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Review of the Genus Cathorops (Siluriformes: Ariidae) from the Caribbean and Atlantic South America, with Description of a New Species

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The New World genus Cathorops in the family Ariidae (Sea Catfishes) includes species that inhabit estuarine and coastal waters as well as freshwaters, playing an important role in Neotropical coastal and estuarine fisheries. The relatively conserved external morphology coupled with the marked sexual dimorphism and ontogenetic variation makes it difficult to recognize and diagnose the species. One of the major problems concerns the nomenclatural and geographical limits of C. spixii, described from tropical Brazil and often treated as the only coastal marine species in the genus from the Western Atlantic. Examination of types of nominal species as well as comprehensive collections of non-types from Caribbean and Atlantic South America, lead us to conclude that spixii is restricted to Brazil and that variolosus are synonyms of nuchalis, spixii, and C. arenatus, respectively. We also describe a new species in the Cathorops mapale species group from Colombia and Venezuela.
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Review of the Genus Cathorops (Siluriformes: Ariidae) from the Caribbean
and Atlantic South America, with Description of a New Species
Alexandre P. Marceniuk
1,2,3
, Ricardo Betancur-R.
4,5
, Arturo Acero P.
6
, and
Janice Muriel-Cunha
3
The New World genus Cathorops in the family Ariidae (Sea Catfishes) includes species that inhabit estuarine and coastal
waters as well as freshwaters, playing an important role in Neotropical coastal and estuarine fisheries. The relatively
conserved external morphology coupled with the marked sexual dimorphism and ontogenetic variation makes it
difficult to recognize and diagnose the species. One of the major problems concerns the nomenclatural and
geographical limits of C. spixii, described from tropical Brazil and often treated as the only coastal marine species in
the genus from the Western Atlantic. Examination of types of nominal species as well as comprehensive collections of
non-types from Caribbean and Atlantic South America, lead us to conclude that C. spixii is restricted to Brazil and that C.
nuchalis is a valid species, ranging from Venezuela to Guyana. The nominal species Arius laticeps,Arius nigricans, and Arius
variolosus are synonyms of C. nuchalis, C. spixii, and C. arenatus, respectively. We also describe a new species in the
Cathorops mapale species group from Colombia and Venezuela.
THE New World genus Cathorops is among the most
abundant and species-rich genera of Sea Catfishes
(family Ariidae). It includes at least 19 species and
ranges from the Gulf of Mexico through southeastern Brazil
in the Western Atlantic, and from Baja California through
northern Peru in the Eastern Pacific (Marceniuk and
Menezes, 2007; Marceniuk and Betancur-R., 2008; Marce-
niuk et al., 2009). While representatives of Cathorops are
most abundant in brackish waters, a few species occur in
marine waters and others are restricted to freshwater
habitats (Marceniuk and Betancur-R., 2008; Betancur-R.,
2010). As compared to other ariids, members of Cathorops
are relatively small, rarely growing beyond 30 cm in total
length.
The genus Cathorops is morphologically well diagnosed
from other Neotropical ariid genera (Marceniuk et al., 2009),
and its monophyly is strongly supported on molecular and
anatomical evidence (Betancur-R. et al., 2007; Marceniuk
and Menezes, 2007; Betancur-R., 2009). In a recent classifi-
cation scheme of New World ariids using mitochondrial,
nuclear, and morphological evidence, Betancur-R. et al.
(2007) hypothesized relationships among 13 species of
Cathorops, recognizing two subgenera, the monotypic Pre-
cathorops for Cathorops dasycephalus, and Cathorops for the
remaining species (see also Betancur-R., 2009; Marceniuk
et al., 2009:fig. 1 and appendix I). While efforts toward
documenting the alpha taxonomy of Cathorops have
increased in the last decade, particularly for Mesoamerican
and Eastern Pacific taxa (Marceniuk, 2007a; Marceniuk and
Betancur-R., 2008; Marceniuk et al., 2009), species differen-
tiation is difficult given their overall similarity in external
morphology and marked sexual dimorphism (e.g., accessory
tooth patches). As a result, the validity of several nominal
species described from the Caribbean and Atlantic coasts of
South America (for the sake of brevity hereafter eastern
South America [ESA]) remains unclear.
With the exception of a few comprehensive revisions (e.g.,
Eigenmann and Eigenmann, 1890; Eigenmann, 1912),
recognition of valid species of Cathorops from ESA is often
based on original descriptions, citation of previous taxo-
nomic accounts, and collections from restricted geographic
locations, without examination of type material (Gu¨nther,
1864; Miranda-Ribeiro, 1911; Gosline, 1945). These difficul-
ties led Taylor and Menezes (1978) to treat all nominal
marine and coastal species of Cathorops from the Western
Atlantic as synonyms of C. spixii, a decision adopted by
several subsequent authors (e.g., Uyeno et al., 1983;
Cervigo´ n, 1992; Castro-Aguirre et al., 1999; Acero P., 2002),
while other more recent accounts have listed species with
uncertain status as species inquirendae (e.g., Arius laticeps,A.
nigricans,A. nuchalis,A. puncticulatus, and A. variolosus;
Marceniuk and Ferraris, 2003; Ferraris, 2007; Marceniuk and
Menezes, 2007). With the aim of clarifying the taxonomic
problems involving ESA species in Cathorops, Betancur-R. and
Acero P. (2005) described C. mapale from Colombia; likewise,
Marceniuk (2007b) redescribed C. agassizii and C. arenatus,
from Brazil and Suriname, respectively.
By comparison of types of species of Cathorops from ESA as
well as a comprehensive collections of non-types from a
wide geographic range, the aims of this paper are to
redescribe Cathorops spixii and redefine its taxonomic and
geographic limits, clarify the status of four names currently
listed as inquirendae, and describe a new species from
Colombia and Venezuela.
MATERIALS AND METHODS
Measurements were made as specified in Marceniuk (2007b),
taken with either a ruler and recorded to the nearest millimeter
or with dial calipers and recorded to the nearest 0.1 mm.
Measurements are given as percents of standard length (SL),
unless stated otherwise. Individual measurements are given for
1
Museu de Zoologia da Universidade de Sa˜o Paulo, Caixa Postal 42494, 04218-970 Sa˜o Paulo, SP, Brazil.
2
Nu´ cleo Integrado de Biotecnologia, Universidade de Mogi das Cruzes, 08780-911 Mogi das Cruzes, SP, Brazil; E-mail: a_marceniuk@
hotmail.com. Send reprint requests to this address.
3
Instituto de Estudos Costeiros, Campus de Braganc¸a, Universidade Federal do Para´, Alameda Leandro Ribeiro, 68600-000 Braganc¸a, Para´,
Brazil.
4
Department of Biological Sciences, Auburn, Alabama.
5
Department of Biological Sciences, The George Washington University, 2023 G Street NW suite 340 Washington, D.C. 20052.
6
Universidad Nacional de Coloˆ mbia, Instituto de Ciencias Naturales, Cerro Punta Betı´n, Apartado 1016 (INVEMAR), Santa Marta, Colombia.
Submitted: 24 December 2010. Accepted: 7 November 2011. Associate Editor: R. E. Reis.
F2012 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-10-202
Copeia 2012, No. 1, 77–97
primary types in tables and meristic values for the type-
specimens are indicated in bold in tables. Whenever possible,
sex of specimens was determined by examination of gonad
morphology under magnification. The dorsomedial groove of
neurocranium referred to herein is formed by the anterior
fleshy portion apposed to the anterior cranial fontanel and
limited by the posterior branches of the mesethmoid and
frontals and a posterior bony portion formed by the mesial
depression of the frontals and the anterior portion of
supraoccipital (Marceniuk et al., 2009). The term ‘‘nuchal
plate’’ refers to the fusion of the anterior and the medial
nuchal plates (Royero, 1987). Vertebral meristics obtained
from radiographs, skeletons, and cleared-and-stained (CS)
specimens include only elements free from the ventral
superficial ossification, counted from the first free vertebra to
the compound vertebra (PU1+U1; as one). While six elements
are associated with the complex vertebra in Cathorops species
(Higuchi, 1982), the number ranges among ariid members
from four to seven (Marceniuk, 2003) and hence were not
included in the counts (i.e., not examined). The term ‘‘rarely’’
in the diagnoses and the dichotomous key refers to uncom-
mon conditions observed in one or two specimens.
A principal component analysis (PCA), obtained with
morphometric and meristic characters, was used to differ-
entiate Cathorops mapale from the new species. The PCA was
performed on the co-variance matrix of the log-transformed
measurements. The values that were constant or appeared to
vary at random, depending on size or sex, were excluded
from the PCA. Thus, PCA results presented here are based on
following variables: gill rakers on first arch (GRFA), gill
rakers on second arch (GRSA), lateral mental barbel length
(LMBL), maxillary barbel length (MBL), mesial mental barbel
length (MMBL), orbital diameter (OD), supraoccipital pro-
cess length (SPL), and width of the supraoccipital process at
the posterior end (WSPPE). Institutional abbreviations are as
listed at http://www.asih.org/node/204.
Mitochondrial cytochrome band ATP synthase subunits 8
and 6 sequences for the new species are available from
GenBank, under accession numbers specified in the material
list (data from Betancur-R. et al., 2007; Betancur-R., 2010).
Cathorops nuchalis (Gu¨nther, 1864)
Orinoco Sea Catfish
Bagre cuinche de rı´o (Spanish)
Figure 1
Arius nuchalis Gu¨nther, 1864:171, figs. Type locality: British
Guiana. Syntypes: BMNH 1975.1.17.3–5 (3), BMNH
2005.5.17.1–2 (2).–Burgess, 1989:169.–Marceniuk and
Ferraris, 2003:450 (as species inquirendae).–Ferraris,
2007:41 (as species inquirendae).–Marceniuk and Me-
nezes, 2007:46 (as species inquirendae).
Arius laticeps Gu¨nther, 1864:171, figs. Type locality: Trinidad
Island and British Guiana. Syntypes: BMNH 1863.6.18.8
(1) Trinidad, 1976.2.25.1 (1) British Guiana.–Burgess,
1989:169.–Marceniuk and Ferraris, 2003:450 (as species
inquirendae).–Ferraris, 2007:41 (as species inquirendae).–
Marceniuk and Menezes, 2007:46 (as species inquirendae).
Tachisurus nuchalis (Gu¨nther).—Eigenmann and Eigenmann,
1888:145 (only name).–Eigenmann and Eigenmann,
1890:86 (British Guiana).
Tachysurus nuchalis (Gu¨nther).Jordan and Evermann,
1896–1898:131, 2782 (British Guiana).
Arius spixii (non Agassiz).—Eigenmann, 1912:145 (British
Guiana); Gosline, 1945:8 (Venezuela, in part).–Lowe-
McConnell, 1962:693 (British Guiana).
Cathorops spixii (non Agassiz).—Schultz, 1944:184 (Venezuela,
Maracaibo basin).–Etchevers, 1978:381–385 (Margarita is-
land).–Taylor and Menezes, 1978 (Belize to Brazil, in part).
–Cervigo´ n, 1991:143–145 (Venezuela, in part).–Cervigo´n,
1992:260 (northern South America, in part).–Gonza´lez-
Bencomo et al., 1997:158 (Venezuela, Lago de Maracaibo).–
Aguilera, 1998:47 (Venezuela, in part).–Marı´n, 2000:68
(Venezuela, in part).–Acero P., 2002:849 (Colombia to Rio
de Janeiro, in part).
Diagnosis.—Cathorops nuchalis can be differentiated from its
subcongeners from the Western Atlantic as follows: from C.
agassizii, from French Guiana to Brazil, by orbital diameter
2.4–3.0 in width of cephalic shield at epioccipital area (vs.
1.6–2.4, rarely 2.4), and wider supraoccipital process at
posterior end (3.0–4.1%SL, rarely 2.9%SL vs. 2.0–2.9%SL;
Fig. 1. Body in lateral view and head in dorsal view. (A) Arius nuchalis, syntype, BMNH 1975.1.17.1-2, 68.5 mm SL, and (B) A. laticeps, syntype,
BMNH 1976.2.25.1, male, 110.5 mm SL.
