Neotropical Ichthyology, 13(3): 499-512, 2015
Copyright © 2015 Sociedade Brasileira de Ictiologia
DOI: 10.1590/1982-0224-20140152
Two new species of the banjo catish Bunocephalus Kner
(Siluriformes: Aspredinidae) from the upper and middle rio São
Francisco basins, Brazil
Tiago P. Carvalho1, Alexandre R. Cardoso2, John P. Friel3 and Roberto E. Reis4
Two new species of banjo catish of the genus Bunocephalus are described from the upper and middle rio São Francisco
basins of Brazil. Bunocephalus hartti is distinguished from all its congeners by the absence of serrations along the anterior
margin of pectoral-in spine in adults (vs. presence of serrations along the anterior margin of the spine). Bunocephalus
minerim can be diagnosed from all congeners, except B. larai, by the absence of an epiphyseal bar between the paired
frontals (vs. presence of the epiphyseal bar at least in adults). Bunocephalus minerim is distinguished from B. larai and other
congeners, except B. chamaizelus, by having nine principal caudal-in rays (vs. 10 principal caudal-in rays).
Duas novas espécies de peixe-banjo do gênero Bunocephalus são descritas para as bacias do alto e médio rio São Francisco
no Brasil. Bunocephalus hartti difere das demais espécies do gênero pela ausência de ganchos ao longo da margem
anterior do espinho da nadadeira peitoral em adultos (vs. presença de ganchos ao longo da margem anterior do espinho).
Bunocephalus minerim pode ser distinguido dos demais congêneres, exceto B. larai, pela ausência da barra epiiseana
entre os frontais (vs. presença da barra epiiseana ao menos nos adultos). Bunocephalus minerim pode ser distinguida de B.
larai e outras congêneres, exceto B. chamaizelus, pela presença de nove raios principais na nadadeira caudal (vs. 10 raios
caudais principais).
Keywords: Biodiversity, Endemism, Neotropical, Systematics, Taxonomy.
omnivores, feeding on terrestrial insects, larvae of aquatic
insects, small ishes, leaves and lowers (Mérigoux &
Ponton, 1998; Melo et al., 2004). Bunocephalus species
are of no commercial interest for food, but several of them
appear regularly in the ornamental ish trade (Friel, 2003).
In an attempt to improve aspredinid classiication,
some species previously included in Bunocephalus (e.g.,
Friel, 2003; Ferraris, 2007) were placed in a new genus,
Pseudobunocephalus Friel, 2008. However, hitherto there
is not an unambiguous diagnosis for Bunocephalus, and
there is no evidence that the remaining species still included
in this genus do form a monophyletic group. Therefore,
species that are included and were recently described in
Bunocephalus lack the synapomorphic characters of other
genera of Aspredinidae (Friel, 1994; Cardoso, 2010).
Introduction
Bunocephalus Kner, 1855 belongs to Aspredinidae, a
group of Siluriformes known as banjo catishes, recognized
by their distinctively depressed head and body, followed by
a slender caudal peduncle, resembling a banjo (Myers, 1960;
Friel, 2003). As currently recognized, the Bunocephalus
contains 11 valid species, distributed through most tropical
river systems in South America, such as the Orinoco,
Amazon, Paraná-Paraguay and rivers in the northwestern
slope of the Andes (Friel, 2003; Cardoso, 2010; Eschmeyer,
2014). Bunocephalus species are benthic, usually found
within leaf litter or buried in the substrate of slowlowing backwaters of creeks and rivers (Leal et al., 2011).
The species of Bunocephalus appear to be generalized
Academy of Natural Sciences of Drexel University, 1900 Benjamin Franklin Parkway, Philadelphia, PA, 19103-1195, USA. carvalho.
ictio@gmail.com (corresponding author)
2
Biosul, Rua dos Soares, 210, Bairro Palermo, 94175-265 Gravataí, RS, Brazil. arcardoso.cardoso@gmail.com
3
Alabama Museum of Natural History, The University of Alabama 119 Smith Hall, Box # 870340, Tuscaloosa, AL, 35487-0340, USA.
john.p.friel@ua.edu
4
Pontifícia Universidade Católica do Rio Grande do Sul - PUCRS, Faculdade de Biociências, Laboratório de Sistemática de Vertebrados,
Caixa Postal 1.429, 90619-900 Porto Alegre, RS, Brazil. reis@pucrs.br
1
499
500
Two new species of Bunocephalus
Furthermore, the genus Amaralia Fowler, 1954 may be
closely related to some species of Bunocephalus (Friel,
1994) and at this moment the monophyly of the genus is
uncertain. There is also a debate as to whether the type
species of Bunocephalus designated by Kner (1855) is
Silurus verrucosus Walbaum, 1792 or Bunocephalus
hypsiurus Kner, 1855 (Ferraris, 1991; 2007; Friel, 2003
contra Mees, 1988; 1996). Final resolution of these issues
may only be possible following future taxonomical changes
as result of phylogenetic analyses within the group, an
ongoing research being performed by the authors.
The rio São Francisco basin has about 181 species of
freshwater ishes with a high level of endemism, about
60% (106 spp.) of the total number of species (Albert et
al., 2011). The irst record of a banjo catish from rio São
Francisco basin was made by Mees (1989) who tentatively
identiied a single specimen from a tributary of the rio das
Velhas as B. larai. As indicated by several authors (Alves
& Pompeu, 2001, 2005; Barbosa & Soares, 2009) the
known populations of Bunocephalus in the São Francisco
basin represent two undescribed species. Here we describe
these two new species as endemics of the upper and middle
portions of the rio São Francisco basin and we comment
about their diagnostic characters.
Material and Methods
Measurements were taken point to point with a digital
caliper. Measurements are expressed as percent of the
standard length (SL), except subunits of head, expressed
as percent of the head length (HL). The measurements
follow those proposed by Friel (1994) and Cardoso (2010),
except for cleithral process length, which was taken from
the anterior margin of the cleithrum on its lateral portion
to the posterior tip of the cleithral process. Vertebral counts
include all preural vertebrae, including the ive vertebrae
modiied into the Weberian Apparatus, plus the PU1+U1
and U2 elements on the caudal skeleton counted as a single
vertebrae, according to Lundberg & Baskin, (1969) and de
Pinna & Ng (2004). Cleared and stained specimens (c&s)
were prepared according to the method described by Taylor
& Van Dyke (1985). Sex was determined by direct inspection
of gonads on dissected specimens. Anatomical illustrations
were made under a stereomicroscope using a camera lucida.
Drawings were digitized and edited using Adobe Photoshop
CC and Adobe Illustrator CC. Photos of pectoral-in spines
were taken using an AxioCam ERc5s camera attached
to a Zeiss Stemi 2000 C steromicroscope. Osteological
descriptions focused on aspects and characters used in
previous phylogenetic studies of the family Aspredinidae
(Friel, 1994; de Pinna, 1996; Cardoso, 2008). Osteological
terminology follows de Pinna (1996), except for the lacrimal,
here treated as antorbital. Institutional abbreviations follow
Sabaj Pérez (2014). Pictures were taken in a photo-tank
following the techniques described by Sabaj Pérez (2009)
with a Nikon D90 and a Nikon D7100 digital SLR.
