roadside and there beside a waterfall. Collecting
was doner mosztly for taxonomic purposes and
main purpose was to detect new species. Usually
nice sites were selected, ignoring a large deal of
other, less attractive habitats. By this way, only a
extremely low, reaching hardly the compensation
part of the bryoflora was registered. To cover all
point. This effect is compensated by (a) higher
habitats in an area, the collections must be done
CO2 (which is produced by the decomposition
systematically, including plantations, secondary
of litter at the forest ground, heavier as air and
forests, river banks etc. Today, the purpose of
thus concentrated at the bottom of the forest, or
collecting is not only taxonomy but also diversity
(b) by higher light intensity as in light gaps.
studies to get an impression of the richness of
Epiphylls are found on all kind of leaves such
different habitats and thus of the value of these
thick leathery leaves, leaves of palms, ferns and
habitats (e.g. with concern to nature
even filmy ferns. Only hairy leaves are not
conservation). Therefore it should be intended
colonized. Usually, the species cannot be
to get complete inventories of study areas. Instead
identified in the field nor properly distinguished.
of collecting here and there in a large area,
Therefore at random collections have to be made
collecting everything in a small site is
to cover most of the species. This means that
recommended.
collection of each 5 leaves with the highest cover
An impression of the alpha-diversity of forest
of epiphylls should be taken from different
types or forests at different elevations can be
phorophyte species and in different
obtained by collecting bryophytes on all available
microhabitats.
substrates (soil, trunks, lianas, shrubs) in a sample
The evaluation of composition, cover,
plot. The sample plot should be homogenous
abundance, frequency etc. of epiphylls on leaves
(avoiding disturbed areas such as roadside
is a special subject, for which special literature
vegetation, mixtures of open and shaded sites)

to allow comparison. It is often recommended to sufficient number so that a statistical evaluation

test the size of the plots with a minimum species will be possible. The outer canopy branches have
area curve. This is, however, only possible for to be sawed off and lowered to the ground by
very experienced collectors who can differentiate ropes for study. If possible and allowed, also large
all species. Based on previous studies, the average branches can be sawed off and studied more
size of the plots is generally at least 25 x 25 m comfortably on the ground. The inventory is
and maximally 100 x 100 m. The plots need not made based on the Johannson-zones (Johannson
to be quadrate but can be rectangular (e.g. 10 x 1974). It is often recommended to study a number
100 m), depending on the topography. It must be of plots of 20 x 20 cm within every Johannson
homogenous. In practice, the size of one hectare zone. This is essential for all further statistics,
has revealed as appropriate. Since many studies since the calculations must be based on a same
are made on a hectare basis, the results can also area. To cover all species, the number of plots
be easily compared. must be quite high. If no statistics are intended,
collecting larger parts of trunks and branches is
Transect studies recommended, because some species may just
Collecting is preferably done along an ecological occur outside the sample plots.
gradient along a transect. Such transects can lead Another method is, to take all bryophytes from a
from a ridge to the bottom of a valley (dry - sample plot of 20 x 20 cm in the lab. This is
moist), or go along a mountain slope (warm - sometimes required, if the wheather or attacks
cool). In this case, the floristic results can be from leeches, ants or mosquitos do not allow to
compared with of measurements of ecological perform longer studies in the field. The removed
parameter und used for correlation analysis to cover can be transported in pizza boxes. In the
see which factors are important for the lab, the samples can be studied under the
composition of bryophytes. Recent studies binocular. This method can be performed only if
preferred altitudinal gradients (e.g. the the epiphyte cover has a certain thickness and
BRYOTROP project) in intervals of 200 m from not for crust epiphytes, but is recommended for
the lowland to the forest line. phytoisocilogical studies.
It is very important for all evaluations that the Depending on the purpose of the study, the
study areas are comparable (not only in size but species are registered as presence/absence (for
also in structure) that not „apples are compared diversity studies) or their cover is estimated (for
with pears“. phytosociological studies), either by percentage
or by Braun-Blanquet indices. Presence/absence
Epiphyte and Canopy studies does nothing say about the different abundance
Special instructions for collecting epiphytic of species and gives no idea of the composition.
bryophytes are given by Gradstein (1996). A A present species can be a tiny liverwort
large part of the diversity of rain forests is interwoven in a moss or a moss covering most
harboured in the crowns of trees. This does also of the branch or trunk.
concern bryophytes, to a smaller content in the For trunk epiphytes, trees with different bark
lowland forests but increasingly in montane structure should be studied. In opposite to trees
forests. Part of the species is even restricted to in temperate regions, the pH of the bark varies
the crown. Trees are usually climbed using the not much and is always low (between 4 and 6).
rope technique or (if possible) the canopy is Species composition is influenced by the structure
accessed by a crane, canopy walk, or balloon. of the bark (smooth, flacy, stripping off) and thre
Rope technique is described by e.g. ter Steege & water stroing capacity of the bark. The higher
Cornelissen (1988). Exceptionally, „destructive the precipitation of an region, the lower is the
collecting technique“ can be used by cutting a influence of bark factors.
tree. Also fallen trees should be studied, which
have come down not too long time ago. Within Cornelissen, J.H.C. & ter Steege, H. 1989. Dis-
the tree, inventories are made of 20 cm2 plots tribution and ecology of epiphytic
within the 5-6 tree zones (Johannson 1974) in a bryophytes and lichens in dry evergreen

102 Frahm
forest of Guyana. J. Trop. Ecol. 5: 131-

150.
Gradstein, S.R. 1996. Corticolous bryophytes.
In: Gradstein, S. R., P. Hietz, R.
Lücking, A. Lücking, H. J. M. Sipman,
H. F. M. Vester, J. Wolf, and E.
Gardette. How to sample the epiphytic
diversity of tropical rainforests.
Ecotropica 2:59-72.
Johannson, D. 1974. Ecology of vascular
epiphytes in West African rain forest.
Acta Phytogeogr. Suecica 59: 1-136.
Ter Steege, H., Cornelissen, J.H. 1988.
Collecting and studying bryophytes in
the canopy of standing rain forest trees.
Pp. 285-290 in: J.M. Glime (ed.)
Methods in Bryology. Nichinan.

APPENDIX II: HERBARIUM MANAGEMENTS
1. Drying
If the collections arrive in the herbarium in wet problematical, since chemicals which poisen
state, they need to be dried up immediately. Some beetles are also harmful for men. Some herbaria
hergaria have special dryer of various modes. still poisen with naphtalene, which causes an
Some consist of boxes, open above, in which the uncomfortable odor. The easiest way is to cool-
plant presses are upright packed. Heat is freeze the specimens. Requirement is that the
generated by light bulbs below. The warm air specimens are absolutely dry. Then they will be
rises through the cardboard between the sealed in plastic bags and stored in a deep freezer
specimens. A similar construction as the field at -20°C for 24 hours. Larger bags also longer.
drier or the field driere itself as described in Ap- By that way, all beetles and larvae, especially
pendix I (4) can also be used in the lab and an tropical ones, are killed. After the freezing, the
electric heater can be used instead of the kerosine are kept in the plastic bags to warm up for again
stove. Other drier consists of a cupboard with 24 hrs.
shelves made from wireand integrated heater,
through which hot air is blown. However,
bryophytes do not need so much care. The 2. Labelling
simplest and easiest way is to spread the
specimens in a room in one layer and wait until Today, labelling is completely done by
they are dried.. computers. Since the beginning of the computer
Bryophytes are rarely attacked by pests. Reason era in the seventies of the last century, biologists
is that they contain antifeeding agents, with which have tried to make use of computers. At first,
they protect themselves against being eaten up punch cards were used to store the collecting
by beetles and snails. Therefore the chance being information. For the first PCs, special labelling
attacked by e.g. museum beetles is low, even in software was designed, and there were numerous
the tropics. But every rule is without exception. different programs with various advantages abut
There are, nevertheless, specialists among beetles also disadvantages.
which can eat bryophytes. I have seen in my Today there exist principally two ways to label
life only one case (in a private herbarium in specimens, (a) by using a label program, (b) to
Germany!). Therefore protective treatments can use a database program. Alternative a is not
be performed. The classic method is chemical recommended, since it only allows to print labels
poisening of the specimens., which is from different localities, which is nothing better

104 Frahm
than copying labels on a xerox machine. The A test can clarify this: a paper sheet with some
advantage of database programs is that all label notes made by different iunk pens is exposed to
information is stored in the computer and can be the sun (at the window) and the notes of non-
searched for. By that way, also evaluations can permanent pens will dissappear soon.
be performed such as: which species were found Also labels must be as waterproof as the ink pens
in the same relevée?, which species were found and therefore the choice of printers is important.
between 6- and 800 m? which species were found Ink printer prints are not waterproof and should
on a special host tree?, in the department XY or be avoided. In contrast, laser printer outprints will
around the village of YZ? And so on. not make any problems.
Any database program can be used for that Beispiele von Labels, Auswertungen
purpose (e.g. MS Access, Filemaker) by
generating fields for
- species 3. Storing
- country
- state/department Traditionally, bryophyte specimens were kept in
- county the same way as flowering plants: the convolutes
- locality were glued upon herbarium sheets, these sheets
- habitat wrapped with an envelope, and the sheets bound
- date to faszicles. This may have made sense in large
- collector herbaria, where the cabinets had the size for her-
- collection number barum sheets. However, herbarium sheets with
- altitude several specimens glued upon one sheet are
- latitude/longitude circumstantial to handle. If the convolutes are
If a mask is generated for the outprint, the label glued upon the sheets, they can hardly be
can directly be printed to the paper sheet from removed (e.g. after a misidentification) without
which the convolute is folded. This avoids to cut damaging the specimen (this is the reason that
and glue the labels. some herbaria used needles to fix the specimens,
Attention has also be paid to the use of paper, other did not glue the specimens but laid them
glue, pens and printers. Formerly, the paper loosely between sheets with the consequence that
(used for herbarium sheets, envelopes or specimen could easily fall out).
convolutes) contained acids which caused that The proper method is today to store the specimens
the paper was easily broken after hundred years like filecards in cardboard boxes. Then they can
or more. This required extensive procedures (re- easily be sorted. The only (small) disadvantage
packing) of specimens in older herbaria. Today, is the upright position of the specimens. The
even xerox paper is acid free that any paper can boxes are placed in metal cabinets. Fumigating
be used for herbarium puposes and no special is usually not necessary.
paper is required.
If labels are used and glued on the specimens,
attention has to be paid to the kind of glue. Self- Bello y Bello, B. 1992. La colección briológica
adhesive labels, although they seem practically, del Herbario Nacional de México
should be avoided since the glue lasts no more (MEXU). Tropical Bryology 6: 35-38.
than 10-20 years. It is absorbed and degenerated Fosberg, F. R., and M.-H. Sachet. 1965. Manual
by the paper with the result that the labels will for tropical herbaria. International
fall off. (Sad own experience). Bureau for Plant Taxonomy. Utrecht,
Problems can also arise by the use of pens to The Netherlands.
mark specimens with collection numbers. The ink
of many ink pens is bleaching under sun light
and disappearing, causing heavy problems with
the identification of specimens. Ink pens should
therefore not only waterproof but also permanent.

Fig. II.1: Herbarium label generated with the

computer program Filemaker. All text is stored
in fields and can be searched for and extracted
for reports or evaluations.
Fig. II.2 (right): Modern storing herbarium speci-

mens in cardboard boxes in a steel cabinet (Her-
barium BONN).
Fig. II.3: Old style of bryophyte herbarium in faszicles designed for flowering plants (Hb.
BONN).
106 Frahm

APPENDIX III: IDENTIFICATION
The amount of available literature varies much Greene, S.W., Harrington, A.J. 1989. The Con-
in the different parts of the tropics. The situation spectus of Bryological Taxonomic Li-
is relatively the best in the neotropics as compa- terature Part 2. Guide to national and re-
red with Africa or tropical Asia. Recently, the gional literature. Bryophytorum Biblio-
British Bryological Society Tropical Bryology theca 37.
Group started to enhance the knowledge of the
bryology of tropical Africa with different activi-
ties. 2.1 Neotropics
A first step is to identify families or genera. This
is possible by generic bryophyte floras such as The neotropics are the bryologically best
that by Gradstein et al. (2001) for the neotropics. surveyed tropical area in the world, which is
A similar flora for tropical Africa is in preparati- documented by a comparably and relatively high
on. number of checklists or even floras. There exists
Next, the identification of species can be even a generic flora for the whole neotropics
(a) taxon-orientated by monographs or revisions (Gradstein et al. 2001). This book is a
of genera. A directory for monographs was pu- breakthrough for bryology in the neotropics. It
blished by Greene & Harrington (1988), which covers a short introduction to the anatomy,
is now slightly outdated. For mosses, all relevant morphology and systematics of bryophytes, a
literature is found in the TROPICOS database at description of the bryophyte regions with rele-
the Missouri Botanical Garden vant literature, a description of habitats with
(www.mobot.org). representative species and genera, a list of floras
(b) flora-orientated. An account of the floristic and a comprehensive bibliography and - as the
literature of the world was given by Grrene & mayor part - keys for and descriptions of hepatic
Harrington (1989). The most relevant tropical and moss genera with illustrations of
floras are listed below. Also checklists may be representative species. A comparable book for
consulted . tropical Africa is under preparation by the
Tropical Bryology Working Group of the British
Greene, S.W., Harrington, A.J. 1988. The Con- Bryological Society.
spectus of Bryological Taxonomic Li-
terature Part 1. Index to monographs and Allen, B. 1994. Moss Flora of Central America
regional reviews. Bryophytorum Biblio- Part 1. Sphagnaceae - Calymperaceae.
theca 35. Missouri Bot. Garden.

108 Frahm
Allen, B. 2002. Moss Flora of central America Group: http://britishbryologicalsociety.org.uk.

Part 2. Encalyptaceae - Orthotrichaceae. Hepatics
Missouri Bot. Garden. • Key to African Caudalejeunea (Van-
Bartram, E.B. 1949. Mosses of Guatemala. Fiel- den Berghen, 1984)
diana (Botany) Chicago. • Key to tropical African Frullania
Buck, W.R. 1998. Pleurocarpous Mosses of the (Vanden Berghen, 1976)
West Indies. Memoirs of the New York • Key to African species of Jungerman-
Botanical garden vol. 82, 400 pp. nia (Vána, 1974-5)
Churchill, S.P. 1994. The Mosses of Amazonian Mosses
Ecuador. AAU reports 35, Aarhus. • Key to families of African pleurocarps
Churchill, S.P., Linares, E.L. 1995. Prodromus (Petit, 1978)
Bryologiae Novo-Granatensis. 2 vols. • Keys to the Hookeriales of Africa
Bogotá. (Demaret & P. de la Varde, 1951-5)
Crum, H.A. and Steere, W.C. (1957). Mosses • Key to African Hypnaceae (Petit,
of Puerto Rico and the Virgin Islands. 1978)
N.Y. Academy of Sciences 7(4). [Useful • Key to genera of African Leucobrya-
for the Caribbean.] ceae (Onraedt, 1976)
Duarte-Bello, P. 1997. Musgos de Cuba. Ma- • Key to Regmatodontaceae of the
drid. 717 pp. world (Eakin, 1975)
Florschütz, P.A. 1964. Flora of Suriname, • Key to African Rhachitheciaceae
Musci, part I. (O’Shea, 1997)
Florschütz-de Waard, J. 1986. Flora of • Key to Rhacocarpaceae of the world
Suriname, Musci , part II. (Frahm, 1996)
Florschütz-de Waard, J. et al. 1996. Flora of • Key to African Rhizogoniaceae
Suriname, Musci , part III. Royal (O’Shea, 1997)
Botanic Gardens Kew. • Generic key to African Sematophylla-
Gradstein, S.R. (1989). A key to the Hepaticae ceae (O’Shea, 1999)
and Anthocerotae of Puerto Rico and • Key to tropical African Breutelia (De
the Virgin Islands. The Bryologist Sloover, 1975)
92(3): 329-348. [A key emphasising • Key to African species of Campylopus
vegetative characters for 237 species in (Frahm & O’Shea, 1996)
92 genera of liverworts and hornworts • Key to tropical African Leptodontium
recorded from Puerto Rico and the (De Sloover, 1987)
Virgin Islands; also useful for other • A key to African Leucophanes (Onra-
parts of tropical America.] edt, 1976)
Griffin, D. and Morales, M.I. (1983). Keys to • Key to African species of Neckera (De
the genera of mosses from Costa Rica. Sloover, 1977)
Brenesia 21: 299-323. [Useful key to • Key to most African species of Raco-
genera of Central America - over 200 mitrium
genera are dealt with.] • Key to tropical African Schoenobryum
Sharp, A.J., Crum, H.A., Eckel, P.M. 1994. The (Bizot & Pócs, 1982)
Moss Flora of Mexico. 2 vols. New
York Bot. Garden. Pócs, T. & O´Shea, B.J. 1991. Quick Reference
List of Basic Literature to Identify
Tropical African Bryophytes. Tropical
2.2 Tropical Africa Bryology 4: 69-854.
Compiled by Tamás Pócs and Brian O’Shea
Various keys are found on the webpage of the

British Bryological Society Tropical Bryology

HEPATICAE and ANTHOCEROTAE Andrewsianthus - Grolle, R. 1963:437; Vána,

J. 1980:228.
Checklists or floras with keys, illustrations or Aneura - Meenks, J.L.D. & Pócs, T. 1985:79.
descriptions.
Anomalolejeunea see under Cheilolejeunea
Arnell, S. 1956. Hepaticae collected by O. Aphanolejeunea - Pócs, T. 1984b:239.

Hedberg et al. on the East African Mountains.
Ark. f. Bot. 3:517-562. Arachniopsis see under LEPIDOZIACEAE/
Arnell, S. 1963. Hepaticae of South Africa. LEPIDOZIOIDEAE
Stockholm,411 pp.
Demaret, F. 1942. Prodrome des bryophytes du Archilejeunea - Vanden Berghen, C.
Congo belge et du Ruanda-Urundi. II.- 1951c:112.
Hepaticae. Bull. Jard. Bot. Bruxelles 16:287.
Grolle, R. 1978. Die Lebermoose der Bazzania - Arnell, S. 1965:66 (Madag-Masc.);
Seychellen. Wiss. Ztschr. Friedrich- Schiller- Jones, E.W. 1975:299, 1980:312; Onraedt, M.
Univ. Jena, Math.-Nat. R. 27:7-17. 1977:139 (Madag-Masc).
Jones, E.W. 1990. African Hepatics XL. An
artificial key to the genera of African Brachiolejeunea see most African species under
Hepatics. J. Bryol. 16:9-40. Frullanoides, but consult van Slageren, M.
Jones, E.W. & Harrington, A.J. 1983. The 1985.
hepatics of Sierra Leone and Ghana. Bull.
Brit. Mus. Nat. Hist. Botany 11:215-289. Bryopteris - Stotler, R.E. & Crandall-Stotler,
Vanden Berghen, C. 1972. Hépatiques et B. 1974:1.
anthocérotales. In Symoens,J.-J. (Ed.):
Résultats scientifiques. Exploration Calycularia - Jones, E.W. 1985b:497.
hydrobiologique du bassin du Lac
Bangweolo et du Luapula. Vol.8, fasc.1. Calypogeia - Bischler, H. 1970:63, Jones,E.W.
Hépatiques et Anthocérotales. 202 pp. 1976b:43.
Vanden Berghen, C. 1978. Hépatiques du
Shaba. Corrections et additions. Bull. Jard. Capillolejeunea - Arnell, S. 1965:69.
Bot. Nat. Belg. 48:367-372.
Caudalejeunea - Vanden Berghen, C.
1948:764; 1972:432; 1984:99; Jones, E.W.
1953c:164.
Reference list according to taxa
Cephalojonesia - Grolle, R. & Vanden
Acanthocolea - Kruijt, R.H.C. 1988:1. Berghen, C. 1970b:764; Jones, E.W. 1987:503.
Acrolejeunea - Gradstein, S.R. 1975:1. Cephalozia - Vána, J. 1988:179.
Adelanthus - Grolle, R. 1972:325. Cephaloziella - Jones, E.W. 1958:430.

(lowland species).
Allisoniella - Vána, J. 1985:86.
Ceratolejeunea - Vanden Berghen, C.
Alobiellopsis - Schuster, R.M. 1969:682. 1951a:61; 1953:279; 1973:381.
Anastrophyllum - Vána, J. 1982:72. Chaetolejeunea see under Rectolejeunea

110 Frahm
Chandonanthus - Vanden Berghen, C. Cololejeunea/Pedinolejeunea - Tixier, P.

1965:135. 1985:15.
Cheilolejeunea - Grolle, R. 1979:343; Jones, Cololejeunea/Taeniolejeunea - Benedix, E.H.

E.W. 1954b:380, 1973:548, 1976b:49, 1982:37. 1953; Jones, E.W. 1968b:569.
Cheilolejeunea/Anomalolejeunea - Vanden Colura - Jones, E.W. & Pócs, T. 1987:495;

Berghen, C. 1951d:364, 1953:278. Jovet-Ast, S. 1953:206; 1954:1; 1956:272;
1958:19; 1976:909; 1980:277.
Cheilolejeunea/Euosmolejeunea - Jones, E.W.
1954a:375, 1973:551; Vanden Bergh-en, C. Conoscyphus - Engel, J.J. 1987:533; Piippo, S.
1965:148. 1985:129.
Cheilolejeunea/Strepsilejeunea - Jones, E.W. Cyathodium - Jones, E.W. 1952:55; Jovet-Ast,

1985a:395, 1988:149; Vanden Berghen, C. S. 1970:57.
1960:133.
Cylindrocolea - Jones, E.W. 1976b:46.
Chiloscyphus - Grolle, R. 1984:505; Jones,
E.W. 1953d:172. Dicranolejeunea - Jones, E.W. 1970a:72.
Chonecolea - Jones, E.W. 1985b:498. Diplasiolejeunea - Jones, E.W. 1973:552;

Tixier, P. 1977:117; 1980:743; 1984:11;
Cladolejeunea - Zwickel, W. 1933:112. 1987:219; Vanden Berghen, C. 1960:119;
1977:216.
Clasmatocolea - Engel, J.J. 1980:1; Grolle, R.
1960:78.; Grolle, R. & Vanden Berghen, C. Diplasiolejeunea/Pellucidae - Jones, E.W.
1970:385; Jones, E.W. 1987b:503. 1954c:393; Tixier, P. 1985:331.
Cololejeunea - Jones, E.W. 1954e:408; Pócs, Drepanolejeunea - Jones, E.W. 1968b:567;

T. 1975:353; 1980:305; 1985:113; Tixier, P. Vanden Berghen, C. 1961:61.
1977:173 (Madagascan spp.); 1979:602.
Drepanolejeunea/Kolpolejeunea - Grolle, R.
Cololejeunea - species with acute leaves - 1976:191.
Vanden Berghen, C. 1977:235.
Euosmolejeunea see under Cheilolejeunea
Cololejeunea - species with dentate leaves -
Jones, E.W. 1953b:158. Evansiolejeunea - Vanden Berghen, C.
1965:151.
Cololejeunea - species with hyaline leaf
margin - Jones, E.W. 1953a:144; Vanden Exormotheca - Schiffner, V. 1942:40.
Berghen, C. 1972:467.
Frullania - Vanden Berghen, C. 1976c:1;
Cololejeunea - species with papillate leaf 1976b:335; 1982:207 (Madagascan spe-cies).
margin - Vanden Berghen, C. 1977:232.
Cololejeunea - species with papillose dorsal Frullanoides - van Slageren, M. 1985. (covers
leaf surface - Vanden Berghen, C. 1977:239. previous African Brachiolejeu-nea); Jones, E.W.
1968b:565; Vanden Berghen, C. 1951b:87;
Cololejeunea/Lasiolejeunea - Tixier, P. 1978:124.
1985:177.
Gongylanthus - Jones, E.W. 1964:649.