78 Copeia 2012, No. 1
Fig. 2A); from C. aguadulce, from Mexico, by lacking fleshy
papillae on lateral and mesial surfaces of first and second gill
arches (vs. with fleshy papillae intercalated with the gill
rakers), 18–23 gill rakers on first arch (vs. 14–16), shorter
snout (5.4–8.9%SL vs. 9.3–11.6%SL), and wider supraoc-
cipital process at posterior end (3.0–4.1%SL vs. 2.5–2.6%
SL); from C. arenatus, from Venezuela to Brazil, by possessing
silvery or whitish coloration on flanks and abdominal
region in life (vs. with yellow coloration), 41–42 free
vertebrae (vs. 39–40; Table 1), and orbital diameter 1.1–1.8
in caudal peduncle height (vs. 1.9–3.0; Fig. 2B); from C.
belizensis, from Belize, by its dorsal-fin spine shorter or as
long as pectoral-fin spine (vs. dorsal-fin spine longer than
pectoral-fin spine), and dorsal-fin spine length 2.1–2.5 in
distance from snout to pelvic fin (vs. 2.8–3.4); from C.
higuchii, from Honduras to Panama, by its densely granu-
lated cephalic shield (vs. with very few granules), and
supraoccipital process length 1.5–2.4 in maxillary barbel
length (vs. 2.5–4.5; Fig. 3A); from C. kailolae, from Mexico
and Guatemala, by lacking fleshy papillae on lateral and
mesial surfaces of first and second gill arches (vs. with fleshy
papillae intercalated with the gill rakers), 18–23 gill rakers
on first arch (vs. 14–16), wider cephalic shield at epioccipital
area (12.4–13.4%SL vs. 11.2–12.3%SL), and wider supraoc-
cipital process at posterior end (3.0–4.1%SL vs. 1.9–2.6%
SL); from C. mapale, from Colombia, by having a wider
cephalic shield at epioccipital area (12.5–13.4%SL, rarely
12.4%SL, vs. 11.5–12.4%SL, rarely 12.5%SL), and caudal
peduncle height 1.7–2.5 (rarely 1.6) in supraoccipital process
length (vs. 1.2–1.6); from C. melanopus, from Guatemala, by
its pale pigmentation on pelvic and pectoral fins (vs. with
intense black pigmentation), and short and inconspicuous
serrations on posterior margin of pectoral-fin spine (vs. with
marked long serrations); from C. spixii, from Brazil, by its
silvery or whitish coloration on flanks and abdominal
region in life (vs. with yellow coloration), wider supraoccip-
ital process at posterior end (3.0–4.1%SL, rarely 2.9%SL, vs.
2.0–2.9%SL; Fig. 2A), and supraoccipital process length 1.5–
2.4 in maxillary barbel length (vs. 2.5–4.5; Fig. 3A); from C.
wayuu, from Colombia to Venezuela, by having snout length
1.6–2.4 in supraoccipital process length (vs. 1.0–1.4), orbital
diameter 1.1–1.7 in snout length (vs. 1.8–2.5, rarely 1.7),
Table 1. Meristic Frequencies of Vertebrae Free from the Ventral
Superficial Ossification for Species of Cathorops from Eastern South
America. Bolded numbers indicate counts for primary types.
39 40 41 42 43
Cathorops agassizii 13
Cathorops arenatus 39
Cathorops mapale 42
Arius nigricans 1
Cathorops nuchalis 52
Cathorops spixii 31
Arius variolosus 1
Cathorops wayuu 1331
Fig. 2. Plot of width of the supraoccipital process at the posterior end
versus standard length for Cathorops agassizii,C. nuchalis, and C. spixii
(A), and orbital diameter versus caudal peduncle height for Cathorops
nuchalis and C. arenatus (B).
Fig. 3. Plot of supraoccipital process length versus maxillary barbel
length for C. nuchalis and C. spixii (A), and supraoccipital process length
versus standard length for Cathorops arenatus,C. nuchalis, and C.
wayuu (B).
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 79
longer supraoccipital process (11.7–18.9%SL vs. 9.1–11.5%
SL; Fig. 3B), and supraoccipital process length 1.0–1.5 in
width of cephalic shield at lateral ethmoid area (vs. 0.6–0.9).
Species of Cathorops nuchalis may be distinguished from its
subcongeners from the eastern Pacific as follows: from C.
fuerthii, from Costa Rica to Panama, by its osseous portion of
dorsomedial groove of neurocranium conspicuous and deep,
with straight margins tapering posteriorly (vs. inconspicu-
ous and remarkably shallow, with irregular margins parallel
along its entire extension), dorsal-fin spine thicker or as
thick as pectoral-fin spine (vs. remarkably thinner), and
narrow caudal-fin lobes, pointed posteriorly (vs. wide and
rounded posteriorly); from C. hypophthalmus, from Panama,
by having 18–23 gill rakers on first arch (vs. 37–40), 16–19
gill rakers on second arch (vs. 37–40), and narrower mouth
(8.5–12.1%SL vs. 13.1–13.3%SL); from C. liropus, from
Mexico, by width of supraoccipital process at posterior end
1.2–1.7 in orbital diameter (vs. 1.8–2.0); from C. raredonae,
from Mexico to El Salvador, by its dorsal-fin spine thicker or
as thick as pectoral-fin spine (vs. remarkably thinner),
caudal-fin lobes narrower and pointed posteriorly (vs. wide
and rounded posteriorly), and 18–23 gill rakers on first arch
(vs. 13–15); from C. manglarensis, from Colombia, by having
narrow caudal-fin lobes, pointed posteriorly (vs. wide and
rounded posteriorly), and 20–24 anal-fin rays (vs. 25–27,
rarely 24); from C. multiradiatus, from Panama to Peru, by
having narrow caudal-fin lobes, pointed posteriorly (vs.
wide and rounded posteriorly), and having 20–24 anal-fin
rays (vs. 25–27); from C. steindachneri, from El Salvador to
Panama, by having fleshy portion of dorsomedial groove
of neurocranium inconspicuous, wide and shallow, not
continuous to the level of posterior nares (vs. conspicuous,
narrow and deep, continuous to the level of posterior nares);
from C. taylori, from Guatemala to El Salvador, by its longer
mesial mental barbel (11.8–17.6%SL vs. 8.3–10.3%SL); from
C. tuyra, from Panama, by having thin lips (vs. lips quite
thick), and accessory tooth plates and posterior expansion of
dentary with small molariform teeth (vs. with large
molariform teeth).
Description.—Counts given in Table 1–4, morphometric
data in Table 5. Head relatively short and depressed,
profile elevated posteriorly, straight at frontal and supra-
occipital area. Snout short and rounded transversely.
Anterior nostril round, with fleshy edge, posterior nostril
covered by flap of skin; nostrils quite close to one another
and moderately close to orbit, but not connected by fleshy
furrow. Eye lateral and large; eyes distant to one another.
Three pairs of moderately long teretiform barbels; maxil-
lary barbel usually extending past base of pectoral-fin
spine, lateral mental barbel reaching margin of gill
membrane, and mesial mental barbel not reaching margin
of gill membrane. Osseous bridge formed by lateral
ethmoid and frontal long and slender, evident under skin.
Cephalic shield exposed, rough and granulated, evident
on postorbital region; shield moderately long and large on
lateral ethmoid, frontal, and supracleithrum areas. Ante-
rior portion of dorsomedial groove of neurocranium
fleshy, conspicuous and continuous to level of posterior
nares; posterior portion of groove osseous, deep and
conspicuous, with straight margins tapering posteriorly.
Supraoccipital process funnel-shaped, long and wide on
posterior portion, and profile straight. Nuchal plate
crescent-shaped, moderately short and wide.
Mouth subterminal to terminal, relatively small; lower jaw
arched. Lips thin, lower lip thinner than upper lip.
Vomerine tooth plates absent. One pair of oval shaped
accessory tooth plates, variable in size, closer to one another
anteriorly, with molariform teeth. Premaxilla relatively long
and narrow, with sharp teeth, sometimes fused at symphy-
sis. Dentary separated at midline, with pronounced poste-
rior expansion and sharp teeth on anterior portion,
molariform teeth on posterior portion and some conical
teeth in between. Gill membranes fused, attached to
isthmus. Eighteen to 23 acicular gill rakers on first arch,
16–19 spike-shaped gill rakers on second arch. Mesial
surfaces of all gill arches with developed gill rakers, lateral
and mesial surfaces of first and second gill arches lacking
fleshy papillae intercalated with gill rakers.
Body width slightly greater than body depth at pectoral
girdle area, progressively more compressed from pectoral fin
to caudal peduncle. Forty-one or 42 free vertebrae from
ventral superficial ossification. Lateral line sloping ventrally
on anterior one-third, extending posteriorly to caudal
peduncle, bending abruptly onto dorsal lobe of caudal fin.
Dorsal-fin spine long and thick; anterior margin with
granules on basal two-thirds, distal one-third with short
serrations; posterior margin with serrations along almost its
entire length. Seven dorsal-fin soft rays. Pectoral-fin spine
long, shorter than dorsal-fin spine; anterior margin with
granules on basal two-thirds and distal one-third with short
serrations; posterior margin straight on basal one-fourth,
distal three-fourths with short serrations. Ten pectoral-fin
soft rays. Posterior cleithral process exposed, smooth and
triangular shaped, short and pointed posteriorly. Pelvic fin
deep and long at base, with six rays. Adipose-fin base
relatively short, less than one-half the length of anal-fin
base, anterior origin at level of anterior half of anal fin. Anal
fin deep and relatively short at base, with 20–24 rays and
distal margin slightly concave. Caudal peduncle low. Caudal
fin forked, dorsal and ventral lobes relatively short,
posteriorly slightly pointed; dorsal lobe longer than ventral
lobe.
Coloration.—In life, grayish to dark brown on dorsum,
silvery on flanks and white on abdomen. In alcohol, dark
brown on dorsum, flanks are brownish to silvery and white
on venter; fins brownish. Maxillary barbel dark, mental
barbel lighter (Fig. 1).
Sexual dimorphism.—Sexual dimorphism was observed in
27 females (117–160 mm SL) and 16 males (100–166 mm
SL). Accessory tooth plates are remarkably broader in
femalesthaninmales(1.51.9%SL vs. 0.7–1.2%SL) and
consequently relatively more distant from one another in
males (4.0–7.2%SL vs. 3.0–4.5%SL). Accessory tooth
plates in males containing relatively fewer and smaller
molariform teeth (Fig. 4A) and nuptial males usually
showing accessory tooth plates covered by epithelial
tissue. Posterior expansion of dentary longer and with
more molariform teeth in females (Fig. 4A). Females have
a relatively longer pelvic fin (12.6–16.5%SL vs. 10.9–
13.3%SL) than males.
Distribution and habitat.—Cathorops nuchalis is distributed
from the Lago de Maracaibo, Venezuela, to Guyana (non-
type specimens examined are all from Venezuela). It
inhabits predominantly freshwaters at lower reaches of
80 Copeia 2012, No. 1
rivers; some specimens were collected in estuarine waters
(Fig. 5).
Size.—The largest examined specimen is 244.7 mm SL
(USNM 121208).
Remarks.—Gu¨nther (1864:171) diagnosed Arius nuchalis and
A. laticeps based on the size of accessory tooth patches and
teeth, as well as other non-informative features. According to
Gu¨nther, in A. nuchalis ‘‘the teeth on the palate are coarsely
granular, and form two subtriangular patches of moderate
extent, which, some times, are subcontinuous with their
anterior angles’’, whereas in A. laticeps, ‘‘the teeth on the
palate are granular and form two small, separated patches
anteriorly on the palate.’’ As discussed by several recent
authors, those differences in fact reflect sexual dimorphism in
Cathorops (e.g., Kailola and Bussing, 1995; Acero et al., 2005;
Marceniuk, 2007a, 2007b). While sexual discrimination
based on gonad examination of the types of the two species
was not feasible, the size and shape of accessory tooth patches
and teeth suggest that Arius nuchalis is a female and A. laticeps
is a male (Fig. 4A). Given that Arius nuchalis and A. laticeps
were described simultaneously, a first reviser is required to fix
precedence on the names. To our knowledge, this study
serves as first reviser by validating A. nuchalis over A. laticeps.
Cathorops nuchalis, distributed from Venezuela through
Guyana, is most similar to the C. agassizii from French
Guyana and Brazil. The two species have large eyes, short
maxillary barbels, and are mainly freshwater inhabitants.
Thus, reports of C. nuchalis from northern Brazil by
Boulenger (1897), Miranda-Ribeiro (1911), and Starks
(1913), likely represent C. agassizii, an abundant species in
the area (Marceniuk, 2007b).