Results
Bunocephalus hartti, new species
urn:lsid:zoobank.org:act:510B1ECB-BF95-4174-B7BC856AB3BE3FCA
Fig. 1, Table 1
Bunocephalus larai non Ihering, 1930. -Mees, 1989: 238 [doubtfully
referred]. -Friel, 2003: 263 [referred as undescribed].
Bunocephalus sp. n. -Alves & Pompeu, 2001: 185 [listed].
Bunocephalus sp. N. 1. -Alves & Pompeu, 2005: 597 [listed as
undescribed].
Bunocephalus sp. A. -Barbosa & Soares, 2009: 162 [listed].
Holotype. MZUSP 62745, 54.8 mm SL, Brazil, Minas
Gerais, Presidente Juscelino, rio Cipó at Fazenda Duas
Barras, 18°41’04”S 43°59’18”W, 1 Jul 2000, C. B. M. Alves
& P. S. Pompeu.
Paratypes. All from Minas Gerais, Brazil, rio São Francisco
basin: CAS 53522, 1, 57.7 mm SL, creek tributary to rio das
Velhas ca. 35 miles north of Belo Horizonte, 2 Feb 1941, T.
D. White and others. MCP 45236, 1, 39.9 mm SL, Augusto
de Lima municipality, rio Curimataí, tributary to rio das
Velhas, 17°59’32”S 44°10’47”W, 9 Jul 2009, C. G. Leal
& D. C. de Carvalho. MCP 48280, 1 c&s, 21.5 mm SL,
Iguatama municipality, ribeirão São Miguel, tributary to
rio São Francisco, 20°12’04”S 45°39’12”W, 26 Sep 2003,
B. P. Nogueira and others. MNRJ 31385, 4 (1c&s), 37.242.6 mm SL, Piumhi municipality, mouth of rio Piumhi,
20°20’31”S 45°59’03”W, 28 Feb 2007, P. L. Gallo, L. H.
Silva, D. L. Z. Kantek & W. A. M. Perez. MZUSP 39443,
1, 45.5 mm SL, rio Formoso, tributary to rio São Francisco,
8 Feb 1988, Y. Sato. MZUSP 39480, 1, 40.0 mm SL, rio
São Francisco at mouth of rio Formoso, approx. 17°26’S
44°57’W, 8-10 Feb 1988, Y. Sato. MZUSP 64227, 3, 36.044.7 mm SL, Juataba municipality, rio Paraopeba between
Juataba and Betim, approx. 19°56’S 44°18’W, 6 Nov 2000,
C. B. M. Alves.
Diagnosis. Bunocephalus hartti is distinguished from all
congeners by the absence of serrations along the anterior
margin of the pectoral-in spine in adults (Fig. 2a; vs.
presence of serrations along the anterior margin in adults,
Fig. 2c). Bunocephalus hartti can be further distinguished
from most congeners, except for B. verrucosus, by having
the last dorsal-in ray completely or almost completely
adnate to the dorsum (vs. dorsal-in ray completely free or
with less than half extension connected to the dorsum).
Description. Morphometric data summarized in Table
1. Maximum body size moderate to small compared to
congeners (maximum observed size 57.7 mm SL). Dorsal,
left lateral and ventral views of body in Fig. 1. Head and
body depressed, lateral proile ascending from tip of
T. P. Carvalho, A. R. Cardoso, J. P. Friel & R. E. Reis
snout to dorsal-in origin, with bony skull ornamentations
in between. Posterodorsal proile of body straight and
descending from dorsal-in origin to near base of caudal
in, becoming slightly convex anterior to caudal-in base.
Ventral body proile convex from mouth to insertion of
pelvic in; concave from this point to anal-in origin,
straight and ascending from anal-in origin to base of caudal
in, slightly concave at caudal-in base. Caudal peduncle
slender, somewhat rounded in cross section, but lattened
dorsally and ventrally, shallowest at midpoint between end
of anal in and caudal-in origin.
Skull ornamentation weakly developed. Eye small and
positioned dorsolaterally. Skin covering eye dense and
pale. Anterior nostril located terminally at tip of snout,
associated with leshy tube projecting beyond upper lip.
501
Posterior nostril without lap, opening anteromedially near
eye. Mouth subterminal, upper lip more prominent relative
to lower lip. All barbels simple, unbranched; maxillary
barbel reaching or slightly surpassing insertion of pectoralin spine, posterolateral mental barbel twice as long as
anteromedial one. Opercular opening reduced to small
valvular slit located just anterior and medially to insertion
of pectoral-in spine. Axial slit pore present, dorsoventrally
inclined underneath posterior cleithral process. Adult
males with digitiform testes. Integument covered with
large unculiferous tubercles, forming series of aligned
longitudinal rows on posterior portion of body. Large and
well-deined rows of tubercles on caudal peduncle, one on
middorsum, and three on lateral of body. Other rows poorly
deined.
Fig. 1. Bunocephalus hartti, MZUSP 62745, holotype, 54.8 mm SL, rio Cipó at fazenda Duas Barras, Presidente Juscelino,
Minas Gerais, Brazil.
502
Two new species of Bunocephalus
Fig. 2. Right pectoral-in spines of cleared and stained
specimens, in dorsal view. a. Adult specimen of Bunocephalus
hartti, MNRJ 31385, 42.2 mm SL. b. Juvenile specimen of
Bunocephalus hartti, MCP 48280, 21.5 mm SL. c. Adult
specimen of Bunocephalus minerim, MCP 28379, paratype,
48.4 mm SL. Scale bar 1 mm.
Osteological description based on two cleared and
stained specimens (21.5-42.2 mm SL, see paratype
list). Anterior margin of mesethmoid slightly concave,
anterolateral projection slightly pronounced (Fig. 3a).
Ethmoid cartilage separate from articular facet of palatine.
Frontal with lateral projections forming dorsal margin of
eye. Frontal posteriorly projected laterally to posterior
cranial fontanel and contacting supraoccipital, epiphyseal
bar present. Supratemporal fossa present at middle portion
of contact between pterotic and supraoccipital bones.
Pterotic with laterally expanded and pointed bony shelf.
Premaxilla with somewhat rectangular shape, bearing
few teeth on its posteromedial margin. Dentary slender,
abutting counterpart at medial portion, symphyseal portion
slightly expanded, teeth present along anterior half of
dorsal margin. Ascending process of Meckel’s cartilage
present. Coronomeckelian bone present. Hyomandibula
associated with preopercle and posterior portion of
mandibular laterosensory canal, supraopercle absent.