Gottschea see under Schistochila Lejeunea/Inflatolejeunea - Vanden Berghen, C.

1965:147.
Gottschelia - Grolle, R. 1968:13.
Lejeunea/Microlejeunea see under
Gymnocoleopsis - Vána, J. 1982:79. Microlejeunea
Gymnomitrion or -um - Vána, J. 1985:88. Lejeunea/Pleurolejeunea - Jones, E.W.

1969:787.
Harpalejeunea - Arnell, S. 1965:75.
Lejeunea/Umbilicatae (species with eplicate
Hygrolejeunea see under Lejeunea perianth) - Jones, E.W. 1967:299.
Inflatolejeunea see under Lejeunea LEJEUNEACEAE - Schuster, R.M. 1963.

(generic key with annotations).
Isotachis - Vána, J. 1982:63; Vanden Berghen,
C. 1965:130. LEJEUNEACEAE/PTYCHANTHOIDEAE -
Gradstein, S.R. 1985:13.
Iwatsukia - Grolle, R. & Onraedt, M. 1974:232;
Vána, J. 1980:233. Lepidozia see under LEPIDOZIACEAE/
Jamesoniella - Grolle, R. 1971:1. LEPIDOZIOIDEAE
Jungermannia - Vána, J. 1974:277: 1975:357. LEPIDOZIACEAE/LEPIDOZIOIDEAE -

Pócs, T. 1984a. (discusses nomenclature & dis
Kurzia see under LEPIDOZIACEAE/ tribution of African Arachniopsis, Kurzia,
LEPIDOZIOIDEAE Lepidozia, Psiloclada, Sprucel-la and Telaranea).
Lejeunea - Jones, E.W. 1979:389; 1984:159; Leptocolea see Cololejeunea

1985a:385.
Leptolejeunea - Vanden Berghen, C. 1961:58;
Lejeunea - L. eckloniana complex - Jones, 1963(1964):49; 1977:213.
E.W. 1974b:77.
Leptoscyphus - Grolle, R. 1962:1; Jones, E.W.
Lejeunea - L. flava complex - Jones, E.W. 1953d:196.
1968a:548.
Leucolejeunea - Jones, E.W. 1973:545.
Lejeunea - L. caespitosa group (small,
monoecious species) - Jones, E.W. 1972:36. Lophocolea see under Chiloscyphus
Lejeunea - (small, dioecious species) - Jones, Lopholejeunea - Vanden Berghen, C.

E.W. 1972:23; 1989:665. 1978:123; 1984a:393.
Lejeunea/Chaetolejeunea see under Lophozia - Vána, J. 1982:80.

Rectolejeunea
Marchantia - Vanden Berghen, C. 1954:37.
Lejeunea/Cladolejeunea see under
Cladolejeunea MARCHANTIALES - Vanden Berghen, C.
1965:166 (generic key).
Lejeunea/Hygrolejeunea - Vanden Berghen, C.
1961:65; 1972:438; 1977:202. Marchesinia - Jones, E.W. 1970a:78; Vanden
Berghen, C. 1976a:926.

112 Frahm
Marsupella - Vána, J. 1985:91. Prionolejeunea - Vanden Berghen, C.

1952:170; 1973:383; 1977:209.
Mastigolejeunea - Vanden Berghen, C.
1948a:49; 1951b:90. (sub Brachiolejeunea Ptychanthus - Vanden Berghen, C. 1951e:64.
nigra).
Ptychocoleus see under Schiffneriolejeunea
Mastigophora - Vanden Berghen, C.
1973:365. Pycnolejeunea - Grolle, R. 1979a:179; Jones,
E.W. 1979:397.
Metzgeria - Arnell, S. 1953:107 (S.Afr. spp.);
Kuwahara, Y. 1973:566; 1986:1 (although deals Radula - Jones, E.W. 1977:461; Yamada, K.
with Neotropic species, many records of African 1975:115.
taxa); Vanden Berghen, C. 1948:187;
1960:111. Rectolejeunea - Jones, E.W. 1974a:71.
Microlejeunea - Jones, E.W. 1969:775; Rectolejeunea/Chaetolejeunea - Jones, E.W.

1979:394; Jovet-Ast, S. 1959:191; Vanden 1969:784.
Berghen, C. 1965:143; 1977:204.
Riccardia - Jones, E.W. 1956:74; Meenks, J.
Nardia - Grolle, R. 1964:297. & Pócs, T. 1985:81.
Nesolejeunea see under Lejeunea Riccia - Jones, E.W. 1957b:210; Gillet, H. &
Jovet-Ast, S. 1957:62; Jovet-Ast, S. 1986:287
Notothylas - Hässel de Menendez, G. 1976:19. (N-Afr).
Odontolejeunea - Vanden Berghen, C. Ricciocarpus - Jones, E.W. 1957b:209.

1963(1964):52; Teeuwer, M. 1989:1.
Scapania - Arnell, S. 1957:26.
Odontoschisma - Grolle, R. 1960b:207.
Schiffneriolejeunea - Jones, E.W. 1954d:396;
Otolejeunea - Grolle, R. 1985:45; Tixier, P. 1982:45; Vanden Berghen, C. 1976a:925.
1980a:607.
Schiffneriolejeunea/Pappeanae - Gradstein,
Oxymitra - Garside, S. 1958:83 (all species S.R. & Vanden Berghen, C. 1985:173.
considered).
Schistochila - Grolle, R. & Zijlstra, G. 1984:87;
Pallavicinia - Grolle, R. 1984:508; Vanden Jones, E.W. 1976a:33.
Berghen, C. 1965:164.
Sprucella - Vanden Berghen, C. 1946:91;
Paraschistochila see under Schistochila 1983:321.
Plagiochasma - Bischler, H. 1978:223. Stenorrhipis - Grolle, R. 1963:441; Jones, E.W.

1968b:565.
Plagiochila - Jones, E.W. 1962:254; 1980:13;
Vanden Berghen, C. 1981:41. Stictolejeunea - Gradstein, S.R. 1985:195;
Jones, E.W. 1976b:50.
Pleurolejeunea see under Lejeunea
Strepsilejeunea see under Cheilolejeunea
Porella - Jones, E.W. 1963:446.

Symbiezidium - Gradstein, S.R. & van Beek, Gradstein, S.R. 1975. A taxonomic monograph
J. 1985:221. of the genus Acrolejeunea (Hepati-cae), with
an arrangement of the genera of
Symphyogyna - Grolle, R. 1980:330; Vanden Ptychanthoideae. Bryoph. Bibl. 4:1-216.
Berghen, C. 1965:156. Gradstein, S.R. 1985. Contribution to a
Monograph of the Lejeuneaceae, Subfam-ily
Syzygiella - Jones, E.W. 1976b:48; Vána, J. Ptychanthoideae. Beih. Nova Hedwigia 80:1-
1985:81. 253.
Gradstein, S.R. & van Beek, J. 1985. A revision
Taxilejeunea - Grolle 1974:93; Jones, E.W. of the genus Symbiezidium Trevis. in
1967:289; 1976b:50. Gradstein, S.R. 1985. Contribution to a
Monograph of the Lej-euneaceae, Subfamily
Thysananthus - Vanden Berghen, C. 1950:35. Ptychanthoideae. Beih. Nova Hedwigia
80:221-253.
Tritomaria - Vána, J. 1982:78. Gradstein, R.S. & Vanden Berghen, C. 1985.
Schiffneriolejeunea sect. Pappeanae en
Tylimanthus - Jones, E.W. 1980:317. Afrique. in Gradstein, S.R. 1985.
Contribution to a Monograph of the
Lejeuneaceae, Subfamily Ptychanthoideae.
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Genus Cheilolejeunea. Trans. Brit. Bryol. Some species of Lejeunea. J. Bryol. 7:23-46.
Soc. 2:380-392. Jones, E.W. 1973. African Hepatics XXIV.
Jones, E.W. 1954c. African Hepatics VIII. Lejeuneaceae: some new or little- known
Diplasiolejeunea albifolia (Taylor) E.W. species and extensions of range. J. Bryol.
Jones comb. nov. Trans. Brit. Bryol. Soc. 7:545-562.
2:393-395.

Jones, E.W. 1974a. African Hepatics XXV. Lejeuneaceae, Diplasiae (supplement). Rev.
Rectolejeunea. J. Bryol. 8:71-76. Bryol. Lichénol. 23:1-22.
Jones, E.W. 1974b. African Hepatics XXVI. Jovet-Ast, S. 1956. Deux Colura nouveaux de
The Lejeunea ecklonia complex. J. Bryol. Madagascar. Rev. Bryol. Lichénol. 25:272-
8:77-92. 276.
Jones, E.W. 1975. African Hepatics XXVII. Jovet-Ast, S. 1958. Localités nouvelle de
Bazzania. J. Bryol. 8:299-327. diverses espèces du genre Colura. Rev. Bryol.
Jones, E.W. 1976a. African Hepatics XXVIII. Lichénol. 27:19-23.
Schistochila Dum. J. Bryol. 9:33- 42. Jovet-Ast, S. 1959. Un Microlejeunea nouveau
Jones, E.W. 1976b. African Hepatics XXIX. de l’Isle de la Réunion. Rev. Bryol. Lichénol.
Some new or little-known species and 27:191-194.
extensions of range. J. Bryol. 9:43-55. Jovet-Ast, S. 1970. Cyathodium africanum Mitt.
Jones, E.W. 1977. African Hepatics XXX. The au Yémen et en Afrique. Rev. Bryol.
genus Radula Dumortier. J. Bryol. 9:461- Lichénol. 37:57-62.
504. Jovet-Ast, S. 1976. Compléments à la
Jones, E.W. 1979. African Hepatics XXXI. Rare connaissance des Colura: espèces et localités
or little-known Lejeuneaceae and extensions nouvelles. Rev. Bryol. Lichénol. 42:909-922.
of range. J. Bryol. 10:387-400. Jovet-Ast, S. 1980. La section Oidocorys S. J.-
Jones, E.W. 1980. African Hepatics XXXII. A. du genre Colura Dum. est- elle âgée de
Some little known species and exten-sions plus de 100 millions d’années? Cryptogamie,
of range. J. Bryol. 11:311-324. Bryol. Lichénol. 1:277-288.
Jones, E.W. 1982. African Hepatics XXXIII. Jovet-Ast, S. 1986. Les Riccia de la région
Some new Lejeuneaceae. J. Bryol. 12:37-49. méditerranéene. Cryptogamie, Bryol.
Jones, E.W. 1984. Notes on the lobule in the Lichénol. 7, Suppl. fasc. 3:287-427.
Lejeuneoideae. Cryptogamie, Bryol. Kruijt, R.H.C. 1988. A Monograph of the
Lichénol. 5:159-172. Genera Dicranolejeunea and Acanthoco-lea.
Jones, E.W. 1985a. African Hepatics XXXIV. Bryoph. Bibl. 36:1-136.
Some little-known or new Lejeuneac-eae. J. Kuwahara, Y. 1973. Further Notes on the
Bryol. 13:385-398. Production of Vegetative Thallus Structures
Jones, E.W. 1985b. African Hepatics XXXV. by Female Involucres of Metzgeria and a
Some new or little-known species and some New Species of Metzge-ria. The Bryologist
noteworthy records. J. Bryol. 13:497-508. 76:566-570.
Jones, E.W. 1987. African Hepatics XXXVII. Kuwahara, Y. 1986. The Metzgeriaceae of the
Some little-known species and extensions of Neotropics. Bryoph. Bibl. 28:1- 254.
range. J. Bryol. 14:503-510. Meenks, J.L.D. & Pócs, T. 1985. East African
Jones, E.W. 1988. African Hepatics XXXVIII. Bryophytes IX. Aneuraceae. Ab-stracta Bot.
Cheilolejeunea subgenus Strepsile-jeunea 9:79-98.
(Spruce)Schust., with special reference to Onraedt, M. 1977. Bryophytes des îles
East Africa. J. Bryol. 15:149-160. mascareno-malgaches et Seychelles III.
Jones, E.W. 1989. African Hepatics XXXIX. Hepaticae: Bazzania. Bull. Jard. Bot. Nat.
Some dioecious species of Lejeunea. J. Bryol. Belg. 47:139-144.
15:665-674. Piippo, S. 1985. Bryophyte Flora of the Huon
Jones, E.W. & Pócs, T. 1987. African Hepatics Peninsula, Papua New Guinea, XII.
XXXVI. Three new species of Colura. J. Geocalycaceae (Hepaticae). Acta Bot. Fenn.
Bryol. 14:495-502. 131:129-167.
Jovet-Ast, S. 1953. Le genre Colura. Pócs, T. 1975. New or little-known epiphyllous
Hépatiques, Lejeuneaceae, Diplasiae. Rev. liverworts I. Cololejeunea from tropical
Bryol. Lichénol. 22:206-312. Jovet-Ast, S. Africa. Acta Bot. Acad. Sci. Hung. 21:353-
1954. Le genre Colura. Hépatiques, 375.

116 Frahm
Pócs, T. 1980. New or little known epiphyllous Endèmisme et spéciation. Nova Hedwigia
liverworts II. Three new Co-lolejeunea from 34:743-767.
East Africa. J. Hattori Bot. Lab. 48:305-320. Tixier, P. 1984. Contribution à l’étude du genre
Pócs, T. 1984a. Synopsis of the African Diplasiolejeunea (Spruce)Schiffner, 4. La
Lepidoziaceae. Proc. Third Meeting Bryol. section Villaumeae P.Tx. (Subgenus
Centr. East Eur.,Praha 107-119 (discusses Diplasiolejeu-nea) sur la côte est de
nomenclature & distribu-tion of African Madagascar. Acta Bot. Hung. 30:11-26.
Arachniopsis, Kurzia, Lepidozia, Psiloclada, Tixier, P. 1985. Contribution a la Connaissance
Sprucella and Telaranea). des Cololejeunoideae. Bryoph. Bibl. 27:1-
Pócs, T. 1984b. New or little known epiphyllous 440.
liverworts III. The genus Aphanolejeunea Tixier, P. 1987. La notion d’espèce chez le genre
Evans in Tropical Africa. Cryptogamie, Diplasiolejeunea 4. Diplasi-olejeunea
Bryol.Lichénol. 5:239-268. symoensii Vand.Bergh. Cryptogamie,
Pócs, T. 1985. East African Bryophytes, VII. Bryol.Lichénol. 8:219-226.
The Hepaticae of the Usambara Rain Forest Vána, J. 1974. Studies über die
Project Expedition, 1982. Acta Bot. Hung. Jungermannioideae (Hepaticae). 5.
31:113-133. Jungermannia Subg. Plectocolea and Subg.
Schiffner, V. 1942. Monographie der Gattung Solenostoma: Afrikanishe Arten. Folia
Exormotheca. Hedwigia 81:40-74. Geobot. Phytotax. 9:277-312.
Schuster, R.M. 1963. An Annotated Synopsis of Vána, J. 1975. Studien über die
the Genera and Subgenera of Lejeuneaceae Jungermannioideae (Hepaticae). 9.
I: Introduction, annotated keys to subfamilies Jungermannia subgen. Plectocolea und
and genera. Beih. Nova Hedwigia 9:1-204. subgen. Solenostoma in Hawaii:
Schuster, R.M. 1969. Studies on Hepaticae Ergänzungen und Synopsis der Gattung
XLVI-XLVII. On Alobiella (Spr.)Schiffn. Jungermannia. Folia Geobot. Phytotax.
and Alobiellopsis Schust. Bull. Nat. Sci. Mus. 10:357-382.
Tokyo 12:659-683. Vána, J. 1980. Some new South and Central
Stotler, R.E. & B.Crandall-Stotler 1974. A American hepatics. J. Hattori Bot. Lab.
Monograph of the Genus Bryopteris 48:225-234.
(Swartz)Nees van Esenbeck. Bryoph. Bibl. Vána, J. 1982. Notes on some African hepatic
3:1-159. genera. 1-5. Fol. Geobot. Phyto-tax. 17:63-
Teeuwer, M. 1989. Nova Hedwigia 48:1-32. 87.
[Odontolejeunea] Vána, J. 1985. Notes on some African hepatic
Tixier, P. 1977a. Espèces nouvelles malgaches genera, 6-9. Fol. Geobot. Phyto-tax. 20:81-
du genre Diplasiolejeunea (Spruce)Schiffn. 99.
(Hepaticae). Lindbergia 4:117-126. Vána, J. 1988. Cephalozia (Dum.)Dum. in
Tixier, P. 1977b. La famille des Africa, with notes on the genus (Notes on
Cololejeuneoideae (Grolle) dans l’Océan some African hepatic genera 10). in Engel,
Indien occidental. Essai monographique. J.J. and Hattori, S. Bryo-logical
Bull. trim. Acad. Malg. 55:173- 247. Contributions Presented in Celebration of the
Tixier, P. 1979. Contribution to the Knowledge Distinguished Schol-arship of Rudolf M.
of the Genus Cololejeunea VIII. Some New Schuster. Beih. N. Hedw. 90:179-198.
Species of Malagasy Cololejeunea. The Vanden Berghen, C. 1946. Note sur Sprucella
Bryologist 82:602-608. succida (Mitt.)Steph. au Congo Belge. Bull.
Tixier, P. 1980a. Deux nouveaux genres de Jard. Bot. Bruxelles 18:91-95.
Lejeunéacées: Otolejeunea Grolle & P.Tx. Vanden Berghen, C. 1948a. Note sur le genre
et Allorgella P.Tx. Nova Hedwigia 32:607- Ptychocoleus Trev. en Afrique tropicale.
622. Bull. Jard. Bot. Bruxelles 19:37-49.
Tixier, P. 1980b. Diplasiolejeunea insigne P.Tx. Vanden Berghen, C. 1948b. Contribution à
à Angaroleky (Tanarive, Madagascar) l’étude des espèces africaine du genre

Metzgeria. Bull. Jard. Bot. Bruxelles 19:187- tanganyikaise. Bull. Soc. Roy. Bot. Belg.
204. 98:129-174.
Vanden Berghen, C. 1948c. Rev. Bryol. Vanden Berghen, C. 1972. Hépatiques
Lichénol. 17:764- [Caudalejeunea]. épiphylles récoltées au Burundi par L.
Vanden Berghen, C. 1950. Le genre Lewalle. Bull. Jard. Bot. Nat. Belg. 42:431-
Thysananthus Lindenb. en Afrique. Rev. 494.
Bryol. Lichénol. 18:35-37. Vanden Berghen, C. 1973. Quelques hépatiques
Vanden Berghen, C. 1951a. Contribution à récoltées au Gabon par G. le Testu. Rev.
l’étude des espèces africaine du genre Bryol. Lichénol. 39:365-385.
Ceratolejeunea (Spruce)Schiffn. Bull. Jard. Vanden Berghen, C. 1976a. Deux Lejeunéacées
Bot. Bruxelles 21:61-81. Holostipées nouvelles pour la flore Africaine.
Vanden Berghen, C. 1951b. Contribution à Rev. Bryol. Lichénol. 42:923-929.
l’étude des espèces africaine du genre Vanden Berghen, C. 1976b. Distributiones
Brachiolejeunea (Spruce)Schiffn. Bull. Jard. Plantarum Africanarum 11:335-372
Bot. Bruxelles 21:87- 94. Vanden Berghen, C. 1976c. Frullaniaceae
Vanden Berghen, C. 1951c. Contribution à (Hepaticae) africanae. Bull. Jard. Bot. Nat.
l’étude des espèces africaine du genre Belg. 46:1-220.
Archilejeunea (Spruce)Sciffn. Rev. Bryol. Vanden Berghen, C. 1977. Hépatiques
Lichénol. 20:112-121. épiphylles récoltées par J.L. de Sloover au
Vanden Berghen, C. 1951d. Note sur quelques Kivu (Zaire), au Rwanda et au Burundi. Bull.
hépatiques récoltées par R.E. & T. Fries en Jard. Bot. Nat. Belg. 47:199-246.
1922, au Mont Kénia. Svensk Bot. Tid. Vanden Berghen, C. 1978. Notes sur quelques
45:362-367. Lejeunéacées holostipées afri-caines. Rev.
Vanden Berghen, C. 1951e. Note sur trois Bryol. Lichénol. 44:123-132.
hépatiques du Congo belge. Bull. Soc. Roy. Vanden Berghen, C. 1981. Le genre Plagiochila
Bot. Belg. 84:61-64. (Dum.)Dum. (Hepaticae) à Mada-gascar et
Vanden Berghen, C. 1952. Note sur quelques aux Mascareignes, principalement d’après les
Lejeunéacées de l’Afrique continen-tale. récoltes de M. Onraedt. Bull. Jard. Bot. Nat.
Bull. Jard. Bot. Bruxelles 22:165-175. Belg. 51:41-103.
Vanden Berghen, C. 1953. Quelques hépatiques Vanden Berghen, C. 1982. Le genre Frullania
récoltées par O. Hedberg sur les montagnes Raddi (Hepaticae) à Madagascar. Récoltes de
de l’Afrique orientale. Svensk Bot. Tid. P. Tixier. Cryptogamie, Bryol. Lichénol.
47:263-283. 3:207-224.
Vanden Berghen, C. 1954. Le genre Vanden Berghen, C. 1983. Lepidozia Dum.
Marchantia L. au Congo Belge. Bull. Jard. emend. Joerg. subgen. Sprucella (Steph.)
Bot. Bruxelles 24:37-50. Vanden Berghen comb. et stat. nov.
Vanden Berghen, C. 1960. Hépatiques récoltées (Hepaticae). Bull. Jard. Bot. Nat. Belg.
par le Dr. J.J. Symoens dans la région péri- 53:321-330.
tanganyikaise. Bull. Soc. Roy. Bot. Belg. Vanden Berghen, C. 1984a. Le genre
92:111-138. Lopholejeunea (Spruce)Schiffn.
Vanden Berghen, C. 1961. Hépatiques récoltées (Lejeuneac-eae, Hepaticae) en Afrique. Bull.
par le Dr. J.J. Symoens dans la région péri- Jard. Bot. Nat. Belg. 54:393-464.
tanganyikaise. Bull. Soc. Roy. Bot. Belg. Vanden Berghen, C. 1984b. Le genre
93:55-74. Caudalejeunea (Steph.)Schiffn.
Vanden Berghen, C. 1963(1964). Lejeunéacées (Lejeuneac-eae, Hepaticae) en Afrique.
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Vanden Berghen, C. 1965. Hépatiques récoltées monograph of the genera Brachiolejeunea
par le Dr. J.J. Symoens dans la région péri- and Frullanoides (Hepaticae). Thesis
published by the author, Utrecht, 309 pp.

