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Two new species of Corydoras are described from the rio Madeira basin, Brazil. The intermediate long-snouted new species can be distinguished from its congeners by presenting the following combination of features: posterior margin of dorsal-fin spine with laminar serrations directed towards the origin of the spine; presence of two longitudinal black stripes on flanks; anterior portion of dorsal fin with sparse black chromatophores, not forming any conspicuous pattern; absence of a conspicuous black marbled coloration pattern on head; black spots on caudal fin, some spots arranged, forming transversal bars; and brownish dorsal-fin spine. The short-snouted new species can be distinguished from its congeners by the following combination of features: short mesethmoid; posterior laminar expansion of infraorbital 2 very reduced, not in contact with compound pterotic; two or three longitudinal black stripes on flanks; absence of an oblique or vertical black blotch across the eye; anterior portion of dorsal fin with sparse black chromatophores, not forming any conspicuous pattern; and ventral surface of trunk naked or covered by sparse platelets.
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Neotropical Ichthyology, 14(1): e150063, 2016 Journal homepage: www.scielo.br/ni
DOI: 10.1590/1982-0224-20150063 Published online: 30 March 2016 (ISSN 1982-0224)
Two new species of Corydoras Lacépède, 1803 (Siluriformes:
Callichthyidae) from the rio Madeira basin, Brazil
Luiz Fernando Caserta Tencatt1 and Willian Massaharu Ohara2
Two new species of Corydoras are described from the rio Madeira basin, Brazil. The intermediate long-snouted new species
can be distinguished from its congeners by presenting the following combination of features: posterior margin of dorsal-
n spine with laminar serrations directed towards the origin of the spine; presence of two longitudinal black stripes on
anks; anterior portion of dorsal n with sparse black chromatophores, not forming any conspicuous pattern; absence of a
conspicuous black marbled coloration pattern on head; black spots on caudal n, some spots arranged, forming transversal
bars; and brownish dorsal-n spine. The short-snouted new species can be distinguished from its congeners by the following
combination of features: short mesethmoid; posterior laminar expansion of infraorbital 2 very reduced, not in contact with
compound pterotic; two or three longitudinal black stripes on anks; absence of an oblique or vertical black blotch across
the eye; anterior portion of dorsal n with sparse black chromatophores, not forming any conspicuous pattern; and ventral
surface of trunk naked or covered by sparse platelets.
Duas espécies novas de Corydoras são descritas da bacia do rio Madeira, Brasil. A espécie nova de focinho longo
intermediário pode ser distinguida de suas congêneres por apresentar a seguinte combinação de características: margem
posterior do espinho da nadadeira dorsal com serrilhas laminares direcionadas para origem do espinho; presença de duas
faixas pretas longitudinais nos ancos; porção anterior da nadadeira dorsal com cromatóforos pretos esparsos, sem formar
nenhum padrão conspícuo; ausência de um padrão de coloração marmoreado de preto conspícuo na cabeça; manchas pretas
na nadadeira caudal, algumas delas alinhadas, formando barras transversais; e espinho dorsal amarronzado. A espécie
nova de focinho curto pode ser distinguida de suas congêneres por apresentar a seguinte combinação de características:
mesetmóide curto; expansão laminar posterior do infraorbital 2 muito reduzida, sem contato com o pterótico composto;
duas ou três faixas pretas longitudinais nos ancos; ausência de uma mancha preta oblíqua ou vertical através do olho;
porção anterior da nadadeira dorsal com cromatóforos pretos esparsos, sem formar nenhum padrão conspícuo; e superfície
ventral do tronco nua ou coberta por plaquetas esparsas.
Keywords: Amazon, Corydoradinae, Mato Grosso, rio Aripuanã, Taxonomy.
1Universidade Estadual de Maringá, Programa de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Av. Colombo,
5790, 87020-900 Maringá, PR, Brazil. luiztencatt@hotmail.com (corresponding author)
2Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494, 04299-970 São Paulo, SP, Brazil. willianmohara@gmail.com
Introduction
Corydoras Lacépède, 1803 is the most species-rich
genus of Siluriformes, comprising approximately 170
valid species (Reis, 2003; Eschmeyer, 2015). The genus is
widely distributed in cis-andean South America and the
largest diversity is found in the Amazon basin, where more
than the half of the known species occurs (see Eschmeyer,
2015). The species inhabit many different environments,
from lakes and streams to large rivers. Typically found in
shallow depths or close to the margins, Corydoras are often
associated with sandy or muddy substrates.
Efforts to understand the taxonomy and systematics
of Corydoras have included Ellis (1913), Gosline (1940),
Nijssen (1970), Nijssen & Isbrücker (1980), Britto (2003)
and Alexandrou et al. (2011). Multiple studies have noted
apparent cases of convergence of color pattern in Corydoras.
Nijssen & Isbrücker (1980) were the rst to implicitly
recognize a convergent color pattern by placing C. arcuatus
Elwin, 1939 and C. narcissus Nijssen & Isbrücker, 198 0, two
species with almost equal color pattern, into two different
morphological groups, and noting that C. narcissus is more
similar to C. acutus Cope, 1872 than to C. arcuatus. Britto
et al. (2009) later described a third species with very similar
color pattern to C. arcuatus and C. narcissus and showed
that all three species belong to distinct clades. Based on
phylogenetic analyses, Britto (2003) noted another case of
apparent color convergence involving Corydoras nattereri
Steindachner, 1876 in the Corydoras clade and C. prionotos
Nijssen & Isbrücker, 1980 in the Scleromystax clade.
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
140
Alexandrou et al. (2011) study of community structure in
Corydoradinae was the rst paper to explicitly hypothesize
an adaptive basis for the convergence of color patterns in
Corydoras. The study was based on a molecular phylogenetic
analysis that established nine well-dened clades – each
with characteristic snout morphology. The authors noted
multiple cases of apparent convergence of color pattern
withi n syntopic representatives of these clades, and provided
evidence to support a hypothesis of Müllerian co-mimicry.
With recent survey efforts undertaken by the project
“Monitoramento e Conservação da Ictiofauna do rio
Madeira” (Assessment and conservation of the ichthyofauna
of the rio Madeira) from 2009 to 2013. Britto (2013)
documented 14 species of Corydoras from the Brazilian
territory of the rio Madeira basin (but mentioned the
occurrence of as many as twenty species). Here we report a
new case of convergent color pattern in Corydoras. During a
recent eld expedition in the rio Aripuanã and its tributaries
in the rio Madeira basin, two new species with similar
color pattern but showing divergent snout morphology
were found. In this paper, we describe these two sympatric
and syntopic species and demonstrate that although they
have similar color pattern, they belong to clearly distinct
morphological groups that correspond to different lineages
sensu Alexandrou et al. (2011).
Material and Methods
Measurements were obtained through digital calipers to
the nearest tenth of millimeter. Morphometric and meristic
data were taken following Reis (1997), with modications
of Tencatt et al. (2013). Morphometrics are reported as
percentages of standard length (SL) and head length (HL).
Homology and terminology of barbels follows Britto &
Lima (2003). For our osteological analysis, some specimens
were cleared and stained (c&s) following the protocol
of Taylor & Van Dyke (1985). Osteological terminology
was based on Reis (1998), except for the use of parieto-
supraoccipital instead of supraoccipital (Arratia & Gayet,
1995), compound pterotic instead of pterotic-supracleithrum
(Aquino & Schaefer, 2002), and scapulocoracoid instead of
coracoid (Lundberg, 1970). Nomenclature of latero-sensory
canals and preopercular pores are according to Schaefer
& Aquino (2000) and Schaefer (1988), respectively. The
supra-preopercle sensu Huysentruyt & Adriaens (2005)
will be treated here as a part of the hyomandibula according
to Vera-Alcaraz (2013). Vertebral counts include only
free centra, with the compound caudal centra (preural 1+
ural 1) counted as a single element. Terminology of snout
external morphology follows Alexandrou et al. (2011). In the
description and diagnosis, the dark longitudinal stripes were
counted only when well dened, continuous and not fused to
the blackened dorsal portion of the sh’s body.
