Neotropical Ichthyology, 14(1): e150062
DOI: 10.1590/1982-0224-20150062
Journal homepage: www.scielo.br/ni
Published online: 07 March 2016 (ISSN 1982-0224)
A new Corydoras Lacépède, 1803 (Siluriformes: Callichthyidae)
from the rio Araguaia basin, Brazil, with comments about
Corydoras araguaiaensis Sands, 1990
Luiz Fernando Caserta Tencatt1 and Marcelo Ribeiro de Britto2
A new species of Corydoras is described from the rio Araguaia basin, Goiás, Brazil. The new species can be distinguished
from its congeners by presenting the following features: infraorbital 1 with very large ventral laminar expansion; infraorbital
2 contacting compound pterotic; lanks densely covered by irregular black spots; and ventral surface of trunk densely covered
by coalescent relatively well-developed platelets. It is also discussed the possibility that C. araguaiaensis comprehends a
complex of species.
Uma espécie nova de Corydoras é descrita da bacia do rio Araguaia, Goiás, Brasil. A espécie nova pode ser distinguida
de suas congêneres por apresentar as seguintes características: infraorbital 1 com expansão laminar ventral muito grande;
infraorbital 2 em contato com o pterótico composto; lancos densamente cobertos por manchas pretas irregulares e superfície
ventral do tronco densamente coberta por plaquetas coalescentes relativamente bem desenvolvidas. Discute-se também a
possibilidade de C. araguaiaensis compreender um complexo de espécies.
Keywords: C65, Corydoradinae, Corydoras sp. “Guaraná”, Goiás State, Taxonomy.
by ecological and phylogenetic aspects, showing relations
and patterns between Müllerian co-mimics of certain
regions. In that work, C. araguaiaensis is proposed as
member of a mimetic group along with C. maculifer and
an undescribed species coded as C122 (see more details
about the “C-number” system and its species in Fuller &
Evers, 2005: 280). The most obvious way to recognize
each mimetic morphotype is the snout morphology. There
are at least three conspicuous types of snouts, the typical
long snout pattern, present in the species of the lineage
1, as C. maculifer, the straight or intermediate long snout
pattern, present in the species of the lineage 8, as C122,
and the short snout pattern, present in the species of the
lineages 4, 5, 6, 7, and 9, as C. araguaiaensis.
Generally, the mimetic pairs or groups are composed by
one representative of each snout pattern (see Alexandrou et
al., 2011: 3, ig. 2). However, since the short snout pattern
is present in conspicuously diferent lineages, it is possible
that two or more short-snouted species may also present
homoplastic color patterns, as the observed for Corydoras
guapore Knaack, 1961, from the lineage 4, and C.
caudimaculatus Rössel, 1961, from the lineage 9, both from
Introduction
The Callichthyidae armored catishes are easily
diagnosed from other Siluriformes by the presence
of two longitudinal series of dermal plates on lanks
(Eigenmann & Eigenmann, 1890; Reis, 1998). The family
currently comprises more than 200 species, including
Corydoras Lacépède, 1803, the most species-rich genus
of Siluriformes, with more than 170 valid species (Reis,
2003; Eschmeyer, 2015). Since the 1940’s, many eforts to
understand the taxonomy and phylogenetic relationships
of the species of Corydoras have been made (e.g., Gosline,
1940; Nijssen, 1970; Nijssen & Isbrücker, 1980; Britto,
2003; Alexandrou et al., 2011), however, the knowledge
of interrelations among Corydoras species is still poorly
known (Britto et al., 2007).
Currently, only three species of Corydoras are
recorded from the rio Araguaia basin: C. araguaiaensis
Sands, 1990, C. cochui Myers & Weitzman, 1954, and
C. maculifer Nijssen & Isbrücker, 1971 (Eschmeyer,
2015). Alexandrou et al. (2011) studied the community
structure of mimetic lineages of Corydoras propitiated
1
Universidade 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)
2
Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Setor de Ictiologia. Quinta da Boa Vista s/n,
São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil. mrbritto2002@yahoo.com.br
1
Neotropical Ichthyology, 14(1): e150062
2e1500622
A new Corydoras from the rio Araguaia basin
the rio Guaporé basin (see Alexandrou et. al., 2011: suppl.
ig. 2). After the analysis of some Corydoras specimens
captured by Hans-Georg Evers during an expedition in
Brazil, a new short-snouted species with similar color pattern
to C. araguaiaensis was revealed, which is described herein.