118 Frahm
Yamada, K. 1975. Notes on Radula from

Tanzania, East Africa. J. Jap. Bot. 50:115-
118.
Zwickel, W. 1933. Zwei neue Gattungen, einige
neue Arten und Umstellung bei den
Lejeuneaceen. Ann. Bryol. 6:105-122.

APPENDIX IV: TECHNIQUES OF TAXONOMIC REVISIONS
Taxonomy is the basis of biology! Even in a collected in different parts e.g. in the neotropics
period when systematics and taxonomy are during different expeditions e.g. in Brazil,
regarded as secondary, it must be understood that Colombia, Bolivia and Mexico, and every time
taxonomic knowledge is necessary. No described as new. Later, bryologists had not the
physiologists or molecular biologist can work knowledge to revise the species worldwide but
without the circumscription of the species he is they were rather specialized on countries or
working on, no ecology is possible without the continents (the French on Africa, the North
knowledge of the species living in an ecosystem. Americans on South America) and a species
The necessary knowledge is obtained by could be known from both continents under two
taxonomic revisions and monographs. They (or more) different names.
differ in the intensity of work. Revisions are
confined to a partricular geographic area, to the The rates of reduction are quite high, as seen from
necessary study of type material and in addition some examples in the Dicranaceae (see bottom
to representative material. Monographs are more page).
detailed and deal (especially in smaller genera)
with all available herbarium material. These reductions have also an effect on the rate
Revisions and monographs have an enormous of endemism, which is also decreasing, e.g. in
importance. Their results are needed to obtain the genus Campylopus.
the exact species number of a genus and they
provide the necessary basis for a The phytogeographical consequences of
phytogeographical research. revisions are enormous, as shown by the
Generally, much more species have been distribution maps of species of Microcampylopus
described than there really exist. This is the result and Campylopodium before and after a revision.
of a) a narrow species concept, especially in the Therefore phytogeographical interpretations of
19th century, when small modifications (e.g in unrevised genera must lead to fatal errors!
colour) were described as new species; b)
theological reasons, since the evolution theory General introductions to taxonomy can be
was not known or later not accepted in the 19th obtained from the existing textbooks (e.g. Stuessy
century and it was believed that God has created 1990). Therefore the script is confined here to
the species anew in every continent and every the techniques of taxonomic work, especially
island; and c) the wide ranges of bryophytes, because there are no instructions available.
which the result that the same species was

120 Frahm
Species described Species accepted Reduction %
Pilopogon 14 8 43
Atractylocarpus 19 9 53
Microcampylopus 27 3 89
Campylopodium 12 2 84
Campylopus
Africa 256 50 81
Neotropics 320 65 80
Subantarctis 74 14 81
Percentage of endemic Campylopus species:
Before Revision After Revision Flowering Plants

Hawaii 70 30 94
New Caledonia 50 0 90
Tab. IV.1: Numbers of species in genera and percentage of endemic species before and after taxono-
mic revisions.
1. Taxonomic revisions taxonomic methods were published by Frahm

(1989).
Revisions are often said to be „old fashioned“ as
they are mostly and primarily based on traditional In the business world, each process is highly
methods. However, we have to realize that we structured and wasted time is minimized.
are working with a group of organisms for which Taxonomic work can also be efficiently
there is no well-founded estimate of the total structured in much the same way. For that reason
number of species world-wide – due to the lack a working scheme has been derived for revisions
of generic revisions on a world-wide basis. prepared by students. This working scheme is
proposed and explained here. It shows that
A major problem is the revisions of genera with taxonomic revisions can be prepared in a
high numbers of species. It is clear that mostly reasonable time with the highest efficiency and
genera with relatively few species have been without delay caused by an incorrect organization
revised so far. Large genera are said to be time- of work.
consuming and difficult because of the large
number of specimens one has to examine, copious There are principally two methods of preparing
literature and taxonomic data. Therefore it is said taxonomic treatments:
that revising a genus of 30-50 species seems to (1) The specimen orientated method and
be impossible to do in a reasonable time, (2) The type specimen orientated method.
especially for students. However, this is a matter The first method starts a study from zilch. All
of efficiency. taxa already described are at first not considered;
God revisions can easily be made by interested only specimens are examined and these are
and motivated students, as is shown by many classified independently. After that, the taxa
publications prepared even as teachers exam or described are taken into account and compared
master‘s theses. The main problem is that a lot with the results of the initial classification.
of time will be wasted if the work is not well Theoretically this seems to be the most indepen-
organized. With this in mind, instructions to using dent way to produce a monograph or revision,

but is has practical disadvantages. It takes an copied in the library. The information on the type
enormous amount of time to obtain one´s ideas material is extracted and added to the taxonomic
of how to classify 3000 or 5000 specimens All file.
specimens from different herbaria must be
continously filed as new and sorted according to 5. Additionally, all available literature from the
the developing ideas. Clearly, using this method important geographical ranges is examined.
requires a great deal of taxonomic experience,
which a student usually does not have. Although 6. All literature is compiled in a literature file.
this method is free without from predisposition
and may give the best results in the end, I cannot 7. When the borrowed herbarium specimens
recommend it for beginners. arrive, all types are studied first. They are
examined and, if possible, permanent microscope
The second method is the type- (or better the type slides are prepared. („If possible“ means that
specimen) orientated method. This concerns sometimes the type specimens are too scanty to
revisionary studies based primarily on the study allow deatching any part of it permanently;
of type material. All other specimens are furthermore, some herbaria exclusively forbid
compared with the species concept derived from detaching and removing any parts of type
the type specimens. This seems to be the easier specimens.) All essential characters are illustrated
of the two methods and it is also the more or photographed and plates with figures are
commonly used. It also gives results in a prepared for each type specimen. If necessary,
reasonable time and is recommended for students. nomenclatural problems are studied, as for
instance lectotypifications.
The following is an outline of a working scheme
for mosses and consists of 12 steps as summarized 8. The labels of the type specimens and all other
in the figure. specimens received on loan are copied and glued
on file cards or the label information is typed into
1. Compile from Index Muscorum and its Sup- the computer (text or database). The file cards
plements a list of accepted species, with are arranged alphabetically. Notes can be added
synonyms on file cards or in a taxonomic file to the files or file cards during study.
(text file or database).
9. All illustrations of the types are arranged for
2. Compose a list of all authors of taxa (see Sayre easy access, for example by pinning them up on
1977 for the locations of herbaria). Then request the wall or a bulletin board. By comparing the
to borrow the necessary type specimens (plus all illustrations, possible synonyms can be detected.
additional material of that genus) from the These initial suspicions can be confirmed by
herbaria. If you are a beginner and your name is comparing the permanent slides.
not well-known (yet), it is advisable to have some
words of recommendation from your supervisor 10. For the species remaining at this stage of the
or professor. The addresses can be obtained from work, an identification key is prepared (again
„Index Herbariorum“, or, for bryophyte herbaria, with the help of illustrations and slides). During
from Vitt et al. (1985). this preparation, further synonyms may be
detected.
3. Look at the distribution data in the Index
Muscorum to determine which additional 11. Using this key, the non-type specimens from
herbaria should be taken into account in the herbaria are studied to determine whether they
requesting to borrow further material. fit into the species concept derived from the types.
In this manner, misidentified specimens can be
4. While waiting to receive the requested loans, revised, specimens belonging to other genera can
copies of the original descriptions (protologues) be excluded, or specimens representing
are ordered through the interlibrary system or undescribed species can be found.

122 Frahm
Fig. IV:1: Working Scheme for taxonomic revisions. From Frahm (1989).

12. In order to prepare the manuscript, the 2. Phenetic and Cladistic analysis
existing files and illustrations are simply merged
together, and descriptions are added. If you are Today, the classical revision is accomplished by
planning to publish the paper in a certain journal, either a phenetic or cladistic analysis, which
you should take into account the journal‘s style makes the interrelationships between taxa more
and format. clear.
A basic revision then consists of the following: A. Phenetics

a) taxonomic data of accepted species and
synonyms (with taxa, citations, and type A phenetic analysis is based on morphological
information) taken from the taxonomic file. and anatomical characters. The resulting
b) descriptions: this is provided with the help of dendrogram reflects the „phenetic distance“, that
the illustrations, microscope slides, and means the affinities between taxa based on overall
examination of the herbarium specimens to cover similarity.
the variation of characters. Special notes are taken In contrast to cladistic programs, no difference
from the comments on the specimen file. is made between apomorphous (derived) and
c) illustrations: this is taken in the case of several plesiomorphous (primitive) character states.
synonyms from the best or most typical one. If Therefore it is often argued that phenetic analysis
there is considerable character variation, at least show morphological similarity but not
some of it should be displayed in the illustrations. evolutionary trends; in other words, the groups
d) lists of specimens examined compiled from achieved through a phenetic analyses are not
the specimen file, and natural, because they do not accurately reflect
e) references, compiled from the literature file. the evolutionary history (speciation) of the group.
In fact, the results of phenetic and cladistic

Other methods used in taxonomy are biometrics, analyses of a given group are often very similar.
chemosystematics, cytology, cultures, SEM The reason is that similarities are often based on
techniques, techniques for illustrations, and homologies and homologhies have a common
knowledge of nomenclatural rules, which are not evolutionary origin. So similarity frequently can
dealt with here. The necessary knowledge can (but must not always) be based on common
be obtained from the general literature because origin.
it is principally very similar to taxonomic work
of other systematic groups, e.g. flowering plants. Phenetic analysis is performed with a cluster
Phenetic and cladistic analyses are discussed in analysis. The data for a cluster analysis have to
the next chapter. be prepared in a table form, in which species (in
rows) and character states (in columns) are listed.
Frahm, J.-P. 1989. Taxonomic methodology. (By the same way, species and relevées can be
Bryol. Times 89: 1-3. used in vegetation analysis)
Sayre, G. 1977. Authors of names of bryophytes
and the location of their herbaria. Bryologist 60: The variables (character expressions) can be
502-521. coded in several ways:
Stuessy, T.F. 1990. Plant taxonomy. The 1. Sequentially. All characters are listed and
systematic evaluation of comparative data. New numbered. The corresponding numbers are
York. entried in the table.
Vitt, D.H., Gradstein, S.R. & Iwatzuki, Z. 2. Presence/absence. For every character state,
1985. Compendium of bryology. Bryophyt. the expression is coded as 0 or 1.
Biblioth. 30: 1-335. The results are the same regardless of the style
of the table.

124 Frahm
Next, the similarity of species 1 against species Systematics“ in German, which was translated
2, 3, 4 and so on, are calculated, then 2 against 1, by others into English. Later, the original German
3, 4, and so on, species 3 against 1,2,4,5, and so manuscript was published. It is different in some
on. The similarity is calculated as percentage respects from the English translation. This first
similarity, various Similarity Indices (Jaccard, cladistic analyses were „hand made“ in contrast
Sörensen etc.), or Euclidean Distance. It is to later computer-assisted analyses.
important to know that Euclidean Distance cannot
be used for data matrices with 0 and 1. The method is principally based on the
Finally, the similarity indices are plotted against assumption of apomorphous (derived) and
each other in a dendrogram. The axis shows the plesiomorphous (primitive) characters states.
value of the similarity index. Most easily Only groups defined by apomorphous character
understandable is the percentage similarity. E.g., states are accepted, since only they are considered
if two species have 9 of 10 characters in common, monophyletic, i.e. sharing the most recent
they are linked in the dendrogram on the 90% common ancestor and including all descendants
level. of that ancestor. Due to recognition of
apomorphous and plesiomorphous characters
This method has the advantage that it is absolutely atates, the method is somewhat subjective,
objective and the resulting dendrogram illustrates although there are principles to determine which
very nicely the morphological similarity of the characters are apomorpous and plesiomorphous,
taxa. and therefore it is hypothetical (as also stated very
strictly by Hennig). Later the computerized
Programs for cluster analysis cladistic analysis was introduced as a „scientific“
approach to systematics.
MVSP (Multivariate Statistical Analysis)
Written by W.L. Kovach. It includes several For a better understanding of cladistic principles,
statistical methods such as correspondence performing a handmade cladistic analysis is
analysis etc. The price is about £ 80. A shareware strongly recommended.
version can be tested free and can be obtained
from the IAB software library by J.-P. Frahm.
This version can only deal with up to 100 varia- List of Computer Programs for Cladistic Analy-
bles. sis
STATISTICA Most of the programs are today outdated, since

A commercial Windows program, which includes these were DOS-programs which do nut run
many statistical features. Price about $900. anymore on an actual PC. The most commonly
used program is PAUP. It was written for Apple
computers and is accordingly easy to use. The
Windows version is not as user friendly; it is more
B. Cladistics or less the DOS version running in a window.
Therefore many labs, although they have
A cladistic analysis shall „reconstruct Windows-PCs, have an Apple computer for
phylogeny“. It is based on a similar data matrix runnning PAUP.
as the phenetic analysis, but the resulting
dendrogram does not show phenetic but Hennig86 (Farris 1988)
„patristic“ distance, that means distance in A relatively simple but very effective program
evolutionary terms. for finding most parsimonious trees with options
for tree manipulating, successive weighting, and
The cladistic method was founded by the Germ- various free statistics. It is a mainframe program
an entomologist Willi Hennig, who worked in compiled for PC users. It runs under DOS and is
the USA. He wrote a textbook on „Phylogenetic relatively out of date concerning the use of

Fig. IV.1: Phenetic, cladistic and molecular analysis of the Campylopodioideae (Dicranaceae). A.
Phenetic analysis (clusater analysis), B. Cladistic analysis using the program Treesearch, C. molecu-
lar analysis of cpDNA (ITS2), maximum parsimony analysis. A-B after Frahm, C. after Stech. All
trees give different results, there is no truth in reconstructing relationsships or phylogeny but diffe-
rent hypotheses.

126 Frahm
memory capacity. The handling of the program COMPONENT Vers. 2.0, 1993
is in the mainframe style and needs input of Written by Roderick M. Page as a Windows Ver-
commands. The data have to be prepared with a sion. It is comparable to MacClade and does not
text editor in a special way. The program is sold generate new trees but investigates and compares
by the author, J.S. Farris, Swedish Museum of trees. Available from the Natural History Muse-
Natural History, S 10405 Stockholm. At least um London at a price of about £ 80.
older versions have a non-graphical outprint. The
compendium by Lipscomb (1994) is highly TREESEARCH (Bruggeman)
recommended as a supplement to the manual. Written by the husband of Ida Bruggeman-
Nannenga for her thesis. A DOS-Program which
PAUP (Phylogenetic Analysis Using Parsimony) is relatively easily menu-operated. The data
An extensive program for finding most matrix must be prepared with a text editor. It has
parsimonous trees with options for morphological no real graphics output and is relatively slow but
and molecular data, tree statistics, bootstrapping, is available free. Copies can be obtained from
various concensus techniques etc. There is an the IAB software library and ordered from J.-P.
earlier version for PC from 1985 and a much more Frahm.
extensive version for Macintosh from 1993. The
old DOS version is not recommended. It needs a Other programs:
math-coprocessor to run. The program was CLINCH
written by D.L. Swofford and was distributed ClaDOS
originally by the Illinois Natural History Survey NONAme and Pee-Wee
at a price of $50, but is now commercially
distibuted by Cambridge University Press at a Of these programs, PAUP is recommended of
higher price, also as a Windows version. The the commercial and Treesearch of the public
windows technique in the Mac and Windows domain programs. The main problem with all
version makes it easy to use and gives high quality problems is that they need a special data format.
outprints of the cladograms. Therefore it is suggested to prepare own data by
„overwriting“ existing files (e.g. the demo files)
MacClade (Maddison & Maddison 1992). to avoid difficulties.
This program does not generate cladistic analyses
but allows to manipulate trees generated with A serious problem and a funny exercise is to
other programs such as PAUP. Only available „feed“ different cladistic programs with the same
for Macintosh computers. It comes with an easy data. The results will not at all be comparable!
data matrix generator. Therefore it is
recommended to use this program in conjunction Farris, J.S. 1983. The logical basis of
with PAUP to generate data matrices and analysis phylogenetic analysis. In: N.I. Platnick
of the trees generated by PAUP. & V.A. Funk (eds.), Advances in
Cladistics 2: 7-36.
Phylip (Phylogenetic Interference Package) Platnick, N.I. 1987 An empirical comparison of
This package contains a variety of phylogenetic microcomputer parsimony programs.
programs for molecular and morphological data, Cladistics 3: 121-144.
inclusively maximum likelihood, parsimony and Sober, E.R. 1983. Parsimony methods in
bootstrapping facilities. Versions for PC (DOS, systematics. In: N.I. Platnick & V.A.
not much recommended because of its difficult Funk (eds.) Advances in Cladistics 2:
operation) and Macintosh. Phylip runs slower 37-47.
than PAUP or Hennig86, but is, however, Wiley, E.G. 1981. Phylogenetics. J. Wiley &
available free of charge from the author J. Felsen- Sons, New York. 439 pp.
stein, Dept. of Genetics SK-50, University of
Seattle, Washington, Seattle, Washington 98195,
USA.

APPENDIX V: VEGETATION ANALYSIS (PHYTOSOCIOLOGAL METHOD AFTER

BRAUN-BLANQUET)
Vegetation analysis (in Europe unfortunately the tree-, herb- and cryptogam layer of boreal
called „plant sociology“) studies the composition forests is dominated by a few species. This
of plant communities. There are different method could, however, hardly be used in Central
approaches concerning the methods and results Europe. Therefore the swiss botanist Braun-
in North America and Europe. In North America, Blanquet developed a method using a modified
usually random plots are studied using statistical scale with 5 units and used character species and
methods. In Europe, estimations of the cover of differential species for classification of vegetation
species are made within a selected plot.The units. Character species are typical of only one
reasons for the different methods are historical. community (which is indicated by this species),
When plant sociology was established in the and differential species are species which can be
1920s, a hundred years long knowledge of the found in more than one vegetation units but
indicator values of the species existed and was differentiate a plant community from others in
common sense. So „typical“ species for a habitat the surroundings. The study plots are not selected
were used to characterize plant communties. At at random but they are chosen according to a
that time, many universities in North America principle. They must be typical, well-developed
did not even exist, and there was no knowledge and homogenous. The so called Braun-Blanquet
of the ecological indicator value of species before, method was said not to work in the tropics. This
on which studies could be based. In addition, is surely true for tropical forests, however, at the
North America is much larger and botanically end of the 1970s it was successfully applied by
much more diverse than Europe. Therefore a Dutch botanists in the páramos of Colombia.
botanist from California could do nothing else
than use statistical methods when he was working Bryophyte communities in the tropics were rarely
in Massachusetts. studied in the past. Jovet-Ast (1949) was the first
who studied epiphytic bryophyte communities in
In Europe there were originally different methods the French Antilles. Giacomini & Ciferi (1950)
for plant sociology developed. In the boreal described a terrestrial bryophyte community from
forests, the Finnish Hult and the Swedish Venezuela with the Braun-Blanquet method. Petit
Sernander estimated the cover of species by a & Simons (1974) described epiphytic bryophyte
scale of 10 units and classified plant communities communities from Burundi. During the
by dominance of species, which was easy, since BRYOTROP project, studies of the epiphytic

128 Frahm
bryophytes were originally made along the Peru Field practice

transect with plastic sheets, which were wrapped
around the trunks. The outlines of bryophyte The methods described are here applied to the
cushions were marked with a pen and later the study of bryophyte communities. Several items
cover and combination of species were evaluated such as phenology or stratification, which are
in the lab (Frahm 1987). However, this method only important for flowering plant communities,
failed in Borneo, since the bryophytes were too are omitted.
humid and water condensed below the plastic.
Therefore the Braun-Blanquet method was used 1. Selection of study plot
(Kürschner 1990) and proved to be very
successful, since tropical bryophyte communities The study plot must be homogenous.
do not differ in their structure from temperate Homogenity can be determined with some
bryophyte communities. Later this method was experience from the aspect or by statistical
used again in Central Africa (Kürschner 1995). methods, e.g. a line taxation or a frequency
analysis.
Frahm, J.-P. 1987. Struktur und Zusammen- The size can also be determined by experience .
setzung der epiphytischen Moos- The right size determines whether all species of
vegetation in Regenwäldern NO-Perus. a community are included or not. If the area is
Beih. Nova Hedwigia 88: 115-141. too small, not all species are included.The size
Giacomini, V. & Ciferri, R. 1950. can be determined by a mininum area curve
Un’associazione crittogamica a (species number per area curve). Species numbers
Polytrichadelphus e Cora (Coreto- are determined on a small scale (e.g. 5 x 5 cm),
Polytrichadelphetum ciferrii) su rocce then the area is doubled and the species numbers
della Foresta delle nebbie in Venezue- are determined again, and so on. The species
la. Atti Ist. Vot. Univ. Pavia, Ser. 5, 9: numbers per area are plotted on a graph. The
211-217. minimum area is reached when the curve
Jovet-Ast, S. 1949. Les groupements des becomes a horizontal line, meaning the species
muscinées épiphytes aux Antilles number is not increasing anymore.
françaises. Revue Bryol. Lichénol. 18, The size for bryophyte communities is between
3-4: 125-146. 5 x 5 cm (epiphyllous communties) and ca. 20 x
Kürschner, H. 1990. Die epiphytischen Moos- 30 cm (epiphytic communities) or even
gesellschaften am Mt. Kinabalu (Nord considerably larger (terrestrial communities). It
Borneo, Sabah, Malaysia). Nova is adapted to the size of the substrate (leaf, branch,
Hedwigia 51: 76-85. trunk). It needs not to be a square but can be a
Kürschner, H. 1995. Wissenschaftliche Ergeb- rectangle of any dimensions. The circumference
nisse der BRYOTROP Expedition nach of the study plot maybe marked with tape or wire.
Zaire und Rwanda 4. Epiphytische
Moosgesellschaften im östlichen 2. Analysis
Kongobecken. Nova Hedwigia 61: 1-64.
Petit, E. & Symoens, F. 1974. Les bryophytes For each analysis, several parameters are noted:
des bois artificiels de Cupressus et date
d’Acacia au Burundi. Analyse description of the locality with exact indication
factorielle de la vegetation of the location
bryophytique. Bull. Jard. Bot. Nat. Belg. elevation
44: 219-247. subsequent number (or a combination letter/digit,
Pócs, T. 1978. Epiphyllous communities and e.g. A1, to indicate different analyses in dif-
their distribution in East Africa. Bryo- ferent forests, different student groups etc.)
phyt. Biblioth. 13: 681-713. size of the study plot (square centimeters or better
length x width)
exposition (use compass)

inclination (if no clinometer is available, a large a) horizontally by arranging all species in the
protractor with a perpendicel can be used) sequence according to their cover, e.g. the species
total cover of bryophytes, eventually also that of with high cover at top,
lichens and phanerogams b) vertically. This arrangement groups similar
special notes can be made concerning other study plots together, e.g. separates different
factors, e.g. the height of the tree, the diameter communities or divides a community in subunits
of the trunk, the forest type, illumination, (e.g. wet and dry expressions, expressions from
density of canopy etc. vertical or inclined trees etc.). The table can also
all species within the study plot are listed, starting be arranged according to ecological parameters
with the most dominant ones. in the head of the table, e.g. pH of bark or soil,
At least, the cover of a single species is elevation, exposition etc.
determined or estimated. The Braun-Blanquet For every species, the frequency (steadiness) can
school uses the following index: be calculated, that means how often it is
represented in all analyses in %. The frequency
r (rare) a single plant is expressed in a scale from I-V:
+ a few plants with a cover less than 1% I = 1-20%
1 1-5% II = 20-20%
2 5-25% III = 40-60%
3 25-50% IV = 60-80%
4 50-75% V = 80-100%
5 75-100%
Based on this table, many statistical studies can
The index 2 has proved to be insufficient, because be made to analyse the community according to
many species have a cover between 5 and 25%. homogeneity, community coefficients etc.
Therefore some authors divided it into 2a (5-
15%) and 2b (15-25%). An easy way to show the interrelationships
The problem of this scale is that it is non-linear between the study plots is to proceed the data in
and cannot be used for computerized evaluations a cluster analysis. The table is then reduced to a
(e.g. a cluster analysis). In this case, the indices two-dimensional matrix of species and study plots
are converted later into means of the indices (e.g. with their values (variables). The resulting
37,5 for 3). It is also possible (although not dendrogram shows the affinities between the
official Braun-Blanquet method) to use study plots. For details of the cluster analysis,
estimations of the percentage cover of each see the Taxonomy section of this paper.
species, which can be directly processed in the
computer.
The Braun-Blanquet method includes the 4. Phytosociological systematics
indication of the „sociability“ of species, a scale
of 5 units (1-5) to indicate whether the plants Communities can be classified by
grow e.g. single, in groups or in mats. Since the a) dominant species, that means the species with
sociability is regarded as part of the life form und the highest cover. The species need, however,
thus species-specific, it is not recommended here not be typical of a community, since they can be
and does not give very much additional weedy species occurring in several communities
information. (and thus having no indicator value),
b) differential species, that means species which
differentiate a community from another one
3. Tablework within a stand (e.g. forest). They cluster in the
table in a block. This is the best way to classify
All separate vegetation analyses are combined communities in the tropics,
in a table (the „raw table“), today by means of a c) by character species. Character species are
spreadsheet program. The table is then sorted indicator species for certain narrow ecological