Cartilaginous contact of hyomandibula with neurocranium
restricted to sphenotic bone. Anterodorsal process of
hyomandibula developed, contacting ventral surface
of sphenotic. Opercular condyle of hyomadibula well
developed. Metapterygoid present, contacting quadrate and
hyomandibula. Endopterygoid present, somewhat triangular
in shape, located underneath contact of palatine and lateral
ethmoid. Posterior margin of palatine cartilaginous and
rounded. Opercle “L” shaped, posterior arm larger than
ventral arm. Interopercle present, triangular in shape and
irmly attached to ventral arm of opercle. Dorsal hypohyal
absent. Anterior ceratohyal with expanded lamina on
anteroventral margin, contacting posterior ceratohyal by
means of cartilage and interdigitated suture. Posterior
ceratohyal with foramen on midventral portion. Interhyal
present. Four branchiostegal rays. Urohyal present,
pointed anteriorly, with foramen and well developed
lateral wings. First and second pharyngobranchials absent;
third and fourth present and ossiied. First hypobranchial
ossiied, second and third cartilaginous. Second and
third basibranchial ossiied, fourth cartilaginous. Third
epibranchial bearing uncinated process. Gill rakers present
in all branchial arches, but limited to few on irst and
second arches. Pharyngeal teeth well developed on upper
tooth plate; about two rows of teeth on ifth ceratobranchial
limited to its medial margin.
Dorsal lamina of Weberian complex reaching dorsal
surface of body, lateral proile of lamina ascending
posteriorly with anterior concavity and bony knob at middle
portion and elevated crest posteriorly. Parapophysis of
fourth vertebra forming broad lamina over swim bladder,
contacting parapophysis of ifth vertebrae extensively.
Parapophysis of ifth vertebra long, extending to lateral
body surface transverse to main body axis. Distal portion of
ifth parapophysis expanded. Total vertebrae 35. Vertebra
bearing horizontal transverse processes from centrum
nine to 31. Hemal spine contacting anal-in pterygiophores
biid. Four to ive pairs of ribs (modally ive), on vertebrae
six to nine or ten. Abdominal vertebrae foramina (hemal
arches) for hemal canal on 6th or 7th and posteriorly on 10th
vertebrae.
Fig. 3. Dorsal view of neurocranium and Weberian complex.
a. Bunocephalus hartti, MNRJ 31385, paratype, 42.2 mm
SL. b. Bunocephalus minerim, MCP 28379, paratype, 43.1
mm SL. ACF anterior cranial fontanel; EX, extrascapula;
EP, epioccital; F, frontal; LAT, lateral ethmoid; SPH,
sphenotic; SOC, supraoccipital; STF supratemporal fossa;
SU, supracleithrum; PCF, posterior cranial fontanel; PTE,
pterotic; VP4 parapophysis of fourth vertebra; and VP5
parapophysis of ifth vertebra. Scale bar = 5 mm.
503
T. P. Carvalho, A. R. Cardoso, J. P. Friel & R. E. Reis
Dorsal in with ive or six rays (modally ive), without
spinelet. First ray unbranched followed by four or ive
branched rays. Membrane of last dorsal-in ray adnate to
dorsum. Anterior nuchal plate absent, middle nuchal plate
contacting posterior nuchal plate laterally. Posterior nuchal
plate not developed laterally, lateral limit not extending
beyond contact with middle nuchal plate. Pectoral in
with one rigid spine and ive branched soft rays. Pectoral
spine curved, feeble serrations present in anterior portion
in juveniles, but absent in adults (Figs. 2a,b). Serrations
along posterior margin of spine increasing in number with
larger body sizes, maximum of 10 serrations on posterior
margin. Two ossiied plus one cartilaginous pectoral-in
radial. Postcoracoid process of pectoral girdle extending
slightly posterior to postcleithral process in lateral view.
Pelvic in with six soft rays, second and third rays longest,
not reaching anal-in origin, irst ray unbranched. Posterior
margin of basipterygium jagged. Lateral cartilage of
basipterygium extending from its anteriormost portion
to contact with last pelvic-in ray. Anal in with seven to
nine rays (modally eight), irst two or three unbranched,
third of length of last anal-in ray extension adnate by
membrane to body. Caudal in with ten principal rays,
ive associated with upper lobe and ive with ventral lobe,
posterior margin of caudal in convex. Lowermost and
uppermost caudal-in rays unbranched, with proximal
expansion and slightly shorter than branched middle rays.
Caudal in with two procurrent rays on upper and lower
lobes, anterior procurrent ray small, triangular in shape,
posterior procurrent ray longer and spine like. Posterior
margin of upper hypural plate extending posteriorly
further than lower hypural plate (hypurals one and two
fused with parhypural). Second ural half-centrum well
developed. Adipose in absent.
Table 1. Morphometric data of holotype and paratypes of Bunocephalus hartti (n=11 including the holotype) and
Bunocephalus minerim (n=17 including the holotype). H= holotype; Max = maximum; Min = minimum; SD = standard
deviation; and unb = unbranched.
Bunocephalus hartti
Standard length
H
Min
54.8
35.8
Bunocephalus minerim
Max
Mean
SD
H
Min
Max
Mean
54.8
41.75
-
37.9
29.5
45.4
38.2
1.92
24.0
22.1
27.3
23.7
SD
-
Percent of standard length
Head length
22.6
21.9
27.7
24.1
1.31
Prepectoral length
20.4
20.4
26.1
23.1
1.58
24.2
21.9
26.0
24.0
1.09
Cleithral width
27.4
26.9
29.2
28.0
0.74
29.8
27.1
31.4
29.7
1.14
Maximum head depth
12.9
12.1
14.6
13.4
0.73
14.7
12.6
15.0
13.8
0.67
Pectoral-spine length
20.2
20.3
25.8
24.1
1.64
24.8
21.0
25.8
23.7
1.53
Distance between coracoid processes
19.9
16.0
23.1
18.5
2.09
21.1
16.6
21.1
19.2
1.31
Coracoid process length
9.8
7.0
9.9
8.7
1.10
10.0
8.8
12.6
11.0
1.06
Distance between cleithral processes
20.9
19.1
21.5
20.1
0.77
22.9
20.6
24.6
22.7
1.12
Cleithral process length
13.8
12.9
15.4
14.1
0.75
15.5
13.0
15.6
14.5
0.69
Predorsal length
43.0
42.3
45.0
43.4
0.77
42.7
40.1
45.8
42.0
1.69
Depth at dorsal-spine insertion
11.7
10.4
13.6
12.1
0.93
14.5
10.4
17.1
14.2
2.02
Dorsal-spine length
12.6
12.4
17.0
14.7
1.44
14.7
14.8
18.2
16.4
1.07
Prepelvic length
43.8
42.0
49.3
45.0
2.26
48.5
40.4
48.5
43.7
2.24
Length of 1st unb. pelvic-in ray
11.3
11.3
13.7
12.3
0.77
12.9
12.6
15.1
13.8
0.92
Preanal length
60.9
60.6
63.4
62.0
0.95
63.8
56.6
63.9
60.0
2.07
Anal-in base length
19.1
14.7
19.2
16.3
1.21
21.6
18.1
23.6
20.6
1.56
Caudal-peduncle length
18.8
18.1
24.6
21.9
2.10
18.4
18.1
23.6
20.6
1.56
Caudal-peduncle depth
4.2
3.9
4.7
4.3
0.27
4.7
3.6
5.1
4.4
0.41
Caudal -in length
15.7
15.7
23.7
20.8
2.24
22.9
21.1
27.3
24.2
1.67
Snout length
29.0
26.5
32.3
29.8
1.49
35.1
26.8
35.2
30.0
2.26
Eye diameter
8.9
7.4
10.4
9.0
0.90
8.7
8.6
12.2
10.4
1.17
Percent of head length
Interorbital width
25.0
23.5
30.2
26.9
2.36
31.8
30.2
35.8
33.0
1.52
Maxillary-barbel length
71.7
64.3
94.8
76.0
9.98
87.9
72.0
111.1
93.4
11.36
Distance between anterior nares
16.9
16.7
21.9
18.7
1.90
21.9
17.3
24.7
22.0
1.90
Distance between posterior nares
27.4
24.0
31.3
28.4
2.46
31.8
27.6
34.5
31.9
1.73
Mouth width
33.8
31.1
41.7
36.2
3.33
36.2
32.2
45.3
38.3
3.38
504
Two new species of Bunocephalus
Nasal canal ossiied and positioned laterally to mesethmoid,
one or two separate tubular ossiications around canal.