Additionally some comments about the taxonomic status of
C. araguaiaensis were also provided.
History, Smithsonian Institution (USNM), Washington, D.C.
Coleção Zoológica de Referência da Universidade Federal
de Mato Grosso do Sul (ZUFMS-PIS), Campo Grande.
Material and Methods
u r n:lsid:zooba n k .org:a ct:541C78B5 -98B6 - 4F BF- 89A 3 6961C9EA74D2
Measurements were obtained using digital calipers to
the nearest tenth of millimeter. Morphometric and meristic
data were taken following Reis (1997) with modiications
of Tencatt et al. (2013). Morphometrics are reported as
percentages of standard length (SL) and head length (HL).
Homology of barbels follows Britto & Lima (2003). Some
specimens were cleared and stained (c&s) according to
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 followed Britto et al. (2009).
Comparative data of Corydoras esperanzae Castro, 1987,
C. evelynae Rössel, 1963, C. leopardus Myers, 1933, C.
melanotaenia Regan, 1912, Corydoras multiradiatus (OrcésVillagomez, 1960), C. orphnopterus Weitzman & Nijssen,
1970, C. rabauti La Monte, 1941, C. schwartzi Rössel, 1963,
and C. xinguensis Nijssen, 1972, were obtained through their
original descriptions and/or high resolution photographs of
type-specimens available from 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: Asociación Ictiológica de
La Plata (AI), La Plata; Academy of Natural Sciences of
Drexel University (ANSP), Philadelphia; Natural History
Museum (BMNH), London; Laboratório de Biologia de
Peixes da Universidade Estadual Paulista “Júlio de Mesquita
Filho” (LBP), Botucatu; Museu de Ciências e Tecnologia
da Pontifícia Universidade Católica do Rio Grande do Sul
(MCP), Porto Alegre; Museum of Comparative Zoology
(MCZ), Harvard University, Cambridge; Museu Nacional,
Universidade Federal do Rio de Janeiro (MNRJ), Rio de
Janeiro; Museu de Zoologia da Universidade de São Paulo
(MZUSP), São Paulo; Naturhistoriska Riksmuseet (NRM),
Stockholm; Coleção Ictiológica do Núcleo de Pesquisas
em Limnologia, Ictiologia e Aquicultura da Universidade
Estadual de Maringá (NUP), Maringá; Royal Ontario
Museum (ROM), Toronto; National Museum of Natural
Results
Corydoras eversi, new species
Figs. 1, 2a, c, 3-5
Corydoras cf. araguaiaensis ‘C65’.-Alexandrou et al., 2011: 22
(supplement).
Corydoras sp. C65.-Fuller & Evers, 2005: 282 [checklist], 309
[illustration].
Corydoras sp. ‘Guarana’ [sic].-Fuller & Evers, 2005: 310
[illustration].
Holotype. MNRJ 43195, 44.5 mm SL, Brazil, Goiás,
Montes Claros de Goiás, unnamed stream tributary to the
rio Araguaia, 15°53’10”S 51°41’34”W, 21 Mar 1998, H.-G.
Evers & P.V. da Silva.
Paratypes. MZUSP 117333, 6, 23.4-35.8 mm SL; NUP
17309, 5, 24.8-42.9 mm SL; NUP 17310, 3 c&s, 42.5- 45.3
mm SL; ZUFMS-PIS 4062, 5, 32.2-39.2 mm SL, same data
as holotype.