Comparative data of the following species were obtained
through their original descriptions and/or high resolution
photographs of type-specimens hosted in the Natural History
Museum, London: Corydoras acrensis Nijssen, 1972, C.
axelrodi Rössel, 1962, C. baderi Geisler, 1969, C. boesemani
Nijssen & Isbrücker, 1967, C. evelynae Rössel, 1963, C.
gomezi Castro, 1986, C. habrosus Weitzman, 1960, C.
haraldschultzi Knaack, 1962, C. isbrueckeri Knaack, 2004,
C. leopardus Myers, 1933, C. loxozonus Nijssen & Isbrücker,
1983, C. noelkempf Knaack, 2004, C. ornatus Nijssen &
Isbrücker, 1976, C. orphnopterus Weitzman & Nijssen,
1970, C. pulcher Isbrücker & Nijssen, 1973, C. robustus
Nijssen & Isbrücker, 1980, C. schwartzi Rössel, 1963, C.
sipaliwini Hoedeman, 1965, C. spectabilis Knaack, 2000, C.
surinamensis Nijssen, 1970 and C. urucu Britto, Wosiacki &
Montag, 2009. Photographs of other pertinent type specimens
were available for examination via the All Catsh Species
Inventory website (Morris et al., 2006).
In the description, numbers between brackets represent the
total number of specimens with those counts. Numbers with
an asterisk refer to the counts of the holotype. Institutional
abbreviations are: AI, Asociación Ictiológica de La Plata,
La Plata; ANSP, Academy of Natural Sciences of Drexel
University, Philadelphia; BMNH, The Natural History
Museum, London; INPA, Instituto Nacional de Pesquisas
da Amazônia, Manaus; LBP, Laboratório de Biologia de
Peixes da Universidade Estadual Paulista, Botucatu; MCP,
Museu de Ciências e Tecnologia da Pontifícia Universidade
Católica, Porto Alegre; MCZ, Museum of Comparative
Zoology, Harvard University, Cambridge; MNRJ, Museu
Nacional, Universidade Federal do Rio de Janeiro, Rio de
Janeiro; MZUSP, Museu de Zoologia da Universidade de
São Paulo, São Paulo; NRM, Naturhistoriska Riksmuseet,
Stockholm; NUP, Coleção Ictiológica do Núcleo de Pesquisas
em Limnologia, Ictiologia e Aquicultura da Universidade
Estadual de Maringá, Maringá; ROM, Royal Ontario Museum,
Toronto; ZUFMS-PIS, Coleção Zoológica de Referência da
Universidade Federal de Mato Grosso do Sul, Campo Grande.
Results
Corydoras brittoi, new species
urn:lsid:zoobank.org:act:3F9B3430-8D4C-4180-9B82-
1E70648E7BD1
(Figs. 1-2a,b, 3a,c, 4, Table 1)
Holotype. MNRJ 43316, 38.1 mm SL, Brazil, Mato Grosso
State, Colniza Municipality, Guariba District, tributary to
the rio Guariba, rio Aripuanã drainage, rio Madeira basin,
09°06’47.4”S 60°25’14.1”W, 15 Jul 2013, W. M. Ohara, D.
B. Hungria & B. Barros.
Paratypes. All from Brazil, Mato Grosso State, Colniza
Municipality, Guariba District, rio Aripuanã drainage,
rio Madeira basin. NUP 17311, 2, 32.4-35.2 mm SL, rio
Aripuanã, 09°21’00”S 59°19’33”W, 16 Jul 2013, W. M.
Ohara, D. B. Hungria & B. Barros; ZUFMS-PIS 4063, 1,
L. F. C. Tencatt & W. M. Ohara
Neotropical Ichthyology, 14(1): e150063, 2016
141
38.1 mm SL, Igarapé Pica-Pau, a tributary to the rio Juma,
09°22’27.2”S 60°02’59.9”W, 16 Jul 2013, W. M. Ohara, D.
B. Hungria & B. Barros. INPA 48032, 4, 34.7-36.7 mm SL;
MCP 48747, 3, 34.5-39.0 mm SL; MNRJ 43573, 4, 32.7-36.8
mm SL; MZUSP 117334, 5, 35.4-37.5 mm SL; NUP 17312,
4, 34.8-37.4 mm SL; NUP 17313, 2 c&s, 31.4-34.1 mm SL;
same data as the holotype.
Diagnosis. Corydoras brittoi can be distinguished from all
of its congeners, with exception of the species from ‘lineage
8’ sensu Alexandrou et al. (2011), by the presence of
posterior margin of dorsal-n spine with laminar serrations
directed towards the origin of the spine (vs. serrations, when
present, conical; directed towards dorsal-n spine tip in the
members of the remaining lineages). Corydoras brittoi can
be distinguished from other members of ‘lineage 8’, with
exception of C. bifasciatus Nijssen, 1972, C. pinheiroi
Dinkelmeyer, 1995 and C. pulcher, by the presence of two
longitudinal black stripes on anks (vs. presence of a single
arched black stripe on dorsal portion of ank in C. arcuatus;
stripe, when well dened, located on midline of ank in
C. gomezi, C. incolicana Burgess, 1993, C. leopardus, C.
orphnopterus, C. robineae Bu rgess , 1983 and C. robustus;
three to four slender longitudinal black stripes on anks
in C. ornatus; presence of three to four longitudinal rows
of black spots on anks, which may be coalescent and
form stripes in some specimens of C. haraldschultzi, C.
isbrueckeri, C. noelkempf and C. spectabilis; absence of
stripes on anks in remaining species). Corydoras brittoi
can be distinguished from C. bifasciatus and C. pulcher by
presenting anterior portion of dorsal n with sparse black
chromatophores, not forming any conspicuous pattern
(vs. conspicuously blackened in C. bifasciatus; hyaline
with conspicuous whitish yellow pigmentation in C.
pulcher); from C. pinheiroi by the absence of a conspicuous
black marbled coloration pattern on head (vs. presence).
Addit i onal ly, Corydoras brittoi can also be distinguished
from C. bifasciatus by the presence of black spots on caudal
n, some spots arranged, forming transversal bars (vs.
spots absent, covered by brown chromatophores); from C.
pulcher by the presence of brownish dorsal-n spine (vs.
conspicuously whitish yellow).
Fig. 1. Corydoras brittoi, holotype, MNRJ 43316, 38.1 mm SL, Brazil, Mato Grosso State, Colniza Municipality, Guariba
District, tributary to the rio Guariba, rio Aripuanã drainage, rio Madeira basin, 09°06’47.4”S 60°25’14.1”W. Dorsal (top),
lateral (middle) and ventral (bottom) views. Photo by Celso Ikedo.
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
142
Description. Morphometric data presented in Table 1. Head
compressed with acutely convex dorsal prole; triangular in
dorsal view. Snout pointed and straight. Head prole nearly
straight from tip of snout to anterior nare; ascending nearly
straight from this point to tip of posterior process of parieto-
supraoccipital. Prole slightly convex along dorsal-n base.
Postdorsal-n body prole nearly straight to adipose-n
spine; concave from this point to caudal-n base. Ventral
prole of body slightly convex from isthmus to base of rst
anal-n ray; concave from this point to caudal-n base.
Body acutely elliptical in cross section at pectoral girdle,
gradually becoming more compressed toward caudal n.
Eye rounded, located dorso-laterally on head; orbit
delimited dorsally by lateral ethmoid, frontal and
sphenotic, ventrally by infraorbitals. Anterior and posterior
nares close to each other, only separated by ap of skin.
Anterior naris tubular. Posterior naris relatively distant to
anterodorsal margin of orbit, separated from it by distance
equal to twice of naris diameter. Mouth small, subterminal,
width nearly equal to bony orbit diameter. Maxillary barbel
relatively large, almost reaching anteroventral limit of gill
opening. Outer mental barbel slightly larger than maxillary
barbel. Inner mental barbel eshy, with base close to its
counterpart. Small rounded papillae covering entire surface
of all barbels, upper and lower lips, and isthmus.
Mesethmoid long; anterior tip well developed, larger
than 50% of bone length (see Britto, 2003: 123, character
1, state 0; g. 1A); posterior portion conspicuously narrow
and entirely covered by a thick layer of skin. Nasal slender,
curved laterally, with very reduced laminar expansion on
its inner margin; mesial border contacting only frontal.
Frontal elongated, narrow, with width slightly smaller than
half of entire length; anterior projection short, size smaller
than nasal length. Frontal fontanel large, oblong; posterior
tip extension slightly entering anterior margin of parieto-
supraoccipital. Parieto-supraoccipital wide, posterior
process long and contacting nuchal plate; region of contact
between posterior process and nuchal plate covered by
thick layer of skin.
Two laminar infraorbitals with minute odontodes;
infraorbital 1 large, ventral laminar expansion poorly
developed; anterior portion with well-developed expansion
(Fig. 2a); infraorbital 2 small, slender; with posterior
laminar expansion well developed; posteroventral margin
contacting posterodorsal ridge of hyomandibula, dorsal
tip contacting sphenotic and compound pterotic (Fig.