130 Frahm
niches. They need not automatically have high

frequencies. This requires, however, knowledge
of the ecology of the species.
All other species are named „associates“. They

have no indicator value or are part of the
community by chance.
The name of the community is given by the name

of the character species or differential species
with the ending -etum, the species name in the
genetive. If there is another species involved, it
is added with the ending -o.
Examples (from BRYOTROP studies in Zaire):

Plagiochiletum terebrantis
Drepanolejeuneo - Microlejeuneetum africanae
Lejeuneo flavae - Plagiochiletum divergentis
The basic community is called association. As in

the systematics of plants, subunits (comparable
to subspecies) are made, here called
subassociations. They are expressed with the
name of the characteristic species with the ending
-etosum, e.g. Marchesionio excavatae -
Plagiochiletum salvadoricae - Plagiochiletosum
praemorsae.
Several associations are grouped to unions (like

species to genera), unions to orders and orders to
classes. Orders have the ending -ion, orders -
etalia, classes -etea.

APPENDIX VI: BRYOPHYTE CHECKLISTS OF TROPICAL COUNTRIES
A. Neotropics
Belize Ecuador
Whittemore, A.T., Allen, B. 1996. The liverworts Steere, W.C. 1948. Contribution to the
and hornworts of Belize. The Bryologist bryogeography of Ecuador I. A review
99: 64-67. of the species of Musci previously
reported. Bryologist 51: 65-167.
Bolivia
Hermann, F.J. 1976. Recopilacion de los musgos Guianas
de Bolivia. Bryologist 79: 123-171. Florschütz-de Waard, J. 1990. A catalogue of the
bryophytes of the Guianas. II. Musci.
Brazil Trop. Bryol. 3: 89-104.
Yano, O. 1981. A checklist of Brazilian mosses. Gradstein, S.R. & Hekking, W.H.A. 1989. A
J. Hattori Bot. Lab. 50: 279-456. catalogue of the bryophytes of the
Guianas. I. Hepaticae and Anthocerotae.
Colombia J. Hattori Bot. Lab. 66: 197-230.
Churchill, S.P. 1989. Nuevo Catalogo de los
Musgos de Colombia. Tropical Mexico
Bryology 1: 95ff. Bourell, M. 1992. A checklist of the bryophytes
Churchill, S.P. & Linares, E.L. 1995. Prodromus of Chiapas, Mexico. Tropical Bryology
Bryologiae Novo-Granatensis. 2 vols. 6: 39-56.
Bogotá. Sharp, A.J., Crum, H.A., Eckel, P.M. (eds.) 1994.
Churchill, S.P., Griffin III, D., Munoz, J. 2000. The moss flora of Mexico. 2 vols. New
A checklist of the tropical Andean York Botanical Garden.
countries. Ruizia 17: 1-203.
Panama
Costa Rica Stotler, R., Salazar Allen, N., Gradstein, S.R.,
Bowers, F.D. 1974. The mosses reported from McGuiness, W., Whittemore, A.,
Costa Rica. Bryologist 77: 150-171. Chung, C. 1998. A Checklist of the
Morales Z., M.I. 1991. Las Hepaticas Hepatics and Anthocerotes of Panama.
communicadas para Costa Rica Tropical Tropical Bryology 15: 167 ff.
Bryology 4: 25-58.

132 Frahm
Paraguay C. Tropical SE Asiatic countries; Mosses

Buck, W.R. 1985. A preliminary list of the (by TIMO KOPONEN)
mosses of Paraguay. Candollea 40: 201-
209. Areas (according to “Index muscorum“) and
countries of east and southeast Asia, and the
Peru Pacific, and their moss checklists.
Menzel, M 1992. Preliminary checklist of the
mosses of Peru. J. Hattori Bot. Lab. 71: As 2:
175-254. China
Redfearn, P. L., Tan, B. C. & He, S. 1996: A
Venezuela newly updated and annotated checklist of Chinese
Moreno, E.J. 1992. Aproximación al mosses. J. Hattori Bot. Lab. 79: 163-357.
conocimiento de las briofitas de Vene- Jiangxi
zuela. Trop. Bryol. 6: 147-156. Fang, Y.-M., Enroth, J. Piippo, S. & Koponen,
Pursell, R.A. 1973. Un censo de los musgos de T. 1998: The bryophytes in Jiangxi Province,
Venezuela. Bryologist 76: 473-500. China: An annotated Checklist. - Hikobia
Hunan
Rao, P.-C., Enroth, J., Piippo, S. & Koponen,
B. Tropical Africa T. 1997: The bryophytes of Hunan Province,
China: An annotated checklist. - Hikobia 12:
Born, S., Frahm, J.-P. & Pócs, T. 1993. 181-203.
Taxonomic Results of the BRYOTROP Hong Kong
Expedition to Zaire and Rwanda 26. A Li, Z. & Lin, P.-C. 1997: A checklist of
new checklist of the mosses of Central bryophytes from Hong Kong. - J. Hattori Bot.
Africa. Tropical Bryology 8:.223-274. Lab. 81: 307-326.
Kis, G. 1984. Checklist of the mosses of the Taiwan
south-east tropical Africa. , J. Vana Kuo, C.M. & Chiang, T.Y. 1987: Index to
(ed.): Proceedings of the 3rd Meeting Taiwan mosses. - Taiwania 32: 119-207.
of the Bryologists from Central and East
Europe. Univ. Karlova, Prague. Japan
Kis, G. 1985. Mosses of south-east tropical Iwatsuki, Z. 1991: Catalog of the mosses of
Africa. An annotated list with Japan. - 182 pp. Hattori Botanical Laboratory,
distributional data. 170 p. , Vacratot Nichinan.
(Ungarn): Institute of Ecology and
Botany of the Hungarian Academy of Korea
Sciences. Choe, D.M. & Choi, H.H. 1980: A list of
O´Shea, B. 1995. Checklist of the mosses of sub- bryophytes of Korea. - Rep. Sci. Education 12:
Saharan Africa. Tropical Bryology 10: 27-55.
91-198. Gao, C. & Chang, K. C. 1983: Bryophytes of
North Korea. - Misc. Bryol. Lichen. 9: 163-170.
By the activities of the British Tropical Bryology
working group, the checklists for Africa are As 3:
available as downloads from the Internet. Bangladesh
O´Shea, B.J. 2003. An overview of the mosses
of Bangladesh, with a revised checklist. J. Hattori
Checklist of Malawi bryophytes Bot. Lab. 93: 259-272.
Checklist of mosses of sub-Saharan Africa Bhutan

Long, D. G. 1994. Mosses of Bhutan II. A
Checklist of liverworts of sub-Saharan Africa checklist of the mosses of Bhutan. - J. Bryol. 18:

339-364. As 4:
Eddy, A. 1988, 1989, 1996: A handbook of
Burma (Myanmar) Malaysian mosses. 1-3. - 1. Sphagnales to
Tan, B. C. & Iwatsuki, Z. 1993: A checklist of Dicranales. - 204 p. 1988; 2. Leucobryaceae to
Indochinese mosses. - J. Hattori Bot. Lab. 74: Buxbaumiaceae. - 256 p. 1989; 3.
325-405. Splachnobryaceae to Leptostomataceae. - 277 p.
1996.
Cambodia (Kampuchea)
Tan, B. C. & Iwatsuki, Z. 1993: A checklist of Indonesia
Indochinese mosses. - J. Hattori Bot. Lab. 74: Borneo
325-405. Touw, A. 1978: The mosses reported form
Borneo. - J. Hattori Bot. Lab. 44: 147-176.
India Java
Gangulee, H. C. 1969-1980: Mosses of eastern Fleischer, M. 1904-1923: Die Musci der Flora
India and adjacent regions. 1-8. - Calcutta. von Buitenzorg, zugleich Laubmoosflora von
Java. 1-4. - Leiden.
Laos
Tan, B. C. & Iwatsuki, Z. 1993: A checklist of Lesser Sunda I.
Indochinese mosses. - J. Hattori Bot. Lab. 74: Touw, A. 1992: A survey of the mosses of
325-405. the Lesser Sunda Islands (Nusa Tenggara),
Indonesia. - J. Hattori Bot. Lab. 71: 289-366.
Nepal Irian Jaya
Noguchi, Z. & Iwatsuki, Z. 1975: Musci. - In: Koponen, T., Norris, D. H. et al. 1983-1997:
Ohashi, H. (ed.) Flora of eastern Himalaya. Third Bryophyte flora of the Huon Peninsula, Papua
report. - Bull. Univ. Mus. Univ. Tokyo 8: 206- New Guinea I-LIX.
242. Schultze-Motel, W. 1963: Vorläufiges
Verzeichnis der Laubmoose von Neuguinea.
Pakistan - Willdenowia 3: 399-549
Nishimura, N. & Higuchi, M. 1993: Mosses from
Pakistan. - In: Nakaike, T. & Malik, S. (eds.),
Cryptogamic flora of Pakistan 2: 275-299. Malaysia
Townsend, C. C. 1993: New records and Peninsular Malaysia
bibliography of the mosses of Pakistan. - J. Bryol. Mohamed, H. & Tan, B. C. 1988. A checklist
17: 671-678. of mosses of Peninsular Malaya and
Townsend, C. C. 1994: A small collection of Singapore. Bryologist 91: 24-44.
mosses from Himalayan Pakistan. - J. Bryol. 18: Borneo
181-185. Touw, A. 1978: The mosses reported form
Townsend, C. C. 1995: Further mosses from Borneo. - J. Hattori Bot. Lab. 44: 147-176.
Himalayan Pakistan. - J. Bryol. 18: 811-814.
Singapore
Sri Lanka Mohamed, H. & Tan, B. C. 1988. A checklist of
Abeywickrama, B. A. & Jansen, M. A. B. 1978. mosses of Peninsular Malaya and Singapore. -
A check list of the mosses of Sri Lanka. - Bryologist 91: 24-44.
UNESCO - Man and Biosphere National
Committee for Sri Lanka Publ. 2: 1-25. Brunei and Sabah
Touw, A. 1978: The mosses reported form
Vietnam Borneo. - J. Hattori Bot. Lab. 44: 147-176.
Tan, B. C. & Iwatsuki, Z. 1993: A checklist of
Indochinese mosses. - J. Hattori Bot. Lab. 325-
405.

134 Frahm
Papua New Guinea Tuvalu and Kiribati

Koponen, T., Norris, D. H. et al. 1983-1999: Tonga
Bryophyte flora of the Huon Peninsula, Papua Vanuatu
New Guinea I-LXIV. – Ann. Bot. Fennici & Acta
Bot. Fennica Austr 1:
Schultze-Motel, W. 1963: Vorläufiges
Streimann, H. & Curnow, J. 1989: Catalogue of
Verzeichnis der Laubmoose von Neuguinea. -
Willdenowia 3: 399-549. mosses of Australia and its external territories. -
Australian Flora Fauna Ser. 10: i-viii, 1-479.
Philippines
Tan, B. C. & Iwatsuki, Z. 1991: A new annotated
Philippine Moss Checklist. - Harvard Papers Bot. Austr 2:
3: 1-64. Fife, A. J. 1995: Checklist of the mosses of New
Zealand. - Bryologist 98: 313-337.
Oc:
Koponen, T., Norris, D. H. et al. 1983-1997:
Bryophyte flora of the Huon Peninsula, Papua Checklists of bryophytes of SE Asiatic
New Guinea I-LIX. countries; Hepatics
Miller, H. A., Whittier, H. O. & Whittier, B. A. SINIKKA PIIPPO
1978: Prodromus florae muscorum Polynesiae.
– Bryophyt. Biblioth. 16: 1- 334. Areas (according to “Index muscorum“) and
countries of east and southeast Asia, and the
Carolines (Federated States of Micronesia) Pacific, and their hepatic checklists
Fiji
As 1:
Marianas
Russia
Hawaii Konstantinova, N. A, Potemkin, A. D. &
Hoe, W. J. 1974. Annotated checklist of Hawaiian Schljakov, R. N. 1992: Check-list of the
mosses. Lyonia 1: 1-45. Hepaticae and Anthocerotae of the former USSR.
Marshalls - Arctoa 1: 87-127.
Marguesas Konstantinova, N. A. & Potemkin, A. D. 1996:
New Caledonia Liverworts of the Russian Arctic: an annotated
Pursell, R. A. & Reese, W. D. 1982: The mosses check-list and bibliography. - Arctoa 6: 125-150.
reported from New Caledonia. - J. Hattori Bot.
As 2:
Lab. 53: 449-482. China
Samoa He, X. -L. 1997: A review and checklist of the
Schulze-Motel, W. 1974: Die Moose der Samoa- Lejeuneaceae in China. - Abstracta Bot. 21(1):
Inseln. - Willdenowia 7: 333-408. 69-77.
Society Islands Piippo, S. 1990: Annotated catalogue of Chine-
se Hepaticae and Anthocerotae. - J. Hattori Bot.
Whittier, H. O. 1976: Mosses of the Society
Lab. 68: 1-192.
Islands. - x, 410 p. The University Presses of Jiangxi
Florida, Gainesville. Fang, Y.-M., Enroth, J. Piippo, S. & Koponen,
Solomons T. 1998: The bryophytes in Jiangxi Province,
China: An annotated Checklist. - Hikobia
Koponen, T., Norris, D. H. et al. 1983-1997: Hong Kong
Bryophyte flora of the Huon Peninsula, Papua Li, Z. & Lin, P.-J. 1997: A checklist of
New Guinea I-LIX. bryophytes from Hong Kong. - J. Hattori Bot.
Lab. 81: 307-326.

Hunan India
Rao, P.-C., Enroth, J., Piippo, S. & Koponen, Kachroo, P. 1969-1977: Hepaticae of India. I-V.
T. 1997: The bryophytes of Hunan Province, - (For details see Greene & Harington, 1989).
China: An annotated checklist. - Hikobia
12:181-203. Laos
Sichuan
Piippo, S., He, X.-L. & Koponen, T. 1997: Nepal
Hepatics from northwestern Sichuan, China, Grolle, R. 1966: Die Lebermoose Nepals.
with a checklist of Sichuan hepatics. - Ann. Khumbu Himal, Ergebnisse des Forschunsunter-
Bot. Fennici 34: 51-63. nehmens Nepal Himalaya 1(4): 262-298.
Taiwan Grolle, R. 1974: Nachtrag zu “Die Lebermoose
Kuo, C.M. & Chiang, T.Y. 1987: Index to Nepals“. Khumbu Himal, Ergebnisse des Forsch-
Taiwan mosses. - Taiwania 32: 119-207. unsunternehmens Nepal Himalaya 6(2): 117-120.
Yunnan Hattori, S. 1975: Anthocerotae, Hepaticae. - In:
Piippo, S., He, X.-L., Koponen, T., Li, X.-J. Ohashi, H. (ed.) Flora of eastern Himalaya. Third
& Redfearn, P., 1998: Hepatics from Yunnan, report. - Bull. Univ. Mus. Univ. Tokyo 8: 206-
China, with a checklist of Yunnan hepatics and 242.
Anthocerotae. - J. Hattori Bot. Lab. 84
Pakistan
Mizutani, M., Furuki, T., Yamada, K. & Higuchi,
Japan M. 1994: Hepatics from Pakistan collected by
Furuki, T. Mizutani, M. 1994: Checklist of the Botanical Expedition of the National Science
Japanese Hepaticae and Anthocerotae, 1993. - Museum, Tokyo in 1990. - Bull. Natn. Sci. Mus.
Proc. Bryol. Soc. Japan. 6: 75-83. Tokyo, Ser. B 20: 143-150.
Mizutani, M. & Furuki, T. 1994: A comparison
list between “Checklist of Japanese Hepaticae and Sri Lanka
Anthocerotae, 1983" and “1993". - Proc. Bryol. Abeywickrama, B. A. & Jansen, M. A. B. 1978.
Soc. Japan. 6: 84-93. A check list of the liverworts of Sri Lanka. -
UNESCO - Man and Biosphere National
Korea Committee for Sri Lanka Publ. 1: 1-10
Choe, D.M. & Choi, H.H. 1980: A list of Onraedt, M. 1981: Bryophytes récoltées à Sri
bryophytes of Korea. - Rep. Sci. Education 12: Lanka (Ceylan). V. - J. Hattori Bot. Lab. 50: 191-
27-55. 216.
Gao, C. & Chang, K. C. 1983: Bryophytes of
North Korea. - Misc. Bryol. Lichen. 9: 163-170. Vietnam
Pócs, T. 1965: Prodrome de la bryoflore du Vi-
As 3: etnam. - Egri Tanárképzö Föiskola Tudományos
Bangladesh Közleményei, N.S. 3: 453-495.
Pócs, T., Tixier, P. & Jovet-Ast, S. 1967: Adatok
Bhutan Észak-Vietnam mohaflórájához. II. Seconde
Long, D. G. 1979: Hepaticae from Bhutan, East contribution à la bryoflore du Nord Vietnam. -
Himalaya. - Lindbergia 5: 54-62. Botanikai Közlemények 54: 27-38.
Long, D. G. & Grolle, R. 1990: Hepaticae of
Bhutan II. - J. Hattori Bot. Lab. 68: 381-440. As 4:
Schiffner, V. 1898: Conspectus Hepaticarum
Burma (Myanmar) Archipelagi Indici. - 382 pp. Batavia, Staats-
druckerei.
Cambodia (Kampuchea)

136 Frahm
Indonesia checklist of Philippine Hepaticae. - J. Hattori Bot.

Borneo Lab. 60: 283-355.
Menzel, M. 1988: Annotated catalogue of the
Hepaticae and Anthocerotae of Borneo. - J. Oc:
Hattori Bot. Lab. 65: 145-206. Koponen, T., Norris, D. H. et al. 1983-1997:
Java Bryophyte flora of the Huon Peninsula, Papua
Schiffner, V. 1900: Die Hepaticae der Flora New Guinea I-LIX.
von Buitenzoerg. I. Band. - 220 p. Leiden, E. Miller, H. A., Whittier, H. O. & Whittier, B. A.
J. Brill. 1983: Prodromus florae hepaticarum Polynesiae
with a key to genera. - Bryophytorum Bibl. 25:
Irian Jaya 1- 423.
Grolle, R. & Piippo, S. Annotated catalogue
of Western Melanesian bryophytes. I. Carolines (Federated States of Micronesia)
Hepaticae and Anthocerotae. - Acta Bot. (see Greene & Harington 1989)
Fennica 125:1-86.
Koponen, T., Norris, D. H. et al. 1983-1999: Fiji
Bryophyte flora of the Huon Peninsula, Pa-
pua New Guinea I-LXIV. – Ann. Bot. Fennici Marianas
& Acta Bot. Fennica
Hawaii
Malaysia Miller, H. A. 1963: Notes on Hawaiian Hepaticae
Peninsular Malaysia V. Collections from recent Swedish expeditions.
Johnson, A. 1958: An account of the thallose - Arkiv för Bot., Ser 2, 5: 489-531.
liverworts found in Malaysia. - Malayan Na-
ture Journal 13: 52-69. Marshalls
Borneo Marguesas
Menzel, M. 1988: Annotated catalogue of the
Hepaticae and Anthocerotae of Borneo. - J. New Caledonia
Hattori Bot. Lab. 65: 145-206. Kitagawa, N. 1983: Hepaticae and Anthocerotae
of New Caledonia. - Proc. Bryol. Soc. Japan 3:
Singapore 119-122.
Brunei and Sabah Samoa

Menzel, M. 1988: Annotated catalogue of the Grolle, R. 1980: Zur Kenntnis der Lebermoose
Hepaticae and Anthocerotae of Borneo. - J. Hat- von Samoa I. - Wiss. Zeitschr. Friedrich-Schil-
tori Bot. Lab. 65: 145-206. ler-Universität, Jena, Math.-Nat. R. 29: 637-648.
Grolle, R. & Schultze-Motel, W. 1972: Vorläu-
Papua New Guinea figes Verzeichnis der Lebermoose von Samoa. -
Grolle, R. & Piippo, S. 19??: Annotated J. Hattori Bot. Lab. 36: 75-89.
catalogue of Western Melanesian bryophytes. I.
Hepaticae and Anthocerotae. - Acta Bot. Fennica Society Islands
125:1-86.
Koponen, T., Norris, D. H. et al. 1983-1997: Solomons
Bryophyte flora of the Huon Peninsula, Papua Grolle, R. & Piippo, S. Annotated catalogue of
New Guinea I-LIX. Western Melanesian bryophytes. I. Hepaticae and
Anthocerotae. - Acta Bot. Fennica 125:1-86.
Philippines Koponen, T., Norris, D. H. et al. 1983-1997:
Tan, B. C. & Engel, J. J. 1986: An annotated Bryophyte flora of the Huon Peninsula, Papua
New Guinea I-LIX.