Diagnosis. Corydoras eversi can be distinguished from
its congeners, with exception of C. aeneus (Gill, 1858),
C. britskii (Nijssen & Isbrücker, 1983), C. diluviatilis
Britto & Castro, 2002, C. eques Steindachner, 1876, C.
garbei Ihering, 1911, C. melanotaenia, C. multiradiatus,
C. rabauti, C. splendens (Castelnau, 1855), and C.
zygatus Eigenmann & Allen, 1942, by the presence of
infraorbital 1 with very large ventral laminar expansion
(vs. poorly-, or moderately-developed ventral expansion);
and infraorbital 2 contacting compound pterotic (vs.
not contacting). The new species can be distinguished
from C. aeneus, C. britskii, C. diluviatilis, C. eques, C.
garbei, C. melanotaenia, C. multiradiatus, C. rabauti, C.
splendens, and C. zygatus by presenting lanks densely
covered by irregular black spots (vs. background color of
lanks orange yellow with dorsolateral body plates entirely
or almost entirely blackened in C. aeneus, C. britskii, C.
melanotaenia, C. multiradiatus, C. rabauti, C. splendens,
and C. zygatus; background color of lanks orange yellow
with lanks almost entirely blackned in C. eques; midline of
lank with a longitudinal series of large rounded blotches,
remaining areas with black spots in C. diluviatilis and C.
garbei); and ventral surface of trunk densely covered by
coalescent relatively well-developed platelets (vs. ventral
surface of trunk naked; or covered by sparse and relatively
small platelets).
Neotropical Ichthyology, 14(1): e150062
L. F. C. Tencatt & M. R. Britto
3
Fig. 1. Corydoras eversi, holotype, MNRJ 43195, 44.5 mm SL, Brazil, Goiás, Montes Claros de Goiás, unnamed stream
tributary to the rio Araguaia, 15°53’10”S 51°41’34”W. Dorsal (top), lateral (middle) and ventral (bottom) views. Photo by
Celso Ikedo.
Neotropical Ichthyology, 14(1): e150062
4e1500624
A new Corydoras from the rio Araguaia basin
Description. Morphometric data presented in Table 1.
Head compressed with convex dorsal proile; triangular
in dorsal view. Snout short and rounded. Head proile
convex from tip of snout to anterior nares; ascending
nearly straight from this point to tip of posterior process of
parieto-supraoccipital; region just posterior to eye slightly
concave in some specimens. Proile slightly convex along
dorsal-in base. Postdorsal-in body proile nearly straight
to adipose-in spine; concave from this point to caudal-in
base. Ventral proile of body slightly convex from isthmus
to pelvic girdle. Proile nearly straight from pelvic girdle
to base of irst anal-in ray; concave until caudal-in base.
Body roughly elliptical in cross section at pectoral girdle,
gradually becoming more compressed toward caudal in.
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 lap 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 moderate in
size, not reaching anteroventral limit of gill opening. Outer
mental barbel slightly larger than maxillary barbel. Inner
mental barbel leshy, with base close to its counterpart.
Small rounded papillae covering entire surface of all
barbels, upper and lower lips, and isthmus.
Table 1. Morphometric data of holotype and 19 paratypes
of Corydoras eversi. SD = standard deviation.
Standard length (mm)
Holotype
Low-High
Mean±SD
44.5
23.4-45.3
-
Percents of standard length
Depth of body
43.1
40.9-45.7
Predorsal distance
49.7
48.9-54.3
42.7±1.1
51.6±1.2
Prepelvic distance
47.4
46.4-53.0
48.7 ±2.0
Preanal distance
83.1
77.7-87.2
82.7±2.0
Preadipose distance
89.0
84.9-98.8
87.9±2.9
Length of dorsal spine
22.9
21.6-30.4
26.7±2.7
Length of pectoral spine
25.4
25.2-32.8
28.7±2.1
Length of adipose-in spine
6.1
5.2-9.3
7.5±1.0
Depth of caudal peduncle
15.1
14.6-17.2
15.9±0.7
Length of dorsal-in base
17.1
16.8-20.1
18.2±0.9
Dorsal to adipose distance
27.0
18.0-27.0
22.0±1.9
Maximum cleithral width
30.1
28.0-31.9
30.0±1.0
Head length
44.3
43.5-48.0
45.5±1.2
14.4
9.6-17.0
12.1±1.9
Length of maxillary barbel
Percents of head length
Head depth
91.9
84.4-94.8
89.0±2.8
Least interorbital distance
34.0
31.4-36.3
33.4±1.3
Horizontal orbit diameter
20.8
19.2-24.3
22.4±1.6
Snout length
35.0
31.5-38.0
35.3±1.6
Least internarial distance
18.8
14.0-18.8
16.2±1.5
Mesethmoid short; anterior tip poorly developed,
smaller than 50% of bone length (see Britto, 2003: 123,
character 1, state 1; ig. 1B); posterior portion relatively
narrow, partially exposed and bearing minute odontodes.