2b). Posterodorsal ridge of hyomandibula close to its
articulation with opercle oblong; exposed, conspicuously
slender; dorsal ridge of hyomandibula between compound
pterotic and opercle covered by thick layer of skin; exposed
Table 1. Morphometric data for Corydoras brittoi and Corydoras pavanelliae. N= number of measured specimens and SD
= standard deviation.
Corydoras brittoi Corydoras pavanelliae
NHolotype Low-High Mean±SD NHolotype Low-High Mean±SD
Standard length (mm) 20 38.1 31.4-39.0 36.1.5±1.8 20 45.1 23.3-45.1 26.6±4.8
Percents of standard length
Depth of body 20 38.6 34.7-40.5 38.7±1.2 20 43.2 35.8-43.2 38.7±1.6
Predorsal distance 20 51.2 49.5-53.3 51.1±0.8 20 54.5 48.6-54.5 52.2±1.6
Prepelvic distance 20 48.8 47.8-51.1 49.1±0.7 20 50.1 45.8-51.4 48.3±1.5
Preanal distance 20 83.2 80.3-85.1 82.1±1.2 20 85.8 78.1-85.8 81.3±1.8
Preadipose distance 20 84.3 83.8-86.5 84.8±0.9 20 88.2 83.5-88.7 85.6±1.5
Length of dorsal spine 19 23.6 23.6-27.2 25.3±1.2 19 30.8 26.7-32.4 30.3±1.4
Length of pectoral spine 20 24.9 24.2-27.2 25.9±0.9 20 31.7 28.7-33.8 31.1±1.3
Length of adipose-n spine 19 8.4 8.0-10.2 9.0±0.6 20 8.6 7.6-9.8 8.7±0.7
Depth of caudal peduncle 20 14.7 13.7-15.7 14.9±0.5 20 15.7 15.0-16.9 15.8±0.6
Length of dorsal-n base 20 19.4 18.6-21.6 20.2±0.8 20 19.3 16.7-19.5 18.1±0.8
Dorsal to adipose distance 20 16.8 14.9-19.1 16.7±1.3 20 18.2 18.0-20.5 19.3±0.8
Maximum cleithral width 20 23.1 22.9-25.2 24.0±0.6 20 29.3 26.5-29.3 27.7±0.8
Head length 20 44.4 42.6-49.0 45.0±1.4 20 47.7 44.2-48.6 46.6±1.3
Length of maxillary barbel 20 17.3 12.1-19.2 16.3±1.5 20 15.1 9.7-15.1 12.3±1.7
Percents of head length
Head depth 20 79.2 71.9-84.3 81.0±3.1 20 84.7 73.3-84.7 79.1±2.9
Least interorbital distance 20 29.2 28.1-32.3 31.0±1.2 20 33.5 29.3-34.0 31.4±1.3
Horizontal orbit diameter 20 22.8 20.9-24.3 22.8±0.8 20 21.4 21.4-27.4 24.9±1.3
Snout length 20 33.2 35.9-43.4 40.7±1.8 20 37.7 32.7-37.8 35.3±1.4
Least internarial distance 20 14.9 10.0-14.5 12.6±1.1 20 16.3 11.8-16.3 14.2±1.1
L. F. C. Tencatt & W. M. Ohara
Neotropical Ichthyology, 14(1): e150063, 2016
143
areas bearing small odontodes. Interopercle covered by
thin layer of skin, somewhat triangular, anterior projection
well-developed. Preopercle slender, elongated, minute
odontodes sparse on external surface. Opercle elongated
dorso-ventrally, width smaller than half of its length;
free margin convex, without serrations and covered by
small odontodes. Anteroventral portion of cleithrum and
posterolateral portion of scapulocoracoid exposed; minute
odontodes sparse on exposed areas. Vertebral count 22 (2);
ribs 7 (2), rst pair conspicuously large; complex vertebra
slender in shape. Neural and haemal spines with pointed
distal tips.
Fig. 2. Lateral view of the head of c&s specimens of Corydoras brittoi, NUP 17313, 34.1 mm SL, showing the poorly-
developed ventral expansion of the infraorbital 1 (a) and infraorbital 2 in contact with compound pterotic (b), and of
Corydoras pavanelliae, NUP 17315, 25.4 mm SL, showing the poorly-developed ventral expansion of the infraorbital 1
(c) and infraorbital 2 not in contact with compound pterotic (d). The dotted lines in (b) and (d) represent the suture between
sphenotic and compound pterotic bones. Abbreviations: io1: infraorbital 1, io2: infraorbital 2, sph: sphenotic, cpt: compound
pterotic. Scale bar = 1.0 mm.
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
144
Four branchiostegal rays decreasing in size posteriorly.
Hypobranchial 2 somewhat triangular, tip ossied and
directed towards anterior portion, posterior margin
cartilaginous; ossied portion well developed, about twice
size of cartilaginous portion. Five ceratobranchials with
expansions increasing posteriorly; ceratobranchial 1 with a
very reduced process on anterior margin of mesial portion;
ceratobranchial 3 notched on postero-lateral margin;
ceratobranchial 5 toothed on postero-dorsal surface, 34
to 36 (2) teeth aligned in one row. Four epibranchials with
similar size; epibranchial 2 slightly larger than others, with
small pointed process on laminar expansion of posterior
margin; epibranchial 3 with somewhat quadrangular
uncinate process on laminar expansion of posterior margin.
Two wide pharyngobranchials (3 and 4), pharyngobranchial
3 with a rippled laminar expansion on posterior margin.
Upper tooth plate oval; 34 to 40 (2) teeth aligned in two
rows on postero-ventral surface.
Lateral-line canal entering neurocranium through
compound pterotic, splitting into two branches before
entering sphenotic: pterotic branch with a single pore;
preoperculomandibular branch conspicuously reduced,
with a single pore opening close to postotic main canal.
Sensory canal continuing through compound pterotic,
entering sphenotic as temporal canal, which splits into
two branches: one branch giving rise to infraorbital canal,
other branch entering frontal through supraorbital canal,
both with single pore. Supraorbital canal not branched,
running through nasal bone. Epiphyseal pore opening at
supraorbital main canal; slightly directed toward frontal
fontanel region. Nasal canal with three openings, rst on
posterior edge, second on posterolateral portion and third
on anterior edge. Infraorbital canal running through entire
second infraorbital, extending to infraorbital 1 and opening
into two pores. Preoperculomandibular branch giving rise
to preoperculo-mandibular canal, which runs through
entire preopercle with three openings, leading to pores 3, 4,
and 5, respectively.
Dorsal n triangular, located just posterior to second
dorsolateral body plate. Dorsal-n rays II,8, posterior margin
of dorsal-n spine 11 to 14 moderately-developed serrations
directed towards dorsal-n spine origin; serrations absent
proximally (Fig. 3a). Nuchal plate moderately developed;
exposed, with minute odontodes; spinelet short; spine
moderately developed, adpressed distal tip reaching to
or slightly surpassing origin of last dorsal-n branched
ray; anterior margin with small odontodes. Pectoral n
triangular, its origin just posterior to gill opening. Pectoral-
n rays I,8 (17), I,9* (3); posterior margin of pectoral-n
spine with 17 to 19 moderately-developed laminar serrations
along its entire length; ser rations directed towards pectoral-
n spine origin (Fig. 3c). Pelvic n oblong, located just
below rst ventrolateral body plate, and at vertical through
rst branched dorsal-n ray. Pelvic-n rays i,5. Adipose
n roughly triangular, separated from base of last dorsal-
n ray by generally six dorsolateral body plates. Anal n
triangular, located just posterior to 13th ventrolateral body
plates, and at vertical through anterior margin of adipose-
n spine. Anal-n rays ii,6. Caudal-n rays i,12,i, generally
ve dorsal and ventral procurrent rays; bilobed; dorsal lobe
generally slightly larger than ventral lobe.
Three laterosensory canals on trunk; rst ossicle tubular,
second ossicle laminar, third lateral-line canal encased
in third dorsolateral body plate. Body plates with minute
odontodes scattered over exposed area, a conspicuous line
of odontodes conned on posterior margins; dorsolateral
body plates 22 (4), 23* (14), 24 (2); ventrolateral body plates
20* (12), 21 (8); dorsolateral body plates along dorsal-n
base 6* (18), 7 (2); dorsolateral body plates between adipose
and caudal ns 6 (7), 7* (12), 8 (1); preadipose platelets 2
(1), 3* (18), 4 (1); small platelets covering base of caudal-
n rays; small platelets disposed dorsally and ventrally
between junctions of lateral plates on posterior portion of
caudal peduncle. Anterior margin of orbit, above lateral
ethmoid, dorsal surface of snout, nasal capsule region, and
upper lip covered by platelets bearing odontodes. Ventral
surface of trunk covered by sparse irregular platelets
bearing odontodes.