Piippo, S. 1993: Hepatics from the Solomon

Islands. 1. - Nova Hedwigia. 56: 355-365.
Tuvalu and Kiribati
Tonga
Vanuatu
Austr 1:
Scott, G. A. & Bradshaw, J. A. 1986: Australian
liverworts (Hepaticae): annotated list of
binomials and check-list of published species
with bibliography. - Brunonia 8: 1-171.

138 Frahm

APPENDIX VII: PRINCIPLES OF PREPARING CHECKLISTS

by TIMO KOPONEN, SINIKKA PIIPPO, JOHANNES ENROTH, PENGCHENG RAO & YIN-
MING FANG
While compiling the bryophyte checklists of the for a checklist including references to all
Hunan (Rao et al. 1997) and Jiangxi (Fang et al. published records of Ceratodon purpureus for
1998) Provinces of China we came across several Finland, while to cite the records of Ceratodon
difficulties. First, some of the records for Hunan for Pakistan is still reasonable. Most useful
and Jiangxi were mere lists of names without checklists are those which give the reasons of
documentation, and second, some of the records the nomenclatural changes, list the synonymies,
are doubtful on the basis of the general known and cite the additions to the flora. This is done
distribution of the taxa. Also, some records cited by citing the papers dealing with the
in earlier checklists were based on literature not nomenclature and floristic publications. Floristic
available to us, or even on unpublished publications cite, or their should cite, the
manuscripts. We feel it necessary to discuss the specimen on which the record is based, and the
methodology of preparing checklists here. museum or herbarium in which this voucher is
being kept. Some checklists include floristic
novelties, Grolle & Piippo (1984) being a good
Checklists are a useful tool of floristics and example of this. The discussion above leads to
phytogeography and a summary of the flora of a the first principle.
certain area. The area dealt with may be a country,
some smaller area such as a province, or, more Principle 1
rarely, a continent or some other larger The basic principle of all checklists is that they
geographical area. Examples of all of these can should be based on herbarium specimens which
be cited and are listed by Lane (1978) and Greene can be traced and the identity of which can be
& Harrington (1989). Basically, two different checked, when or if necessary.
kinds of lists have been published. Some
checklists are mere lists of the flora existing in The basis of any checklist is the floras of the area
the area dealt with, while some others carefully dealt with, if such works exist. Fortunately,
document all the knowledge by citing published bryologists of earlier generations were careful in
records. The method and layout partly depend their documentation. Works such as Fleischer’s
on the area dealt with. In many European (1904-1923) flora of Java, Brotherus’ (1923)
countries, North America, and Japan the Fennoscandian flora, Bartram’s (1939) Philippine
checklists may be mere lists without any further flora, as well as some more modern floras
information. This is easily understood; for (Gangulee’s1969-1977, Whittier 1976, Magill
instancce, it might be difficult to find a publisher 1981, 1987, Allen 1994, Bai & Zhao 1997) cite
specimens. This tradition should be continued

140 Frahm
when the floras of imperfectly known areas are be given directly in the list, and not as numerical
published. This can be done very shortly e.g. by code (compare Piippo 1990 and Redfearn et al.
citing the specimens from which the figures of 1996). Miller et al. (1978) contains a vast body
the work were drawn, or by citing widely of knowledge of the floristics of the Pacific area.
distributed exsiccata (e.g. Noguchi 1987-1994). The distribution of taxa is readily available, but
We have found it most unfortunate that Eddy finding the references for a certain taxon within
(1988-1996) did not continue this tradition. a certain group of islands is laborious.
Nomenclatural and floristic additions can be cited
The lack of documentation in flora works creates in notes (e.g. Koponen et al. 1977, Corley et al.
problems to checklist writers. How to deal with 1981).
records which are highly doubtful on the basis
of the general distribution of the taxon? Here is Principle 4
one example. Chang (1978), in her flora, Information in checklists should be given in a
published a description and an illustration of form easily available.
Mnium venustum (= Plagiomnium venustum) and
cited it for many Chinese provinces. However, Acknowledgements. Grants nos. 10134229 and
P. venustum is an endemic of the Pacific coast of 153706 from Academy of Finland to Timo
North America; the illustration in Chang’s flora Koponen are cordially acknowledged.
does not represent it; and no Chinese specimens
have been available for study (Koponen 1981,
Koponen & Lou 1982). Redfearn et al. (1966) References
noted these references and did not accept P.
venustum as reliably recorded for China. Allen, B. 1994: Moss flora of Central America.
Part 1. Sphagnaceae-Calymperaceae. - 242 pp.,
Missouri Botanical Garden, St. Louis.
Principle 2
Flora works lacking documentation should be Bai, X.-L. & Zhao, Z.-T. 1997: Flora
used very critically as basis of checklists. bryophytarum intramongolicarum. - 541 pp.
Typis Universitatis Intramongolicae.
The records published in popularizing books, or
in works dealing with forestry or describing Bartram, E. B. 1939: Mosses of the Philippines.
vegetation, are often not documented. These - Philipp. J. Sci. 68: 1-437.
records can be doubted on the basis of the general
distribution of species, although all distributions Brotherus, V. F. 1923: Die Laubmoose
are not yet known in detail. Checklists should Fennoskandias. - Flora Fennica 1:1-635.
not be based on unpublished manuscripts, (Facsimile 1974, Koenigstein.)
undocumented mimeographed lists, or data files
not readily accessible, either. Chang M.-S. 1978. Bryophyta in Flora
Tsinglingensis. Tom 3. Bryophyta (Par. 1).
Principle 3 Science Publisher, Beijing. 329 pp.
Popularizing books and plant sociological papers
lacking documentation should be omitted or used Corley, M. F. V., Crundwell, A. C., Düll, R., Hill,
very critically as basis of checklists. Records in M. O. & Smith, A. J. E. 1981: Mosses of Europe
unpublished manuscripts and data files not and Azores; an annotated list of species, with
generally available should not be used as basis synonyms from recent literature. - J. Bryol. 11:
of checklists. 608-689.
Well-documented checklists are useful sources Eddy, A. 1988, 1989, 1996: A handbook of
of information and using them should not be time- Malaysian mosses. 1-3. - 1. Sphagnales to
consuming. The references to the literature should Dicranales. - 204 p. 1988; 2. Leucobryaceae to

Buxbaumiaceae. - 256 p. 1989; 3. 1978: Prodromus florae muscorum Polynesiae. -

Splachnobryaceae to Leptostomataceae. - 277 p. Bryophytorum Bibl. 16: 1- 334.
1996.
Noguchi, A. 1987-1994: Illustrated moss flora
Fang, Y.-M., Enroth, J. Piippo, S. & Koponen, of Japan 1-5. - Hattori Botanical Laboratory,
T. 1998: Annotated checklist of the bryophytes Nichinan.
in Kiangsi Province, China. - Hikobia
Piippo, S. 1990: Annotated catalogue of Chinese
Fleischer, M. 1904-1923: Die Musci der Flora Hepaticae and Anthocerotae. - J. Hattori Bot. Lab.
von Buitenzorg, zugleich Laubmoosflora von 68: 1-192.
Java. 1-4. Leiden.
Rao, P.-C., Enroth, J. Piippo, S. & Koponen, T.
Gangulee, H. C. 1969-1977?: Mosses of eastern 1997: Annotated Checklist of the bryophytes in
India and adjacent regions. 1-6. Calcutta. Hunan Province, China. - Hikobia
Greene, S. W. & Harington; A. J. 1989: The Redfearn P.L. Jr., B.C. Tan & S. He, 1996: A
conspectus of bryological literature. Part 2. Guide newly updated and annotated checklist of Chinese
to national and regional literature. - mosses. - J. Hattori Bot. Lab. 79: 163-357.
Bryophytorum Bibl. 37: 1-321.
Whittier, H. O. 1976: Mosses of the Society
Grolle, R. & Piippo, S.: Annotated catalogue of Islands. - x, 410 p. The University Presses of
Western Melanesian bryophytes. I. Hepaticae and Florida, Gainesville.
Anthocerotae. - Acta Bot. Fennica 125: 1-86.
1984.
Koponen, T. 1981: A synopsis of Mniaceae

(Bryophyta). VI. Southeast Asian taxa. - Acta
Bot. Fennica 117: 1-34.
Koponen, T., Isoviita, P. & Lammes, T. 1977:

The bryophytes of Finland: An annotated
checklist. - Flora Fennica 6: 1-77.
Koponen, T. & Lou, J.-S. 1982: Miscellaneous

notes on Mniaceae (Bryophyta). XII. Revision
of specimens in the Institute of Botany, Academia
Sinica, Beijing, China. - Ann. Bot. Fennici 19:
67-72.
Lane, D. 1978: A geographical guide to

bryophyte floras of the world 1. - Miscellanea
Bryol. Lichenol. 8: 15-34.
Magill, R. E. 1981, 1987: Flora of southern

Africa. Bryophyta. 1, 2. - 1, Sphagnaceae-
Grimmiaceae, i-xv, 1-291 p. 1981. - 2,
Gigaspermaceae-Bartramiaceae, i-ix, 292-443. -
Government Printer, Pretoria.
Miller, H. A., Whittier, H. O. & Whittier, B. A.

142 Frahm

APPENDIX VIII: METHODOLOGY OF RELEVÉE STUDIES
Bryology is not only collecting bryophytes. Alt- Base for such studies can be hectare plots (see
hough collecting was and is the main motive or appendix II) or trees for epiphyte studies.
even urge for many bryologists and the only sti- Most important is that all parameters, especially
mulus for many amateur bryologists, it is just the the size of the study sites, are comparable. Com-
necessary base for further scientific evaluations. parisons of species numbers are only possible if
In the begin of bryology, collecting was done for based on the same area. There were publications
taxonomy (discovery and description of new spe- comparing species numbers of national parks of
cies). At that time, species were collected like very different size, in different altitudes and by
stamps in most complete sets. This period is over collections from different periods (for one park
now. Althrough there is still a chance to detect even collections from 90 years ago included). The
new species in the tropics, this chance is small, advantage of hectare plot studies is that the base
requires much taxonomic knowledge and is so- for comparison is given. There are also examp-
mething for specialists. The next step was to de- les in which the species numbers of trees are com-
scribe the distribution of known species. This has pared without regard to their size. Species num-
been done by many single floristic studies, in bers can only be compared for the same area,
Europe also by systematic mapping of species. and this concerns also studies of epiphytes, were
There is still a need for such floristic studies in the calculationhas to be based on for instance a
the tropics, especially in undercollected areas, square meter around the tree trunk.
however, this can not be all. Therefore some in-
structions are given here to complete and inten- 1. Calculation of alpha diversity (species per
sify the bryological knowledge of tropical coun- area). For that purpose, the area (study plot, rele-
tries. vée) must have the same size and should have a
Usually, at random collecting makes scientifically minimum size to cover all species of a vegetati-
not much sense and is something for hobby-bryo- on type. The minimum size can be determined
logists. Important is to gather data. For the eva- by a species-area curve: with measuring tape,
luation of data, a comparable base is required. an area of 1 square meter is marked and all spe-

144 Frahm
cies in this area are counted. It is not necessary (c) The vegetation belts differ in mountains of
that the species can be named but need only to different altitude: they go higher up in large
be differentiated. Next, the side length is doub- mountains but are lower in small mountains
led to 2m (4 m2), then to 4, 16, 32 and 64 m („Massenerhebungseffekt“).
(4096m2) and again all species are counted (or The main ecological factor which shall be ex-
those in addition to the previous area). Finally, pressed by the altitude is the temperature., which
the species number is plotted against the area. decreases by 0.6-1°C per 100 m altitude. Instead
The resulting curve shows at first a steep increa- of giving the altitude, it is possible to refer to the
se, but will get suddenly flat, when the the spo- mean annual temperature. Usually, mean annual
ecies number is saturated and ends asymptotic. temperatures can only be determined by long
Minimum area is the area where the curves flat- series of temperature measurements, to cover the
tens. different temperatures in different seasons (sea-
sonally climate). They can also be determined
2. After collecting and identifying the species, by the temperature of sping water (except ther-
the data can be evaluated: mal springs, of course), since the water has a con-
- by writing a species list by families stant temperature over the year. Measurements
- by calculating the ratio hepatics (incl. horn- over a year are, however, not necessary in the
worts) : mosses, inner tropics , where the daily fluctuations of the
- by calculating the number of species by fami- temperature are higher than the annual fluctuati-
lies and displaying the results in a bar chart or ons (daily climate). The mean annual tempera-
pie-chart. ture is measured in the soil below 30 cm depth,
- by calculating the number of species on diffe- where the influence of the day/night fluctuations
rent substrates (soil, rocks, rotten wood, epiphy- are no more expressed. To determine the soil tem-
tic). perature in 30 cm depth, a solid metal stick of
this length is required, which is bent 90° at the
3. Calculation of beta-diversity. After the inven- top. The stick is pushed into the soil to form a
tory of several relevées along an ecological gra- hole of 30 cm depth. In this hole, a thermometer
dient (see appendix II), the relevées can be com- is lowered. It can be a normal thermomter fixed
pared. For the calculation, diversity indices such by a cord or an electronic thermometer lowered
as the Sörensen Index can be used: by its cable. There are special electronic soil ther-
mometer with stick-like electrode.
Species common in both relevées2 By this way, the mean annual temperature can
----------------------------------------- be used as reference instead of the altitude.
Species number relevée A + B
5. Determination of cover. The cover of bryo-
The species numbers of different relevées can be phytes varies much between lowland, montane
visualized in a line chart. and subalpine forests. It can easily be determi-
ned even without knowing any species but gives
4. Determination of the mean annual tempe- nevertheless good results for the determination
rature. For altitudinal transects and comparison of altitudinal zones, because bryophytes reflect
of relevées, the elevation is used as parameter. best the different ecological conditions in diffe-
This has, however, important disadvantages: rent altitudes. The cover is estimated as average
(a) The vegetation belts vary in altitude by lati- percentage on the ground and on trees. Determi-
tude; they are higher situated in equatorial latitu- nation of cover is different from that of species
des but decrease towards the poles. richness, since both factors are not correlated. A
(b) The exposition varies the vegetation. At the few certain species can form large masses. By
same altitude, we may have different vegetation this way, the species numbers can be sometimes
types on N- and S- exposed slopes. Therefore a higher in areas with low cover and vice versa. It
N-exposed relevée in 2000 m cannot be compa- is also recommended to differentiate between the
red with a S-exposed one at the same altitude. cover of hepatics and mosses.

For epiphytes, the thickness of epiphyte cover cylinder is also 1m and the cylinder has an area
can also be determined. In the lowlands, epiphy- of 1 m2. If the tree has a circumference of 50 cm,
tes are usually found only as crusts of liverworts, the hight must be 2 m to give an area of 1 m2. If
whereas in the subalpine forests hugh, thick the trees are too small that the length of the trunk
masses of epiphytes cover trunks and branches. area gets too long, or if the amount of bryophy-
The measurements are taken with a ruler. tes is too much that it will get difficult to handel,
also half a square meter may be taken. All bryo-
6. Determination of life forms. As expressed in phytes are scratched off the trunk with a knife
chapter 5.1, the different life forms are direct ex- and collected in a bag. The material is air dried
pressions of different ecological conditions be- and the weight is determined by a balance. Any
cause they are adaptations to these conditions. pieces of bark, litter, accompanying ferns, filmy
The main life forms are determined: ferns, other epiphytes or dirt has to be removed
- crusts: predominantly liverworts (Lejeuneaceae, before. By this way, the phytomass may be de-
certain Frullania and Radula species) creeping termined on several representative trees and an
on bark of trees). average can be calculated. Statistics recommen-
- cushions ded the use of random trees, however, in this case
- tufts (Dicranum, Campylopus-species) the number of tress must be quite high. Often,
- mats (most pleurocarps) the epiphytic cover on trees is very uniform and
- wefts (Lepidozia, Bazzania, Teleranea) in this case, even the result of the determination
- turfs (Polytrichaceae) of the phytomass of one representative tree co-
- fans (Porotrichum, some Plagiochilas) mes the result of the determination of the phyto-
- pendants (predominantly Meteoriaceae) mass of numerous trees chosen at random very
Some life forms listed here may not be found, close.
others my be differentiated by themselves if it With some reservation, the phytomass of epiphy-
seems appropriate (e.g. creeping mats of tic bryophytes can be calculated for a hectare.
Macromitrium or similar, dendroid turfs). For First it is estimated how many times the open
each life form, the percentage is noted. area on the trunk, where the material was scrat-
ched off, fits on the whole tree (say 12 times).
7. Determination of phytomass. The amount of Then the number of trees with a similar cover
phytomass of bryophytes is not only an expres- are counted in an area which can be overlooked
sion of the „mossiness“ of a habitat but determi- (e.g. 20 x 20 m, say 5). This would mean that 60
nes the water storing capacity of the forest. In times the weight of the phytomass per square-
tropical forests, it concerns usually the phytomass meter is found on 20 x 20 m and approximately
of epiphytes due to the lack of larger amounts of 25 times as much on a hectare. This estimation
bryophytes on other substrates (except of subal- gives a rough idea of the phytomass per hectare
pine forests). and may not be interpreted too exact. It is only
Some bryologists have determined the phytomass an approximation but gives an idea whether the-
of a complete tree, which had to be cut for that re are 20 kg or a ton of bryophytes to be expec-
purpose, and all bryophytes had to be remove, ted on a hectare.
dried and weighted. This requirtes much effort
and is not always possible. Therefore an alterna- 8. Determination of water storing capacity.
tive method has been developed based on mea- From the dry weight of bryophytes per square
surements of the phytomass per squaremeter. For meter, the amount of water can be determined
that purpose, one square meter is marked on a which is stored in the bryophytes. For that pur-
tree trunk. The trunk resembles a cylinder (2πrh). pose, the whole mass of bryophytes taken from
First the circumference of the trunk is measured one m2 is soaked in water. Water is now stred
with a measuring tape (2πr). Next the height of within the bryopyhtes as well as between the bry-
the cylinder is determined by dividing 1 square- ophytes. The latter, the so called interception
meter by the circumference. Example: If a tree water, is difficult to determine. If one tries to put
has a circumference of 1 m, the height of the the bryophytes on a balance, the interception

146 Frahm
water will run off. Therefore the surplus water result (+/- 0.2 values).
is carefully shaken off and the wet bryophytes
are weighted. The experience shows that the bry- 10. Determination of light intensity. Light in-
ophytes normally store water between 2,5 and tensity, humidity and temperature depend on per-
3,5 of their dry weight. An exception are sam- manent fluctuation. Therefore single measure-
ples from lowland forests with much Leucobrya- ments cannot be taken into account. An excpeti-
ceae and Calymperaceae, which store more (5x). on is the relative light intensity. It indicates the
Roughly, the dry weight may be multiplied by percentage of light in an habitat, e.g. different
three to get the water storing capacity. The same parts of the forest floor, as compared to the light
can be done with the hectare values. in the open. Therefore the relative light intensity
of e.g. 5% can be determined at any time of the
9. Determination of pH of substrates. Species day or during cloudy or sunny wheather.
composition is generally much influenced by the Although the intensity of certain wavelengths
pH of the substrate, but not as important in the relevant for phyotosynthesis (PAR) is measured-
wet tropics. The higher the precipitation, the lo- for physiological purposes, the cheaper lux me-
wer is the influence of the substrate. All measu- ter with a sensibility for a different light spec-
rements of barks of trees in the tropics revealed trum can be used to characterize different light
acidic conditions, ranging betwen 4 and 6, which and dark habitats.
can , however, be different on different host trees Measurements are taken with the light sensor
and therefire explain different species compoisi- exposed to the sky in different parts of a forest.
tion (beside bark texture and chemistry). Bark nstead of making numerous at random readings,
pH is measured by cutting off small pieces of the it is recommended to measure the lightest and
outernmost bark, filling them in a 50 ml plastic the darkest spot which are inhabited by bryoph-
bottle and adding 2.5 times as much distilled ytes to get the span of light intensity in a forest
water. The bottles can be shaken and measured stand. Alternatively, the light can be measured
immediately or kept for 24 hours and measured for habitats of different species to get the photo-
then. Important is, that always the same methods philous and skiotolerant species differentiated.
is used (same amount of bark, water, time). Mea- Shortly before or afterwards, the light is measu-
surements are taken with an electronic pH-me- red in the open (on a road, clearing, spot of a
ter. If the pH-meter is new or has not be used for fallen tree, open trail). The reading from the open
a longer time, it is important (a) to soak the elec- is divided by the values received in the forest to
trode in water (no distilled water!) before and to get the percentage light. Complications arise only
calibrate it with two buffers. The electrode has if there are wandering clouds. In this case atten-
to be stored all the time between he measure- tion has to be paid that the measurements have
ments in water. Recommende are electrodes only be made at moments with open sky. Equal-
which are sealed or filled with gel which need ly mats of clouds are not a problem.
not to be refilled. There are special flat head elec-
triodes for direct measurements on bark. It is also 11. Determination of relative humidity and
the only way to measure the pH of bare rock. temperature. For these factors, momentary mea-
The surface of a plane piece of bark (which can surements make rarely sense. Only if data log-
be removed from the tree for that pupose) or a gers are not available, measurements in minimal
plane part of a rock is covered with distilled wa- hours intervalls can provide daily curves of both
ter and the elctrode is pressed upon it. There must factors. For both purposes, electronic devices are
be direct contact with water between surface and recommended.
electrode. Humidty was originally measured with psychro-
Soil samples are treated in the same way: soil is meters. These were two thermometers, of which
filled in a plastic bottle and diluted with 2.5 ti- one had a „sock“ covering the bulb. The sock
mes of distilled water. The treatment (shaking or was wettened with water, causing a decrease of
not, leaving it for short or longer time before the temperature by evaporation. The lower the air
measurement) has only a small influence on the humidity the higher was the evaporation and the

higher the difference between the „dry“ and the propgata over wide distances? An analysis of the
„wet“ thermometer. The relative humidty (rh) life strategies already gives rough results. Also a
could then be interpolated from a table. It is still calculation of the mode and frequency of vege-
possible to prepare two thermometers in such a tative or sexual propagation. Detailed analyses
way for that puropse as the cheapest way of a how the species propagate (by gemmae, tubers,
psychrometer. Only the table is required, which fragments, leaves, or sexually by spores, how
can be copied from ecology textbooks. Later, large are the spores, how long are they presumab-
electronic psychrometers could calculate the va- ly distributed, can be helpful.
lue, but were still operating with two thermome-
ters. Some time ago, a substance has been dico-
vered which changed the resistance of electrici-
ty in dry and wet state. Therefore simple modi-
fied micro-voltmeter could be used for measur-
ring humidity with this new electrode similar to
an electronic thermometer.
Long term measurements of temperature and
humidity are possible with data loggers. Such
instruments were extremely costly and large in
the past but as all electronic equipment got small
and cheap within the time. Recent dataloggers
have the size of a credit card or a match box. The
latter cost less than 130$ and can store up to 1700
readings in any intervall. They are connected to
a computer to initialize and download the data,
which are displayed in curves and can be expor-
ted to MS Excel.
Humidity data loggers give an idea how long
the bryophytes are in wet state in different habi-
tats. Bryophytes are poicilohydric and therefore
metabolic active only when turgescent. So they
do not suffer from any temperature or humidity
in dry state (which is called anabiosis). Important
is, how long they can be photosynthetic active in
different habitats. For that purpose, there must
be sufficient light and an air humidity above 80%.
Below 80% rH, the bryophytes dry up (some
hygrophytes even sooner). Since phytomass is a
direct result of net photosynthesis, the „mossi-
ness“ of an area is a result of photosynthetic ac-
tive phases (except for lowland forests with high
rates of respiration) which can be determined by
a datalogger.
12. Determination of life strategies and pro-

pagation methods. This answers the question,
how is the bryoflora composed, of species which
are sterile, or propagate vegetatively and are thus
less flexible against destruction or disturbance
and are candidates to get likely locally extinct by
human impact or by fertile spcies which easily