Nasal slender, curved laterally, with inner margin laminar;
mesial border contacting frontal and mesethmoid. 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 parietosupraoccipital. 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 well
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.
2c). Posterodorsal ridge of hyomandibula close to its
articulation with opercle oblong; exposed, relatively
thick; dorsal ridge of hyomandibula between compound
pterotic and opercle exposed; exposed areas bearing
small odontodes. Interopercle entirely or almost entirely
covered by thick layer of skin, somewhat triangular,
anterior projection well-developed. Preopercle relatively
thick, elongated, minute odontodes sparse on external
surface. Opercle conspicuously 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 20 (2), 21 (1);
ribs 6 (3), irst pair conspicuously large; complex vertebra
compact in shape. Neural and haemal spines with expanded
in distal tips.
Four branchiostegal rays decreasing in size posteriorly.
Hypobranchial 2 somewhat triangular, tip ossiied and
directed towards anterior portion, posterior margin
cartilaginous; ossiied 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, 37
to 46 (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 irregular laminar expansion on posterior margin.
Upper tooth plate oval; 34 to 43 (3) teeth aligned in two
rows on postero-ventral surface.
Neotropical Ichthyology, 14(1): e150062
L. F. C. Tencatt & M. R. Britto
5
Fig. 2. Lateral view of the head of c&s specimens of (a, c) Corydoras eversi, paratype, NUP 17310, 42.5 mm SL, showing
the well-developed ventral expansion of the infraorbital 1 and the platelets on the snout (a) and infraorbital 2 in contact with
compound pterotic (c); and (b, d) Corydoras araguaiaensis, MZUSP 87155, 31.8 mm SL, showing the moderately-developed
ventral expansion of the infraorbital 1 (b) and infraorbital 2 not in contact with compound pterotic (d). The dotted lines in
(c) and (d) represent the suture between sphenotic and compound pterotic. Abbreviations: io1: infraorbital 1, io2: infraorbital
2, sph: sphenotic, cpt: compound pterotic. Scale bar = 1.0 mm.
Neotropical Ichthyology, 14(1): e150062
6e1500626
A new Corydoras from the rio Araguaia basin
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. Nasal canal with three openings, irst 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 or three 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 in triangular, located just posterior to second
dorsolateral body plate. Dorsal-in rays II,7 (1), II,8* (19),
posterior margin of dorsal-in spine with ive to 14 very
reduced serrations directed towards tip of spine; serrations
absent proximally. 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-in branched ray;
anterior margin with small odontodes. Pectoral in triangular,
its origin just posterior to gill opening. Pectoral-in rays I,7
(10), I,8* (10); posterior margin of pectoral spine with 23 to
32 small serrations along its entire length; serrations directed
towards tip of spine (Fig. 3). Pelvic in oblong, located just
below irst ventrolateral body plate, and at vertical through
irst branched dorsal-in ray. Pelvic-in rays i,5. Adipose in
roughly triangular, separated from base of last dorsal-in
ray by generally seven dorsolateral body plates. Anal in
triangular, located just posterior to 12th ventrolateral body
plates, and at vertical through anterior margin of adipose-in
spine. Anal-in rays ii,5 (6), ii,6* (13). Caudal-in rays i,7,i
(1), i,11,i* (1), i,12,i (18), generally four dorsal and ventral
procurrent rays; bilobed, dorsal lobe generally slightly larger
than ventral lobe.