Fig. 3. Dorsal- and pectoral-n spines of (a, c) Corydoras
brittoi, NUP 17313, 34.1 mm SL, showing serrations
directed towards the origins of the spines on posterior
margins of the (a) dorsal-n spine (8.6 mm long) and of
the (c) right pectoral-n spine (8.9 mm long), and of (b,
d) Corydoras pavanelliae, NUP 17315, 26.8 mm SL,
showing serrations directed towards the tips of the spines
on posterior margins of the (b) dorsal-n spine (7.9 mm
long) and of the (d) right spine (7.6 mm long).
L. F. C. Tencatt & W. M. Ohara
Neotropical Ichthyology, 14(1): e150063, 2016
145
Color in alcohol. Overall color of body in Fig. 1. Ground
color of body pale yellow, with top of head and snout
dark brown; interorbital region with more intense brown
pigmentation; nuchal plate and border of process of parieto-
supraoccipital contacting rst dorsolateral body plate light
yellow. Medial portion of opercle and cleithrum, posterior
portion of compound pterotic, preopercle, infraorbitals 1
and 2, upper lip, maxillary barbel and anterior portion of
outer mental barbel covered by brownish chromatophores.
First, second and third dorsolateral body plates almost
entirely blackened. Dorsal portion of body irregularly black
pigmented, forming diffuse longitudinal black stripe; black
stripe larger on dorsal-n base region becoming narrow
towards caudal peduncle. Body with two conspicuous
longitudinal black stripes. Midventral half of dorsolateral
body plates blackened, forming broad longitudinal stripe
along ank. Medial portion of ventrolateral body plates
anterior to anal-n last branched ray region blackened,
forming slender longitudinal black stripe along ank.
Dorsal n covered by sparse black chromatophores;
chromatophores slightly more concentrated on region of
rst and second dorsal-n rays, including membranes;
chromatophores generally absent close to dorsal-n base;
dorsal-n spine brownish. Pectoral-n rays with sparse
black chromatophores; pectoral-n spine brownish. Pelvic
n hyaline. Adipose n covered by black chromatophores
on its medial portion; black pigmentation more intense,
for ming an ir regular con s picuous elongated black blotch in
some specimens; adipose-n spine dorsal half blackened.
Caudal-n base blackened; caudal-n lobes with black
spots; some spots arranged in one to four transversal black
bars.
Color in life. Very similar to preserved specimens but with
ground color of body whitish yellow. Region of contact
between process of parieto-supraoccipital and nuchal plate
bright yellow. Body covered by greenish yellow iridescent
coloration, more concentrated on lower half of opercle,
infraorbitals, cleithr um and on region of two longitudinal
lateral stripes (Fig. 4).
Sexual dimorphism. Except for the presence of lanceolate
genital papilla in males, which occurs in all Corydoradinae
(see Nijssen & Isbrücker, 1980; Britto, 2003), no other
sexually dimorphic feature was observed.
Distribution. The new species is known from the rio
Aripuanã basin, Mato Grosso State (Fig. 5).
Ecological notes. The type locality of Corydoras brittoi
is located at 110 meters above sea level, and is a small
clear water stream, with 2-3 m width and 0.5-2 m depth,
with preserved riparian vegetation, swift water current,
and bottom composed mainly of sand and dead leaves.
Specimens of C. brittoi were observed at night during
capture at shallow portions of the stream in small groups
(5-15 individuals), and sometimes associated with a other
new species described below.
Et y mology. Corydoras brittoi is named in honor of
Marcelo Ribeiro de Britto, a dear friend and mentor, for his
extensive contributions to the taxonomy and systematics of
the Corydoradinae. A genitive.
Conservation status. Corydoras brittoi is so far known
only from the type-locality and its conservation status is
uncertain based on the limited knowledge of its geographic
distribution. However, considering that important threats to
the species were not detected in the area, and that it occurs in
a protected area (Reserva Extrativista Guariba Roosevelt),
Corydoras brittoi would be classied 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. Specimen of Corydoras brittoi photog raphed in life, showing the iridescent greenish yellow coloration all over the body.
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
146
Corydoras pavanelliae, new species
urn:lsid:zoobank.org:act:3DDD5672-0B60-49C4-8720-
E7DA1013CDE3
(Figs. 2c,d, 3b,d, 6, Table 1)
Holotype. MNRJ 43317, 45.1 mm SL, Brazil, Mato Grosso
State, Colniza Municipality, Guariba District, tributary to
the rio Guariba, rio Aripuanã drainage, rio Madeira basin,
09°06’47.4”S 60°25’14.1”W, 15 Jul 2013, W. M. Ohara, D.
B. Hungria & B. Barros.
Paratypes. INPA 48033, 5, 22.5-23.2 mm SL; MCP 48748,
5, 19.0-23.5 mm SL; MZUSP 117335, 10, 21.8-25.4 mm SL;
NUP 17314, 17, 20.8-30.7 mm SL; NUP 17315, 3 c&s, 24.7-
26.8 mm SL; ZUFMS-PIS 4064, 8, 20.0-22.9 mm SL; same
data as the holotype.
Diagnosis. Corydoras pavanelliae can be distinguished
from its congeners, with exception of species from lineages
4’, ‘5’, ‘6’, ‘7’ and ‘9’ sensu Alexandrou et al. (2011) by
the presence of a short mesethmoid, with anterior portion
smaller than 50% of the bone length (vs. long, equal or
larger than 50% of the bone length). Corydoras pavanelliae
can be distinguished from species of lineages ‘4’, ‘5’ and
‘7’ sensu Alexandrou et al. (2011) by the absence of contact
between infraorbital 2 and compound pterotic (Figs. 2c,d)
(vs. presence (see Tencatt & Pavanelli (2015: 291, g. 3))).
Corydoras pavanelliae can be distinguished from lineages
‘6’ and ‘9’ sensu Alexandrou et al. (2011), with the exce ption
of C. axelrodi, C. evelynae, C. loxozonus, C. parallelus
Burgess, 1993, C. schwartzi and C. surinamensis, by the
presence of two or three longitudinal black stripes on anks
(vs. a longitudinal black stripe along midline of ank in
C. acrensis, C. baderi, C. boesemani, C. bondi Gosline,
1940, C. coppenamensis Nijssen, 1970, C. habrosus, C. julii
Fig. 5. Map showing the type-locality (red circle) of Corydoras brittoi and Corydoras pavanelliae, a tributary to the rio
Guariba, Mato Grosso. The blue triangle represents the Igarapé Pica-Pau, also a tributary to the rio Guariba, and the yellow
triangle represents the rio Aripuanã, both representing additional records of C. brittoi.
L. F. C. Tencatt & W. M. Ohara
Neotropical Ichthyology, 14(1): e150063, 2016
147
Steindachner, 1906, C. nattereri, C. sipaliwini, C. trilineatus
Cope, 1872; a longitudinal series of large black blotches
along midline of ank in C. diphyes Axenrot & Kullander,
2003, C. ehrhardti Steindachner, 1910, C. aveolus Ihering,
1911, C. longipinnis Knaack, 2007, C. paleatus (Jenyns,
1842), C. reynoldsi Myers & Weitzman, 1960 and C.
tukano Britto & Lima, 2003; anks densely covered by
small rounded black spots, which can be diffuse or absent
in some specimens, in C. albolineatus Knaack, 2004 and
C. potaroensis Myers, 1927; a single oblique black stripe
from dorsal-n base region descending to base of caudal
peduncle in C. melini Lönnberg & Rendahl, 1930; four to
six longitudinal rows of black spots on anks, which may be
coalescent and form stripes in some specimens of C. sterbai
Knaack, 1962; presence of a single arched black stripe on
dorsal portion of ank in C. urucu; absence of stripes on
anks in remaining species). Corydoras pavanelliae can be
distinguished from C. axelrodi, C. evelynae, C. loxozonus,
C. parallelus, C. schwartzi and C. surinamensis by the
absence of an oblique or vertical black blotch across the eye
(vs. presence); from C. axelrodi, C. evelynae, C. loxozonus,
C. parallelus and C. surinamensis by presenting anterior
portion of dorsal n with sparse black chromatophores,
not forming any conspicuous pattern (vs. entirely or almost
entirely conspicuously blackened); from C. evelynae and
C. schwartzi by having ventral surface of trunk naked
or covered by sparse platelets (vs. densely covered by
coalescent platelets).