Material examined.—BMNH 1863.6.18.8, 1, syntype of Arius
laticeps, 159.7 mm SL, Trinidad; BMNH 1975.1.17.3–5, 2,
syntypes of Arius nuchalis, 100.1–122.3 mm SL, British
Guiana; BMNH 1976.2.25.1, 1, syntype of Arius laticeps,
110.5 mm SL, British Guiana; INVEMAR-PEC uncatalogued,
2 (tissue #CO0622, CO0623), 161–208 mm SL, Venezuela,
Delta Amacuro; LBP 6061, 1, 183 mm SL, Venezuela, Isla
Margarita, mouth of Rı´o Nova Esparta; USNM 121206, 9,
117–160 mm SL, Venezuela, Rı´o de Los Pajaros, 3 km above
Lago de Maracaibo; USNM 121207, 3, 128–135 mm SL,
Venezuela, mouth of Cano De Sagua, 25 km north of
Sinamaica; USNM 121208, 2, 160.3–244.7 mm SL, Vene-
zuela, Rı´o Agua Caliente, 2 to 3 km above Lago de
Maracaibo; USNM 121209, 1, 125.6 mm SL, Venezuela,
Lago de Maracaibo at Yacht Club, Maracaibo; USNM
121210, 4, 110–137 mm SL, Venezuela, Lago de Maracaibo,
1 km off Pueblo Viejo.
Cathorops spixii (Agassiz, 1829)
Madamango Sea Catfish
Bagre cuinche amarilo (Spanish)
Uricia-amarela, bagre amarelo (Portuguese)
Figure 6A
Pimelodus spixii Agassiz in Spix and Agassiz, 1829:19. Type
locality: Equatorial Brazil. Neotype: MZUSP 49345, by
present designation.
Arius nigricans Valenciennes, 1834:no p., pl. 3 (fig. 3). Type
locality: Rı´o de la Plata, Montevideo, Uruguay. Holotype:
MNHN 0000–0176.
Arius spixii (Agassiz).—Cuvier and Valenciennes, 1840:76
(copied).–Fisher, 1917:406 (Brazil, Maceio).–Gosline,
1945:8 (Venezuela, French Guiana, Brazil, in part).
Table 2. Meristic Frequencies of Anal-Fin Rays for Species of Cathorops from Eastern South America. Bolded numbers indicate counts for
primary types.
19 20 21 22 23 24 25
Cathorops agassizii 113 27 9
Cathorops arenatus 323 14 4
Cathorops mapale 128121
Arius nigricans 1
Cathorops nuchalis 14231
Cathorops spixii 925 21
Arius variolosus 1
Cathorops wayuu 253533
Table 3. Meristic Frequencies of Gill Rakers on First Arch for Species of Cathorops from Eastern South America. Bolded numbers indicate counts for
primary types.
15 16 17 18 19 20 21 22 23 24
Cathorops agassizii 520 18 33
Cathorops arenatus 12101515 11
Cathorops mapale 27393
Arius nigricans 1
Cathorops nuchalis 11 4221
Cathorops spixii 21418 2
Arius variolosus 1
Cathorops wayuu 12 75 41
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 81
Table 4. Meristic Frequencies of Gill Rakers on Second Arch for Species of Cathorops from Eastern South America. Bolded numbers indicate counts
for primary types.
15 16 17 18 19 20 21 22
Cathorops agassizii 315 19 7 5
Cathorops arenatus 216 23 22
Cathorops mapale 18 11 1 1
Arius nigricans 1
Cathorops nuchalis 2135
Cathorops spixii 122013 1
Arius variolosus 1
Cathorops wayuu 2576
Table 5. Morphometric Data for Cathorops nuchalis. Standard length is expressed in millimeters, and all other measurements are expressed in
percents of standard length.
Cathorops nuchalis
Syntypes
Arius laticeps
Syntypes
Arius nuchalis n Mean Range
Standard length (mm) 110.5–159.7 100.1–122.3 26 100.1–244.7
Head length 25.4–25.6 24.7–25.5 14 25.9 24.7–28.9
Snout length 7.9–8.2 7.4–8.9 15 7.0 5.4–8.9
Distance between anterior nostrils 4.0–4.3 4.4 13 4.7 4.0–5.5
Distance between posterior nostrils 4.7–4.8 5.5 22 5.4 3.9–7.1
Distance between anterior nostril and orbit 5.0–5.5 6.7 22 6.0 3.5–7.3
Distance between posterior nostril and orbit 8 4.9 4.4–5.2
Orbital diameter 4.8–6.0 5.3–6.5 22 5.2 4.3–6.6
Interorbital distance 11.0–12.6 10.4–11.7 13 12.6 10.4–14.4
Maxillary barbel length 27.7–30.2 33.4–40.7 22 28.1 19.7–40.7
Lateral mental barbel length 20.0–21.7 22.2 13 19.7 17.5–22.8
Mesial mental barbel length 12.9–15.5 17.6 13 13.6 11.8–17.6
Mouth width 10.4–11.0 9.6–12.1 22 10.3 8.5–12.1
Width of cephalic shield at lateral ethmoid area 11.6–12.6 11.9 23 12.6 10.5–14.8
Width of cephalic shield at frontals area 7.0–7.0 8.3 22 8.3 7.0–10.0
Width of cephalic shield at epioccipital area 9 12.8 12.4–13.4
Width of cephalic shield at supracleithrum area 19.6–20.0 21.3 22 18.8 16.5–21.3
Supraoccipital process length 12.6–13.8 16.9 23 14.6 11.7–18.9
Supraoccipital process width at posterior end 3.3–3.8 3.3 20 3.3 2.9–4.1
Nuchal-plate length 8 7.2 6.6–7.9
Nuchal-plate width 17 7.6 6.5–8.8
Body depth 10 18.6 16.6–19.9
Body width 10 20.6 17.1–22.3
Distance from snout to pectoral fin 19.3–19.9 19.7 13 22.9 19.3–26.0
Distance from snout to dorsal fin 37.9–38.2 37.9 13 37.4 34.6–38.7
Distance from snout to pelvic fin 50.2–51.2 52.2 13 51.5 49.9–53.4
Distance from snout to adipose fin 75.0–79.9 74.7 13 76.6 74.5–79.9
Distance from snout to anal fin 63.9–67.1 67.2 13 67.2 63.9–70.4
Caudal-peduncle height 7.6–7.9 7.3 21 7.6 6.6–8.6
Pectoral-fin spine length 19.6–19.9 18.6–20.9 12 19.1 17.2–20.9
Dorsal-fin spine length 21.4–21.8 23.6–24.5 19 21.7 16.9–24.5
Pelvic-fin base length 3.3–3.5 4.9 13 4.2 3.3–4.9
Pelvic-fin height 12.3–14.3 10.9–15.1 14 13.7 10.9–17.1
Adipose-fin base length 10 7.0 6.0–7.8
Adipose-fin height 8 3.9 3.3–4.4
Anal-fin base length 18.6–19.9 17.8 13 18.0 14.5–19.9
Anal-fin height 18.5–19.2 19.7 13 15.5 12.3–19.7
Caudal-fin dorsal lobe length 33.6 33.4 12 33.1 29.8–36.1
Caudal-fin ventral lobe length 29.9–30.6 30.0 13 29.7 25.0–32.2
82 Copeia 2012, No. 1
Tachisurus spixii (Agassiz).—Eigenmann and Eigenmann,
1888:146 (Para´ , Maranha˜o, Bahia, Rio de Janeiro, Sa˜o
Paulo, in part).–Eigenmann and Eigenmann, 1890:88
(Para´ , Maranha˜o, Bahia, Rio de Janeiro, Sa˜o Paulo, in
part).
Tachysurus spixii (Agassiz).—Jordan and Evermann, 1896–
1898:131, 2783 (French Guiana to Brazil, Para´ to Santos,
in part).–Miranda-Ribeiro, 1911:337, 339 (Suriname to
Santos, in part).–Fowler, 1942:140 (Brazil, Para´ to Santos,
in part).–Fowler, 1951:449 (Brazil, in part).–Tommasi,
1965:9 (Brazil, in part).
Cathorops spixii (Agassiz).—Taylor and Menezes, 1978 (Belize
to Brazil, in part).–Acero P., 2002:849 (Colombia to Rio de
Janeiro, in part).–Marceniuk and Ferraris, 2003:449 (Ve-
nezuela to Brazil, in part).–Marceniuk, 2005:97 (French
Guiana to Parana´ , in part).–Ferraris, 2007:40 (Caribbean
to Brazil, in part).–Marceniuk and Menezes, 2007:45
(Brazil, Para´ to Rio de Janeiro).
Diagnosis.—Cathorops spixii is distinguished from its sub-
congeners from the Western Atlantic (excluding C. arenatus)
by its yellowish coloration on flanks and abdominal region
in life (vs. silvery or whitish coloration). The species can be
further differentiated as follows: from C. agassizii, from
French Guiana to Brazil, by having the maxillary barbel
reaching or surpassing base of pectoral-fin spine (vs. not
reaching base of pectoral-fin spine), orbital diameter 5.7–
10.0 in maxillary barbel length (vs. 3.2–5.2; Fig. 7A), and
orbital diameter 4.1–7.4 in external mental barbel length
(vs. 1.9–3.8); from C. aguadulce, from Mexico, by lacking
fleshy papillae on lateral and mesial surfaces of first and
second gill arches (vs. with fleshy papillae intercalated with
the gill rakers), and 17–20 gill rakers on first arch (vs. 14–16);
from C. arenatus, from Venezuela to Brazil, by having 41–42
free vertebrae (vs. 39–40; Table 1), orbital diameter 1.8–2.7
in width of cephalic shield at lateral ethmoid area (vs. 2.8–
4.3, rarely 2.7; Fig. 7B), and orbital diameter 2.8–4.1 in
width of cephalic shield at supracleithrum area (vs. 4.2–6.0,
rarely 4.1); from C. belizensis, from Belize, by its dorsal-fin
spine shorter or as long as pectoral-fin spine (vs. dorsal-fin
spine longer than pectoral-fin spine), and dorsal-fin spine
length 2.0–2.7 in distance from snout to pelvic fin (vs. 2.8–
3.4); from C. higuchii, from Honduras to Panama, by its
densely granulated cephalic shield (vs. with very few
granules), and width of supraoccipital process at posterior
end 1.6–2.5 (rarely 1.5 ) in orbital diameter (vs. 1.0–1.5);
from C. kailolae, from Mexico and Guatemala, by lacking
fleshy papillae on lateral and mesial surfaces of first and
second gill arches (vs. with fleshy papillae intercalated with
the gill rakers), 17–20 gill rakers on first arch (vs. 14–16), and
longer lateral mental barbel (19.5–33.9%SL vs. 15.9–18.6%
SL); from C. mapale, from Colombia, by having 17–19 (rarely
20) gill rakers on first arch (vs. 21–24, rarely 20; Table 3);
from C. melanopus, from Guatemala, by its pale pigmenta-
tion on pelvic and pectoral fins (vs. with intense black
pigmentation), short and inconspicuous serrations on
posterior margin of pectoral-fin spine (vs. with marked long
serrations), and 22–25 anal-fin rays (vs. 19–20); from C.
nuchalis,fromVenezuelatoGuyana,byitsnarrower
Fig. 4. Tooth plates of Cathorops nuchalis (A), C. spixii (B), and C.
wayuu (C). Abbreviations: atp–accessory tooth plates; Den–dentary;
pe–posterior expansion; PMtp–premaxillary tooth plate. Scale bar 5
2 mm.
Fig. 5. Sampling localities of examined material. Filled symbols
represent type specimen localities; open symbols represent non-type
specimen localities. Some symbols indicate more than one locality or
lot of specimens.
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 83
supraoccipital process at posterior end (2.0–2.9%SL vs. 3.0–
4.1%SL, rarely 2.9%SL; Fig. 2A), and supraoccipital process
length in 2.5–3.7 maxillary barbel length (vs. 1.5–2.4;
Fig. 3A); from C. wayuu, from Colombia to Venezuela, by
having orbital diameter 1.8–2.7 in width of cephalic shield
at lateral ethmoid area (vs. 2.8–3.9, rarely 2.7; Fig. 7B).