Antorbital present, with anterior limb pointed, extending
anterior to anterior margin of premaxilla. Antorbital mesial
limb rounded and associated with laterosensory canal.
Infraorbital canal present with three tubular ossiications,
canal exiting antorbital, passing below eye margin and
entering neurocranium through sphenotic. Mandibular canal
interrupted, with two tubular ossiications lateral to posterior
portion of dentary, and two tubular ossiications near to
contact with preopercle. Extrascapular present. Lateral line not
associated with fourth parapophyses, anterior portion running
just aside margin of parapophyses. Lateral line complete,
extending variably to caudal peduncle, formed by simple
tubes, median portion presenting small inconspicuous hooks.
Color in alcohol. Head and body light brown dorsally,
ventral portions lighter brown to yellowish pale. Four
saddles of dark coloration on dorsal surface of body. First
dark saddle at level of dorsal in, second at anal in vertical,
third at middle caudal peduncle and fourth at origin of
caudal in. Second, third and fourth saddles sometimes not
connected at middorsal line. Dorsal in mostly dark brown
with light distal margin; pectoral in whitish cream to
hyaline with small light brown spots, pelvic and anal ins
mostly hyaline, without conspicuous dark spots. Caudal in
hyaline with dark blotch at proximal portion and scattered
black spots on distal third sometimes forming band.
Distribution. Known from several tributaries of the
upper and middle rio São Francisco basins including
the das Velhas, Paraopeba and Formoso rivers in Minas
Gerais State, Brazil (Fig. 4).
Etymology. The epithet hartti is a patronym honoring
Charles Frederick Hartt, a Canadian-American geologist,
and first professor of Geology at Cornell University.
Hartt worked extensively in Brazil, and a few of his
notable accomplishments include the publication of
“Geology and physical geography of Brazil” (Hartt,
1870), and serving as the founder and director of the
section of geology at the Museu Nacional of Brazil from
1866 to 1867.
Conservation status. Bunocephalus hartti is known
from an Extent of Occurrence (EOO) of approximately
34,000 km 2 , and despite some areas within its range
suffer continuing decline in habitat quality because
of contamination from the city of Belo Horizonte and
also mining and agriculture, there is no evidence of
its habitat being severely fragmented or occurring
extreme fluctuations in range or number of individuals.
Considering that no specific threats to the species were
detected, B. hartti is categorized as Least Concern (LC)
according to the International Union for Conservation of
Nature (IUCN) categories and criteria (IUCN Standards
and Petitions subcommittee, 2014).
Fig. 4. Distribution map of the new species of Bunocephalus in the rio São Francisco basin, Brazil. Circles represent
Bunocephalus hartti and squares represent B. minerim, solid symbols indicate the type-localities.
T. P. Carvalho, A. R. Cardoso, J. P. Friel & R. E. Reis
Bunocephalus minerim, new species
u r n:lsid:zooba n k.org:act:41D3FB3D -FA F6 - 4894 -A8C95EC56878B8FB
Figs. 5, 6, Table 1
Bunocephalus sp. N. 2. -Alves & Pompeu, 2005: 597 [listed as
undescribed].
Bunocephalus sp. B. -Barbosa & Soares, 2009: 162 [listed].
Bunocephalus sp. -Leal et al., 2011: 148 [ecology]. -de Carvalho et
al., 2011: 85 [barcoded].
Holotype. MCP 47087, 37.9 mm SL, Brazil, Minas Gerais,
Guarda-Mor municipality, córrego Guarda-Mor, near the
town of Guarda-Mor on highway BR-364, 17°44’52”S
47°05’40”W, 21 Jan 2012, R. E. Reis, E. H. L. Pereira & P.
C. Lehmann A.
Paratypes. All from Minas Gerais, Brazil, rio São
Francisco basin: ANSP 172080, 1, 24.4 mm SL, Curvelo
municipality, rio Picão approx. 20 km NE of Curvelo,
18°36’14”S 44°17’06”W, 11 Jul 1993, S. A. Schaefer and
others. MCP 16742, 1, 36.8 mm SL, Curvelo municipality,
rio Picão at road from Curvelo to Monjolos, 18°36’14”S
44°17’06”W, 11 Jul 1993, R. E. Reis, E. L. Pereira &
S. A. Schaefer. MCP 28378, 1, 45.3 mm SL, Paracatu
municipality, stream at highway BR-040 from Paracatu to
João Pinheiro, 17°18’15”S 46°46’16”W, 24 Jan 2001, C. A.
S. Lucena, J. F. P. Silva, E. L. Pereira & A. R. Cardoso. MCP
28379, 2 c&s, 43.1-48.4 mm SL, Guarda-Mor municipality,
córrego Macaúba tributary to rio Claro on road between
Coromandel and Guarda-Mor, 17°58’57”S 47°06’41”W,
24 Jan 2001, C. Lucena, J. F. P. Silva, E. H. L. Pereira &
A. R. Cardoso. MCP 34665, 3 (1 c&s), 29.8-32.9 mm SL,
Iguatama municipality, ribeirão São Miguel tributary to
rio São Francisco, 20°12’00”S 45°39’09”W, 26 Sep 2003,
B. P. Nogueira. MCP 45156, 10, 36.2-42 mm SL, Augusto
de Lima municipality, rio Curimataí, tributary to rio das
Velhas, 17°59’34”S 44°10’48”W, 9 Dec 2009, C. G. Leal
& D. C. Carvalho. MCP 45242, 23 (2 c&s), 11.2-41.7 mm
SL, Augusto de Lima municipality, rio Curimataí, tributary
to rio das Velhas, 17°59’34”S 44°10’48”W, 9 Dec 2009,
C. G. Leal & D. C. Carvalho. MCP 45759, 1, 42.4 mm SL,
Augusto de Lima municipality, rio Curimataí, tributary to
rio das Velhas, 17°59’34”S 44°10’48”W, 24 May 2009, F. A.