148 Frahm

APPENDIX IX: THE USE OF COMPUTERS IN THE FIELD
Computers got irreplaceable tools for scientific cameras. It has to be observed that the converter
work, also for the fieldwork. They can be used has enough watt. For instance, if the notebook
for normal computer work as well as devices for charger has 220V and 1,8A, it needs 400 W. This
measuring instruments or for navigation. Many can easily be calculated by Ohm´s formula.
computer work can already done in the field and Possible applications for notebooks in the field
many time can be saved by using the computer are wordprocessing, use of spreadsheets (e.g. for
during fieldtrips. There are, however, limitations. data from ecological measurements of light,
The computer must be portable, and therefore temperature, humidity etc.), use of databases e.g.
only notebooks, handheld PCs or Pocket PCs are for collecting information, download of data
in question. from data loggers and navigation.
1. Notebooks do almost everything a desktop Navigation is possible by connecting a GPS to
computer can do, but have lower size and weight. the notebook. It requires special software such
They can be synchronized with desktop as FUGAWI (see internet). If maps are loaded
computers and by this way all current projects into the computer, the position is indicated. This
and files can be carried around. The main facilitates travels in the tropics, showing always
restriction is still, that the batteries last only 2-3 the own position. The software also allows to
hrs. Therefore a use abroad is very much calibrate scanned maps or aerial photographs.
confined if the notebook cannot be recharged, By that way, also maps of a national park or the
except for the use in fieldstations. On trips by study site of a field station can be used. By
car, the notebook can be recharged from the car walking through a study site, the exact position
battery. Necessary is a special 12V charging is indicated and even single trees can be located
cable (usually for this special notebook and thus and marked in the map. Beside, notebooks allow
expensive). An alternative is the use of a 12V – access to the internet (webpages, e-mails)
110/220V converter. This converter allows to use through the cell phone in areas which are covered
all electric devices in the car (up to an limit of by cell phone networks. Else the use of satellite
100 – 400 W depending on the model), including cell phones is possible (which is, however,
electric shaver, charger for cell phones, batteries expensive).
for measuring instruments, cameras and video

150 Frahm
So called subnotebooks are smaller (A5 size) but

also run with normal computer systems and have
similar limitations concerning battery life. The
battery life is limited by use for the screen, for
the hard disk and a CD ROM.
2. Handheld and Pocket PCs differ from

notebooks in their size (pocket format) and the
feature that they have all programs and data in
RAM, which costs hardly any electric power.
They have no power consuming disk drives and
most of the energy is used to illuminate the small
screen. By this way, they last 10-40 hrs. Some
older models use 1,5V batteries, which can easily
be exchanged and no charging is required.
Handhelds have a tiny keyboard and are easier
to use for data input, Pocket PCs are used with a
small pointer. There are 3 different systems of
such small PCs: Palm, Psion and Windows CE.
They all require that the handheld or pocket PCs
are connected with a desktop computer or
notebook from which programs or data can be
downloaded. The programs on the small
computers are different from the large PCs and
the files must be converted. Nevertheless, these
small computers allow the use of Word, Excel,
Filemaker or Access files in he field. Accessing
the internet is equally possible by a cell phone
which is connected by cable or IR device. By
the same connection, sending of SMS messages
or even pictures taken with a digital camera is
possible if the cameras uses flashcards and the
Pocket PC has a flash card slot (not in Palmtops).
They can also be used for downloads of data
logger data (much easier than with the heavy
notebook), e.g. Orion dataloggers with Palm
handhelds (special connecting cable required).
They can also be used for navigation using the
right software and by connecting the GPS to the
handheld (special cable required). So much of
the work such as writing notes, drafts of
manuscripts, collecting data, measurement data
can already be done in the field. The PC can be
worn in a pocket, has a weight of 150 – 300 g
and can be used up to 40 hrs without recharging.

APPENDIX X: PHOTOGRAPHING IN THE FIELD
Photographing during collecting trip and projects Of course, not all digital cameras can be used
in the field is a nice way to illustrate the results for our purposes. Not as important is the
for publications and presentations (posters, resolution. All cameras with 2 megapixels and
lectures). For bryologists, knowledge of as well more are sufficient. The latter allows prints of
as equipment for close up photography is half letter size in photo quality. In opposite, too
required. So far, photographing was mainly done high resolution causes too large picture files.
with single lens reflex cameras. For close up Important is, that they have
photography, special equipment such as close - a shortest distance of 1-4 cm (many have only
up lenses, macro objectives, extension tubes, 20 or 40 cm),
bellows, special flash lights (ring or macro - high ASA (can be varied up to 800, even 1600
flashes) and tripods were needed, a proper ASA) to allow pictures in the dark understorey
storage inclusive, which could sum up to several of forests (many have only 100 ASA),
kilograms which had to be carried around.. The - good optics (digital cameras of optical
films had to be developed, framed, and today companies have better optics than those of
usually digitised for Powerpoint presentations, computer companies such as Epson etc.)
beamer slide shows or incorporation in reports - possibility to shut off the flash
or publications. - possibility to over- or underexposure the picture
Today, digital cameras allow to get digital - possibility for spot measurement
pictures directly; they are much smaller and have - possibility to shift the focus/exposure
a lower weight, can be carried around in a pocket, - a common storing device such as a compact
some need not close up equipment, have a built- flash card.
in flash, are much easier to use, cost less than a - a filter screw at the objective
SLR equipment, have a monitor on which the - zoom without elongating the objective.
pictures can be controlled, and bad pictures can So all digital snapshot cameras are excluded.
be erased and cost no money. They are highly Among the remaining cameras, there is a
recommend and therefore photographing with recommendation for Nikon Coolpix (900, 950,
classical cameras is no more treated here. The 990, 4500) cameras. These have developed to
disadvantage that they are sensitive to high „cult cameras“ amongst biologists. The cameras
humidity is shared with 99% of all SLR cameras, allow a shortest distance of 2 cm, giving a picture
which are also operated electronically (there are size of 12 x 17 mm. If this is not enough, a
only very few manual cameras left). handlens can be mounted before the objective

152 Frahm
(a 10x handlens gives pictures of 5x7 mm size).

Most attractive is the use of a 10x lightlens. The
objective can be turned around in all directions,
allowing pictures in difficult positions without
laying in the mud or even from below (a fern
leaf). The camera has all possibilities of the
Nikon SLR cameras such as time or focus
automatic, shift of exposure/focus, spot matrix
measurement, over and under exposure etc. The
built in flash can be used for close ups (from
model 990 on, it can be directed to the object by
a fingertip). These cameras can also easily be
used at the microscope. There are expensive
microscope adapter available, but an ocular, in
which the 28 mm screw of the objective is drilled,
does the same. The monitor can be turned 90°
and allows a control of the picture when sitting
before the microscope. Unfortunately, the newest
models all have a built in battery and no
exchangeable batteries anymore, which was
easier to use in the field.
In general, the highest compression of the picture
file can be used, which results in a size of 500
kb with a resolution of 3 megapixels. A compact
flash card of 48 BM stores thus about 100
pictures.

A B
C D
E F
Fig. X.1: Examples of bryophyte pictures taken in the field with a a pocket size digital camera
(Nikon Coolpix 990) without any close up accessories. A. Tayloria magellanica, B. Hypopterygium
arbuscula, C. Chorisodontium aciphyllum, D. Schistochila spec., E. Symphyogyna spec., F. Bartra-
mia sp.

154 Frahm

APPENDIX 11: A GUIDE TO COLLECTING BRYOPHYTES IN THE TROPICS

by
Brian O’Shea*
1. Introduction
This guide is intended for any individual or often saw their herbaria as collections of ‘one of
group going to collect bryophytes in the tropics. everything’, rather than as reference collections
Experience has shown that general purpose which could be used to assist in future
expeditions are often not successful in making identification, in distributional studies, or for
useful collections of bryophytes: often only the taxonomic studies. Some plants, including
commonest and most conspicuous specimens are bryophytes, have been collected out of existence
collected. Octoblepharum albidum was called in the UK, and it is said that there is more material
‘missionary moss’ by the eminent bryologist of some very rare plants in herbaria than there is
H.N. Dixon because of the frequency with which in the wild. The UK flora is very well known,
it was collected by the inexperienced, and all and is thus particularly susceptible to such
the ‘bryophytes’ of one university expedition threats.
proved to be lichens. Even experienced In the tropics, the position is rather different.
bryologists from temperate regions do not always Very few areas are well enough known to be
make good collections unless they are prepared able to predict where rare or endangered
to expend effort in familiarising themselves with bryophytes are to be found and thus deliberate
the tropical flora before they go. This guide is over-collecting is unlikely to be a problem. Over
produced with the intention of providing the collecting is also not a problem in areas that are
necessary guidance to maximise the scientific endangered by external threats to the
value of tropical collecting expeditions. environment, such as dams or logging, and
There is a strong feeling against collecting indeed for some collecting, for instance for
amongst conservation-minded botanists, perhaps chemical analyses, quite significant quantities
because historically there has been a ‘stamp may be needed. In most areas of the tropics, so
collecting’ mentality amongst botanists. They little collecting has been done that almost any
collecting is likely to be useful. The exception
is the main tourist areas at locations such as
Luquillo Mountains (Puerto Rico), Kilimanjaro
* Reprint from British Bryological Society (Kenya) and Kinabulu (Sabah) which have been
Special Volume 3, 1989, with permission of quite well documented so that there will be little
the author. benefit in the non-expert making collections.

156 Frahm
(Permits are also required for collecting in such bryophytes appear to play a vital rôle in nutrient
places.) cycling and buffering in forests, as well as in
water relations. Evidence about this is only just
But why is collecting bryophytes from the tropics emerging, and this knowledge can only be gained
important? There are two main reasons. The by field investigation and the information
first is the pursuit of scientific knowledge, which derived from collecting.
should need no explanation or justification to
anyone going on a scientific expedition. The This guide does not cover the planning or
majority of the world’s bryophytes exist in the equipping of an expedition in any detail, only
tropics, yet they remain the least known. There how to go about preparing for your trip and what
is often a problem in identifying plants from the to do when you get there to make sure your
tropics because of the poor state of tropical collecting trip is worthwhile for both you and
bryophyte taxonomy (and the lack of relevant science. The tropics may be an exotic place for
literature - see later). Consequently attempts to you, but it is part of everyday life to the people
understand tropical ecology are limited because who live there, so your attitude to them is
of the problems of being certain about what plant important. Show your care and concern for both
is being studied. In many tropical environments, the local people and their environment to make
bryophytes form a significant part of the flora, your trip truly worthwhile.
but the study of their ecology is restricted to the If you have any comments on the material in the
few who can actually identify them. As most guide, please let the author know, and also get
tropical ecologists are not bryologists, this in touch if you have any queries - we have no
diminishes the relevance of their work. The intention of providing an expedition planning
more information we can get about bryophytes, service, but we may be able to point you to the
the more it will possible to help those attempting information you need.
a synthesis of knowledge about the tropics.
The second reason is one of survival. About a 2. Preparation

third of all tropical forest has been felled or
degraded already, and the rate is being 2.1 The tropics In deciding to make a trip to
maintained at about 20 to 40 hectares per minute. the tropics, you should already have a good idea
Most of this disappearing forest is botanically about what you can expect to find when you get
unknown, and thus potentially important there, but you should make sure you have seen
information about specific plants, and also more such publications as Richards (1975), Jacobs
general information of relevance to other areas, (1981), Longman and Jenik (1987) or Whitmore
has gone without trace. Bryophytes are (1984) for tropical forests, Walter (1971) for the
important in two major ways. Firstly, they have non-forested tropics, and Myers (1984) or
specific properties that make them important as Caulfield (1985) for the tropics in general, and
the source of various chemical products of great Pócs (1982) and Richards (1984) for bryophyte
potential value to mankind. This has been habitats. For more specific information about a
known for some time about flowering plants, but particular country or area see Davis et al (1986),
increasingly it is becoming clear that bryophytes and Greene and Harrington (1989) for available
also possess a number of biologically-active bryological literature.
substances, such as anti-carcinogenic
compounds, anti-microbial substances etc. and 2.2 Planning There are several excellent
that the greatest variety of these occur in tropical books and booklets about planning an
climates (see Ando and Matsuo (1984) for a expedition (which also contain pointers to more
summary of these and other uses). The better specialised books), so this is not covered here in
understanding we have of the floristics of these any detail. Examples are Gifford (1983),
plants, the easier it will be to exploit knowledge Blashford-Snell and Ballantyne (1977), and the
of bryophyte biochemistry. Secondly, publications of the Royal Geographical Society’s

Expedition Advisory Centre (EAC), such as If the weather is seasonal and you can choose a
Chapman (1988). Anyone planning a trip should time of year, go shortly after the rainy season,
contact the EAC at 1 Kensington Gore, London when the bryophytes will be at their best.
SW7 2AR (01-581 2057). Two short papers
(Mori and Holm-Nielson, 1981; Delgadillo, You may find throughout your trip that dealings
1987) on botanical/bryological trips to the with officialdom are eased if you travel as a
tropics, although aimed primarily at professional tourist rather than as a scientific expedition.
botanists, are nevertheless of great value to any
collecting trip in looking at both opportunities The Embassy or Tourist Bureau of your
and constraints. Books on backpacking and destination country should also be able to
‘travelling on a shoe-string’ will also be useful provide information, as might specialist travel
if you are not well-financed (e.g. Lonely Planet agencies - but all this should be found in more
Guides, Moon Guides, Frommer ‘Dollarwise’ general guides such as those mentioned at the
guides). Hatt (1985) is a cheap and useful start of this section.
general book about travelling in the tropics.
If you can, talk to your local experts about which
Research the trip thoroughly, and know what to bryophyte groups or geographical areas need
expect in the tropics in general, and in your target particular attention - this may provide a focus
country (geography, language, customs, for your expedition. Try the local bryological
currency, weather etc.). Make sure all society (the BBS in the UK - address on the front
participants are taking it seriously and are page of this guide) (O’Shea, 1985), or the local
similarly prepared, work out costs in detail and herbarium (see below). The wet tropics, rich in
allow a generous contingency, find out what both quantity and variety of bryophytes, may
clothing is appropriate, consider medical appear the most attractive area to visit, but drier
requirements, get injections organised well in areas are much less well-worked, and the plants
advance, organise your visas and discover what are different and of interest because less well-
limitations there are likely to be on tourists. known.
Work out the logistics of travel (how to get about
with your team and equipment) making any 2.3 Overseas contacts, invitations and
bookings before you go if you can - it may take collecting permits The most valuable source of
days arranging a trip across country when you assistance in making your trip worthwhile is
are there. If you need to hire porters or guides, likely to be a contact in the area you are visiting
make sure you know how to go about doing this - but you must be able to demonstrate the
and how much the going rate is. This is likely seriousness of your intent before somebody who
to be much more successful if it is arranged doesn’t know you will make any effort to help.
locally for you, so a contact in the area is very This means amongst other things having
useful. This will also help you over local laws, someone concentrating almost solely on
especially about collecting and travelling in bryophytes, and that person being able to
remote or conservation areas. It should also demonstrate some expertise. If you do not
alert you to any particular dangers including already have a contact, you could try looking in
insects, snakes and other animal life, guerrilla the International Association of Bryologists’
activity etc. - be aware of the dangers and know Compendium (Vitt, Gradstein and Iwatsuki,
how to deal with them, but don’t panic. 1985), which lists bryologists and herbaria in
most countries in the tropics, or write to the
Accommodation should be arranged before Botany Department at the University nearest to
you go. Using tents or ‘dormobile’ type your destination. You will find that an offer to
vehicles gives flexibility, but makes it less easy collect for a tropical herbarium can be used to
to get specimens dried. Missionaries are usually solicit a written invitation from that country,
very welcoming, and it may be possible to use a which is almost essential for fund raising. You
mission as a base, or just for a night or two. may not even be allowed to collect if there is no

158 Frahm
local collaboration. A local contact will also „cryptogamic“ herbarium at the British Museum
advise where collecting permits are essential (Natural History) in London, Edinburgh
(National Parks, Nature Reserves, etc.). Botanical Garden, the National Museum of
Wales at Cardiff, or possibly at a local museum
Many South American countries nowadays or herbarium . Telephone beforehand to make
require you to have a permit (to be obtained via sure that it is convenient to visit, and that there
a local herbarium) for collecting anywhere in is someone there to show you where things are
the country. Moreover, part of each collection (and for local museums, to check that they have
needs to be deposited in a local herbarium. appropriate tropical collections/literature). Their
Some local herbaria may require you to deposit experienced staff will be able to provide advice
labelled specimens before you leave the country. and information on many parts of the world. It
It is obviously essential to be aware of such rules can also be very useful to visit herbaria while
before embarking on a collecting trip in these you are in the tropics, and indeed may
countries. See Delgadillo (1987) for more sometimes be obligatory (see 2.3 above).
details.
2.8 Types of expeditions Two-person trips are
2.4 Funding The Directory of Grant-Making probably best, organised specifically to meet
Trusts (published by the Charities Aid bryological needs. However, it is sometimes
Foundation) and Grants Register (Macmillan) are useful to join a larger, interdisciplinary
very useful, but take a lot of going through and expedition (e.g. a Royal Society expedition)
result in a huge amount of letter writing - most where you may be relieved of some or all of the
of which is likely to be fruitless. The EAC have organisation. This may involve some
a small booklet listing the most useful sources compromising on areas to be visited, which may
of funds, and private companies are also worth not include the best ones for bryology.
trying.
2.5 Maps It is useful to have good large-scale 3. How much do you need to know about
maps, and Stanford’s (12-14 Long Acre, London bryophytes ?
WC2E 9LP) is the best source. They also stock
a wide range of travel guides. Map libraries are This document does not tell you how to
also useful, such as that at the Royal distinguish bryophytes from other plants: if you
Geographical Society (RGS). Look also at are unfamiliar with bryophytes, you are probably
geological maps. This will give some idea of not yet ready for a tropical collecting trip.
which areas might be promising to visit. Collecting bryophytes in the tropics will be much
more profitable if you are already familiar with
2.6 Expedition reports Writing up your your local plants, particularly with the range of
expedition is essential. It is a good idea to look variation that can be expected, so that you know
at a range of expedition reports to see what others what distinguishes one bryophyte taxon from
have done, as not only will they give you an idea another. If your trip is still some time away, and
of what is required, but they may give you you have time to get to know bryophytes before
guidance on how other expeditions have you go, first of all look at the standard texts:
prepared themselves. Botanical Gardens such Watson (1981) and Smith (1978) in the UK, and
as Kew or Edinburgh will have these, as does also Schofield (1985) for a more general account.
the library of the RGS/EAC. If this doesn’t put you off (it takes some time to
become familiar with even the common ones
2.7 Herbaria/libraries It is important to visit a amongst the thousand or so taxa in the UK - so
bryophyte herbarium before a tropical trip, to think of the problems of coping with a flora that
survey relevant literature and to look at could be even larger) attend some meetings of
specimens from the region to be visited. These your local bryological society (O’Shea, 1985)
are usually part of the „lower plants“ or or find a local bryologist who will help you. You

should certainly be able to distinguish between although they are a somewhat neglected habitat,
the main bryophyte groups, such as acrocarpous and there are interesting (and probably new) taxa
and pleurocarpous mosses, the Metzgeriales, to be found, as well as a few plants that are
Jungermanniales and Marchantiales in the confined to these habitats. A rather more
hepatics, and the hornworts, as well as having interesting flora is found in mountain streams.
examined some of the commoner tropical In running water bryophytes will always be
bryophytes. Usually the large and conspicuous attached to rocks, tree roots or the stream bed,
plants are well-known; the dull and the but some will almost always be submerged,
inconspicuous may ultimately be more whilst others will only occasionally be
profitable. Even a little knowledge will make a submerged. In standing water, a few liverworts
big difference to your effectiveness, but I must may be found floating free, and there is often an
emphasise that your collection will be interesting ephemeral flora around fluctuating
significantly more useful if you are already ponds, growing on bare mud. Although some
familiar with bryophytes, and no expedition bryophytes can tolerate brackish conditions or
should set with the main object of studying occasional salt spray, the coast is not usually a
bryophytes unless its members already have productive area, although a few bryophytes
some reasonable knowledge of bryophytes. usually can be found on old mangrove trees.
4.2 Trees Probably no trip to the tropics will

4. Where to look omit the forest habitat, but cultivated land is also
productive because of the greater light, and also
Most mosses and liverworts prefer moist, shaded because the trees are more accessible.
places, although there are a few species restricted
to deserts and other dry habitats. Normally, On trees in dense forests, the height above the
however, the moister the situation, the greater is base of the tree at which particular species of
the development of the bryophyte population, bryophytes grow depends on light and humidity.
and in tropical areas they may compose the Light intensity increases from the ground
greatest part of the visible vegetation. Wherever upwards, but relative humidity decreases (and
clouds and moist winds strike mountainsides wind movement also increases), so that inside a
through the greater part of the year, the forest forest, bryophytes such as Frullania and
may be so overgrown and draped with mosses Macromitrium spp. which require good light and
that the trees can hardly be seen, and a tree trunk are tolerant of low humidities, are found mainly
may actually be only half its apparent diameter. in the tree canopy. On isolated trees in clearings,
In other forests (particularly lowland forest), the parks, savannas, orchards etc., the canopy
degree of shade may be sufficient to severely species grow much lower down, often nearly to
inhibit bryophyte growth, with most of the flora ground level. A rich bryophyte flora can often
being in the forest canopy. If you are visiting a be found near the ground for instance on old
drier area (see also para. 2.1) choose the wet cocoa trees in plantations, but these are often
season to visit. Generally, providing there is fairly common species. Different kinds of tree
enough light, the warmth and wetness provide have different bryophyte floras, although the
good conditions for bryophytes throughout the species of tree is probably less important than
tropics. The ecology of tropical bryophytes is such things as bark acidity and chemistry, and
covered well in Pócs (1982) and Richards (1984). on the quality of the bark as a substrate: smooth
and flaky bark will usually have a poorer flora
It is worth considering four main tropical than rough bark. When collecting bryophytes
substrates in more detail: water, trees, soil and from trees, a sharp knife is essential, and a scrap
rock. of bark in the packet may add to its scientific
value, (although there are simply too many tree
4.1 Water In general, lowland tropical taxa to be able to identify them from pieces of
streams and rivers are not rich in bryophytes, bark). Fallen branches and twigs should always