Cathorops spixii is distinguished from its subcongeners
from the eastern Pacific as follows: from C. fuerthii, from
Costa Rica to Panama, by its osseous portion of dorsomedial
groove of neurocranium conspicuous and deep, with
straight margins tapering posteriorly (vs. inconspicuous
and remarkably shallow, with irregular margins parallel
along its entire extension), dorsal-fin spine thicker or as
thick as pectoral-fin spine (vs. remarkably thinner), and
narrow caudal-fin lobes, pointed posteriorly (vs. wide and
rounded posteriorly); from C. hypophthalmus, from Panama,
by having 17–20 gill rakers on first arch (vs. 37–40), 15–19
gill rakers on second arch (vs. 37–40), and narrower mouth
(7.9–11.7%SL vs. 13.1–13.3%SL); from C. liropus, from
Mexico, by having orbital diameter 5.7–10.0 in maxillary
barbel length (vs. 3.9–5.7, rarely 5.8), and orbital diameter
4.1–7.4 in external mental barbel length (vs. 2.1–4.1, rarely
4.2); from C. raredonae, from Mexico to El Salvador, by its
dorsal-fin spine thicker or as thick as pectoral-fin spine (vs.
remarkably thinner), caudal-fin lobes narrower and pointed
posteriorly (vs. wide and rounded posteriorly), and 17–20
gill rakers on first arch (vs. 13–15); from C. manglarensis,
from Colombia, by having narrow caudal-fin lobes, pointed
posteriorly (vs. wide and rounded posteriorly), and 22–25
anal-fin rays (vs. 25–27, rarely 24); from C. multiradiatus,
from Panama to Peru, by having narrow caudal-fin lobes,
pointed posteriorly (vs. wide and rounded posteriorly), and
having 22–25 anal-fin rays (vs. 25–27); from C. steindachneri,
from El Salvador to Panama, by having fleshy portion of
Fig. 6. Body in lateral view and head in dorsal view. (A) Cathorops spixii, neotype, MZUSP 49345, female, 119.0 mm SL, (B) C. arenatus, non-type,
MZUSP 104013, female, 125.5 mm SL, and (C) profile of osseous portion of dorsomedial groove. Abbreviation: SOp–supraoccipital process.
Fig. 7. Plot of orbital diameter versus maxillary barbel length for
Cathorops agassizii and C. spixii (A), and orbital diameter versus width
of cephalic shield at lateral ethmoid area for Cathorops arenatus,C.
spixii, and C. wayuu (B).
84 Copeia 2012, No. 1
dorsomedial groove of neurocranium inconspicuous, wide
and shallow, not continuous to the level of posterior nares
(vs. conspicuous, narrow and deep, continuous to the level
of posterior nares); from C. taylori, from Guatemala to El
Salvador, by its longer lateral mental barbel (19.5–33.9%SL
vs. 13.2–19.3%SL); from C. tuyra, from Panama, by having
thin lips (vs. lips quite thick), and accessory tooth plates and
posterior expansion of dentary with small molariform teeth
(vs. with large molariform teeth).
Description.—Counts given in Table 1–4, morphometric data
in Table 6. Head relatively short and depressed, profile
elevated posteriorly, slightly convex at frontal and supraoc-
cipital area. Snout short and rounded transversely. Anterior
nostril round, with fleshy edge, posterior nostril covered by
flap of skin; nostrils quite close to one another and
moderately close to orbit, but not connected by fleshy
furrow. Eye lateral and moderately large; eyes distant to one
another. Three pairs of long teretiform barbels; maxillary
barbel reaching to middle of pectoral-fin spine, lateral
mental barbel surpassing margin of gill membrane, and
mesial mental barbel reaching margin of gill membrane.
Osseous bridge formed by lateral ethmoid and frontal long
and slender, evident under skin. Cephalic shield exposed,
rough and granulated, evident on postorbital region; shield
relatively short and narrow on lateral ethmoid, frontal, and
supracleithrum areas. Anterior portion of dorsomedial
groove of neurocranium fleshy, conspicuous and continu-
ous to level of posterior nares; posterior portion of groove
osseous, deep and conspicuous, with straight margins
tapering posteriorly, rarely tapering toward the middle from
both anterior and posterior ends (Fig. 6C). Supraoccipital
Table 6. Morphometric Data for Cathorops spixii. Standard length is expressed in millimeters, and all other measurements are expressed in percents
of standard length.
Cathorops spixii
Holotype
Arius nigricans
Neotype
Cathorops spixii n Mean Range
Standard length (mm) 142 119 39 62–226
Head length 23.2 25.5 39 25.6 22.7–29.5
Snout length 7.0 6.6 39 7.1 5.3–8.6
Distance between anterior nostrils 3.8 4.4 39 4.7 3.8–5.6
Distance between posterior nostrils 4.4 5.3 39 5.9 4.2–7.2
Distance between anterior nostril and orbit 6.5 39 6.4 5.4–7.4
Distance between posterior nostril and orbit 3.2 4.4 39 4.3 3.2–5.2
Orbital diameter 4.6 4.4 39 4.7 4.2–5.5
Interorbital distance 10.1 12.8 39 12.7 10.0–13.7
Maxillary barbel length 29.0 37.8 36 36.4 28.3–45.0
Lateral mental barbel length 19.9 26.9 38 25.6 19.5–33.9
Mesial mental barbel length 14.3 16.2 37 16.8 10.2–25.3
Mouth width 8.7 9.2 39 9.9 7.9–11.7
Width of cephalic shield at lateral ethmoid area 11.2 11.4 39 11.6 9.8–12.9
Width of cephalic shield at frontals area 6.7 6.7 39 6.7 5.9–8.1
Width of cephalic shield at epioccipital area 11.5 37 11.5 8.9–13.1
Width of cephalic shield at supracleithrum area 15.9 16.6 39 17.3 15.5–18.9
Supraoccipital process length 9.1 11.9 39 12.3 9.1–15.0
Supraoccipital process width at posterior end 2.3 2.5 38 2.5 2.0–2.9
Nuchal-plate length 6.1 7.2 39 7.0 5.9–8.1
Nuchal-plate width 6.5 7.1 39 6.9 5.9–8.1
Body depth 17.5 38 17.2 15.6–18.8
Body width 20.3 38 20.7 19.4–21.9
Distance from snout to pectoral fin 18.3 21.3 39 21.2 18.1–26.1
Distance from snout to dorsal fin 33.8 34.5 39 35.0 31.5–38.6
Distance from snout to pelvic fin 49.3 50.6 39 49.5 45.0–53.1
Distance from snout to adipose fin 71.1 73.5 39 73.4 67.0–78.9
Distance from snout to anal fin 64.1 63.9 39 65.7 61.8–70.8
Caudal-peduncle height 8.1 9.0 39 8.7 7.2–10.0
Pectoral-fin spine length 15.5 19.7 39 18.7 14.7–21.2
Dorsal-fin spine length 17.9 22.7 30 21.1 17.7–25.1
Pelvic-fin base length 4.0 4.2 39 4.1 3.1–5.0
Pelvic-fin height 13.0 13.5 37 13.7 9.8–18.0
Adipose-fin base length 6.4 38 7.5 4.5–13.0
Adipose-fin height 4.5 38 4.2 3.0–5.6
Anal-fin base length 17.4 18.2 39 19.6 16.7–22.4
Anal-fin height 14.6 15.9 36 14.7 11.8–17.5
Caudal-fin dorsal lobe length 33.6 26 34.2 27.5–39.0
Caudal-fin ventral lobe length 29.7 35 31.7 24.9–35.7
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 85
process funnel-shaped, moderately long and narrow on
posterior portion, straight in lateral view, rarely convex
(Fig. 6A). Nuchal plate crescent-shaped, moderately long
and narrow.
Mouth subterminal to terminal, relatively small; lower
jaw arched. Lips thin, lower lip thinner than upper lip.
Vomerine tooth plates absent. One pair of oval shaped
accessory tooth plates, variable in size, closer to one another
anteriorly, with molariform teeth. Premaxilla relatively long
and narrow, with sharp teeth, sometimes fused at symphy-
sis. Dentary separated at midline, with pronounced poste-
rior expansion and sharp teeth on anterior portion,
molariform teeth on posterior portion and some conical
teeth in between. Gill membranes fused, attached to
isthmus. Seventeen to 20 acicular gill rakers on first arch,
15–19 spike-shaped gill rakers on second arch. Mesial
surfaces of all gill arches with developed gill rakers, lateral
and mesial surfaces of first and second gill arches lacking
fleshy papillae intercalated with gill rakers.
Body width greater than body depth at pectoral girdle
area, progressively more compressed from pectoral fin to
caudal peduncle. Forty-one or 42 free vertebrae from ventral
superficial ossification. Lateral line sloping ventrally on
anterior one-third, extending posteriorly to caudal pedun-
cle, bending abruptly onto dorsal lobe of caudal fin. Dorsal-
fin spine long and thick; anterior margin with granules on
basal two-thirds, distal one-third with short serrations;
posterior margin with short serrations along almost its
entire length. Seven dorsal-fin soft rays. Pectoral-fin spine
relatively short, shorter than dorsal-fin spine; anterior
margin with granules on basal two-thirds and distal one-
third with short serrations; posterior margin straight on
basal one-fourth, distal three-fourths with short serrations.
Ten or 11 pectoral-fin soft rays. Posterior cleithral process
exposed, smooth and triangular shaped, short and pointed
posteriorly. Pelvic fin deep and long at base, with six rays.
Adipose-fin base relatively short, less than one-half the
length of anal-fin base, anterior origin at level of anterior
half of anal fin. Anal fin moderately deep and long at base,
with 19–25 rays and distal margin slightly concave. Caudal
peduncle high. Caudal fin forked, dorsal and ventral
relatively long, posteriorly pointed; dorsal lobe slightly
longer than ventral lobe.
Coloration.—In life, gray on dorsum, yellow on flanks and
abdomen. In alcohol, gray on dorsum and flanks, venter
region some times yellow; anal, caudal, and pelvic dark on
distal portion, dorsal and adipose fin dusky. Maxillary and
mental barbel blackish (Fig. 6A).
Sexual dimorphism.—Sexual dimorphism was observed in 22
females (80–226 mm SL) and 13 males (64–166 mm SL).
Accessory tooth plates in males containing relatively fewer
and smaller molariform teeth (Fig. 4B) and nuptial males
usually showing accessory tooth plates covered by epithe-
lial tissue. Posterior expansion of dentary longer and with
more molariform teeth in females (Fig. 4B). Females have a
relatively longer pelvic fin (13.0–18.0%SL vs. 11.3–
13.9%SL).
Distribution and habitat.—Cathorops spixii is distributed from
the Amazon River delta (State of Para´) through Rio de
Janeiro (Fig. 5). Inhabits mostly estuarine areas, but can
occur in lower reaches of rivers as well.
Size.—The largest examined specimen is 226 mm SL (MNRJ
825).
Remarks.—The type-specimens of Pimelodus spixii were
collected sometime between 1817 and 1820, when the
botanist Carl von Martius and the zoologist Johann Baptiste
von Spix carried out a collecting expedition in Brazil. The
description of the species was initiated by Spix, but not
concluded due to Spix’s death in 1826. Louis Agassiz was
invited by Martius to complete Spix’s manuscript on
Brazilian Fishes, which was published in 1829 and 1831.
Within the original manuscript, Spix had included a
description and illustration for a species bearing the name
Pimelodus albidus. Agassiz noted the prior use of P. albidus by
Lesueur and assigned the name Pimelodus spixii to the
species described by Spix. The two available names,
published simultaneously as Pimelodus albidus Spix and
Pimelodus spixii Agassiz, were based on at least two
specimens measuring 177.8 mm and 203.2 mm TL from
equatorial Brazil, between Marajo´ Bay (Para´ state) and Sa˜o
Marcos Bay (Maranha˜o state).
Spix’s name is preoccupied by Pimelodus albidus Lesueur,
1819 (p. 148) and considered a junior primary homonym,
and therefore invalid according to articles 11, 12, and 16 of
the International Code of Zoological Nomenclature (ICZN),
as pointed out by Whitehead and Myers (1971), leaving
Pimelodus spixii as the valid name for the species. The
destruction of the type-specimens deposited in the Zoolo-
gische Staatssammlung Mu
¨nchen during World War II
(Terofal, 1983; Kottelat, 1988; Neumann, 2006), makes it
difficult to access the identity of the nominal species. As
mentioned in the present study, three species of Cathorops
are found in equatorial Brazil, and the name P. spixii might
therefore be applied to any one of those species. The length
of the maxillary barbel, described in Spix’s account of
Pimelodus albidus as ‘‘cirrhi 6, maxillares longiores usque ad
pinnarum pectoralium apicem producti’’, indicates the speci-
mens studied could not be referred to C. agassizii, which is
characterized by a short maxillary barbel that never reaches
the base of the pectoral spine (Marceniuk, 2007b). Never-
theless, even the combined descriptions of Spix (P. albidus)
and Agassiz (P. spixii) are not sufficient to unambiguously
assign the name to either of the two remaining species, one
of which, C. arenatus, is a species well defined in Marceniuk
(2007b).