Sampaio, F. Suzuki & R. Couto. MCP 45763, 1, 38.8 mm SL,
Augusto de Lima municipality, rio Curimataí, tributary to
rio das Velhas, 17°59’34”S 44°10’48”W, 24 May 2009, F. A.
Sampaio, F. Suzuki & R. Couto. MNRJ 26500, 3, 30.6-36.6
mm SL, Iguatama municipality, rio São Miguel at Fazenda
Doce de Leite, 20°11’59”S 45°39’09”W, 21 Dec 2003, B.
P. Nogueira, G. A. Pereira & R. E. S. Hojo. MNRJ 31414,
3, 23.1-37.6 mm SL, Pains municipality, ribeirão Moendas,
tributary to rio São Miguel, 20°26’31”S 45°40’00”W, 26 Aug
2007, P. A. Buckup, M. R. Britto & U. Jaramillo. MZUSP
39444, 2, 27.8-30.3 mm SL, rio Formoso, tributary to rio
505
São Francisco, 8 Feb 1988, Y. Sato. MZUSP 73800, 5, 34.442.4 mm SL, Augusto de Lima municipality, rio Curimataí
at fazenda Vitória, 18°05’43”S 44°16’15”W, 16 Aug 2001,
C. B. M. Alves & P. S. Pompeu. UFRGS 11273, 2, 40-40.9
mm SL, Unaí/Palmeirinha municipalities, stream at road
between Unaí and Palmeirinha, 16°09’22”S 46°44’48”W,
11 Sep 2009, G. Frainer, F. R. Carvalho & V. A. Bertaco.
UFRGS 11374, 1, ixed and preserved in ethanol 95%, 42.4
mm SL, Unaí/Palmeirinha municipality, stream at road
between Unaí and Palmeirinha, 16°09’22”S 46°44’48”W,
11 Sep 2009, G. Frainer, F. R. Carvalho & V. A. Bertaco.
Sequences. Genseq-2 COI: Published sequences from
a barcode study of the fauna of the São Francisco River
basin (de Carvalho et al., 2011) identiied as Bunocephalus
sp. are available of paratypes MCP 45156, MCP 45763 and
MCP 45759. These correspond to genseq-2 COI following
the nomenclature of Chakrabarty et al. (2013). GenBank
accession numbers for these sequences are: HM405074 and
HM405075 for MCP 45156; HM405076 for MCP45763; and
HM405077 for MCP45759.
Diagnosis. Bunocephalus minerim can be diagnosed
from all congeners, except B. larai, by the absence of an
epiphyseal bar between the paired frontals (vs. presence of
the epiphyseal bar at least in adults). Bunocephalus minerim
is distinguished from B. larai and other congeners, except
B. chamaizelus, by having 9 principal caudal-in rays (vs. 10
principal caudal-in rays).
Description. Morphometric data summarized in Table 1.
Maximum body size small compared to congeners (maximum
observed size 48.8 mm SL). Dorsal, left lateral and ventral
views of body in Figs. 5-6. Head and body depressed, lateral
proile almost straight and ascending from supraoccipital tip
to dorsal-in origin, bony humps at posterior end of Weberian
lamina and at middle nuchal plate. Posterodorsal proile of
body straight and descending from origin of dorsal in to
caudal-in base. Ventral body proile convex from mouth
to insertion of pelvic in; concave from this point to analin origin, straight and ascending from anal-in origin to
base of caudal in, slightly convex at caudal-in base. Caudal
peduncle slender, round in cross section, shallowest at
midpoint between end of anal in and caudal-in origin.
Skull ornamentations weakly developed, conspicuous
bony knobs absent in adult specimens. Eye small and
positioned dorsolaterally. Skin covering eye dense and
pale. Anterior nostril located terminally at tip of snout,
associated with leshy tube projecting beyond upper lip.
Posterior nostril without lap, opening anteromedially near
eye. Mouth subterminal, upper lip more prominent relative
to lower lip. All barbels simple, unbranched, maxillary
barbel slightly surpassing insertion of pectoral-in spine,
posterolateral mental barbel twice as long as anteromedial
one. Opercular opening reduced to small valvular slit
located just anterior and medially to insertion of pectoral-
506
Two new species of Bunocephalus
in spine. Axial slit pore present, dorsoventrally inclined
underneath posterior cleithral process. Some female
specimens with eggs attached to lateral and ventral surface
of the body and pectoral, pelvic and anal ins (Fig. 6). Adult
males with digitiform testes. Integument covered with
small unculiferous tubercles, those on posterior portion
of body larger and forming series of aligned longitudinal
rows. Large and well-deined rows of tubercles on caudal
peduncle, especially in juveniles, one on middorsum and
three on lateral of body. Other rows poorly deined.
Fig. 5. Bunocephalus minerim, MCP 47087, holotype, 37.9 mm SL, córrego Guarda-Mor near the town of Guarda-Mor on
highway BR-364, Minas Gerais, Brazil.
T. P. Carvalho, A. R. Cardoso, J. P. Friel & R. E. Reis
507
Fig. 6. Paratype specimens of Bunocephalus minerim. a. Dorsal view of MCP 34665, 31.5 mm SL; showing the saddled
pattern of a distinct color morph. b. Left lateral view of a mature female, MCP 45242, 41.4 mm SL, showing the eggs
adhered and marks of previously attached eggs on the lateral and ventral portions of body and ins.
Osteological descriptions based on ive cleared and
stained specimens (32.6-47.6 mm SL). Anterior margin
of mesethmoid straight, anterolateral projections absent.
Ethmoid cartilage separate from articular facet of palatine.
Frontal with lateral projections forming dorsal margin of
eye. Frontal posteriorly projected laterally to posterior
cranial fontanel and contacting supraoccipital, epiphiseal
bar absent. Supratemporal fossa present at middle portion
of contact between pterotic and supraoccipital bones.
Pterotic with laterally expanded and pointed bony shelf.