160 Frahm
be examined, as should fallen trees - it may be Lowland forests have always been assumed to
the only way of seeing what grows at the top of have a rather small bryophyte flora, in contrast
the tree, unless you have tree climbing apparatus to the variety of phanerogams, but recent
(see below). Trees are perhaps the most evidence suggests that lowland forests,
complex of bryophyte habitats in the tropics, particularly canopies are much richer than
because the base, different levels of the trunk, previously thought. Montane forests (especially
the crotches, the branches and the twigs may all ‘cloud’ or ‘elfin’ forests) are, though, much
have different species. richer in both variety and quantity.
In addition, in a moist forest, many bryophytes 4.3 Soil Bryophytes growing on soil are
are epiphyllous (growing on leaves), and the particularly important in seasonally wet habitats
leaves of small trees and shrubs will be covered in climates with dry seasons, such as savanna
with bryophytes - mostly small liverworts - as etc., especially on banks of streams. They show
will buttresses, if there is enough light. You the same intolerance of variation in environment
will usually have to collect whole leaves in this as bryophytes on trees, and the resulting narrow
case, as it is easier to collect a leaf rather than selection of habitat is characteristic of these
trying to scrape the epiphylls from the leaf plants. Some species will be more or less
surface. It is also best to collect whole twigs restricted to heavy, water-holding clays, whereas
where these are covered with pendulous others are always on sand or gravel. Whether
bryophytes, despite the problems they bring in the soil is acidic or basic is extremely important
fitting them into a herbarium. See the to bryophytes, and as this may be characteristic
paragraphs on collecting for further details. for a species, it is necessary to be able to
recognise this either by testing or through
In dense forest, much of the forest life exists in knowledge of the rest of the flora. Prior research
the canopy. There are now several books on the geology of the area will help. The amount
describing the „enchanted canopy“ (e.g. of moisture available at one season or throughout
Mitchell, 1986; Perry, 1986), and Perry in the year, the amount of shade or sunlight, and
particular spends some time describing in outline whether the ground is flat or inclined are all
the mechanics of getting into the canopy and factors which determine directly the species of
moving about in it, but this requires a set of skills mosses and liverworts which occupy any habitat.
and equipment that you may not have with you. An excellent illustration of this intolerance is that
However, several bryologists have managed to we can expect, and find, totally different moss
climb trees successfully and safely (e.g. ter populations in open clay fields and on deeply
Steege and Cornelissen, 1988), usually by using shaded clay banks in a ravine. The clay is the
a bow to shoot a light cord over a high branch, same, but the other environmental factors are
and then hauling up a climbing rope. The rope radically different.
is then climbed using „ascenders“, which is hard
work and takes a long time, but is safe and In open clay fields, dried up ponds or on mud
straightforward provided you have someone with flats along rivers, very small mosses, thalloid
you who has climbing experience and is familiar liverworts and hornworts may be found which
with ropes, slings, karabiners and ascenders. are often overlooked, and could thus be very
Where conservation is not important (see para. interesting. Old termite mounds are a good
1) it may be easier to have trees felled to get to habitat for some bryophytes, such as Fissidens
the canopy bryophytes. In general, however, and Dicranella species. Man-made habitats are
fallen branches and recently felled trees will be worth examining.
sufficient.
Within lowland forests, bryophytes rarely grow
It is not safe to climb trees if you are alone, on the ground. Where there is exposed ground
whether to collect bryophytes or to see where it is often acid, even on basic rocks. The type
you are. of soil which most markedly affects the

bryophyte flora is the white sand found in secondly, during and after rain, the drought-
Amazonia, the Guianas, Borneo etc., which has resistant bryophytes become green and
a different flora from the usual clayey red or conspicuous: mosses suddenly appear
yellow lateritic soils. Another factor is altitude: prominently on rocks, walls and trees which
there are more bryophytes on banks of trails and previously seemed free of them.
roadside cuttings at higher elevations.
Light can be limiting in lowland tropical moist
4.4 Rock A single large rock may form the forest, where only about 2% of the light at the
home for a dozen different bryophytes. Just as canopy reaches the forest floor. It is also
on trees, those at the base will be moisture loving, important in its relation to evaporation, and
and those on top will be less so, depending on bryophytes in bright places are likely to be
the exposure of the rock. The relative amount drought-resistant, unless growing in permanently
of light will have a large effect, so different sides damp or wet places. Although some live in very
of an isolated rock will have different amounts low light intensities (e.g. in caves), light is still
of bryophytes as well as different species. As necessary for photosynthesis.
with soils, the composition of the rock is
extremely important, and alkaline rocks such as Altitude is an important factor in determining
limestone will support an entirely different the distribution of bryophytes as well as other
population from acidic rocks such as granite or plants, as you can see as you ascend a mountain.
quartzite. Be prepared to look for very small In the tropics, bryophytes are always more
species, particularly of liverworts; you will need abundant and varied in mountainous areas,
a knife to scrape these from the rocks. The particularly above 1000m-1500m, and in the
ceilings and walls of caves into which daylight ‘mossy’ forests (2000m-2500m). The effect of
enters should also be examined. altitude is complex, and includes amongst others:
changes in humidity, more rapid radiation of
Look as well at man-made habitats, not only heat, decrease in average temperature, decrease
where natural rock is used, but also on brick in atmospheric pressure, and increase in
(particularly on the mortar) and concrete. ultraviolet radiation. Depending on the
Culverts may combine dampness and decaying physiology of particular plants one or more of
concrete, so may have interesting species these factors will apply more decisively in
growing in them. controlling distribution. Many mosses of high,
exposed places are tinged with red, purple or
Without doubt, the most significant factor brown.
affecting bryophyte growth is the amount of
moisture available. Mosses growing on tree
trunks well above the ground or on exposed 5. How to collect
rocks will depend almost entirely on rain, dew
or fog for whatever moisture they obtain. A simple rule for collecting is to assume that if
Consequently they must be types that tolerate a two plants look different they are different; it is
level of drought. Most of them can reduce their better to collect one species several times than
evaporating area and at the same time protect to pass over a related species entirely - and the
the growing point of the plant by the rolling or range of variation may in itself be a very valuable
curving in of the leaves when they begin to dry. addition to a herbarium. Some species grow
When the plants are moistened, even after many intermingled, especially liverworts, and are
years of drought (for instance in your moss difficult or impossible to distinguish with a hand
packet), the leaves swell, become green and lens. It is also the case that bryophytes often
spread out again. Practical advantage of this have very strict ecological and habitat
can be taken in two ways when collecting: firstly, preferences, and apparently similar plants in
moistening these bryophytes often results in a different habitats may well be different species.
very rapid uncurling which aids identification; So use your instinct and rely on a plant’s „look“

162 Frahm
- a distillation of all of its characteristics: colour, Having found your bryophyte, there is no one
size, manner of growth, shape of leaf, branching, way to deal with it - different collectors favour
habitat and so on. You will notice that different methods. It is necessary first to collect
experienced bryologists usually have a good idea and packet it in some way and to record
what a plant is before they look through their collection information, then dry it (and keep it
hand lens - and part of this skill is purely dry) before packing it to take away. This topic
observational, based on a knowledge of variation is covered from a professional herbarium
amongst bryophyte taxa. Once you have „got viewpoint in Fosberg and Sachet (1965)
your eye in“ you will find this an invaluable skill.
5.1 Equipment The most important items of
In temperate climates, most bryophytes that collecting equipment are a hand lens, a knife a
produce sporophytes do so in spring or late pencil or waterproof marker, and packets. The
summer to autumn, but in more equable tropical hand lens ideally should have a magnification
climates sporophytes can be seen throughout the of x20 with a fairly wide field of view, but this
year, although each species has its own season. may not admit enough light in forests, so also
Sporophytes are often useful or even essential take a x10 or x12. (You may find in a moist
for identification, and they should therefore be forest that your lens is of little use, being
searched for and collected, notably in the constantly misted up.) Always have your lenses
Meteoriaceae, which fruit very sparingly, so any tied round your neck. Even then it is possible
fruiting stems should be picked out and packeted to lose them, so take a spare (as it may be
separately. Nevertheless, do not neglect to impossible to buy another one locally). The
collect a specimen merely because you cannot knife should be sharp and is needed for scraping
see a sporophyte. The type of male and female off or digging up your specimen; it may also be
sex organs is sometimes crucial for required to cut off pieces of bark, and scrape the
identification, so look for these also. surface of soft rock. For entering virgin forest
a machete (large knife) may be needed, unless
You should also make sure you collect the whole you go with a guide. These can usually be
plant. Many tropical epiphytic mosses have bought locally very cheaply.
conspicuous, pendant secondary stems, and
inconspicuous primary stems running along the 5.2 Packeting. Supplies of paper for packets is
bark. The latter may be vital for identification a very important detail, which should not be left
of the material. to chance. One way is to packet your collections
in envelopes made on the spot from local
Some parts of the identification process are easier newspapers, collection data being written on the
with fresh material, so even if you don’t know envelope in water-insoluble ink. Unfortunately,
the identity of a plant, it will still be useful to newspapers can be difficult to obtain in some
look for sporophytes or perianths in the field. countries, and expensive to buy, and in this case
With hepatics, it may be essential to look for oil it may be much better to take thin paper along,
bodies and to measure, count and describe them such as old computer paper. Strong brown paper
before they disappear on drying. This of course bags (as used for vegetables) are another option,
presumes the availability of a microscope in the but not if the bryophytes are wet and the gum
field. (Sometimes the local herbarium, field holding the bags together is water-soluble. If
station or botanical laboratory may be able to the mosses are very wet, gently compress them
provide a microscope.) If you do this, keep the to remove most of the water, and pack in double
shoot that was studied separate in a small packet or treble thickness packets. This method is
within the main packet. (This will guard against described in use in Cameroun by Edwards
the problems associated with a mixed collection.) (1986). Wet paper will disintegrate if not
handled carefully, so carry wet packets with care
in the field and try to dry them as soon as
possible, or transfer the bryophytes to dry

packets at the end of the day. The packets should Some bryophytes growing on soil are only very
be about 12cm x 8cm (about the size of 3in x loosely attached to the substrate, for example in
5in index cards), and can be folded as follows: marshy areas, but others may be very closely
adhering, and can only be removed with a piece
of soil. As the collection dries, this will turn
into dust, and produce a fine powder to decorate
your herbarium whenever you open the packet.
The plants are apt to break up if separated too
roughly from fresh soil, so one strategy is to wash
off excess soil gently at the earliest opportunity
- although this may well also wash away any
gemmae etc. if the washing is too vigorous. It
is probably best to pare away as much surplus
earth as possible from beneath the crust without
breaking up the specimen, and packet in stiff
paper, possibly using double packeting. Avoid
bending the packet, but press lightly. Liverwort
thalli in particular are prone to break into bits
when the crust on which they are growing cracks.
Any particularly small specimens should be Try to avoid the use of plastic bags for collecting,
wrapped in an appropriately sized piece as: they make it difficult to get the specimens
of paper before being put in a normal size packet. dry; species can get very mixed up over a period
This also applies to sporophytes or fertile parts of time; plants are likely to go mouldy or
of plants where these are in short supply, and etiolated; they are more difficult to label
have been isolated in the field - but always keep properly. Plastic bags are acceptable only if
them attached to part of the gametophyte to avoid you are going to transfer the specimens to paper
later confusion. packets within 24 hours, but this is not always
easy to do if you have collected a lot, or if the
Earlier collectors used cloth bags (10cm x 20cm), weather slows down the processing of your day’s
with collection data written on the outside in specimens. Plastic bags are likely to be less of
black lead pencil or indelible pencil, or on a slip a problem in the cool mountains than in the hot
of paper inside. A dozen or so of the smaller lowlands - but they may force ripe hepatic
bags are then put into a larger cloth sack, such sporophytes to shoot within a day.
as a 10 kilo flour or sugar sack, but this would One circumstance where you will need to use
now probably be regarded as unacceptable plastic is if you are bringing live bryophytes back
because the collections would get badly mixed. home, although you are likely to need a permit
The tendency now is to packet species separately for this. You should use very sturdy bags, not
as much as possible. This can however be time the thin sort that tend to stick to the plant.
consuming, and perhaps also difficult and Remove most of the moisture from the plants
inconvenient if the weather or light is bad, but is (squeeze them out) and there must be some air
probably easier in the field when the material is in the bag. Keep them as cool as possible e.g.
still fresh than when jumbled together in the by using a hotel room refrigerator, and reduce
packet later. Generally, though, it is best to the time kept in plastic, for instance be collecting
maximise your collecting time, and sort and them the day before departure.
document your specimens at the end of the day.
You must use your common sense - after reading
this guide you should be able to recognise good
practice, so you must balance expediency with
the scientific value of you collection.

164 Frahm
5.3 How much to collect. This can be an specimen may prove to be inadequate. This
emotive issue. In the past, large quantities were happens even with experienced collectors, for
collected, but today’s more conservation-minded instance with the liverwort families
approach suggests that you should collect more Plagiochilaceae and Lejeuneaceae.
modest quantities - enough to fill a 12cm x 8cm
packet should be plenty for one collection. (This • if a plant is in any way unusual, it may be
packet size is only approximate - choose a size important to distribute samples to a much wider
that fits the way you wish to store them.) audience.
However, you may find that you will need more
than one collection of a specimen, for: Nevertheless, even if you need five or six
collections, do not over collect, in respect of the
• your own herbarium privilege of being allowed to collect, and in
• the local herbarium, so they have samples of deference to the importance of the tropics in the
everything that you collect (in some countries ecology of the world. It is saddening to see
this may be a prerequisite of any collecting trip, torn tufts of moss hanging from trees and rocks
for instance in Brazil, but is in any case an following the depredations of the careless and
essential courtesy) wanton collector.
• if you need to send the plant to an expert for
identification, it is courteous to allow him 5.4 Labelling First of all, each packet should
to keep part or all of the specimen be uniquely identified, clearly and indelibly.
• you may wish to have specimens to exchange Most collectors use sequential numbers for this.
with other bryologists (this is a recognised Many just start with collection number 1 and
practice in botany - herbaria and collectors keep going, and this is perhaps the simplest and
frequently swap specimens to help build up least confusing way. Others elaborate this
representative collections) slightly by adding a date or a site (e.g. 8806/1 or
• a national herbarium 880601/1 or site1/1 or a combination such as 88/
site1/1) and so on, but it is important whatever
Sometimes it may only be possible to collect a method is used that mistakes in numbering do
small specimen, but remember that your not occur - and mistakes are easy when writing
collection won’t be much use as a specimen if hurriedly and in bad conditions, and especially
there is insufficient material to allow when different nationalities are involved in the
identification. Particularly for hepatics there are expedition. Pre-printed collecting notebooks
reasons why your collection should be more than with tear-off labels make it easy to maintain the
just a few scraps: sequence accurately. The numbering scheme
should also be versatile - you may have to add
•there may be a need to search a large amount extra numbers later (for instance where one
of material in order to find the rare mixed collection yields several specimens). The
inflorescence, or the perfect peristome, or to get ink from ball-point pens is liable to fade in the
a feel for the amount of intraclonal variation - tropics, so use ordinary or indelible pencil or
often very important with hepatics. waterproof markers.
• the really important plant in a collection may In field conditions, numbering sequences are not
not be what the collector saw, but some always easy to maintain (you may accidentally
minute plant mixed with it. For a taxonomist use the same number twice, for instance, or
working with such a collection it is maddening forget the last number you used), and you may
to have two stems of a possible new species, in any case prefer to sort out packet numbering
which are insufficient for adequate diagnosis. at the end of the day. In this case, temporary
Similarly, what the collector thought was one field numbers are best, such as taking the date
species may be two or three that are difficult to and time from a watch and writing this on the
distinguish, and what appeared to be an adequate packet, so that there is no chance of getting the

packets out of sequence. (Incidentally, even and have access to identification manuals. As
supposedly waterproof watches seem to leak in it gets dark about 18.00hrs in the tropics, much
the wet tropics, and it is best to use a fairly cheap packeting and documentation has to be done by
watch in a polythene bag with silica gel crystals.) artificial light. It is worthwhile taking a good
Another method is to use pre-numbered tickets, lamp.
such as cloakroom tickets or menu pads, with
the top copy going in the packet, the duplicate The general rule should be to document each
remaining as a record. It is also possible to write specimen as fully as the collector is able. A
on many living phanerogam leaves with a ball- few well-documented specimens may be much
point pen or marker, for instance when collecting more useful than a mass of poorly documented
epiphylls. ones.
If it is necessary to collect as much as possible Keep a notebook throughout your trip, and keep
in a limited time, you can put all specimens it up to date daily. If you are recording written
(packeted) from one habitat at one site in a bag data in the field, it is a good idea to use a different
together, and label the bag - but this method is notebook in the field, as this may get wet or lost,
obviously error prone. and transfer data to you formal notebook at the
end of each day. Provided it is not lost, the field
It will also be necessary to record the details notebook also provides a useful back-up to the
needed for the final herbarium packet, where formal notebook. This should contain details of
these are not constant for the site, or for the day, all the places you visit - description, latitude and
e.g. substrate (rotting log, rock etc.) and identity, longitude or other map reference, altitude (take
even if only known to family level (this saves a an altimeter) and so on, followed by the
lot of time later). The type of rock, soil or tree collection numbers for each site and habitat. You
is also very important, if you can identify, or will need this information to write out the final
describe it. For epiphytic specimens, indicate packets, but also so that you can produce lists
the precise location on the host (e.g. height on a for the sites you visit and write an account of
trunk, side of a branch) and the host species if your trip.
known; for epiphylls, indicate the host species
(again, if known, although this is often Notebooks containing waterproof paper are
impossible), which side of the leaf, and so on. available from Hawkins and Manwaring,
Deep or light shade could be added, colour when Westborough, Newark, Notts.
fresh (specimens may turn brown on drying).
This information can be written directly on the Make sure you have enough information to write
packet, or preferably in a notebook, or better still (as a minimum) the following on a label:
spoken into a dictaphone, which is quicker but
needs transcribing to a notebook at the end of • collection number
the day; tapes should be kept as a back-up in • date
case of loss/soaking of the field notebook. • identification (even if only to a broad grouping)
Remember spare tapes and batteries, and also • locality - description, latitude/longitude or grid
Murphy’s law - if a piece of equipment can go reference
wrong, it will. If you don’t have a dictaphone • habitat - aspect, substrate, vegetation type
(or if it is affected by the humidity, or goes • altitude (if possible)
wrong), it is less tedious to write directly on the • other observations - associated plants, whether
packet, and only use the notebook for less fruiting, abundance or rarity etc.
frequent notes, such as when you arrive at a new
site or habitat, but this is also more risky as paper In some places you may only have a large-scale
packets can disintegrate. If you make field map (if any), and it is then very desirable to draw
identifications, always check them again at the detailed sketch maps of collecting locations,
end of the day, when you are under less pressure together with measured distances to landscape

166 Frahm
features. The description of each location should only solution. Gentle pressing can also be an
be sufficiently detailed that it can be refound, if advantage with some thallose hepatics, but heavy
necessary. pressing and roasting can make it impossible to
recover the original cross-section.
5.5 Drying Drying is essential, otherwise fungi
will attack your collections. If you are in a dry In summary, dry as quickly as possible, at as low
area, it should be possible to air dry them over a a temperature as possible.
few days (either spread in a single layer in the
sun, or in your tent or room), but in a wet area, 5.6 Keeping them dry When specimens are
this may be a major problem and preoccupation. thoroughly dried, they should be placed carefully
For those with an unlimited budget see Croat in polythene bags to prevent them from picking
(1979) - take a pick-up truck with you, equipped up moisture from the air again. It is a good idea
with a professional herbarium drier heated by to bundle packets together with elastic bands
butane. More practical is some arrangement (about 10-12 per bundle) for ease of handling.
such as described in Frahm and Gradstein Greene (1986) recommends adding activated
(1986), using a metal frame holding a wire mesh crystals of silica gel to the plastic bags, which
shelf, surrounded by a cotton curtain, and heated are brought into the field in cotton bags, dried
from below by one or two kerosene stoves. The with the specimens and then put in one or two
apparatus weighs about 2.5 kg, but it is essential paper packets at the top of each bag. This also
to have a reliable method of drying your makes sure that any moisture that might get into
specimens, otherwise, as Frahm and Gradstein the bag due to incomplete sealing or drying will
point out, your specimens may become of more be adsorbed by the crystals and not the
interest to a mycologist. (Do not use the strong bryophytes. Mothballs are equally useful.
heat used for vascular plants, unless you hang Sturdy bags should be used to provide protection
the bryophytes above the frame. ‘Cooking’ during transport. The bags should be sealed
bryophytes will distort cell structures, preventing tightly with sticky tape. Keep a close watch on
reconstitution on re-wetting and use in later the sealed bags, and open immediately any
growth studies.) Another alternative is to use showing condensation, and re-dry the packets.
net shopping or vegetable bags, hung up over A little dichlorbenzene amongst the packets will
heaters or on a line in the sun. Don’t use a heavy deter cockroaches and other pests, if the
plant press, as this can damage sporophytes and collection is to be stored for some time. Always
some critical morphological features of the pack the notebook separately from the
specimen, although leaves with epiphylls should specimens.
be pressed lightly when drying to keep them flat,
as should very large ‘hanging’ moss 5.7 Packing for transport The physical volume
(Meteoriaceae etc.). Use dry paper in this of bryophytes that you collect may be
process, such as botanical drying paper, or thick surprisingly large, and you should think about
wads of newspaper, and change daily until dry how you are going to get them home before you
and don’t mix with bulky flowering plants. The start collecting. The stages will be:
most comfortable method, especially in the
humid tropics, is to hire an air-conditioned hotel • pack them carefully (see ‘Keeping them dry’
room - your specimens, if laid out in a single above) and tightly toavoid shaking about during
layer, should be dry within 24 hours. transport.
Delicate hepatics in particular will blacken with • be able to carry them to wherever they are to
collapse of tissue if they are not dried carefully, be despatched from (which means if you are
and cells that have collapsed in this way do not backpacking you may need to make periodic
recover when soaked. This can be a real trips back to a post office).
difficulty with epiphylls, when the phanerogam • make arrangements to pack them properly -
leaf dries slowly, and prevents the hepatic from preferably in wooden crates (such as tea
drying. A thin press, mentioned above, is the chests) that do not allow the plastic bags to be

damaged, or in strong cardboard boxes, different herbaria. If a specimen is correctly

preferably with strengthened corners. labelled it does not matter if it remains
• be aware of the official requirements for custom unidentified for years. Sorting if possible should
declarations, and the local regulations for the be done by the specialist who is studying the
export of plants, if they are to be sent directly collection. Separation into duplicates before
back home. See ‘Customs’ below. Dried plants this has been done is risky, as it may remove
can be regarded for customs purposes as evidence of plant associates, and also the ability
technically dead. to search through the material for some
• have enough money to pay for the crates and taxonomically important component.
the postage.
• agree with the local herbarium whether you 5.10 Photography Don’t underestimate the
will return duplicate specimens from home value of photography in collecting, both for
following identification, or whether you will general habitat views and for individual plant
send the specimens straight to the herbarium, and studies. A great deal of skill is necessary for
send a list later. Send them from home if at all close-up studies of bryophytes, but Edwards
possible, so you can be absolutely sure what is (1986) discusses some of the problems of
in the packet (see 5.9 below). photographing bryophytes in the tropics, and
• don’t expect the crates to get home for several provides useful advice on equipment,
weeks or possibly months. magnifications etc. The camera will provide a
valuable record, and should be used as a
5.8 Customs It may be necessary to get written supplement to your other methods of recording
permission to export specimens from the host information. An Eastman Kodak publication
country. This can sometimes be counter- on tropical photography (1986) is included as
productive, and local advice should be sought. an appendix in Chapman (1988).
If you have made contact with a local herbarium,
this will solve many of the problems, as they may 6. Suggested research topics
be able to help with the transport back home,
and will be aware of how to deal with customs, Although the purpose of this guide is to assist
postal services, etc. - but don’t expect this sort collecting, there may also be opportunity for
of help to be available unless you have arranged other types of research.
it specifically in advance. For entry into the
UK, herbaria here have special Customs 6.1 Floristics Producing a list of collections
‘Privilege Labels’ to avoid import problems. is useful in itself, and may merit publication.
These are especially useful if specimens are to This will publicise the material for possible use
be posted. Customs are increasingly opening by others, and may add significantly to our
such packages if not so labelled and can damage knowledge of phytogeography.
specimens. Discuss this with your local
herbarium. See note in para. 2.1 about dealing 6.2 Taxonomic revisions It will quickly become
with officialdom. apparent while trying to identify tropical
bryophytes how difficult this is, and the great
5.9 When you get home Once you get them need for regional and world-wide revisions. The
home, you will need to transfer them to latter will often be a very big undertaking,
permanent packets, either as you identify them, requiring access to a good library, a rich
or perhaps as a one-off exercise when you get herbarium, and especially access to a loan facility
back to check that they have all arrived safely. for borrowing material, particularly type
The danger is that packets and collection data specimens. Nevertheless, concentrating your
become separated, which is particularly likely if collections on one particular group or genus may
collections remain unsorted. It is important to provide the foundation for a contribution to such
sort, re-packet, fully label (typed labels and a revision.
photocopied duplicates) and split into sets for