Fig. 6. Corydoras pavanelliae, holotype, MNRJ 43317, 45.1 mm SL, Brazil, Mato Grosso State, Colniza Municipality,
Guariba District, tributary to the rio Guariba, rio Aripuanã drainage, rio Madeira basin, 09°06’47.4”S 60°25’14.1”W. Dorsal
(top), lateral (middle) and ventral (bottom) views. Photo by Celso Ikedo.
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
148
Description. Morphometric data presented in Table 1.
Head compressed with convex dorsal prole; triangular in
dorsal view. Snout short and slightly pointed. Head prole
convex from tip of snout to anterior nares; ascending
nearly straight from this point to tip of posterior process of
parieto-supraoccipital; region just anterior to nares slightly
concave in some specimens. Prole slightly convex along
dorsal-n base. Postdorsal-n body prole nearly straight
to adipose-n spine; concave from this point to caudal-n
base. Ventral prole of body slightly convex from isthmus
to pelvic girdle. Prole nearly straight from pelvic girdle to
base of rst anal-n ray; abruptly concave from this point
to caudal-n base. Body roughly elliptical in cross section
at pectoral girdle, gradually becoming more compressed
toward caudal n.
Eye rounded, located dorso-laterally on head; orbit
delimited dorsally by lateral ethmoid, frontal and sphenotic,
ventrally by infraorbitals. Anterior and posterior nares
close to each other, only separated by ap of skin. Anterior
naris tubular. Posterior naris close to anterodorsal margin of
orbit, separated from it by distance equal to naris diameter.
Mouth small, subterminal, width nearly equal to bony orbit
diameter. Maxillary barbel moderate in size, not reaching
anteroventral limit of gill opening. Outer mental barbel
slightly larger than maxillary barbel. Inner mental barbel
eshy, base slightly separated to its counterpart; insertion
of barbel in middle of lower lip. Small rounded papillae
covering entire surface of all barbels, upper and lower lips,
and isthmus.
Mesethmoid short; anterior tip moderately developed,
smaller than 50% of bone length (see Britto, 2003: 123,
character 1, state 1; g. 1B); posterior portion relatively
wide and entirely covered by thick layer of skin. Nasal
slender, curved laterally, with inner margin laminar; mesial
border contacting only frontal. Frontal elongated, narrow,
with width slightly smaller than half of entire length;
anterior projection short, size smaller than nasal length.
Frontal fontanel large, oval; posterior tip extension slightly
entering anterior margin of parieto-supraoccipital. Parieto-
supraoccipital wide, posterior process long and contacting
nuchal plate; region of contact between posterior process
and nuchal plate covered by thick layer of skin.
Two laminar infraorbitals with minute odontodes;
infraorbital 1 large, ventral laminar expansion poorly
developed; moderately developed in larger specimens
(see Britto, 2003: 128, g. 5B); anterior portion with
well-developed expansion (Fig. 2c); infraorbital 2 small,
slender; with posterior laminar expansion very reduced;
posteroventral margin contacting posterodorsal ridge
of hyomandibula, dorsal tip contacting only sphenotic
(Fig. 2d). Posterodorsal ridge of hyomandibula close to
its articulation with opercle oblong; slender, exposed and
bearing small odontodes; dorsal ridge of hyomandibula
between compound pterotic and opercle covered by
thick layer of skin. Interopercle entirely covered by thick
layer of skin, somewhat triangular, anterior projection
well-developed. Preopercle slender, elongated, minute
odontodes sparse on external surface. Opercle elongated
dorso-ventrally, width smaller than half of its length; free
margin smoothly convex, without serrations and covered
by small odontodes. Anteroventral portion of cleithrum and
posterolateral portion of scapulocoracoid exposed; minute
odontodes sparse on exposed areas. Vertebral count 21 (3);
ribs 6 (3), rst pair conspicuously large; complex vertebra
compact in shape. Neural and haemal spines with pointed
distal tips.
Four branchiostegal rays decreasing in size posteriorly.
Hypobranchial 2 somewhat triangular, tip ossied and
directed towards anterior portion, posterior margin
cartilaginous; ossied portion well developed, about
twice size of cartilaginous portion. Five ceratobranchials
with expansions increasing posteriorly; ceratobranchial 1
with small process on anterior margin of mesial portion;
ceratobranchial 3 notched on postero-lateral margin;
ceratobranchial 5 toothed on postero-dorsal surface, 33
to 39 (3) teeth aligned in one row. Four epibranchials with
similar size; epibranchial 2 slightly larger than others, with
small pointed process on laminar expansion of posterior
margin; epibranchial 3 with curved mesially uncinate
process on laminar expansion of posterior margin. Two
wide pharyngobranchials (3 and 4), pharyngobranchial
3 with small triangular laminar expansion on posterior
margin. Upper tooth plate oval; 41 to 43 (3) teeth aligned in
two rows on postero-ventral surface.
Lateral-line canal entering neurocranium through
compound pterotic, splitting into two branches before
entering sphenotic: pterotic branch with a single pore;
preoperculomandibular branch conspicuously reduced,
with a single pore opening close to postotic main canal.
Sensory canal continuing through compound pterotic,
entering sphenotic as temporal canal, which splits into
two branches: one branch giving rise to infraorbital canal,
other branch entering frontal through supraorbital canal,
both with a single pore. Supraorbital canal not branched,
running through nasal bone. Epiphyseal pore opening at
supraorbital main canal. Nasal canal with three openings,
rst on posterior edge, second on posterolateral portion and
third on anterior edge. Infraorbital canal running through
entire second infraorbital, extending to infraorbital 1 and
opening into two pores. Preoperculomandibular branch
giving rise to preoperculo-mandibular canal, which runs
through entire preopercle with three openings, leading to
pores 3, 4, and 5, respectively.
Dorsal n triangular, located just posterior to second
dorsolateral body plate. Dorsal-n rays II,8, posterior margin
of dorsal-n spine with nine to 12 reduced serrations directed
towards tip of spine; serrations absent proximally (Fig. 3b).
Nuchal plate moderately developed; exposed, with minute
odontodes; spinelet short; spine relatively large, adpressed
distal tip surpassing last dorsal-n branched ray origin;
anterior margin with small odontodes. Pectoral n triangular,
its origin just posterior to gill opening. Pectoral-n rays I,8
L. F. C. Tencatt & W. M. Ohara
Neotropical Ichthyology, 14(1): e150063, 2016
149
(17), I,9* (3); posterior margin of pectoral-n spine with
17 to 22 small serrations along its entire length; serrations
directed towards pectoral-n spine tip (Fig. 3d). Pelvic n
oblong, located just below rst ventrolateral body plate, and
at vertical through rst branched dorsal-n ray. Pelvic-n
rays i,5. Adipose n roughly triangular, separated from base
of last dorsal-n ray by generally six dorsolateral body plates.
Anal n triangular, located just posterior to 12th ventrolateral
body plates, and at vertical through anterior margin of
adipose-n spine. Anal-n rays ii,6. Caudal-n rays i,12,i,
generally four dorsal and ventral procurrent rays; bilobed;
dorsal lobe generally slightly larger than ventral lobe.
Three laterosensory canals on trunk; rst ossicle
tubular, second ossicle laminar, third canal encased in third
dorsolateral body plate. Body plates with minute odontodes
scattered over exposed area, a conspicuous line of odontodes
conned on posterior margins; dorsolateral body plates 22
(4), 23* (14), 24 (2); ventrolateral body plates 20* (12), 21
(8); dorsolateral body plates along dorsal-n base 6* (18),
7 (2); dorsolateral body plates between adipose and caudal
ns 6 (7), 7* (12), 8 (1); preadipose platelets 2 (1), 3* (18),
4 (1); small platelets covering base of caudal-n rays; small
platelets disposed dorsally and ventrally between junctions
of lateral plates on posterior portion of caudal peduncle.
Anterior margin of orbit, above lateral ethmoid, covered
with small platelets bearing odontodes. Ventral surface of
trunk covered by sparse platelets; naked in some specimens.
Color in alcohol. Overall color of body in Fig. 6. Ground
color of body light yellow, with top of head and snout
dark brown; parieto-supraoccipital entirely dark brown.