Premaxilla with somewhat rectangular shape, bearing teeth
on its posteromedial margin. Dentary slender, abutting
counterpart at medial portion, symphyseal portion slightly
expanded, teeth present along most of dorsal margin of
dentary. Ascending process of Meckel’s cartilage present,
contacting main portion of this cartilage. Coronomeckelian
bone present. Hyomandibula associated with preopercle
and posterior portion of mandibular laterosensory
canal, supraopercle absent. Cartilaginous contact of
hyomandibula with neurocranium restricted to sphenotic
bone. Anterodorsal process of hyomandibula developed,
contacting ventral surface of sphenotic. Metapterygoid
present (bilaterally absent in some specimens), small in
size and ventrally contacting dorsal margin of quadrate.
Endopterygoid present, somewhat squared in shape,
located underneath contact of palatine and lateral ethmoid.
Posterior margin of palatine cartilaginous and rounded.
Opercle “L” shaped, posterior arm larger than ventral arm.
Interopercle present, triangular in shape and irmly attached
to ventral arm of opercle. Dorsal hypohyal absent. Anterior
ceratohyal with expanded lamina on anteroventral margin,
contacting posterior ceratohyal by means of cartilage and
508
Two new species of Bunocephalus
interdigitated suture. Posterior ceratohyal with foramen
on midventral portion. Interhyal present. Typically ive
branchiostegal rays (four on one side in few specimens).
Urohyal present, pointed anteriorly, without foramen.
Lateral wings and dorsal keel of urohyal reduced or
absent. First and second pharyngobranchials absent, third
present and ossiied, fourth present and variably ossiied.
First hypobranchial ossiied, second variably ossiied and
third cartilaginous. Second basibranchial ossiied, third
variably ossiied. Third epibranchial bearing uncinated
process. Gill rakers absent on irst and second branchial
arches, small and scattered on third and fourth arches.
Pharyngeal teeth well developed on upper tooth plate;
about two rows of teeth on ifth ceratobranchial.
Dorsal lamina of Weberian complex reaching dorsal
surface of body, lateral proile of lamina ascending
posteriorly with anterior concavity and bony knob at
middle portion (Fig. 3b). Parapophysis of fourth vertebra
forming broad lamina over swim bladder presenting hookshaped process at posterior lateral margin. Parapophysis of
fourth vertebra contacting parapophysis of ifth vertebrae
extensively. Parapophysis of ifth vertebra long, extending
to lateral body surface transverse to main body axis. Distal
portion of ifth parapophysis expanded in adult specimens.
Total vertebrae 34-37 (modally 34). Vertebrae bearing
horizontal transverse processes from 7th to penultimate
centrum. Hemal spines simple, those contacting anal-in
pterygiophores not biid. Four pairs of ribs, on vertebrae six
to nine. Abdominal vertebrae foramina (hemal arches) for
hemal canal on 6th or 7th and posteriorly on 11th vertebrae.
Dorsal in with four or ive rays (modally ive), without
spinelet. First ray unbranched followed by three or four
branched rays. About half length of last dorsal-in ray
adnate to dorsum by membrane. Anterior nuchal plate
absent, middle nuchal plate contacting posterior nuchal
plate laterally. Posterior nuchal plate not developed
laterally, lateral extension passing slightly beyond contact
with middle nuchal plate. Pectoral in with one rigid spine
and ive branched soft rays, last one variably branched.
Pectoral spine curved, bearing recurved serrations
along ¾ of its anterior and posterior margins. Serrations
increasing in number with larger body sizes, to maximum
of 10 serrations on anterior and posterior margin of
pectoral spines. A single ossiied plus one cartilaginous
pectoral-in radial, one specimen with two ossiied and
one cartilaginous radial. Postcoracoid process of pectoral
girdle extending slightly posterior to postcleithral process
in lateral view. Pelvic in with six soft rays, second and
third rays longest, just reaching to anal-in origin, irst
ray unbranched. Basipterygium with reniform shape, its
posterior margin jagged and not bearing cartilaginous tip
in adults. Lateral cartilage of basipterygium extending
from anteriormost portion of bone to contact with last
pelvic-in ray. Anal in with seven to nine rays (modally
nine), irst three to ive unbranched. Caudal in with nine
principal rays, ive associated with upper lobe and four
with ventral lobe, posterior margin of caudal in convex.
Lowermost and uppermost principal caudal-in rays
unbranched with proximal expansion and slightly shorter
than branched middle rays. Caudal in with two procurrent
rays on upper and lower lobes, anterior procurrent circular
to rectangular in shape, posterior procurrent ray longer
and spine-like. Posterior margin of upper hypural plate
extending posteriorly further than lower hypural plate.
Second ural half-centrum well-developed. Adipose in
absent.
Nasal canal ossiied positioned laterally to mesethmoid,
variably one or two tubular ossiications around canal.
Antorbital present, with anterior limb pointed, extending
anterior to anterior margin of premaxilla. Antorbital mesial
limb rounded and associated with laterosensory canal.
Infraorbital canal present with three tubular ossiications,
canal exiting antorbital, passing below eye margin and
entering neurocranium through sphenotic. Mandibular
canal interrupted, with one tubular ossiication lateral
to posterior portion of dentary, and one or two tubular
ossiication near to contact with preopercle. Extrascapular
present. Lateral line not associated with fourth
parapophysis, anterior portion running just ventrally to
margin of parapophysis. Lateral line complete; extending
variably to caudal peduncle, with simple ossiied tubes
presenting variably few inconspicuous hooks on anterior
portion.
Color in alcohol. Pigmentation variable with two distinct
color morphs of overall dark and light patterns (Figs. 6a-b).
Dark morph with dorsal portion of head and body dark
brown with three poorly deined and variably present dark
saddles, irst beneath dorsal in, second at vertical through
middle of anal in and third at end of caudal peduncle
(Fig. 5). Light morph have similar pigmentation pattern
but with light brown dorsal surface contrasting with
clearly deined dark brown saddles; lateral portion of body
with irregularly mottled dark blotches (Fig. 6a). Ventral
surface of body overall light brown with dark pigment
concentrated in unculiferous tubercles. Pectoral, ventral
and anal ins with scattered dark pigment. Dorsal and
caudal ins overall dark with light distal portions. Caudal
in sometimes bearing clear patch at proximal portion.
Distribution. Known from several tributaries of the upper
and middle rio São Francisco basins including the das
Velhas, Formoso, Paraopeba and Paracatu rivers in Minas
Gerais State, Brazil (Fig. 4).
Etymology. The speciic epithet, minerim, refers to the
tipically regional manner of pronouncing the Portuguese
word “mineirinho”, diminutive of “mineiro”, which refers
to a person that comes from the State of Minas Gerais. The
name is an allusion to the region where it is found and also
to its relative small size in comparison with other species
of Bunocephalus. A noun in apposition.