Three or four laterosensory canals on trunk; irst ossicle
tubular, second ossicle laminar, third and fourth lateral-line
canals, if present, encased in third and fourth dorsolateral
body plates, respectively. Body plates with minute odontodes
scattered over exposed area, a conspicuous line of odontodes
conined on posterior margins; dorsolateral body plates 21
(2), 22 (9), 23* (9); ventrolateral body plates 18 (1), 19 (8), 20
(10), 21* (1); dorsolateral body plates along dorsal-in base
5* (6), 6 (14); dorsolateral body plates between adipose- and
caudal-in 6 (10), 7* (9), 8 (1); preadipose platelets 2 (3), 3 (16),
4* (1); small platelets covering base of caudal-in 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, snout,
region surrounding ventral margin of preopercle, and upper
lip covered with relatively large platelets bearing odontodes
(Fig. 2a). Ventral surface of trunk covered by coalescent
relatively well-developed platelets (Fig. 4); region between
pelvic and anal ins with platelets in some specimens.
Fig. 3. Right pectoral spine (11.4 mm long) of Corydoras
eversi, paratype, NUP 17310, 45.3 mm SL, showing the
serrations on posterior margin.
Fig. 4. Ventral surface of the trunk of Corydoras eversi,
paratype, NUP 17310, 44.4 mm SL, showing the relatively
large coalescent platelets covering the area between pectoral
and pelvic girdles. Scale bar = 1.0 mm.
Color in alcohol. Overall color of body in Fig. 1. Ground
color of body brownish yellow, with top of head and snout
dark brown. Ventral margin of orbit, above infraorbital
1, blackened. Maxillary barbel and anterior portion of
outer mental barbel covered by black chromatophores.
Dorsal and lateral portion of body, with exception of small
region of ventrolateral body plates surrounding pelvic in,
densely covered by irregular black spots; spots arranged
longitudinally on lanks, forming irregular stripes.
Ventral region of body close to pectoral-in spine origin
with black spots in some specimens. Ground color of ins
grayish brown. All ins covered by irregular black spots;
pectoral, pelvic, anal, and caudal ins with black spots
difuse or absent in some specimens. Caudal in generally
with scattered spots, not forming transversal black bars;
with faded spots arranged in up to six transversal slender
black bars in few specimens.
Color in life. Very similar to preserved specimens but
with ground color of body intense orange; body covered by
greenish yellow iridescent coloration (Fig. 5).
Neotropical Ichthyology, 14(1): e150062
L. F. C. Tencatt & M. R. Britto
7
Fig. 5. Aquarium specimen of Corydoras eversi, nearly 37.0 mm SL, showing its peculiar bright orange color pattern. Photo
by Hans-Georg Evers.
Sexual dimorphism. Presence of lanceolate genital papilla
in males, which occurs in all Corydoradinae (see Nijssen &
Isbrücker, 1980; Britto, 2003). The males of the new species
present irst and second branched dorsal-in rays slightly
larger than in females. Additionally, the males of Corydoras
eversi present pointed oblong pelvic in, while in females
the pelvic in is rounded. In Corydoras females, the rounded
pelvic in is generally used to make a ventral pouch to hold
the eggs during spawning (H.-G. Evers pers. comm.).
Distribution. Corydoras eversi is known from its type
locality, an unnamed stream tributary to the rio Araguaia
(Fig. 6).
Ecological notes. The specimens examined herein were
captured in a clear water stream, with sandy ground and fast
lowing. After capture the specimens are light orange. This
color remains in aquarium when the water conditions are
adequate to its biological aspects. In the type locality of the
new species no other congener were observed in syntopy
(H.-G. Evers pers. comm.).
Etymology. Corydoras eversi is named in honor of
Hans-Georg Evers, a dear friend and great enthusiast
in the ishkeeping hobby, especially in the breeding of
Corydoras species. Hans collected the specimens of C.
eversi that apparently originated all the stock present in
the hobby until the present day and also the specimens
used herein for the description.
Fig. 6. Map showing the type locality (red star) of Corydoras
eversi, an unnamed stream tributary to the rio Araguaia,
Goiás State.