Maxillary barbel and proximal region of outer mental
barbel covered by black chromatophores. Anterior portion
of body with irregular black blotches. Posterior margin of
rst and second dorsolateral body plates blackened. Dorsal
portion of body irregularly black pigmented, forming
diffuse slender longitudinal black stripe, more conspicuous
from dorsal-n base to adipose-n origin. Body with two
or three conspicuous longitudinal black stripes. Medial
portion of dorsal half of dorsolateral body plates with black
blotches anteriorly to adipose n; blotches aligned forming
short and narrow longitudinal stripe below dorsal-n base
in some specimens. Midventral half of dorsolateral body
plates blackened, forming broad longitudinal stripe along
ank. Dorsal half of medial portion of ventrolateral body
plates blackened, forming slender longitudinal black stripe
along ank; stripe absent or diffuse posteriorly to anal-
n last branched ray region. Dorsal n covered by sparse
brown chromatophores, more concentrated on its base;
upper half of rst and second dorsal-n rays, including
membranes, with more concentrated black chromatophores;
dorsal-n spine brownish. Pectoral-n rays with sparse
black chromatophores; pectoral-n spine brownish. Pelvic
n hyaline. Adipose n with posterior margin brownish or
blackened; adipose-n spine dorsal half blackened. Caudal
n with scattered brownish chromatophores; black spots
arranged in one to four transversal black bars; bars diffuse
or absent in some specimens; caudal-n base blackened;
region just posterior to caudal n base hyaline and with
light yellow pigmentation.
Sexual dimorphism. Except for the presence of lanceolate
genital papilla in males, which occurs in all Corydoradinae
(see Nijssen & Isbrücker, 1980; Britto, 2003), no other
sexually dimorphic feature was observed.
Distribution. The new species is known from its type-
locality, a tributary to the rio Aripuanã, Mato Grosso State
(Fig. 5).
Ecological notes. The only known specimens of Corydoras
pavanelliae were collected among C. brittoi specimens. For
notes on its ecology, see Ecological notes of C. brittoi.
Et y mology. Corydoras pavanelliae is named in honor of
Carla Simone Pavanelli, advisor of the rst author and dear
friend , for her extensive contribut ions to the knowledge of the
ecology and taxonomy of the Neotropical shes. A genitive.
Conservation status. The known specimens of Corydoras
pavanelliae are relatively numerous and were collected
among C. brittoi specimens. For this reason, we also suggest
that C. pavanelliae would be classied as Least Concern
(LC) according to the International Union for Conservation
of Nature (IUCN) categories and criteria (IUCN Standards
and Petitions Subcommittee, 2014).
Discussion
In general, Corydoradinae species with convergent color
pattern can be clearly distinguished from each other by
snout morphology (see Alexandrou et al., 2011: 2, g. 1).
Corydoras pavanelliae presents the general morphological
features common to lineages ‘6’ and ‘9’ sensu Alexandrou
et al. (2011), both characterized mainly by (I) the presence
of a short mesethmoid; (II) posterior margin of the pectoral-
n spine with serrations generally directed towards the tip
of the spine; (III) infraorbital 1 generally with poorly to
moderately developed ventral laminar expansion; and (IV)
infraorbital 2 generally not contacting compound pterotic.
Despite both lineages sharing some general external
morphology, Alexandrou et al. (2011) and Vera-Alcaraz
(2013) found evidence that these two clades do not form
a monophyletic group. However, most of Alexandrou et
al.’s (2011) phylogenetic inferences are based on molecular
characters, which are not yet available for C. pavanelliae.
Therefore, it is as yet not possible to clearly determine which
lineage C. pavanelliae belongs to.
On the other hand, Corydoras brittoi presents the
typical intermediate long snout patter n, which is present in
most species from the ‘lineage 8’ sensu Alexandrou et al.
(2011). The species from the ‘lineage 8’, with exception of
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
150
C. diuviatilis Britto & Castro, 2002 and C. garbei Ihering,
1911, can be distinguished from the other congeners by the
presence of dorsal-n spine with serrations directed towards
the origin of the spine. Other features common to species
of the lineage 8 are (I) the presence of long mesethmoid;
(II) laminar serrations on posterior margins of dorsal- and
pectoral-n spines; (III) infraorbital 1 generally with poorly
to moderately developed ventral laminar expansion; and
(IV) infraorbital 2 contacting compound pterotic. Despite
the fact that C. diuviatilis and C. garbei are recognized
as belonging to lineage 8 according to Alexandrou et al.
(2011), they display a morphological pattern much closer to
the species of ‘lineage 7’, which can be characterized by
(I) the presence of rounded snout; (II) serrations on dorsal-
and pectoral-n spines, when present, poorly developed
and directed towards the tip of the spines; (III) ventral
laminar expansion of infraorbital 1 very large; and (IV)
infraorbital 2 contacting compound pterotic. Therefore, the
phylogenetic positions of C. diuviatilis and C. garbei, may
require further investigation. Notwithstanding these two
exceptions, C. brittoi shares the same serration pattern of
the dorsal-n spine observed in all of the species from the
lineage 8 sensu Alexandrou et al. (2011).
Twe nt y-t w o Corydoras species have previously
been described from the rio Madeira drainage: C.
albolineatus Knaack, 2004, C. bilineatus Knaack, 2002, C.
caudimaculatus Rössel, 1961, C. cervinus Rössel, 1962, C.
cruziensis Knaack, 2002, C. geryi Nij s se n & Isb r ücke r, 19 83,
C. gossei Nijssen, 1972, C. gracilis Nijssen & Isbrücker,
1976, C. guapore Knaack, 1961, C. haraldschultzi, C.
isbrueckeri, C. latus Pearson, 1924, C. mamore Knaack,
2002, C. noelkempf, C. negro Knaack, 2004, C. paragua
Knaack, 2004, C. pinheiroi, C. pygmaeus Knaack, 1966,
C. sarareensis Dinkelmeyer, 1995, C. seussi, C. similis
Hieronimus, 1991, C. spectabilis, and C. sterbai.
Seven more Corydoras species were recorded from the
rio Madeira basin by Britto (2013): C. armatus (Günther,
1868), Corydoras cf. ambiacus Cope, 1872, Corydoras
aff. bondi, Corydoras aff. griseus Holly, 1940, Corydoras
aff. melanistius Regan, 1912, Corydoras cf. polystictus
Regan, 1912 and Corydoras cf. trilineatus. Also, with
the recent survey efforts, ten additional species were
recorded: Corydoras acutus, Corydoras aeneus (Gill,
1858), Corydoras cf. arcuatus, C. narcissus, C. ourastigma
Nijssen 1972, C. splendens (Castelnau, 1855) and four
morphotypes that represent possibly new species. A key
of the Corydoras species collected until 2011 in the rio
Madeira basin (Brazilian territory) and information about
them, are available in Britto (2013).
With the two new species described here, a total of
42 species are known from the rio Madeira basin. This
represents 24.7% of all Corydoras diversit y. Corydoras
brittoi and C. pavanelliae can be distinguished from all of
their congeners in the rio Madeira basin, with the exception
of C. bilineatus, C. pinheiroi and C. pygmaeus, by the
presence of two to three (up to three only in some specimens
of C. pavanelliae) longitudinal black stripes on anks (vs. a
single longitudinal black stripe in C. acutus, Corydoras aff.
bondi, Corydoras cf. trilineatus; three to four longitudinal
rows of black spots on anks, which may be coalescent
and form stripes in some specimens of C. haraldschultzi,
C. isbrueckeri, C. noelkempf, C. spectabilis, C. sterbai; a
single black stripe on dorsal portion of ank in C. arcuatus,
C. gracilis and C. narcissus; conspicuous stripes absent in
the remaining species). Both new species presented herein
can be distinguished from C. bilineatus and C. pinheiroi
by the absence of a black marbled coloration on anterior
portion of the body (vs. presence), and from C. pygmaeus by
the presence of contact between nuchal plate and posterior
process of the parieto-supraoccipital (vs. absence).