T. P. Carvalho, A. R. Cardoso, J. P. Friel & R. E. Reis
Conservation status. Bunocephalus minerim is known
from an Extent of Occurrence (EOO) of approximately
75,000 km2, and despite some areas within its range
suffer continuing decline in habitat quality because
of contamination from the city of Belo Horizonte and
also mining and agriculture, there is no evidence of its
habitat being severely fragmented or occurring extreme
luctuations in range or number of individuals. Considering
that no speciic threats to the species were detected, B.
minerim is categorized as Least Concern (LC) according to
the International Union for Conservation of Nature (IUCN)
categories and criteria (IUCN Standards and Petitions
subcommittee, 2014).
Discussion
The new species described herein in Bunocephalus
share three apomorphic features proposed by Friel (1994) to
a clade composed by Amaralia and Bunocephalus: middle
nuchal plate ornamentation well developed; posterior
margin of basipterygium jagged and lateral line ossicles
with small hooks, this last feature being variable in B.
minerim. The new species do not share most of the seven
apomorphic features of Amaralia (Friel, 1994), except for
the presence of four branchiostegal rays and absence of
serration on anterior portion of pectoral-in spine in B.
hartti. With the limited information at hand, we prefer to
include these new species in Bunocephalus since reviewing
the composition and the monophyly of these two genera are
beyond the scope of the present paper.
The two new species of Bunocephalus can be diagnosed
among other species of the genus by apomorphic characters,
which can be related to morphological changes during
ontogeny. Bunocephalus hartti is unique within the species
of the genus by the absence of serrations on the anterior
margin of the pectoral-in spine in adults (Fig. 2a). Feeble
serrations are observed in the pectoral-in spines of juveniles
of Bunocephalus hartti (Fig. 2b), and during ontogeny these
serrations seem to be absorbed by the growth of the anterior
portion of the spine. Within Aspredinidae, serrations
on the anterior margin of the spine are absent in most
hoplomyzontines, Xyliphius Eigenmann, 1912, Amaralia
and seem to have been lost multiple times within the family
(Friel, 1994).
Most species of Bunocephalus possess the plesiomorphic
condition within aspredinids of having an epiphyseal bar
between the paired frontals. The only exceptions are B.
larai, endemic from the upper rio Paraná basin, and the new
species herein described B. minerim. The presence of the
epiphyseal bar can be related to the degree of ontogenetical
development of the specimens: a developmental series of B.
verrucosus (INPA 4395) contains a juvenile specimen (38.4
mm SL) with incomplete medial contact between the frontals,
whereas the bar is completely formed in all individuals above
68 mm SL. According to Friel (1994, 2008) the absence of
an epiphyseal bar is also observed in some members of the
509
genus Pseudobunocephalus and at least in Xyliphius lepturus
Orcés, 1962. Therefore the loss of the epiphyseal bar in
adults seems to have evolved independently at least three
times within aspredinids, perhaps as retention of a juvenile
feature. Within Bunocephalus the absence of the epiphyseal
bar may suggest a close relationship between B. larai and B.
minerim.
An interesting feature observed in Bunocephalus minerim
is the presence of eggs directly attached to the skin surface of
some females (e.g., MCP 45242, MCP 45156, MZUSP 73800
and UFRGS 11273). Such parental care in the form of physical
attachment to developing embryos has previously been
reported in some aspredinids such as Pterobunocephalus,
Platystacus, Aspredo, and Aspredinichthys (Friel, 2003). In
Pterobunocephalus, like in B. minerim, the eggs are directly
attached to the body, whereas in Platystacus, Aspredo, and
Aspredinichthys eggs are attached to leshy stalks, called
cotylephores (Wetzel et al., 1997). Within Bunocephalus this
feature is rarely observed. From an extensive examination of
museum specimens of Bunocephalus (Friel, 1994; Cardoso;
2008; lots listed herein) only three other specimens of a
Bunocephalus cf. coracoideus from French Guiana were
observed carrying adhesive eggs (CUMV 81970). However,
the eggs in these specimens are more supericially attached
to body in comparison with B. minerim, in which depressions
on the skin surface are observed (Fig. 6b).
Comparative Material Examined. Amaralia hypsiura: Brazil:
INPA 32338, 3 (1 c&s), 70-79 mm SL, Porto Trombetas, Pará.
Amaralia sp.: Paraguay: UMMZ 207818, 2 (1 c&s), 84-118 mm
SL, Río Aquidaban at Paso Horqueta, Concepción. Bunocephalus
aloikae: French Guiana: ZMA 102.229, 62.5 mm SL (holotype of
Bunocephalus amaurus aloikae Hoedeman, 1961), Rivière Litany
near Aloiké village. Bunocephalus aleuropsis: Brazil: MCP
34142, 2, 36.3-37.4 mm SL, rio São João, Ribeirão Cascalheira,
Mato Grosso. MCP 35744, 1, 46.3 mm SL, Bujari, rio Riozinho
do Andirá, Bujari, Acre. UNT 2038, 1, 45.3 mm SL, Ipueiras,
rio Tocantins, Tocantins. UNT 2039, 1 c&s, 66.8 mm SL, rio
Tocantins, Tocantins, Brazil. Bunocephalus amaurus: Guyana:
FMNH 53121, 55.8 mm SL (holotype of Bunocephalus amaurus
Eigenmann, 1912), Konawaruk. Suriname: ZMA 102.228, 70.6
mm SL (holotype of Bunocephalus amaurus sipaliwini Hoedeman,
1961), bordering Paru Savannah, Sipaliwini. Bunocephalus
colombianus: Colombia: FMNH 56038, 71.2 mm SL (holotype
of Bunocephalus colombianus Eigenmann, 1912), Raspadura,
Choco. FMNH 56668, 1, 112.7 mm SL (paratype of Bunocephalus
colombianus Eigenmann, 1912), Raspadura, Choco. CAS 35249, 1,
84.7 mm SL (paratype of Bunocephalus colombianus Eigenmann,
1912), río Quito into río Atrato, Quibdó, Choco. Bunocephalus
chaimazelus: Guyana: FMNH 53122, 26.7 mm SL (holotype of
Bunocephalus chamaizelus Eigenmann, 1912), Erukin. FMNH
53123, 1 of 2 35.2 mm SL (paratype of Bunocephalus chamaizelus
Eigenmann, 1912), Erukin. FMNH 53125, 2, 22.3-28.1 mm SL
(paratypes of Bunocephalus chamaizelus Eigenmann, 1912), Gluck
Island. FMNH 7370, 1, 23.3 mm SL (paratype of Bunocephalus
chamaizelus Eigenmann, 1912) Tumatumari, lower Potaro River.
510
Two new species of Bunocephalus
ROM 91314, 5 (1c&s), 17.5-30.3 mm SL, Imbaima Creek tributary
to Kuribrong River, Potaro-Siparuni. ROM 91386, 4 (1c&s), 18.829 mm SL, Creek tributary to upper Kuribrong River, PotaroSiparuni. Bunocephalus coracoideus: Peru: ANSP 138984, 2 (1
c&s), 49.2-80.0 mm SL, Río Nanay at vinicity of Iquitos, Loreto.