Conservation status. At the time of the capture of
Corydoras eversi, the type locality presented high degree of
deforestation and presence of nearby cattle farming. There
was an encampment of the ʻMovimento dos Trabalhadores
Rurais Sem Terra - (MST)ʼ just a few kilometers away
from the collection site (Hans-Georg Evers pers. comm.).
Despite the type locality presents, at least in the year of
1998, nearby human and cattle occupation and severe
deforestation, in addition to the scarcity of the known
material of C. eversi, apparently, no additional collecting
Neotropical Ichthyology, 14(1): e150062
8e1500628
A new Corydoras from the rio Araguaia basin
trip has been performed in the region since 1998. Therefore,
the current situation of the type locality is unknown and,
with the available data, it is not possible to determine if any
of the aforementioned potential threats has directly afected
the new species. Additionally, it is very possible that the
new species may be further spread than the type locality
region, potentially occurring in other creeks and streams
of the region. Thus, it seems like the most appropriate
conservation status according to the International Union
for Conservation of Nature (IUCN) categories and criteria
(IUCN Standards and Petitions Subcommittee, 2014), is
Least Concern (LC).
Remarks. Corydoras eversi is known by exporters and in
the ishkeeping hobby as Corydoras sp. “Guaraná” due to
the intense and shiny orange pigmentation displayed by live
specimens, which is very similar to the Brazilian lemonade.
Evers (1998) coded the new species as C65, according to the
“C-number” system proposed by Evers (1993).
Discussion
The new species present a color pattern similar to C.
albolineatus Knaack, 2004, C. apiaka Espíndola, Spencer,
Rocha & Britto, 2014, C. araguaiaensis, C. haraldschultzi
Knaack, 1962, C. maculifer, C. multimaculatus
Steindachner, 1907, C. polystictus Regan, 1912, and C.
xinguensis Nijssen, 1972. Corydoras eversi can be clearly
diagnosed from C. haraldschultzi and C. maculifer by the
presence of short mesethmoid (vs. long) and serrations on
posterior margin of pectoral spine directed towards the tip
of the spine (vs. directed towards pectoral-spine origin).
From C. albolineatus, C. apiaka, C. araguaiaensis and C.
polystictus, the new species can be distinguished by the
presence of infraorbital 2 in contact with compound pterotic
(Fig. 2c) (vs. not in contact (Fig. 2d)). The new species
difers from C. multimaculatus and C. xinguensis by the
presence of ventral laminar expansion of infraorbital 1 well
developed (vs. moderately developed) and ventral surface
of the trunk densely covered by coalescent relatively well
developed platelets (vs. covered by sparse relatively small
platelets).
Corydoras eversi presents osteological pattern similar
to the species from clade IX sensu Britto (2003). The
presence of a well-developed ventral laminar expansion
in the infraorbital 1 corresponds to the state 3 of character
15 of Britto’s (2003) phylogeny, a synapomorphy of the
species located in clade IX. This clade is composed of
species previously allocated in Brochis Cope, 1871, in
addition to C. aeneus, C. eques, C. rabauti, and C. zygatus.
Corydoras melanotaenia also appeared correlated with
the aforementioned species in the phylogenetic hypothesis
conducted by Alexandrou et al. (2011). As presented in the
diagnosis, the species mentioned above can be promptly
distinguished from C. eversi by their peculiar general color
pattern, with uniform background color and dark dorsal
region, clearly diferent from the densely spotted pattern
of the new species. Additionally, C. coppenamensis, C.
diluviatilis, C. garbei, and C. lymnades also present a welldeveloped ventral laminar expansion in the irst infraorbital.
The new species can be clearly distinguished from these
species by the presence of ventral surface of trunk densely
covered by relatively large coalescent platelets (vs. ventral
surface of trunk naked; or covered by sparse and relatively
small platelets).
The presence of relatively well-developed coalescent
platelets in the ventral surface of the trunk of Corydoras
eversi (Fig. 4) is an uncommon condition only displayed by
few congeners, such as C. araguaiaensis, C. esperanzae,
C. evelynae, C. fowleri Böhlke, 1950, C. julii Steindachner,
1906, C. leopardus, C. orphnopterus, C. schwartzi,
and C. trilineatus Cope, 1872. The new species can be
distinguished from C. evelynae, C. orphnopterus, and C.
schwartzi by the absence of a concentration of a transversal
black stripe (“mask”) across the eye (vs. presence).