Comparative material examined. Corydoras acutus: Peru:
Unknow n depart ment: MNRJ 3985, 2, 47.1-54.8 mm SL, Sansho-
Caño. Corydoras adolfoi: Brazil: Amazonas: MZUSP 26641, 1,
32.5 mm SL, holotype of Corydoras adolfoi Bur ge ss , 1982,
tributary to the upper rio Negro. Corydoras ambiacus: Peru:
Loreto: MCP 26178, 1, 42.5 mm SL, rio Pacaya; MCP 26209, 10
of 19, 25.0-33.3 mm SL, Caño Yarina. Ucayali: MZUSP 26053, 2,
41.8-47.2 mm SL, Iamiriacocha. Corydoras approuaguensis:
French Guyana: Cayenne: MZUSP 27895-6, 2, 43.0-46.1 mm
SL, paratypes of Corydoras approuaguensis Nijs s e n & Is br ü c ker,
1983, rio Approuague. Corydoras araguaiaensis: Brazil: Mato
Grosso: MZUSP 87155, 4 of 33, 24.9-46.7 mm SL, 2 c&s, 27.6-
31.8 mm SL, Corixo da Saudade. Corydoras areio: Brazil: Mato
Grosso do Sul: ZUFMS-PIS 1314, 15, 34.4-41.9 mm SL, 2 c&s,
38.1-38.5 mm SL, Periquito stream. Corydoras armatus: Brazil:
Amazonas: MZUSP 49567, 1, 45.3 mm SL, rio Acre. Corydoras
aurofrenatus: Paraguay: Concepción: NRM 23529, 10 of 33,
31.4-45.7 mm SL, Arroyo Laguna Penayo where it crosses the
road Concepción-Paso Barreto. Corydoras bifasciatus: Brazil:
Pará: MZUSP 38976, 16, paratypes, 23.6-30.0 mm SL, creek at
left bank of the rio Cururu. Corydoras blochi: Brazil: Roraima:
MZUSP 8580, 3, 31.0-42.6 mm SL, paratypes of Corydoras
blochi Nijssen , 1971, Igarapé on Fazenda Canadá, tr ibutar y to the
rio Uraricoera. Corydoras bondi: Guyana: Barima-Waini: ROM
66202, 7 of 134, 33.8-39.9 mm SL, 3 c&s of 134, 36.7-38.6 mm
SL, Waikerebi Creek. Corydoras brevirostris: Ve n e z uela:
Bolívar: LBP 3080, 10, 23.8-27.7 mm SL, 3 c&s, 25.8-27.9 mm
SL, Río Orinoco. Corydoras britskii: Brazil: Mato Grosso do
Sul: ZUFMS-PIS 862, 12, 72.0-78.0 mm SL, marginal lagoon to
rio Vermelho. Corydoras carlae: Brazil: Paraná: NUP 711, 1,
47.9 mm SL, rio Tormenta; NUP 4425, 1 c&s, 45.0 mm SL, rio
Tormenta. Corydoras cervinus: Brazil: Mato Grosso: LBP
10097, 2, 38.9-39.7 mm SL, tributary to rio Guaporé. Corydoras
cochui: Brazil: Gos: MZ USP 89055, 6, 18.7-23.6 mm SL, rio do
Peixe II. Tocantins. MZUSP 35838, 4 of 6, 16.1-18.5 mm SL, rio
Javaés. Corydoras condiscipulus: French Guyana: Cayenne:
MZUSP 38957, 7, 34.1-40.3 mm SL, paratypes of Corydoras
condiscipulus Nijssen & Isbrücker, 1980, Cumuri Creek.
Corydoras coppenamensis: Suriname: Saramacca: MZUSP
13995-99, 5, 28.2-34.9 mm SL, paratypes of Corydoras bondi
coppenamensis Nijssen, 1970, rio Coppename. Corydoras
L. F. C. Tencatt & W. M. Ohara
Neotropical Ichthyology, 14(1): e150063, 2016
151
crimmeni: Brazil: Uncertain state: MZUSP 52490, 1, 36.1 mm
SL, holotype of Corydoras crimmeni Grant, 1998, aquarium
specimens said to be from near the town of Boa Vista, Roraima,
possibly from the rio Branco. Corydoras davidsandsi: Braz il:
Amazonas: MZUSP 110066, 4 of 40, 36.0-41.9 mm SL, 2 c&s of
40, 40.9-42.1 mm SL, rio Inambú. Corydoras diuviatilis:
Brazi l: São Paulo: MZUSP 75268, 1, 39.8 mm SL, holotype of
Corydoras diuviatilis Britto & Castro, 2002, Paulicéia stream.
Corydoras diphyes: Paraguay: Alto Paraná: ANSP 169756, 2,
40.7-43.1 mm SL, drainage ditches north of km 250 (2 km east of
Juan E. O’Leary on route 7). Corydoras ehrhardti: Brazil:
Paraná: NUP 11255, 15, 36.5-46.8 mm SL, rio São Pedro.
Corydoras elegans: Peru: Ucayali: MZUSP 26017, 6, 25.9-28.3
mm SL, Lobococha. Corydoras ephippifer: Brazil: Amapá:
MZUSP 31605, 2, 44.9-49.1 mm SL, rio Cupixi. Corydoras
eques: Brazil: Amazonas: MCZ 8204, 4 of 12, 37.6-44.4 mm SL,
paratypes of Corydoras eques Steindachner, 1876, rio Amazonas
at Codajás. Corydoras aveolus: Brazil: São Paulo: MZUSP
424, 1, 33.4 mm SL, holotype of Corydoras aveolus Ihering,
1911, tributaries to the rio Piracicaba. Corydoras fowleri: Peru:
Loreto: LBP 12462, 9, 44.3-59.9 mm SL, 1 c&s, 50.4 mm SL,
tributary to the rio Ampiyacu. Corydoras garbei: Brazil: Minas
Gerais: MNRJ 18089, 14, 19.2-25.3 mm SL, 2 c&s, 25.9-27.4 mm
SL, Perta-Pé lagoon. Corydoras geoffroy: Suriname: Marowijne:
MZUSP 38984, 2, 38.7-45.2 mm SL, paratypes of Corydoras
octocirrus Nijssen, 1970, fall in the rio Oelamari. Corydoras
gossei: Brazil: Rondônia: MZUSP 38977, 6, 48.4-53.4 mm SL,
paratypes of Corydoras gossei Nijssen, 1972, Igarapé do 13,
tributary to the rio Mamoré. Corydoras griseus: Guyana:
Potaro-Siparuni: MZUSP 108896, 4 of 13, 31.5-36.2 mm SL, 2
c&s of 13, 30.6-34.5 mm SL, Igarapé tributary to the rio
Kuribrong. Corydoras guapore: Brazil: Mato Grosso: ZUFMS-
PIS 4000, 5, 26.9-33.6 mm SL, 2 c&s, 28.8-29.2 mm SL, rio
Guaporé. Corydoras gryphus: Brazil: Paraná: MNRJ 40770, 1,
32.3 mm SL, holotype of Corydoras gryphus Tencatt, Britto &
Pavanelli, 2014, rio Paraná (near Ponte da Amizade). NUP 14676,
3 c&s, 27.7-32.4 mm SL, paratypes of Corydoras gryphus
Tencatt, Britto & Pavanelli, 2014, rio Paraná (near Ponte da
Am izade). Corydoras hastatus: Brazil: Mato Grosso: NUP
6862, 116, 13.1-20.7 mm SL, baía Caiçara. Corydoras incolicana:
Brazi l: Amazonas: MZUSP 45717, 1, 47.6 mm SL, holotype of
Corydoras incolicana Burgess, 1993, rio Içana. Corydoras julii:
Brazi l: Piau í: NUP 16225, 1, 46.8 mm SL, rio At alaia. Corydoras
kanei: Brazil: Uncertain state: MZUSP 52489, 1, 36.6 mm SL,
holotype of Corydoras kanei Grant, 1998, aquarium specimens
said to be from near the town of Boa Vista, Roraima, possibly
from the r io Branco. Corydoras lacrimostigmata: Brazil: Pa r a n á :
MNRJ 40725, 1, 31.8 mm SL, holotype of Corydoras
lacrimostigmata Tencatt, Britto & Pavanelli, 2014, rio Maria
Flora; NUP 14657, 3 c&s, 30.9-34.5 mm SL paratypes of
Corydoras lacrimostigmata Tencatt, Britto & Pavanelli, 2014, rio
Nestor. Corydoras longipinnis: Argentina: Santiago del Estero:
AI 221, 1, 59.5 mm SL, holotype of Corydoras longipinnis
Knaack, 2007, río Sali. Tucumán: NUP 14440, 2 c&s, 29.9-33.4
mm SL, Pampa-Mayo stream. Corydoras lymnades: Brazil:
Minas Gerais: MNRJ 15765, 6, 15.8-17.7 mm SL, 2 c&s, 18.1-18.4
mm SL, rio Peruaçu; MNRJ 40186, 1, 29.7 mm SL, holotype of
Corydoras lymnades Tencatt, Vera-Alcaraz, Britto & Pavanelli,
2013, rio Guarda-Mor. Corydoras maculifer: Brazil: To c a n t i ns :
NUP 8970, 2, 42.0-46.0 mm SL, ribeirão Xambioazinho.