Taking into account the complex nomenclatural history
of C. spixii, in order to clarify the taxonomic status and for
the sake of nomenclatural stability of this important
nominal species, we decide to define it objectively with
the designation of a neotype in accordance to Article 75
and qualifying conditions (Article 75.3.) of the Interna-
tional Code of Zoological Nomenclature (ICZN, 1999). The
neotype chosen is a female, 119 mm SL, from Marajo´Bay,
Para´ state (MZUSP 49345) within the Brazilian equatorial
region, the type-locality referred originally by Spix (Brasilia
aequatoriali). In fact, it could have been chosen from any
estuarine or coastal area between the bays of Marajo´(Para´
state) and Sa˜o Marcos (Maranha˜o state). The decision to
chose a specimen from Marajo´ Bay was made following the
detailed itinerary description included in Spix and Martius
(1938) diary (‘‘Viagem ao Brasil’’, vol. 2, chapter five,
Portuguese version), where it is mentioned that they
arrived at the equatorial region (Marajo´ bay, immediately
next to Vigia city) on 25 July 1819. Examination of the
86 Copeia 2012, No. 1
material collected in August of 1865, by L. Agassiz in the
Brazilian equatorial region, shows that C. spixii is much
more common in Para´ state (six specimens MCZ 7683, MCZ
7709, MCZ 7663), whereas only two specimens of C.
arenatus were collected from Maranha˜o state in Brazil
(MCZ 7685). Additionally our observations indicate that
C. arenatus is predominantly found along the Brazilian
marine coastal region and more rarely in estuarine waters,
where the type-specimens of Cathorops spixii were probably
caught.
We recognize Arius nigricans (Fig. 8A; Table 6) as junior
synonym of C. spixii based on examination of the holotype.
The type-locality of that species, Rı´o de la Plata, Montevideo,
Uruguay, is probably wrong considering the southern extent
of the distribution of any species of Cathorops species is
Parana´ state, southeastern Brazilian region (Marceniuk and
Menezes, 2007). The uncertainty about the type locality of A.
nigricans prevents us from designating its holotype as the
neotype of C. spixii; that designation would imply a southern
range extension for C. spixii based on no reliable data.
Material examined.—MCZ 7663, 1, 154 mm SL, Brazil, Para´,
Bele´ m; MCZ 7678, 1, 153 mm SL, Brazil, Rio de Janeiro; MCZ
7683, 4, 118–144 mm SL, Brazil, Para´, Bele´m; MCZ 7699, 3,
97–98 mm SL, Brazil, Bahia, Salvador; MCZ 7709, 1, 140 mm
SL, Brazil, Para´ , Bele´m; MNRJ 825, 1, 226 mm SL, Brazil,
Bahia, barra da Areia; MZUSP 104088, 1, 155 mm SL, Brazil,
Bahia, Porto Seguro, fish market; MZUSP 107931, 2, 105–
113 mm SL, Brazil, Para´ , baı´a de Marajo´ ; MZUSP 22683, 1,
125 mm SL, Brazil, Rio de Janeiro, Atafona; MZUSP 22783, 1,
121 mm SL, Brazil, Rio de Janeiro, lagoa Marica; MZUSP
24374, 106–106 mm SL, Brazil, Bahia, Maragogipe, mouth of
Rio Paraguac¸u, near Maragogipe; MZUSP 24487, 3, 95–
112 mm SL, Brazil, Rio de Janeiro, Macae´ , Portinha; MZUSP
37214, 1, 64 mm SL, Brazil, Para´ , Vigia; MZUSP 37215, 1,
88 mm SL, Brazil, Maranha˜o, Sa˜o Luiz, Estreito do Coqueiro,
Paje´ , ilha de Sa˜o Luiz; MZUSP 37236, 8, 127–183 mm SL,
Brazil, Sergipe, Rio Sergipe; MZUSP 48526, 2, 166–181 mm
SL, Brazil, Rio de Janeiro, baı´a de Sepetiba; MZUSP 49345, 1,
119 mm SL, neotype of Pimelodus spixii (by present
designation), Brazil, Para´ , baı´a de Marajo´; MZUSP 49348, 1,
62 mm SL, Brazil, Para´ , Ponta Fina, baı´a de Marajo´; MZUSP
49349, 1, 97 mm SL, Brazil, Paraı´ba, Lucena; MZUSP 49350,
1, 132 mm SL, Brazil, Paraı´ba, Lucena; MZUSP 49351, 1,
123 mm SL, Brazil, Paraı´ba, Lucena; MZUSP 49353, 4, 104–
149 mm SL, Brazil, Rio de Janeiro, Atafona Pontal; MZUSP
49357, 2, 117–119 mm SL, Brazil, Sergipe, Pirambu, Rio
Japaratuba; MZUSP 49363, 1, 109 mm SL, Brazil, Sergipe, Rio
Sergipe; MZUSP 49411, 1, 80 mm SL, Brazil, Para´ , Bele´m,
Coroinha, baı´a de Marajo´ ; MZUSP 72982, 1, 117 mm SL,
Brazil, Maranha
˜o, Sa˜o Luis, Rio Curuca, ilha de Sa˜o Luis;
ZUFES 130023, 3, 137–153 mm SL, Brazil, Espı´rito Santo,
baia de Camburi.
Cathorops wayuu, new species, Betancur-R., Acero P.,
and Marceniuk
Wayuu Sea Catfish
Bagre cuinche (Spanish)
Figure 9
Arius spixii (non Agassiz).—Gosline, 1945:8 (Venezuela, in
part).
Cathorops spixii (non Agassiz).—Taylor and Menezes,
1978:35 (Colombia and Venezuela, in part).–Valdez and
Aguilera, 1987:51 (Venezuela).–Cervigo´ n, 1991:143–145
(Venezuela, in part).–Cervigo´ n, 1992:260 (Septentrional
South America, in part).–Aguilera, 1998:47 (Venezuela, in
part).–Marı´n, 2000:68 (Venezuela, in part).–Acero P.,
2002:849 (Colombia and Venezuela, in part).
Cathorops sp.—Betancur-R. and Acero P., 2005:57 (Colom-
bia).–Betancur-R. et al., 2010 (Colombia and Venezuela).
Holotype.—INVEMAR-PEC 8247, male (tissue #LF267),
229 mm SL, Colombia, La Guajira, Camarones at lagoon,
Fig. 8. Body in lateral view and head in dorsal view. (A) Arius nigricans, holotype, MNHN 0000-0176, 142 mm SL, and (B) A. variolosus, holotype,
MNHN 0000-4169, 142 mm SL.
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 87
purchased from local fisherman, approximate location
11u259430N, 73u049150W, R. Betancur-R. and A. Acero P.,
18 December 2010.
Paratypes.—All purchased from local fishermen unless
otherwise indicated. AUM 42758, 4 (tissue #VEN0506–08),
171–204 mm SL, Venezuela, Nueva Esparta, Isla Margarita,
hook and line, R. Betancur-R. and M. I. Sierra, 12 February
2005, GenBank accessions GQ982433–35, GQ982463–65;
AUM42760,2,(tissue#VEN037), 159–265 mm SL,
Venezuela, Sucre, Golfo de Paria, Irapa, R. Betancur-R. and
M. I. Sierra, 16 February 2005, GenBank accessions
GQ982440, GQ982470; AUM 42846, 4 (tissue #VEN0516–
17), 144–205 mm SL, Venezuela, Sucre, Carupano, R.
Betancur-R. and M. I. Sierra, 14 February 2005, GenBank
accessions GQ982436–37, GQ982466–67; AUM 42848, 2
(tissue #VEN0544–45), 139–218 mm SL, Venezuela, Car-
abobo, Puerto Cabello, R. Betancur-R. and M. I. Sierra, 20
February 2005, GenBank accessions GQ982441–42,
GQ982471–72; AUM 44227, 4, (tissue #VEN0526–27),
225–314 mm SL, Venezuela, Sucre, Golfo de Paria, Guiria,
R. Betancur-R. and M. I. Sierra, 14–15 February 2005,
GenBank accessions GQ982438–39, GQ982468–69; INVE-
MAR-PEC 5734, 2 (tissue #494–495), 230–251 mm SL,
Colombia, La Guajira, NE Riohacha at fish market (nearby
mouth of Rı´o Rancherı´as), R. Betancur-R. et al., 13 December
2003, GenBank accessions DQ990470 (cytochrome b),
DQ990642 (ATP synthase subunits 8 and 6), GQ982432
(cytochrome b), GQ982462 (ATP synthase subunits 8 and 6);
INVEMAR-PEC 5735, 4 (tissue #481, 482, 484, 486), 183–
246 mm SL, Colombia, La Guajira, off Camarones, R.
Betancur-R. et al., 11 December 2003, GenBank accessions
GQ982427–30, GQ982457–60; INVEMAR-PEC 8246, 4
(tissue #487), 180–246 mm SL, Colombia, La Guajira,
Camarones at lagoon, approximate location 11u259430N,
73u049150W, R. Betancur-R. et al., 12 December 2003,
GenBank accessions GQ982431, GQ982461; INVEMAR-PEC
8248, 6, 194–249 mm SL, collected with holotype; INVE-
MAR-PEC 8249, 8 (skeleton preparations), 229–242 mm SL,
Colombia, La Guajira, NE Riohacha at fish market (nearby
mouth of Rı´o Rancherı´as), R. Betancur-R. and A. Acero P., 17
December 2010.
Diagnosis.—Cathorops wayuu differs from its subcongeners
from the Western Atlantic as follows: from C. agassizii, from
French Guiana to Brazil, by having the maxillary barbel
reaching or surpassing base of pectoral spine (vs. not
reaching base of pectoral spine), orbital diameter 1.8–2.5
(rarely 1.7) in snout length (vs. 0.7–1.7), orbital diameter
5.2–7.8 in maxillary barbel length (vs. 3.2–5.1, rarely 5.2),
and orbital diameter 4.0–5.7 in external mental barbel
length (vs. 1.9–3.8); from C. aguadulce, from Mexico, by
lacking fleshy papillae on lateral and mesial surfaces of first
and second gill arches (vs. with fleshy papillae intercalated
with the gill rakers), 17–20 gill rakers on first arch (vs. 14–
16), and shorter supraoccipital process (9.1–11.5%SL vs.
11.6–16.3%SL, rarely 11.5%SL); from C. arenatus, from
Venezuela to Brazil, by its silvery or whitish coloration on
flanks and abdominal region in life (vs. with yellowish
coloration); 41–43 (rarely 40) free vertebrae (vs. 39–40;
Table 1), and shorter supraoccipital process (9.1–11.5%SL
vs. 11.6–18.9%SL, rarely 11.5%SL; Fig. 3B); from C.
belizensis, from Belize, by its dorsal-fin spine shorter or as
long as pectoral-fin spine (vs. dorsal-fin spine longer than
pectoral-fin spine), and dorsal-fin spine length 1.05–1.3 in
pectoral-fin spine length (vs. 0.97–1.0); from C. higuchii,
from Honduras to Panama, by its densely granulated
cephalic shield (vs. with very few granules), and orbital
diameter 5.2–7.8 in maxillary barbel length (vs. 7.9–12.4,
rarely 7.8); from C. kailolae, from Mexico and Guatemala, by
lacking fleshy papillae on lateral and mesial surfaces of first
and second gill arches (vs. with fleshy papillae intercalated
with the gill rakers), and 17–20 gill rakers on first arch (vs.
14–16); from C. mapale, from Colombia, by possessing 16–20
(rarely 21) gill rakers on first arch (vs. 21–24, rarely 20;
Table 3), 16–19 gill rakers on second arch (vs. 19–22, rarely
18; Table 4), and eight fixed mitochondrial substitutions
(see Table 7); from C. melanopus, from Guatemala, by its pale
Table 7. Summary of Eight Fixed Substitutions Differentiating Cathorops mapale (17 specimens) and C. wayuu (17 specimens) at the Mitochondrial
Level (Data from Betancur-R. et al., 2011). Numbers above character states are nucleotide positions as compared to the mitochondrial genome of
Ictalurus punctatus (GenBank accession number NC003489).
ATP synthase subunits 8 and 6 (842 bp) Cytochrome b(1095 bp)
9079 9302 9305 9416 9653 15479 15567 16250
Cathorops mapale AGGT T AGC
Cathorops wayuu GAACCCA T
Fig. 9. Body in lateral view and head in dorsal view. (A) Cathorops wayuu, holotype, INV-PEC 8247, 232.8 mm SL (fresh specimen).
88 Copeia 2012, No. 1
pigmentation on pelvic and pectoral fins (vs. with intense
black pigmentation), short and inconspicuous serrations on
posterior margin of pectoral-fin spine (vs. with marked long
serrations), and lower caudal-peduncle (6.2–8.3%SL vs. 8.7–
9.0%SL); from C. nuchalis, from Venezuela to Guyana, by
the snout length 1.0–1.4 in supraoccipital process length (vs.
1.6–2.4), orbital diameter 1.8–2.5 (rarely 1.7) in snout length
(vs. 0.7–1.7), shorter supraoccipital process (9.1–11.5%SL vs.
11.7–18.9%SL; Fig. 3B), supraoccipital process length 0.7–
0.9 in width of cephalic shield at lateral ethmoid area (vs.