168 Frahm
6.3 Ecology Taxonomic knowledge is a 8. Literature on identification

prerequisite for ecological work. There are
many possibilities here, and you will either need You may not find it easy to find books (‘floras’)
to have a very good idea in advance about what to help with the identification of your bryophyte
you are going to do, or have someone on the collection. There are not even lists of known
expedition who is able to give support and bryophytes for many tropical countries. This
advice. You will need to take any equipment position is slowly changing, despite the present
you need with you: it is unlikely to be easily lack of knowledge about the tropics. Some areas
available at your destination. Possible areas of have been worked much better than others,
investigation are: particularly the richer floras such as those of the
Caribbean and South East Asia. Floras covering
• what grows in/at a given area, altitude, habitat, quite wide areas are now becoming available,
tree, etc. for instance Eddy (1988) covers the mosses of
• what habitats are occupied by a given species the whole of S.E. Asia, and Bartram (1949) is
or genus still the most useful book for the mosses of
• estimates of bryomass at different sites or Central America. For the liverworts of tropical
altitudes America, Gradstein’s key for Puerto Rico
(Gradstein, 1989) can be used. In other parts of
6.4 Reproductive biology the tropics the position is not as good, but there
are projects underway or being considered in
• distribution of sporophytes of a given species most of the tropics, although they may take many
in different habitats years to complete. A very useful general survey
• characteristics of sporophyte production of available literature for both individual
between different species in the same habitat countries and for genera (Greene and Harrington,
• seasonal distribution of sporophytes (difficult 1988, 1989) is now available. These documents
unless you are there long enough) should give you pointers to the more general
papers on the country in which you are
interested. It may still be necessary to chase up
7. How to distinguish between different references to more detailed, local papers, and the
bryophytes British Museum (Natural History) Botany
Library may be able to provide you with
Many hints have been provided already on how photocopies (see below for details). At a higher
to distinguish tropical bryophytes sufficiently level, you will find Schofield (1985) very useful
well to be able to predict the value of collecting for deciding to which group a plant belongs, and
them. There are some characteristics of there are also generic floras for some parts of
bryophytes that have more importance and the world. Look through the bibliography for
significance for identification in the tropics than these and other books that cover your area.
in temperate regions, and vice versa, but a There are much fewer books on liverworts and
separate paper is planned on this, and so it is not hornworts than there are for mosses.
dealt with here. Although you should use the If there is no local flora in existence, you may
sources mentioned in the ‘Literature on be venturing into the area of taxonomic research.
identification’ section below as a source of It will be then that you benefit from the quality
information before you go, only collecting in the and frequency of your collections, both to get a
field, and work at the end of the day on picture of the variability of the taxa, and also to
identifying your collections, will give you a feel search for perianths, peristomes, etc., which may
for this. be necessary for identification and study.

At a major library such as the British Museum as erosion controllers and bioindicators) and
(Natural History), you will find several journals biochemical.]
concerned with bryology, such as Journal of
Bryology, Lindbergia, Journal of the Hattori Bartram, E.B. (1939). Mosses of the
Botanical Laboratory, The Bryologist and Philippines. Philipp. J. Sci. 68. Reprint
Cryptogamie, as well as many journals devoted Hbk:DM135. [A useful book for Malaysia as
to particular areas of the tropics. These will well as the Philippines. The recent reprint is
often give leads to useful papers about the area unfortunately very expensive.]
you are visiting.
Bartram, E.B. (1949). Mosses of Guatemala.
Fieldiana: Botany 25. Chicago Natural History
9. Acknowledgements Museum. [A useful book for all parts of tropical
America, well-written and easy to use, with good
I am very grateful for the contributions (some keys, descriptions and illustrations. Still
quite substantial) made to the text of this guide available quite cheaply via the Missouri
by Jeff Duckett, Jan-Peter Frahm, Rob Gradstein, Botanical Garden, although the nomenclature is
Alan Harrington, Eustace Jones, David Long, somewhat outdated.]
Royce Longton, Angela Newton, Paul Richards,
Cliff Townsend, and Martin Wigginton. Thanks Blashford-Snell, J. and Ballantyne, A. (1977).
to Martha Newton for the illustration. Expeditions, the Experts’ Way. London. [A
standard ‘how to do it’ book.]
10. Bibliography Breen R.S. (1963). Mosses of Florida.

University of Florida Press, Gainesville. [Florida
The following bibliography contains references has a relatively small number of tropical mosses,
to publications some of which you may find it but this book has good short descriptions and
difficult to locate. The British Museum (Natural quite good illustrations of most of the common
History) may be able to provide (quite expensive) mosses of the Caribbean region.]
photocopies of some of the scientific articles
(contact the Library on 01-938 9123), but copies Brotherus, V. (1924). Musci (Laubmoose). In:
of most items are held by (or have been seen by) Die natürlichen Pflanzenfamilien, Bd. 10, 11.
the author of this guide, who will be willing to Engelmann, Leipzig. Reprint, 1978: DM394. [A
help where possible. world-wide summary of all moss families and
genera, with brief keys to species.]
Where prices are quoted, these are as at June
1988 or later. Chapman, R. (1988). Tropical Forest
Expedition Manual. (3rd ed.). Expedition
Abeywickrama, B.A. (1960). The genera of Advisory Centre (1 Kensington Gore, London
mosses of Ceylon. Ceylon J. Sci. (Bio. Sci.) SW7 2AR). Pbk. £5. [The EAC is jointly
3(1):42-122. [Like other generic keys (e.g. administered by the Royal Geographical Society
Griffin and Morales (1983), Van der Wijk (1958), and the Young Explorer’s Trust, and provide an
Van der Wijk and Chopra (1966)), this is not easy information and training service. This manual
to use, as it requires knowledge of ‘phylogenetic’ is designed for those with limited expedition
characters that are often difficult to observe.] experience, who are intending to visit a relatively
unexplored tropical rain forest area. It covers
Ando, H and A. Matsuo. (1984). Applied dress and equipment, movement and navigation,
Bryology. Advances in Bryology: 2. J. Cramer. camping and cooking, local assistance, air
[Covers all known uses and potential uses of supply, and some guidance on research topics,
bryophytes, including medical, ecological (both as well as appendices on photography and
reference sources.]

170 Frahm
Croat, T.B. (1979). Use of a portable propane keys, descriptions, illustrations and habitat and
gas oven for field drying plants. Taxon 28: 573- distribution data.]
580. [Of more use to those with well-financed
logistics.] Edwards, S.R. (1986). Bryophyte collecting
and plant photography. University of Hull
Crum, H.A. and Steere, W.C. (1957). Mosses Department of Geography Miscellaneous Series
of Puerto Rico and the Virgin Islands. N.Y. 30: 65-72; 102-108. 2 fig. [Describes collecting
Academy of Sciences 7(4). [Useful for the and photography in Cameroun.]
Caribbean.]
Fleischer, M. (1902-1922). Die Musci der Flora
Caulfield, C. (1985). In the Rainforest. von Buitenzorg. 4 vols. E.J. Brill, Leiden.
Heinemann, London (Hbk:£10.95) and Pan (Reprinted in two volumes, 1976. Hbk:DM400).
(Pbk:£3.95). [A journalist’s account of the [This is strictly speaking a moss flora of Java,
nature and fate of tropical rainforest, with but it covers a large part of the Eastern tropics
emphasis on man’s assault on the forest; very and is (according to P.W. Richards) „by far the
well written, most informative, and hard-hitting. best tropical moss flora ever written, though now
Like Myers, this provides plenty of evidence to of course somewhat out of date“.]
justify a scientific expedition to the tropics.]
Florschütz, P.A. (1964). Musci of Suriname,
Davies, S.B. et al. (1986). Plants in danger : Part 1. (Flora of Suriname, Vol. 6 Part 1). E.J.
what do we know. IUCN (International Union Brill, Leiden. Pbk:DM48. [This is one of the
for the Conservation of Nature and Natural best tropical moss floras, and includes most of
Resources), Gland, Switzerland and Cambridge, the acrocarpous mosses likely to be met with in
U.K. Pbk:£15. [This is a mine of information the lowlands of the Guianas and Amazonia.]
on the flora and vegetation of each country in
the world, as a background to conservation Florschütz-De Waard, J. (1986). Musci, Part
requirement, although bryophytes don’t get a 2. (Flora of Suriname, Vol. 6 Part 1.) Pbk:DM38.
mention. It will provide a useful starting point [Continuation of Florschütz (1964). Includes
for selecting a country and finding pointers to three families of pleurocarps, including
further information.] Hookeriaceae.]
Delgadillo M., C. (1987). Additional Frahm, J.-P. and Gradstein, S.R. (1986). An
recommendations for bryologists visiting the apparatus for drying bryophytes in the field.
tropics. Taxon 36: 289-291. [Particularly aimed Bryological Times 38: 5. [Describes a home-
at professional botanists, and their relations with made, portable aluminium frame with a wire
tropical colleagues during visits, but of relevance mesh shelf, weighing 2.5 kg in total, heated with
to anyone who wants to collect. An addendum a kerosene stove, used in Borneo and South
to Mori and Holm-Nielson (1981).] America.]
Eastman Kodak. (1986). Tropical
Photography. Kodak Publication C-24. [Seven Gifford, N. (1983). Expeditions and
pages of advice on care of equipment and Exploration. Macmillan, London. [A book with
materials, exposures, processing and dealing lots of lists (e.g. what to take, what to put in your
with fungus. This item is also included as an medical kit etc.), and advice from those who have
Appendix to Chapman (1988).] planned expeditions or gone on them. An
extensive bibliography.]
Eddy, A. (1988). A Handbook of Malesian
Mosses Volume 1: Sphagnales to Dicranales. Gradstein, S.R. (1989). A key to the Hepaticae
British Museum (Natural History). Pbk:£15. and Anthocerotae of Puerto Rico and the Virgin
[The first of five parts. Indispensable, containing Islands. The Bryologist 92(3): 329-348. [A key
emphasising vegetative characters for 237

species in 92 genera of liverworts and hornworts Herzog, T. (1926). Geographie der Moose. G.
recorded from Puerto Rico and the Virgin Fischer, Jena. Reprinted 1975, DM120. [This
Islands; also useful for other parts of tropical is out of date, but is still a good introduction to
America.] floristics, including the tropics, for anyone who
can read German.]
Greene, S.W. (1986). Keeping them dry.
Bryological Times 38:6. [Describes how to keep Jacobs, M. (1988). The Tropical Rain Forest.
dried specimens dry by adding silica gel to the Springer-Verlag. Pbk:£20.65. [Original Dutch
polythene bags of dried specimens, on a trip to edition published 1981. A well presented and
Chilean rain forests.] useful general book on the structure, ecology,
physiology etc. of lowland tropical rain forests
Greene, S.W. and Harrington A.J. (1988). The with an emphasis on S.E. Asia. This sort of book
Conspectus of Bryological Taxonomic Literature is essential as a source book for planning an
- 1: Index to monographs and regional reviews. educational expedition to such areas. It
(Bryophytorum Bibliotheca 35). J. Cramer in specifically excludes other tropical areas,
der Gebrüder Borntraeger Verlagsbuchhandlung, including upland forests, where bryophytes are
Berlin and Stuttgart. Pbk:DM120 [A world- likely to be in greater abundance and variety.]
wide list of taxonomic literature, presented
alphabetically by genus and family.] Long, D.G. (1982). Collection and preservation
of bryophytes in Arabia. Bull. Emirates N. H.
Greene, S.W. and Harrington A.J. (1989). The Gp (Abu Dhabi) 18: 18-19. [A brief guide to
Conspectus of Bryological Taxonomic Literature recognising bryophytes, collecting, packeting,
- 2: Guide to national and regional literature. drying and labelling.]
(Bryophytorum Bibliotheca 37). J. Cramer in Longman, K.A. and Jenik, J. (1987). Tropical
der Gebrüder Borntraeger Verlagsbuchhandlung, forest and its environment. 2nd ed. Longman,
Berlin and Stuttgart. Pbk:DM120 [The England. Hbk:£17.50 [Excellent account of
indispensable source for the main bryological rainforest ecology in a small comprehensive
literature of countries, regions and islands of the format; some emphasis on Africa.]
world.]
Mitchell, A.W. (1986). The Enchanted Canopy.
Griffin, D. and Morales, M.I. (1983). Keys to Fontana/Collins. Pbk:£9.95. [Subtitled ‘secrets
the genera of mosses from Costa Rica. Brenesia from the rainforest roof’. Mainly describes
21: 299-323. [Useful key to genera of Central animal life, and bryophytes don’t get a mention,
America - over 200 genera are dealt with.] but lots of beautiful photographs.]
Fosberg, F.R. and Sachet, M.-H. (1965). Mori, S.A. and Holm-Nielson, L.B. (1981).
Manual for tropical herbaria. (Regnum Recommendations for botanists visiting
Vegetabile 39). International Association of Plant neotropical countries. Taxon 30: 87-89. [Aimed
Taxonomy, Utrecht. [A very detailed account of at professional botanists, with the items of more
herbarium management in the tropics. Covers general interest already mentioned in this guide.
collecting and labelling as well as herbarium See also Delgadillo (1987) for elaboration of the
techniques, procedures and administration.] more specifically bryological points.]
Hatt, J. (1985). The Tropical Traveller. Pan, Myers, N. (1984). The Primary Source: Tropical
London. Pbk:£3.95. [Preparation, equipment, Forests and our Future. Norton and Co. [An
money problems, health, exploring, etc. A very important and influential book about the
cheap way to get a feel for the problems.] importance of tropical forests to the world. The
book contains all the information and arguments
you will need to justify a trip to the tropics.]

172 Frahm
O’Shea, B.J. (1985). Bryological Societies and Smith, A.J.E. (1978). The Moss Flora of Britain
Working Groups. Bryological Times 31: 7-8. [A and Ireland. Cambridge University Press.
list of all known bryological societies and Pbk:£22.50. [The standard UK text on
working groups, giving basic information and identifying mosses, covering the whole flora.
contact points.] See also Watson (1981).]
Perry, D. (1986). Life Above the Jungle Floor. Steere, W.C. (1944). Instructions to naturalists
Simon and Schuster. [A popular account of a in the Armed Forces for botanical field work:
biologist’s discoveries in the tree tops of a Costa No. 3 The collecting of mosses and liverworts.
Rican jungle, including creating a tree top Supplement to Company D Newsletter. 1-13.
platform and a network of ropes from which he Company D, 3651 S.U. Department of Botany,
hung to observe the wildlife.] University of Michigan, Ann Arbor, USA.
[Covers how to collect, where to look, how to
Pócs, T. (1982). Tropical forest bryophytes. In: document etc. A previous (and quite successful)
A.J.E. Smith (ed.), Bryophyte Ecology. Pp. 59- attempt to cover a similar area to that of this
104. Chapman and Hall, London (Hbk:£50). guide, but now out of date in its approach to
[This is an excellent account of tropical collecting.]
bryophyte ecology, that fills out the more general
picture provided by Jacobs (1981) and Whitmore ter Steege, H. and Cornelissen, J.H.C. (1988).
(1984). It gives some idea of what species can Collecting and studying bryophytes in the
be expected in different habitats in different parts canopy of standing rain forest trees. In J.M.
of the tropics. Along with Richards (1984), Glime (ed.) Methods in Bryology, pp. 285-290.
essential.] Hattori Botanical Laboratory, Nichinan. [Briefly
reviews possible methods of tree climbing and
Richards, P.W. (1952, 5th reprint with gives details of the method the authors used in
corrections 1975). The Tropical Rain Forest. French Guiana - using various rope climbing
Cambridge University Press. Hbk:£40; techniques, which are described. Several
Pbk:£22.50. [The classic text - and written by a bryophyte species new to Guyana were found in
bryologist - but a more academic approach than the canopy.]
Jacobs (1981); still indispensable.]
Van der Wijk, R. (1958). Precursory studies
Richards, P.W. (1984). The ecology of tropical on Malaysian Mosses II. A preliminary key to
forest bryophytes. In: R.M. Schuster (ed.), New the moss genera. Blumea 9: 142-186. [Usefully
Manual of Bryology Vol. 2, pp. 1233-1270. generally for tropical Asia.]
Nichinan. [Complements Pócs (1982), based
on wider geographical area, and with a more Van der Wijk, R. and Chopra, R.S. (1966). A
detailed review of epiphylls.] preliminary key to the genera of Indian mosses.
Schofield, W.B. (1985). Collecting bryophytes Res. Bull. Panjab Univ. (N.S.) 17:149-191. [See
and processing for study. Appendix A (pp 387- comments on Abeywickrama (1960) regarding
391) of Introduction to Bryology. Macmillan. generic keys.]
Hbk:£35. [A useful general summary of where
and when to collect, collecting tools and Vitt, D.H., Gradstein S.R. and Iwatsuki Z.
methods, observations on fresh material, (1985). Compendium of Bryology.
labelling, packeting, filing and storage. The (Bryophytorum Bibliotheca Bd. 30). Verlag J.
book itself is excellent, and particularly useful Cramer, Braunschweig. [A world listing of
for the circumscriptions of each family, which herbaria, collectors, bryologists and current
may help in preliminary identifications.] research - based on data gathered in 1983-4.]

Walter, H. (1971). Ecology of tropical and

subtropical vegetation. Ed. J. Burnett, Oliver
and Boyd, Edinburgh. [This book deals with
high mountains, savannas, deserts, etc., which
are not dealt with in Jacobs, Richards and
Whitmore.]
Watson, E.V. (1981). British Mosses and

Liverworts. Cambridge University Press.
Pbk:£27.50. [A less comprehensive text than
Smith (1978) for mosses (only abbreviated
descriptions of the uncommon species), but
includes liverworts (unlike Smith). Essential for
the beginner who is serious about getting to know
British bryophytes.]
Whitmore, T.C. (1984). Tropical Rain Forests

of the Far East. (2nd ed.) Clarendon Press,
Oxford. Pbk:£25; hbk:£50. [Of general, world-
wide interest, despite the title. Rather more
academic than Jacobs (1981), and more up to
date on literature than Richards (1952).]
Whittier, H.O. (1976). Mosses of the Society

Islands. University of Florida Presses,
Gainesville. [Good for Pacific Island genera,
even if the species are different.]

174 Frahm

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2 tab.

Research needs and priorities
(1-4 modified from Gradstein (1995)
(1) Taxonomic revisions im important genera to allow identification of

specimens of difficult groups.
(2) As complete as possible inventories of different forest stands of dif-
ferent forest types.
(3) Comparisons of disturbed and undisturbed forests to assess the hu-
man impact.
(4) Studies on the ecology and reproductive biology of common and
rare species.
(5) Prepation of checklists for countries or provinces as a base of all
bryological studies.
(6) Preparation of registers of collections.
(7) Mapping of the registered data to obtain first ideas about the vertical
and horizontal distribution.
(8) Mapping of the diversity of bryophyte species per square unit (1 ha,
10 x 10 km, a province) in different parts of a country to get an estimate
of the location of hot spots and species rich areas, which can be pro-
posed for conservation.
(9) Preparation of red lists of endangered species for countries and pro-
vinces to be submitted to conservation authorities.
(10) Start of bioindication studies in urban areas or human influenced
regions.
Gradstein, S.R. 1995. Biodiversity of non-vascular epiphytes in tropi-

cal rain forests. Fibal Scientific report, Second international ESF-work-
shop on tropical canopy research, Schloß Reisensburg, 27.-30.July 1995.

196 Frahm

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Bonn, April 2003 Jan-Peter Frahm

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Research needs and priorities...........................................................................................................195

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Ecosystems. Ecosystems of the World Textbooks

Sociedad Latinoamericana de Briología.

Sastre, I., Tan, B.C. 1995. Directory of grants

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2. DIVERSITY OF BRYOPHYTE SPECIES IN THE TROPICS

2.1. Global diversity

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Congo (Brazzav.) 49 76 unusual - that one of them was properly

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An annotated catalogue. Bryophytorum 2.2.3 Tropical SE-Asia

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Tab. 2.1: Number of mosses in SE-Asia

No. of taxa No. of endemics % endemics Ref.

Bangladesh 183 3 1.6 O´Shea 2003a

Malaysian Tropics. Bryologist 56(2): of Bangladesh, with a revised checklist.

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1. in different types of forests (primary - Allen, N. S., Gradstein, S. R. & Churchill, S.

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3. ORIGIN AND AGE OF TROPICAL BRYOPHYTES

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Fig. 3.3: ranges of moss species found in Dominican amber.

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northern Andes. Mscr. Univ. Amster- 1. Presence of a central strand

The use of life forms as indicators of environ- 4. Water sacks