Corydoras melanistius: Guyana: Unknown region: BMNH
1864.1.21.86, 1, 35.0 mm SL, lectotype of Corydoras melanistius
Regan, 1912, designated by Nijssen & Isbrücker, 1967, rio
Essequibo. Corydoras melini: Brazil: Amazonas: MZ USP 81163,
2, 37.0-45.0 mm SL, rio Tiquié. Corydoras multimaculatus:
Brazi l: Minas Gerais: MCP 29025, 2, 20.1-25.4 mm SL, rio
Peruaçu. Corydoras napoensis: Peru: Loreto: MZUSP 26341, 1,
27.8 mm SL, paratype of Corydoras napoensis Nijssen &
Isbrücker, 1986, Moronacocha. Corydoras nattereri: Brazil:o
Paulo: MZUSP 110255, 4 of 31, 32.0-32.8 mm SL, 2 c&s of 31,
32.3-34.4 mm SL, rio Paraitinga. Corydoras paleatus: Uruguay:
Canelones: NRM 54230, 1, 53.5 mm SL, Sarandí stream.
Corydoras panda: Peru: Huánuco: ROM 55815, 6, 26.5-39.7 mm
SL, unknown stream somewhere above Panguana in Llullapichis
drainage. Corydoras pantanalensis: Brazil: Mato Grosso: NUP
10188, 1 c&s, 46.4 mm SL, Baía Sinhá Mariana. Mato Grosso do
Sul. NUP 12593, 21, 38.7-51.2 mm SL, tributary to the rio
Miranda. Corydoras parallelus: Brazil: Amazonas: MZUSP
45716, 1, 47.4 mm SL, holotype of Corydoras parallelus Burgess,
1993, rio Içana. Corydoras pinheiroi: Brazil: Ronnia: MZUSP
48099, 1, 54.3 mm SL, holotype of Corydoras pinheiroi
Dinkelmeyer, 1995, stream tributary to the rio Ribeiro, at
Guajará-Mirim. Corydoras potaroensis: Guyana: Potaro-
Siparuni: ROM 61526, 3 of 15, 35.0-44.8 mm SL, 2 c&s of 15,
32.6-35.1 mm SL, rio Potaro. Corydoras pygmaeus: Peru: Lor et o:
MZUSP 26344, 4, 13.5-20.0 mm SL, Moronacocha. Corydoras
reticulatus: Peru: Ucayali: MZUSP 28752, 3, 37.3-45.1 mm SL,
Iamiriacocha masisea. Corydoras robineae: Brazil: Amazonas:
MZUSP 27175, 1, 33.7 mm SL, holotype of Corydoras robineae
Burgess, 1983, rio Aiuana. Corydoras sarareensis: Brazil: Mato
Grosso: MZUSP 48100, 1, 40.9 mm SL, holotype of Corydoras
sarareensis Dinkelmeyer, 1995, rio Sararé. Corydoras seussi:
Brazi l: Rondônia: MZUSP 49323, 10, 44.3-54.0 mm SL,
paratypes of Corydoras seussi Dinkelmeyer, 1996, small stream
tributary to the rio Pacas-Novos (= Pacaás Novos), near Guajará-
Mirim. Corydoras similis: Brazi l: Acre: LBP 10648, 7, 21.4-34.3
mm SL, rio Iquiri. Corydoras splendens: Brazil: Gos: NUP
1299 0, 1, 43.7 mm SL, tr ibut ary to the rio Ar a g u aia. Mat o Grosso.
NUP 10195, 1 c&s, 54.6 mm SL, Pai Caetano lake. Corydoras
stenocephalus: Brazil: Amazonas: MNRJ 3625, 3, 31.2-62.3 mm
SL, rio Javari. Corydoras sterbai: Brazil: Mato Grosso: MZUSP
94998, 1, 39.9 mm SL, rio Guaporé. Corydoras treitlii: Brazil:
Maranhão: NUP 16224, 3, 21.5-45.6 mm SL, rio Medonho.
Corydoras trilineatus: Brazil: Acre: MZUSP 30857, 3 of 25,
40.9-44.1 mm SL, 2 c&s of 25, 44.2-43.8 mm SL, rio Tarauacá.
Corydoras tukano: Brazil: Amazonas: MZUSP 82100, 40.9 mm
SL, holotype of Corydoras tukano Britto & Lima, 2003, rio
Tiquié. Corydoras xinguensis: Brazil: Mato Grosso: MZUSP
38987, 1, 34.5 mm SL, paratype of Corydoras xinguensis Nijssen,
1971, Igarapé upstream Porori village. Corydoras zygatus:
Brazi l: Acre: MZUSP 30858, 4 of 15, 41.7-47.3 mm SL, rio
Tara u a cá.
Two new Corydoras from the rio Madeira basin
Neotropical Ichthyology, 14(1): e150063, 2016
152
Acknowledgements
The Núcleo de Pesquisas em Limnologia, Ictiologia
e Aquicultura (Nupélia) of the Universidade Estadual de
Maringá and the Laboratório de Zoologia da Universidade
Federal de Mato Grosso do Sul provided logistical support.
We are grateful to Bruno Barros (Naturae), Diogo Hungria
(GIA), Ariana Ribeiro, Fabíola Vieira (UFRO), and Cintia
Oliveira (FSL) for help during the eldwork. To Carolina
Doria and Mariluce Messias (UNIR) for the donation of the
specimens used in the description of both new species. This
work is part of the project “Monitoramento e Conservação
da Ictiofauna do rio Madeira”, a partnership involving the
Universidade Federal de Rondônia, Instituto de Estudos e
Pesquisas do Agronegócio e Organizações Sustentáveis and
Santo Antônio Energia (2009-2012). The authors are grateful
to Carlos Lucena (MCP), Cláudio Oliveira (LBP), Mário de
Pinna, Aléssio Datovo and Osvaldo Oyakawa (MZUSP),
Mariluce Messias and Ângela Araujo (UFRO) and Otávio
Froehlich (in memoriam) (ZUFMS-PIS) for hosting museum
visits, curatorial assistance and loaning of material. We also
thank Hernán López-Fer nández, Don Stacey and Erling Holm
(ROM), Jorge Casciotta and Adriana Almirón (AI), Juan
Mirande (Fundación Miguel Lillo), Marcelo Britto (MNRJ)
and Sven Kullander (NRM) for the loaning and/or donation
of several specimens analyzed in this paper. To Andressa
Oliveira, Francisco Severo-Neto and Thomaz Sinani
(ZUFMS-PIS), Carlos Lucena and Héctor Vera-Alcaraz
(MCP), Cláudio Oliveira, Ricardo Britzke, Fábio Roxo,
Bruno Melo and Gabriel Silva (LBP), Osvaldo Oyakawa and
Túlio Teixeira (MZUSP) for gently welcome LFCT during
museum visits. To Marcelo Britto (MNRJ) for kindly share
his knowledge about Corydoradinae with LFCT, and for
sending several photographs of types hosted in BMNH. To
Robert McLure for the pleasant late-night talks on Corydoras
and for kindly reviewing the English language of this paper.
To William Crampton and four anonymous reviewers for
the valuable suggestions on this manuscript. To Celso Ikedo
for taking the photos used in gures 1 and 6. To Fernando
Paiva and Lucas Blanco by permitting the use and by the
assistance in the image capture laborator y of the Universidade
Federal de Mato Grosso do Sul. The Conselho Nacional de
Desenvolvimento Cientíco e Tecnológico (CNPq) provided
fellowships to LFCT (process #141061/2014-6) and the
Fundação de Amparo à Pesquisa do Estado de o Paulo
(FAPESP) provided grants to WMO (process #2013/22473-8).
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Submitted May 11, 2015
Accepted January 04, 2016 by William Crampton
... Despite the wide geographic distribution of Corydoras in cisandean South America, its representatives predominantly occur in the Amazon basin, in which more than the half of the known species can be found (Tencatt & Ohara, 2016b). The rio Madeira basin, which flows into the rio Amazonas, is the world's richest drainage regarding the number of fish species (Jézéquel et al., 2020;Torrente-Vilara et al., 2013), and currently harbours 44 species of Corydoras, representing one quarter of the total species of the genus (Ohara et al., 2016;Tencatt & Evers, 2016;Tencatt & Ohara, 2016a, 2016b. ...
... Despite the wide geographic distribution of Corydoras in cisandean South America, its representatives predominantly occur in the Amazon basin, in which more than the half of the known species can be found (Tencatt & Ohara, 2016b). The rio Madeira basin, which flows into the rio Amazonas, is the world's richest drainage regarding the number of fish species (Jézéquel et al., 2020;Torrente-Vilara et al., 2013), and currently harbours 44 species of Corydoras, representing one quarter of the total species of the genus (Ohara et al., 2016;Tencatt & Evers, 2016;Tencatt & Ohara, 2016a, 2016b. ...
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