1.0–1.5); from C. spixii, from Brazil, by having silvery or
whitish coloration on flanks and abdominal region in life
(vs. with yellowish coloration); and orbital diameter 2.8–3.9
(rarely 2.7) in width of cephalic shield at lateral ethmoid
area (vs. 1.8–2.7).
The new species may be distinguished from its subcon-
geners from the eastern Pacific as follows: from C. fuerthii,
from Costa Rica to Panama, by its osseous portion of
dorsomedial groove of neurocranium conspicuous and deep,
with straight margins tapering posteriorly (vs. inconspicu-
ous and remarkably shallow, with irregular margins parallel
along its entire extension), dorsal-fin spine thicker or as
thick as pectoral-fin spine (vs. remarkably thinner), and
narrow caudal-fin lobes, pointed posteriorly (vs. wide and
rounded posteriorly); from C. hypophthalmus, from Panama,
by having 18–23 gill rakers on first arch (vs. 37–40), 16–19
gill rakers on second arch (vs. 37–40), and narrower mouth
(8.9–11.7%SL vs. 13.1–13.3%SL); from C. liropus, from
Mexico, by its smaller orbital diameter (3.5–4.9%SL vs. 5.0–
6.4%SL), and wider cephalic shield at supracleithrum area
(16.6–18.2%SL vs. 18.3–19.3%SL); from C. raredonae, from
Mexico to El Salvador, by its dorsal-fin spine thicker or as
thick as pectoral-fin spine (vs. remarkably thinner), caudal-
fin lobes narrower and pointed posteriorly (vs. wide and
rounded posteriorly), and 16–21 gill rakers on first arch (vs.
13–15); from C. manglarensis, from Colombia, by having
narrow caudal-fin lobes, pointed posteriorly (vs. wide and
rounded posteriorly), and 19–24 anal-fin rays (vs. 25–27,
rarely 24); from C. multiradiatus, from Panama to Peru, by
having narrow caudal-fin lobes, pointed posteriorly (vs.
wide and rounded posteriorly), and having 19–24 anal-fin
rays (vs. 25–27); from C. steindachneri, from El Salvador to
Panama, by having fleshy portion of dorsomedial groove of
neurocranium inconspicuous, wide and shallow, not con-
tinuous to the level of posterior nares (vs. conspicuous,
narrow and deep, continuous to the level of posterior nares),
and shorter supraoccipital process (9.1–11.5%SL vs. 11.7–
15.9%SL); from C. taylori, from Guatemala to El Salvador, by
its longer mesial mental barbel (10.8–16.7%SL vs. 8.3–
10.3%SL); from C. tuyra, from Panama, by having thin lips
(vs. lips quite thick), and accessory tooth plates and
posterior expansion of dentary with small molariform teeth
(vs. with large molariform teeth).
Description.—Counts given in Table 1–4, morphometric data
in Table 8. Head relatively long and depressed, profile
slightly elevated posteriorly, straight or concave at frontal
and supraoccipital area. Snout long and rounded or slightly
blunt transversely. Anterior nostril round, with fleshy edge,
posterior nostril covered by flap of skin; nostrils quite close
to one another and moderately closer to orbit, not
connected by fleshy furrow. Eye lateral and small; eyes
distant to one another. Three pairs of relatively short
teretiform barbels; maxillary barbel reaching/surpassing
(smaller individuals) or falling short (larger individuals) of
pectoral-fin spine; lateral mental barbel reaching/surpassing
(smaller individuals) or falling short (larger individuals) of
margin of gill membrane, and mesial mental barbel reaching
margin of gill membrane only in small individuals. Osseous
bridge formed by lateral ethmoid and frontal long and
slender, evident under skin. Cephalic shield exposed, rough
and granulated, evident on postorbital region; shield
moderately long and large on lateral ethmoid, moderately
narrow on frontal and supracleithrum areas. Anterior
portion of dorsomedial groove of neurocranium fleshy,
conspicuous and continuous to level of posterior nares;
posterior portion of groove osseous, deep and conspicuous,
with straight margins tapering posteriorly. Supraoccipital
process funnel-shaped, short and moderately wide on
posterior portion, and profile straight or slightly convex;
nuchal plate crescent-shaped, short and relatively narrow.
Mouth subterminal to terminal, relatively small; lower jaw
arched. Lips thin, lower lip thinner than upper lip.
Vomerine tooth plates absent. One pair of oval shaped
accessory tooth plates, variable in size, closer to one another
anteriorly, with molariform teeth. Premaxilla relatively long
and narrow, with sharp teeth, sometimes fused at symphy-
sis. Dentary separated at midline, with pronounced poste-
rior expansion and sharp teeth on anterior portion,
molariform teeth on posterior portion and some conical
teeth in between. Gill membranes fused, attached to
isthmus. Sixteen to 21 acicular gill rakers on first arch, 16–
19 spike-shaped gill rakers on second arch. Mesial surfaces of
all gill arches with developed gill rakers, lateral and mesial
surfaces of first and second gill arches lacking fleshy papillae
intercalated with gill rakers.
Body width greater than body depth at pectoral girdle area,
progressively more compressed from pectoral fin to caudal
peduncle. Forty to 43 free vertebrae from ventral superficial
ossification. Lateral line sloping ventrally on anterior one-
third, extending posteriorly to caudal peduncle, bending
abruptly onto dorsal lobe of caudal fin. Dorsal-fin spine
relatively short and thick; anterior margin with granules on
basal two-thirds, distal one-third with short serrations;
posterior margin with short serrations along almost its entire
length. Seven dorsal-fin soft rays. Pectoral-fin spine relatively
short, shorter than dorsal-fin spine; anterior margin with
granules or short serrations along entire length; posterior
margin straight on basal one-fourth, distal three-fourths with
short serrations. Ten pectoral-fin soft rays. Posterior cleithral
process exposed, smooth and triangular shaped, short and
pointed posteriorly. Pelvic fin deep and long at base, with six
rays. Adipose-fin base relatively short, less than one-half the
length of anal-fin base, anterior origin at level of anterior half
of anal fin. Anal fin moderately deep and relatively short at
base, with 19–24 rays and distal margin slightly concave.
Caudal peduncle low. Caudal fin forked, dorsal and ventral
lobes moderately long, posteriorly pointed or slightly
rounded; dorsal lobe longer than ventral lobe.
Coloration.—In life, gray on dorsum, silvery white on sides
and abdomen; fins dusky often with red tinges; maxillary
barbel dark, mental barbel lighter; adipose and other fins
darker than venter (Fig. 9). In alcohol, brown on dorsal and
lateral portions, light beige to white ventrally.
Sexual dimorphism.—Sexual dimorphism was observed in 12
females (159–214 mm SL) and six males (191–225 mm SL).
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 89
Males with anterior portion of body relatively longer than in
females, as evidenced by distance from tip of snout to
pectoral fin (22.0–27.4%SL vs. 19.7–24.8%SL). Accessory
tooth plates in males containing relatively fewer and smaller
molariform teeth (Fig. 4C) and nuptial males usually
showing accessory tooth plates covered by epithelial tissue.
Posterior expansion of dentary longer and with more
molariform teeth in females. Females have longer pelvic
fin (14.4–18.1%SL vs. 12.9–14.6%SL) than males.
Distribution and habitat.—Cathorops wayuu occurs northeast-
ward to the Santa Marta Massif in northern Colombia, from
La Guajira to Golfo de Paria in Venezuela. Inhabits brackish
and coastal marine waters (Fig. 5).
Size.—The largest examined specimen is 314 mm SL (AUM
44227).
Remarks.—The Cathorops mapale species group comprise the
sister species Cathorops mapale from Colombia and C. wayuu
from Colombia and Venezuela (Betancur-R. and Acero P.,
2005; Betancur-R. et al., 2010). Meristic and morphometric
analyses preclude unambiguous differentiation of the two
species (Betancur-R. et al., 2010). Although the number of
gill rakers on first and second arches (see Diagnosis) and the
bivariate plot maxillary barbel/posterior internarial distance
(4.8–7.9, mean 6.1 6SD 0.8 in C. mapale; 3.6–6.0, mean 4.4
6SD 0.8 in Cathorops wayuu) are the best morphological
discriminators, overlap exists at the univariate/bivariate
level. In addition, while a principal component analysis
on six morphometric and two meristic variables shows two
major multivariate clusters, two specimens of C. wayuu
overlap with C. mapale (Fig. 10). As indicated by analyses of
mitochondrial sequences (ATP synthase 8/6 and cytochrome
b) the two species are, however, unambiguously differenti-
ated at the molecular level, separated by eight fixed
substitutions (Betancur-R. et al., 2010; Table 7). Further-
more, they are reciprocally monophyletic and allopatric,
segregated by the Santa Marta Massif in northern Colombia
(see Betancur-R. et al. [2010] for a discussion on the
biogeography).
Whereas mitochondrial distances estimated from protein-
coding genes range from 1.5–2.8%among sister-species pairs
in the genus Cathorops (see Betancur-R. et al., 2010:table 3),
divergence between C. mapale and C. wayuu is only 0.7–
1.2%. Furthermore, molecular clock estimates for the split of
Table 8. Morphometric Data for Cathorops wayuu. Standard length is expressed in millimeters, and all other measurements are expressed in
percents of standard length.
Cathorops wayuu
Holotype nMean Range
Standard length (mm) 232.8 22 144–314
Head length 28.1 22 27.1 24.2–31.3
Snout length 8.6 22 8.5 7.1–10.5
Distance between anterior nostrils 4.7 22 4.7 4.2–5.2
Distance between posterior nostrils 6.6 22 6.1 5.4–7.2
Distance between posterior nostril and orbit 4.5 22 4.7 4.1–5.4
Orbital diameter 4.4 22 4.1 3.5–4.9
Interorbital distance 12.0 22 13.1 11.6–14.3
Maxillary barbel length 25.3 22 27.0 21.8–33.4
Lateral mental barbel length 17.8 22 19.4 15.9–24.6
Mesial mental barbel length 12.5 22 13.5 10.8–16.7
Mouth width 10.5 22 10.3 8.9–11.7
Width of cephalic shield at lateral ethmoid area 13.0 22 13.2 12.2–14.4
Width of cephalic shield at frontals area 5.8 22 6.9 5.8–7.9
Width of cephalic shield at supracleithrum area 17.1 21 17.3 16.6–18.2
Supraoccipital process length 10.4 22 10.3 9.1–11.5
Supraoccipital process width at posterior end 2.8 22 2.9 2.4–3.3
Nuchal-plate width 6.5 22 7.2 6.5–8.2
Body depth 16.8 21 19.1 16.1–21.8
Body width 17.4 22 20.0 17.4–21.8
Distance from snout to pectoral fin 24.4 22 22.5 19.7–27.4
Distance from snout to dorsal fin 38.1 22 37.0 34.4–40.8
Distance from snout to pelvic fin 53.4 22 51.0 48.8–54.3
Distance from snout to adipose fin 77.8 22 75.6 72.0–77.9
Distance from snout to anal fin 69.7 22 67.9 64.6–70.7
Caudal-peduncle height 6.9 22 7.6 6.2–8.3
Pectoral-fin spine length 16.5 13 17.8 16.3–20.0
Dorsal-fin spine length 19.6 12 20.3 16.8–23.2
Pelvic-fin base length 3.8 22 4.1 3.5–4.6
Pelvic-fin height 13.4 22 15.5 12.9–18.1
Adipose-fin base length 6.4 22 6.4 5.0–8.6
Adipose-fin height 7.7 21 9.2 7.5–11.9
Anal-fin base length 16.3 22 16.1 14.3–18.9
90 Copeia 2012, No. 1
the two lineages (,0.9 my; Betancur-R. et al., 2010) are
slightly lower than generally inferred times for allopatric
speciation in fishes (2.3–1.0 my; McCune and Lovejoy,
1998). In light of the above, Betancur-R. et al. (2010)
concluded that C. mapale and C. wayuu represent a case of
incipient speciation. Nevertheless, in a taxonomic frame-
work, we opt to recognize the specific status of both
lineages. This is particularly relevant considering that they
play an important role in artisanal fisheries for coastal
populations along Colombia and Venezuela.
Etymology.—The new species is described after the Wayuu
native American ethnic community from La Guajira penin-
sula (type locality) in northern Colombia and Venezuela.
The specific epithet is a noun in apposition.