Corydoras eversi difers from C. julii, C. leopardus, and
C. trilineatus by the absence of a conspicuous longitudinal
black midlateral stripe bordered dorsally and ventrally by
an unpigmented area (vs. presence). From C. araguaiaensis,
the new species difers by the presence of infraorbital 1
with ventral laminar expansion well developed (Fig. 2a) (vs.
moderately developed (Fig. 2b)) and relatively large platelets
bearing odontodes on snout (vs. plates generally absent;
when present, plates are scarce and restrict to the anterior
margin of orbit, above lateral ethmoid). From C. fowleri, it
can be distinguished by having short and rounded snout (vs.
long and conspicuously pointed). Finally, C. eversi can be
distinguished from C. esperanzae by the presence of black
spots on the body (vs. absence).
Despite the great general resemblance between
Corydoras eversi and C. araguaiaensis, both species
present clearly diferent morphology, mainly in osteological
features, as mentioned above. However, even the similar
color pattern presents some conspicuous diferences. The
spots are smaller and more numerous in the new species
(Fig. 1) whereas C. araguaiaensis presents larger and
less numerous spots (Fig. 7). The caudal-in color pattern
is difuse in C. eversi, with faded spots (Fig. 1), forming
difuse conspicuously slender transversal black bars only
in few specimens, contrary to C. araguaiaensis which
presents conspicuous and thicker transversal black bars on
caudal in (Fig. 7).
Evers (1993) proposed a coding system for the
Corydoradinae to avoid the creation of nomina nuda by
using trade names. Some potentially new species from the
rio Araguaia basin with similar color pattern to Corydoras
araguaiaensis were coded in this system, including C.
eversi. Additionally, a species from the rio Cristalino
basin, Mato Grosso State, very similar to C. araguaiaensis
but with larger spots and more intense black pigmentation
on anterior portion of dorsal in, was considered new and
coded as C45 (Fig. 8). Initially, C45 appears to be easily
Neotropical Ichthyology, 14(1): e150062
L. F. C. Tencatt & M. R. Britto
distinguished from C. araguaiaensis by its color pattern,
but after a deep analysis the general morphology proved
to be very similar, with mixed morphotypes, making all
the aforementioned potential diagnostic features present
some degree of overlap. By this reason, a clear diagnosis
between C. araguaiaensis and C45 was not possible and
they were treated herein as a single taxon. However,
since the original description of C. araguaiaensis lacks
many important information, mainly skeletal, beyond the
existence of two diferent morphotypes, the possibility
that C. araguaiaensis comprehends a complex of species
is reasonable. A complete revisionary study of C.
araguaiaensis, including a molecular approach, is needed
to elucidate its taxonomic status.
9
Comparative material examined. Corydoras acutus: Peru:
Unknown department. MNRJ 3985, 2, 47.1-54.8 mm SL, SanshoCaño. Corydoras adolfoi: Brazil: Amazonas. LBP 6863, 2, 27.5-31.7
mm SL, igarapé Puranga. LBP 6871, 2, 32.2-32.5 mm SL, unnamed
igarapé. Corydoras aeneus: Trinidad-Tobago: Unknown
department. USNM 1116, lectotype of Hoplosoma aeneum Gill,
1858, 38.2 mm SL, Island of Trinidad, West Indies. 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 Nijssen
& Isbrücker, 1983, rio Approuague. Corydoras araguaiaensis:
Brazil: Mato Grosso. MCP 40230, 2, 35.3-44.2 mm SL, rio Paciguara;
Fig. 7. Corydoras araguaiaensis, MZUSP 87155, 45.8 mm SL, showing the general color pattern of the species in lateral
view. Photo by Celso Ikedo.
Fig. 8. Uncatalogued aquarium specimen of Corydoras araguaiaensis attributed to C45, showing the general color pattern
of the species in lateral view. Photo by Hans-Georg Evers.