French Guiana: CUMV 81970, 4, 81.6-84.5 mm SL, Rivière
Sinnamary. Brazil: MNHG 255032, 6 of 10, 70.6-84.0 mm SL,
igarapé Cuxiú, near Ourém, Pará. MHNG 2551.015, 1, 82.9 mm
SL, rio Tocantins, ca. 3 km of São Félix, Tocantins. Bunocephalus
doriae: Brazil. MCP 13176, 7 (1 c&s), 36.0-50.9 mm SL, rio
Uruguai, São Nicolau. MCP 14267, 6 (1 c&s) 61.5-73.2 mm SL, rio
Negro on road between Bagé and Aceguá, Bagé, Rio Grande do
Sul, Bunocephalus erondinae: Brazil: MCP 40877, 82.9 mm SL
(holotype of Bunocephalus erondinae Cardoso, 2010), canal São
Gonçalo, Pelotas, Rio Grande do Sul. MCP 40878, 17 (1 c&s), 56.676.9 mm SL (paratypes of Bunocephalus erondinae Cardoso, 2010).
MCP 43519, 2 (1 c&s), 33.5-61.1 mm SL, rio das Antas at mouth,
Cotiporã, Rio Grande do Sul. Bunocephalus knerii: Ecuador:
FMNH 99481, 1, 50.7 mm SL, stream tributary to Río Payamino,
Capihuara, Napo. NMW 10976, 1 50.7 mm SL (syntype of
Bunocephalus kneri Steindachner, 1882), Canelos. Bunocephalus
larai: Brazil: MCP 28376, 2, 48.6-53.9 mm SL, rio Paranaíba,
córrego tributary of the rio Paranaíba, Rio Paranaíba, Minas
Gerais. MCP 28377, 3 (1 c&s), 51.1-55.6 mm SL, ribeirão de Fora on
road from Rio Paranaíba to Serra do Salitre, Rio Paranaíba, Minas
Gerais. MZUSP 22614, 2, 36.0-37.9 mm SL, rio Paraná in front of
Jupiá, São Paulo. MZUSP 23092, 7, 26.0-31.3 mm SL, rio Paraná, at
Ilha Solteira, São Paulo. Bunocephalus verrucosus: Brazil: MCP
35743, 5 (2 c&s) 55.8-82.7 mm SL, igarapé Maninguari on highway
BR-364, Bujari, Amazonas. MCP 29811, 1, 98.4 mm SL, lago Tefé
near Tefé, Amazonas. INPA 4395, 5 (2 c&s), 27.3-38.8 mm SL, rio
Uraricoera, Roraima. Guyana: MHNG 2281.52, 1, 92.7 mm SL,
afluent du Rupununi, Rupununi, Annai. Peru: MHNG 2395.58,
2, 27.1-29.7 mm SL, tributary of Río Ucayali (Pachitea), Ucayali.
Pseudobunocephalus amazonicus: Bolivia: ZMA 109.246, 37 mm
SL (holotype of Dysichthys amazonicus Mees, 1989), creek near
Todos los Santos, Cochabamba. Brazil: MCP 35751, 6 of 15 (2 c&s)
23.7-29.3 mm SL, rio Ribeirão in the highway BR-425 ca. 62 km S of
highway BR-364, Nova Mamoré, Rondônia. Pseudobunocephalus
biidus: Peru: CAS 35106, 4, 32.6-42.4 mm SL (paratypes of
Bunocephalus biidus Eigenmann, 1942), Yurimaguas creek,
Loreto. Brazil: MCP 32772, 1 c&s, 31.2 mm SL, rio Iquirí,
tributary to rio Ituxi, Amazonas. Pseudobunocephalus iheringii:
Argentina MCP 13377, 7 (1 c&s) 23.3-48.0 mm SL, Arroio
Chimiray, 5 km of Azara, tributary to Río Uruguay, Missiones.
Pseudobunocephalus lundbergi: Venezuela: ANSP 168817,
28.4 mm SL, (holotype of Pseudobunocephalus lundbergi Friel,
2008), caño Barranca, ca. 1.25 hours downstream from Jubillal
(opposite bank) on Río Caura, Bolivar. Pseudobunocephalus
quadriradiatus: Peru: MHNG 21570.21, 31.4 mm SL, (holotype
of Dysichthys quadriradiatus Mees, 1989), Chinguito [= Cocha
Shinguita or Shirguita], Loreto. MHNG 2430018, 4, 27.1-32.1 mm
SL, (paratypes of Dysichthys quadriradiatus Mees, 1989), NRM
15142, 28 (2 c&s) of 66, 19.0-21.7 mm SL, Río Samiria drainage
left bank of stream halfway between Hamburgo and Santa Elena,
Peru. Pseudobunocephalus rugosus: Brazil: MCP 15540, 3 (2
c&s), 19.8-23.9 mm SL, rio Paraguai at Cáceres and outskirts, Mato
Grosso. Pterobunocephalus depressus: Ecuador: ANSP 130606,
1 c&s, Río Conejo at Santa Cecilia, Napo. Pterobunocephalus
dolichurus: Brazil: MCP 35745, 1 c&s, 73.3 mm SL, rio Caeté
at highway BR-364, Acre. Xyliphius lepturus: Venezuela: MCNG
5547, 1 c&s, 107.8 mm SL, Río Bocono, Portuguesa/Barinas.
Acknowledgements
We are grateful to the following people for loan of
specimens and hospitality while visiting museums: M.
Sabaj Pérez, K. Luckenbill, J. Lundberg (ANSP), D. Catania
(CAS), M. Rogers (FMNH), L. Rapp Py-Daniel (INPA);
C. Oliveira and F. Roxo (LBP), Donald Taphorn and Otto
Castillo (MCNG), C. Lucena and M. Lucena (MCP), S.
Fisch-Müller and R. Covain (MHNG), P. Buckup and M.
Britto (MNRJ), M. de Pinna, A. Datovo and O. Oyakawa
(MZUSP), S. Kullander (NRM), H. Wellendorf (NMW), H.
López-Fernández (ROM), P. Lucinda (UNT). Thanks to C.
Leal and D. Carvalho for information on DNA barcoded
specimens of Bunocephalus minerim. Thanks also to C.
B. M. Alves for information regarding these new species.
Thanks to C. Lucena for helping with photos of the pectoralin spines. We also thank John Lundberg for discussions on
catish morphology. This research was partially supported
by the Conselho Nacional de Desenvolvimento Cientíico e
Tecnológico (CNPq) (process number 229355/2013-7 to TPC,
and processes number 305180/2010-0 and 207038/2013-9 to
RER), and Fundação de Amparo à Pesquisa do Estado do
Rio Grande do Sul - FAPERGS (process number 11/0936-5
to RER). The authors are grateful to the All Catish Species
Inventory (NSF DEB-0315963).
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Submitted October 13, 2014
Accepted June 03, 2015 by Francisco Langeani
Published September 25, 2015
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