Taxonomic status of other species of Cathorops from ESA.—
Here, six of the 12 nominal species from ESA ascribed to
Cathorops are recognized as valid and taxonomic status of
other nominal species is defined (see Table 9). In addition to
the species validated and synonymized above, the identity
of Arius variolosus (Fig. 8B) is clarified as a result of this
study. This nominal species was described from a single
specimen collected in Cayenne, French Guiana, and was
recently listed as species inquirendae by Marceniuk and
Ferraris (2003) and Ferraris (2007). Arius variolosus is
conspecific with C. arenatus (Table 10), as redescribed by
Marceniuk (2007b), based on the following features: 39 free
vertebrae (unique among ESA species of Cathorops, vs. 42–43
in C. agassizii and C. mapale, 41–42 in C. nuchalis and C.
spixii, 41–43 [rarely 40] in C. wayuu [Table 1]), orbital
Fig. 10. Scatterplots of principal components PC1 and PC2, obtained from the analysis of six morphometric and two meristic variables for Cathorops
mapale (M–holotype, m–paratypes) and C. wayuu (W–holotype, w–paratypes). Factor loadings on the first and second principal components.
Table 9. Summary of Nominal Species in Cathorops from ESA and Their Current Status and Country Distribution. Two letter country codes follow ISO-
3166.
Nominal species Type locality Status Distribution
Arius arenatus SR Valid as C. arenatus
a
VE, GY, SR, GF, BR
Arius fissus SR Synonym of C. arenatus
a
Arius laticeps TT, GY Synonym of C. laticeps
b
Arius nigricans ´o de la Plata, UR (likely erroneous) Synonym of C. spixii
b
Arius nuchalis GY Valid as C. nuchalis
b
VE, TT, GY
Arius pleurops Marajo, BR Synonym of C. agassizii
a
Arius puncticulatus Buenos Aires, AR (likely erroneous) Species inquirenda
c,b
Cathorops mapale Santa Marta, CO Valid as C. mapale
b
CO
Cathorops wayuu La Guajira, CO Valid as C. wayuu
b
CO, VE
Pimelodus albidus Equatorial BR Invalid, preoccupied by Lesueur
d
Pimelodus spixii Equatorial BR Valid as C. spixii
b
BR
Tachysurus agassizii Rio Grande do Sul, BR (likely erroneous) Valid as C. agassizii
a
GF, BR
a
Marceniuk (2007)
b
This study
c
Marceniuk and Ferraris (2003)
d
Whitehead and Myers (1971)
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 91
diameter 7.6 in maxillary barbel length (vs. 3.2–5.2 in C.
agassizii), orbital diameter 1.9 in caudal peduncle height (vs.
1.1–1.85 in C. nuchalis; Fig. 2B), orbital diameter 3.2 in
width of cephalic shield at lateral ethmoid area (vs. 1.8–2.7
in C. spixii; Fig. 7B), and shorter supraoccipital process
(11.6%SL vs. 9.1–11.5%SL in C. wayuu; Fig. 3B).
Unfortunately, the taxonomic status of Arius puncticulatus,
described from Buenos Aires and listed as species inquirendae
in Cathorops (Marceniuk and Ferraris, 2003; Ferraris, 2007),
remains uncertain. The single type specimen of A. puncticu-
latus, deposited at the Muse´ um National d’Histoire Naturelle
in Paris, appears to be lost (J. Gregorio, pers. comm.; R.
Betancur-R., pers. obs.). Furthermore, given that Parana´ state
in Brazil represents the southern limit of Cathorops distribu-
tion and hence the type locality is likely erroneous, the
original description by Valenciennes (in Cuvier and Valenci-
ennes, 1840) provides insufficient detail to accurately assign
the name to any of the known ESA species of Cathorops.
The specimens of Cathorops examined from the freshwa-
ters of Rio Doce and Rio Paraı´ba do Sul, in Espı´rito Santo
state, and those collected to the south of Ilha Grande, in Rio
de Janeiro state, are not included in the list of material
examined of C. spixii in the present study. Those specimens
have morphological characteristics that differ from those
observed in C. spixii (as redescribed here), and a more
thorough investigation of the identity of these specimens is
required, based on a broader sampling of individuals and
morphological data. It is herein suggested that the specimens
from those localities be designated as Cathorops cf. spixii.
DISCUSSION
Among the 36 families in the Order Siluriformes, Ferraris
(2007) reported the highest number of species inquirendae
in the Ariidae (56 as of 2007), followed by the Bagridae (33).
Recently, the status of four of the 56 ariid species has been
Table 10. Morphometric Data for Cathorops agassizii,C. arenatus,C. mapale, and holotype of Arius variolosus. Standard length is expressed in
millimeters and all other measurements are expressed in percents of standard length.
Cathorops agassizii Cathorops arenatus Cathorops mapale Arius variolosus
(n=51) (n=67) (n=23) (n=1)
Standard length (mm) 76–219 91–182 118–256 141
Head length 22.8–31.6 22.6–29.2 23.7–29.9 27.7
Snout length 5.0–9.0 5.3–8.3 6.0–8.7 7.8
Distance between anterior nostrils 3.3–5.7 4.2–5.8 3.6–5.1 5.4
Distance between posterior nostrils 4.7–7.5 5.3–7.3 4.2–6.8 6.0
Distance between anterior nostril and orbit 5.2–7.5 5.8–7.6 6.6–7.3
Distance between posterior nostril and orbit 3.3–5.6 3.8–5.3 4.0–5.4 4.3
Orbital diameter 4.7–7.1 3.1–4.5 3.7–5.2 4.4
Interorbital distance 10.6–16.8 12.4–14.7 10.7–14.8 12.8
Maxillary barbel length 18.8–29.0 27.3–39.3 26.9–39.3 33.1
Lateral mental barbel length 11.2–22.1 18.4–36.0 17.5–26.4 22.2
Mesial mental barbel length 3.9–15.6 11.6–25.3 11.7–17.8 14.6
Mouth width 7.6–12.2 8.8–12.7 8.3–10.8 12.7
Width of cephalic shield at lateral ethmoid area 10.8–14.1 11.6–14.3 11.4–14.1 14.1
Width of cephalic shield at frontals area 5.0–7.2 6.4–8.0 5.5–7.4 8.0
Width of cephalic shield at epioccipital area 10.3–12.6 11.1–13.9 11.5–12.5
Width of cephalic shield at supracleithrum area 16.0–18.9 17.1–20.0 15.8–18.6 17.7
Supraoccipital process length 9.2–15.0 11.5–15.0 10.0–12.8 11.7
Supraoccipital process width at posterior end 2.1–2.9 2.0–3.9 2.2–3.3 2.6
Nuchal-plate length 6.0–8.2 5.9–7.7 6.8–7.2
Nuchal-plate width 5.9–7.6 6.5–8.5 6.3–7.8 6.9
Body depth 14.1–20.2 14.4–21.1 17.5–19.4
Body width 18.8–22.3 20.0–24.3 17.3–21.7
Distance from snout to pectoral fin 18.4–28.2 18.6–25.8 19.9–25.5 21.3
Distance from snout to dorsal fin 31.5–39.7 32.8–41.1 33.4–38.9 39.0
Distance from snout to pelvic fin 48.8–55.5 46.9–53.3 46.4–53.0 51.8
Distance from snout to adipose fin 68.9–81.0 66.7–78.9 71.3–76.7 75.9
Distance from snout to anal fin 61.9–70.6 62.0–69.3 63.3–69.2 67.4
Caudal-peduncle height 6.3–9.2 7.9–10.6 6.9–8.5 8.3
Pectoral-fin spine length 15.7–20.5 16.4–24.6 16.7–19.1 19.7
Dorsal-fin spine length 17.5–22.0 17.8–27.0 18.5–26.5 21.7
Pelvic-fin base length 2.7–4.9 3.5–5.2 3.0–4.5 3.9
Pelvic-fin height 10.9–18.3 113–17.2 12.2–18.6 13.0
Adipose-fin base length 4.3–8.3 5.0–9.8 5.4–10.4
Adipose-fin height 2.6–5.0 3.0–5.0 3.6–4.3
Anal-fin base length 16.0–21.4 16.2–22.7 15.6–20.6 18.7
Anal-fin height 10.9–17.0 10.5–18.1 13.1–15.4 18.1
Caudal-fin upper lobe length 27.4–34.9 30.9–39.2 31.6–35.6
Caudal-fin lower lobe length 23.8–32.6 24.3–34.6 23.1–31.5
92 Copeia 2012, No. 1
elucidated, including two (out of seven) in the genus
Cathorops (Betancur-R. et al., 2008; Marceniuk et al., 2009).
This study provides clarification for the taxonomic status for
four additional species inquirendae in Cathorops from ESA
(see details in Table 9) and redefines the geographic limits of
the ESA species in Cathorops formerly recognized as
Cathorops spixii sensu lato by Taylor and Menezes (1978),
making a significant contribution toward the alpha taxon-
omy of the genus and ultimately the Ariidae.
Cathorops spixii is redescribed and its geographic range
restricted to Brazil, southeastward to the Amazon River
mouth, with a northern distributional limit around the Para´
state. Cathorops nuchalis, previously listed as species inquir-
endae (Marceniuk and Ferraris, 2003; Ferraris, 2007), is
revalidated based on type and non-type specimens collected
from Venezuela through Guiana. Arius laticeps and Arius
nigricans, previously listed as species inquirendae, are syno-
nyms of C. nuchalis and C. spixii, respectively. Lastly, C. wayuu
is formally described as a new species in the Cathorops mapale
species group, ranging from Colombia through Venezuela.
As a result of the nomenclatural changes, the identity of
the species of Cathorops from northern South America for
non-taxonomy purposes (e.g., fisheries, life history, and
ecology) requires urgent clarification (e.g., Etchevers, 1978;
Arias de Diaz and Bashirullah, 1988; Tijaro et al., 1998).
Particularly, Cathorops spixii is often utilized as a model
species in a variety of studies on pollution bioindication
(e.g., Azevedo et al., 2009a, 2009b), nutritional character-
ization (de Oliveira et al., 2008), population dynamics (e.g.,
Dantas et al., 2010), feeding ecology (e.g., Melo and
Teixeira, 1992), among others. While C. spixii co-occurs
with C. agassizii and C. arenatus in northern Brazil, the latter
two species are rarely recognized as valid. Notably, there are
major differences in the habitat preferences of the three
species. Whereas C. agassizii inhabits chiefly clear waters
with high freshwater influence, C. spixii is most commonly
found on estuaries with low salinity, muddy bottoms, and
turbid waters, and C. arenatus is frequently associated with
hard substrates in open sandy areas and high salinity waters.
Finally, most species of Cathorops play an important role
in artisanal fisheries in ESA. Given that overfishing at several
locations has led to a progressive reduction in reported catch
size below that of the minimum maturation size in recent
years (e.g., Narva´ ez-Barandica et al., 2008), the fishery may
require conservation and management in the immediate
future.
KEY TO THE SPECIES OF CATHOROPS FROM THE WESTERN
ATLANTIC
All estuarine and/or marine, unless otherwise stated.
1a. Lateral and mesial surfaces of first and second gill
arches with fleshy papillae intercalated with the
gill rakers; 14–16 gill rakers on first arch; 13–16 gill
rakers on second arch
__________________________________________________________
2
1b. Lateral and mesial surfaces of first and second gill
arches without fleshy papillae intercalated with
the gill rakers; 17–24 (rarely 16 in C. wayuu) gill
rakers on first arch; 16–22 (rarely 15 in C. spixii) gill
rakers on second arch
__________________________________________________________
3
2a. Snout length 9.3–11.6%SL; distance from tip of
snout to dorsal-fin origin 39.0–40.7%SL
____________________
______________________________________________________________________________
C. aguadulce
(Mexico; freshwater)
2b. Snout length 6.0–8.6%SL; distance from tip of snout
to dorsal-fin origin 33.1–38.0%SL
________________
C. kailolae
(Mexico and Guatemala; freshwater)
3a. Pelvic and pectoral fins with intense black pigmen-
tation; posterior margin of pectoral-fin spine with
conspicuous and long serrations
____________
C. melanopus
(Guatemala; freshwater)
3b. Pelvic and pectoral fins with pale pigmentation;
posterior margin of pectoral-fin spine with incon-
spicuous and short serrations
___________________________________________
4
4a. Dorsal-fin spine as long as pectoral-fin spine
___________
_______________________________________________________________________________
C. belizensis
(Belize)
4b. Dorsal-fin spine longer than pectoral-fin spine
_______
5
5a. Cephalic shield with very few granules
______
C. higuchii
(Honduras to Panama)
5b. Cephalic shield densely granulated
_______________________________
6
6a. Short barbels, maxillary barbel not reaching base
of pectoral spine; eye large, orbital diameter 3.2–
5.2 in maxillary-barbel length
__________________________________________
7
6b. Barbels relatively long, maxillary barbel reaching
or surpassing base of pectoral spine; eyes rather
small, orbital diameter 5.3–10.5 (rarely 5.2 in C.