Neotropical Ichthyology, 14(1): e150062
10
e15006210
A new Corydoras from the rio Araguaia basin
MCP 40243, 3 of 5, 33.9-38.1 mm SL, unnamed stream tributary to
the ribeirão Gameleira; MCP 40270, 1, 27.0 mm SL, ribeirão Santana;
MZUSP 87155, 4 of 33, 24.9-46.7 mm SL, 2 c&s, 27.6-31.8 mm SL,
Corixo da Saudade. Corydoras cf. araguaiaensis (C45): MZUSP
86269, 63, 24.1-49.3 mm SL, 2 c&s, 29.9-36.3 mm SL, Corixão do
Meio. 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 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, tributary 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: Venezuela: 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 cochui: Brazil: Goiás. MZUSP 89055, 6, 18.723.6 mm SL, rio do Peixe II. Tocantins. MZUSP 35838, 4 of 6, 16.118.5 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
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: Brazil: 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 diluviatilis: Brazil: São Paulo. MZUSP 75268,
1, 39.8 mm SL, holotype of Corydoras diluviatilis 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 laveolus: Brazil: São Paulo. MZUSP 424, 1, 33.4 mm SL,
holotype of Corydoras laveolus 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 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 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 Amizade). 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 hastatus: Brazil: Mato Grosso. NUP 6862, 116,
13.1-20.7 mm SL, baía Caiçara. Corydoras incolicana: Brazil:
Amazonas. MZUSP 45717, 1, 47.6 mm SL, holotype of Corydoras
incolicana Burgess, 1993, rio Içana. Corydoras julii: Brazil: Piauí.
NUP 16225, 1, 46.8 mm SL, rio Atalaia. 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 rio Branco.
Corydoras lacrimostigmata: Brazil: Paraná. 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: Tocantins. 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 multimaculatus:
Brazil: Minas Gerais. MCP 29025, 2, 20.1-25.4 mm SL, rio Peruaçu.
Corydoras nattereri: Brazil: São 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: Rondônia. 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, 35.044.8 mm SL, 2 c&s of 15, 32.6-35.1 mm SL, rio Potaro. 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: Brazil: 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: Brazil: Acre. LBP 10648, 7, 21.434.3 mm SL, rio Iquiri. Corydoras splendens: Brazil: Goiás. NUP
12990, 1, 43.7 mm SL, tributary to the rio Araguaia. Mato 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 treitlii: Brazil: Maranhão. NUP 16224, 3, 21.5-
Neotropical Ichthyology, 14(1): e150062
L. F. C. Tencatt & M. R. Britto
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 zygatus: Brazil: Acre. MZUSP 30858, 4
of 15, 41.7-47.3 mm SL, rio Tarauacá.
Acknowledgments
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.
The authors are grateful to Carlos Lucena (MCP), Cláudio
Oliveira (LBP), Mário de Pinna, Aléssio Datovo, and
Osvaldo Oyakawa (MZUSP) and Otávio Froehlich (in
memoriam) (ZUFMS-PIS) for hosting museum visits and
loaning of material. We also thank Hernán López-Fernández,
Don Stacey and Erling Holm (ROM), Jorge Casciotta and
Adriana Almirón (AI), Juan Mirande (Fundación Miguel
Lillo), 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 VeraAlcaraz, 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 Fernando Paiva and Lucas Blanco by
permitting the use and by the assistance in the image capture
laboratory of the Universidade Federal de Mato Grosso do
Sul. To Hans-Georg Evers by sending the type-specimens
and general information about Corydoras eversi. To Robert
‘Rob’ McLure for the pleasant late-night talks on Corydoras
and for kindly reviewing the English language of this paper.
To Celso Ikedo for taking the photos used in the igures 1
and 7. Financial support to LFCT was provided by Conselho
Nacional de Desenvolvimento Cientíico e Tecnológico
(CNPq, process #141061/2014-6), and MRB by Fundação
Carlos Chagas Filho de Amparo a Pesquisa do Estado do
Rio de Janeiro (FAPERJ, process #E-111.268/2014).
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Submitted May 11, 2015
Accepted September 20, 2015 by Fernando Carvalho