wayuu) in maxillary-barbel length
__________________________________
8
7a. Width of the supraoccipital process at the posterior
end 2.1–2.9%SL (Fig. 2A); orbital diameter 1.6–2.4
in width of cephalic shield at epioccipital area
_______
_________________________________________________________________________________
C. agassizii
(French Guiana to Brazil; mainly freshwater)
7b. Width of the supraoccipital process at the posterior
end 3.0–4.1%SL (rarely 2.9%SL; Fig. 2A); orbital
diameter 2.5–3.0 (rarely 2.4) in width of cephalic
shield at epioccipital area
_____________________________
C. nuchalis
(Venezuela to Guyana)
8a. Body short, 39–40 free vertebrae; osseous dorso-
medial groove of neurocranium tapering toward
the middle from both anterior and posterior ends
(condition rarely observed in C. spixii; Fig. 6C);
supraoccipital process convex in lateral view
(condition rarely observed in C. spixii; Fig. 6)
__________
9
8b. Body elongate, 41–43 free vertebrae; osseous
dorsomedial groove of neurocranium with straight
posterior margins, progressively widening anteri-
orly (Fig. 6C); supraoccipital process straight in
lateral view (Fig. 6A)
__________________________________________________________
10
9a. Orbital diameter 2.8–4.3 in width of cephalic
shield at lateral ethmoid area (Fig. 7B); orbital
diameter 4.2–6.0 (rarely 4.1) in width of cephalic
shield at supracleithrum area
____________________
C. arenatus
(Venezuela to Brazil)
9b. Orbital diameter 1.8–2.7 in width of cephalic
shield at lateral ethmoid area (Fig. 7B); orbital
diameter 2.8–4.1 in width of cephalic shield at
supracleithrum area
_______________________________________________
C. spixii
(Brazil)
10a. Caudal peduncle height 1.7–2.5 (rarely 1.6 in C.
nuchalis) in supraoccipital process length
_________________
11
10b. Caudal peduncle height 1.1–1.7 in supraoccipital
process length
________________________________________________________________________
13
11a. Flanks and venter with yellow coloration in life;
caudal peduncle height 3.0–4.4 (rarely 2.9) in width
of the supraoccipital process at the posterior
end
__________________________________________________________________________________
C. spixii
(Brazil)
Marceniuk et al.—Genus Cathorops from the Caribbean and Atlantic South America 93
11b. Body coloration white and silvery or brownish but
never yellow; caudal peduncle height 1.6–2.9 in
width of the supraoccipital process at the posterior
end
______________________________________________________________________________________________
12
12a. Supraoccipital process length 11.7–18.9%SL
(Fig. 3B); supraoccipital process length 1.0–1.5 in
width of cephalic shield at lateral ethmoid area
_______
___________________________________________________________________________________
C. nuchalis
(Venezuela to British Guiana, mainly freshwater)
12b. Supraoccipital process length 9.1–11.5%SL
(Fig. 3B); supraoccipital process length 0.6–0.9 in
width of cephalic shield at lateral ethmoid area
_______
_______________________________________________________________________________________
C. wayuu
(Colombia to Venezuela)
13a. Orbital diameter 1.8–2.6 (rarely 2.7 in C. spixii)in
width of cephalic shield at lateral ethmoid
(Fig. 7B)
_____________________________________________________________________________________
14
13b. Orbital diameter 2.7–3.9 in width of cephalic shield
at lateral ethmoid (Fig. 7B)
______________________________________________
15
14a. Flanks and venter yellow in life; 17–19 (rarely 20)
gill rakers on first arch (Table 3)
_______________________
C. spixii
(Brazil)
14b. Body coloration white and silvery or brownish but
never yellow; 21–24 (rarely 20) gill rakers on first
arch (Table 3)
_________________________________________________________
C. mapale
(Colombia)
15a. 21–24 (rarely 20) gill rakers on first arch; 19–22
(rarely 18) gill rakers on second arch (Table 4)
_________
______________________________________________________________________________________
C. mapale
(Colombia)
15b. 17–20 (rarely 21) gill rakers on first arch; 16–19 gill
rakers on second arch (Table 4)
______________________
C. wayuu
(Colombia to Venezuela)
MATERIAL EXAMINED
Counts given in Tables 1–4, and morphometrics data given
in Table 10. Material examined from Eastern Pacific, see
Marceniuk et al. (2009).
Cathorops agassizii: BMNH 1897.7.17.7, 1, holotype of Arius
pleurops, 158 mm SL, Brazil, Para´ , Marajo´; MCZ 7667, 1,
113 mm SL, Brazil, Ceara´ , Fortaleza; MCZ 7670, 1, holotype
of Tachisurus agassizii, 187 mm SL, Brazil, Rio Grande do Sul
(probably wrong); MZUSP 25169, 1, 186 mm SL, Brazil, Para´,
Vigia; MZUSP 37228, 1, 148 mm SL, Brazil, Maranha˜o, ilha
de Sa˜o Luı´s, estreito do Coqueiro; MZUSP 37230, 3, 102–
148 mm SL, Brazil, Alagoas, Maceio´ ; MZUSP 37232, 6, 119–
176 mm SL, Brazil, Alagoas, Maceio´ , lagoa Mundau´; MZUSP
37234, 1, 116–180 mm SL, Brazil, Alagoas, Maceio´, lagoa
Mundau´ ; MZUSP 37237, 2, 149–151 mm SL, Brazil, Sergipe,
Rio Sergipe; MZUSP 37241, 1, 123 mm SL, French Guiana,
Pointe des Roches; MZUSP 49341, 1, 148 mm SL, Brazil,
Alagoas, Maceio´ , lagoa Mundau´; MZUSP 49342, 1, 144 mm
SL, Brazil, Alagoas, Maceio´ , lagoa Mundau´; MZUSP 49343, 2,
107–113 mm SL, Brazil, Alagoas, Maceio´ , lagoa Mundau´;
MZUSP 49344, 1, 147 mm SL, Brazil, Maranha˜o, iIha de Sa˜o
Luı´s, Rio Curuc¸a´; MZUSP 49346, 3, 154–157 mm SL, Brazil,
Para´ , Cajueiro, baı´a de Marajo´ , ilha de Mosqueiro; MZUSP
49347, 2, 201–219 mm SL, Brazil, Para´ , Vigia, fish market;
MZUSP 49354, 1, 166 mm SL, Brazil, Sergipe, mouth of Rio
Pomonga´ , near of Aracaju´; MZUSP 49355, 3, 147–159 mm
SL, Brazil, Sergipe, Pirambu, Rio Japaratuba; MZUSP 49356,
1, 172 mm SL, Brazil, Sergipe, Pirambu, Rio Japaratuba;
MZUSP 49359, 6, 96–185 mm SL, Brazil, Sergipe, Rio Sergipe;
MZUSP 49412, 1, 84 mm SL, French Guiana, Barrieˆra.
Cathorops aguadulce: FMNH 4678, 1, holotype of Galeichthys
aguadulce, 227 mm SL, Mexico, Rı´o Tesechoaca´ n at Pe´rez,
Veracruz, Rı´o Papaloapan basin; UMMZ 97483, 5, 134–
187 mm SL, Mexico, Veracruz, Rı´o Papaloapan, 2 miles W of
San Cristobal; UMMZ 186482, 1, 190 mm SL, Mexico,
Veracruz, Estero tributary to Bahia Tecolutla, on Tecolutla-
Nautla hwy 2.8 mi SSE Tecolutla (ferry landing, S side) at
Rancho, Rı´o Tecolutla basin; UMMZ 210777, 1, 152 mm SL,
Mexico, Oaxaca, Rı´o Papaloapan, ca 100 m below bridge to
Papaloapan.
Cathorops arenatus: MCZ 7656, 1, 100.8 mm SL, British
Guiana; MCZ 7685, 2, 175–195 mm SL, Brazil, Maranha˜o,
Sa˜o Luis; MNHN 0000-4169, holotype of Arius variolosus,1,
141 mm SL, French Guiana; MZUSP 37215, 1, 91 mm SL,
Brazil, Maranha˜o, ilha de Sa˜o Luis, Page´ , estreito de
Coqueiro; MZUSP 37241, 1, 141 mm SL, French Guiana,
Pointe des Roches; MZUSP 48523, 1, 155 mm SL, Brazil, Para´,
Jubim, baı´a de Marajo
´ ; MZUSP 49364, 1, 172 mm SL, Brazil,
Para´ , fish market; MZUSP 104081, 4, 139–194 mm SL, Brazil,
Para´ , Braganc¸a, fish market; MZUSP 104082, 3, 135–183 mm
SL, Brazil, Para´ , Curuc¸a´ , Rio Curuc¸a´ estuary; MZUSP 104083,
4, 138–156 mm SL, Brazil, Para´, Salino´ polis, Mac¸arico beach;
MZUSP 104084, 3, 171–195 mm SL, Brazil, Maranha˜o, Sa˜o
Luı´s, beach next to Raposa port; RMNH 3036, holotype of
Arius fissus, 1, 139 mm SL, Suriname; RMNH 3099, 1,
holotype of Arius arenatus, 122 mm SL, Suriname; USNM
66101, 1, 136 mm SL, British Guiana, fish market of
Georgetown; USNM 225446, 3, 101–106 mm SL, Suriname,
Nickerie District, Corantijn River at Clara Creek, 05u559N,
57u059W; USNM 233482, 5, 110–133 mm SL, Venezuela,
mouth of Rı´o Orinoco, 08u569N, 60u119W; USNM 286394, 3,
130–158 mm SL, Brazil, Amapa´, 03u179N, 50u129W; USNM
286395, 2, 92–94 mm SL, Suriname, 06u049N, 54u519W;
USNM 286458, 1, 185 mm SL, Brazil, Amapa´, 02u559N,
49u449W; USNM 286459, 1, 150 mm SL, Brazil, Amapa´,
03u169N, 50u039W; USNM 286461, 1, 126 mm SL, French
Guiana, 04u599N, 51u589W; USNM 286463, 2, 159–169 mm
SL, Brazil, Amapa´, 03u279N, 50u259W; USNM 286465, 1,
130 mm SL, French Guiana, 04u439N, 51u299W; USNM
286471, 3, 115–144 mm SL, Brazil, Amapa´, 01u269N,
48u149W; USNM 286472, 1, 125–147 mm SL, Brazil, Amapa´,
00u249N, 47u329W; USNM 286473, 3, 125–146 mm SL, French
Guiana, 04u479N, 51u379W; USNM 286509, 2, 163–171 mm
SL, Brazil, Amapa´, 02u169N, 48u479W; USNM 286750, 2, 142–
182 mm SL, Brazil, Amapa´, 01u049N, 48u069W.
Cathorops belizensis: FMNH 77783, 4, 168–218 mm SL,
Belize, 500 yards off Belize City; FMNH 95978, 3, 207–
248 mm SL, collected with holotype; USNM 286399, 1,
holotype of Cathorops belizensis, 175 mm SL, Belize, Belize
City, mangrove swamps St. John’s college; USNM 385181, 8,
147–240 mm SL, collected with holotype.
Cathorops higuchii: UF 16243, 1, 180 mm SL, Costa Rica,
Limo´ n, Tortuguero; USNM 38645, 3, 144–165 mm SL,
Panama, Colo´ n, Aspinwall; USNM 79345, 1, 140 mm SL,
Panama, Toro Point, canal zone; USNM 79346, 2, 162–
168 mm SL, Panama, Colo´ n, Mindi Reef, Mindi, canal zone;
USNM 79348, 1, 163 mm SL, Panama, Colo´ n, Fox Bay;
USNM 79355, 2, 127–143 mm SL, Panama, Toro Point, canal
zone; USNM 79363, 1, holotype of Cathorops higuchii,
129 mm SL, Panama, Colo´ n Reef; USNM 79365, 1, 138 mm
SL, Panama, Toro Point, canal zone; USNM 79366, 1,
172 mm SL, Panama, Toro Point, canal zone; USNM
79415, 1, 134 mm SL, Panama, Mindi Reef, Mindi, canal
zone; USNM 286763, 1, 140 mm SL, Honduras, Rı´o Cruta,
94 Copeia 2012, No. 1
Caratasca lagoon, 15u269N, 83u419W; USNM 286764, 3, 130–
140 mm SL, Honduras, Caratasca lagoon to Rı´o Cruta,
15u269N, 83u419W; USNM 286766, 2, 97–141 mm SL,
Honduras, Rı´o Cruta, Caratasca lagoon, 15u219N, 83u349W;
USNM 385182, 3, 125–192 mm SL, collected with holotype.
Cathorops kailolae: AMNH 35074, 3, 106–135 mm SL,
Guatemala, Izabal, Rı´o Cienaga, 1 km above mouth into
´o Dulce; ANSP 142725, 1, 215 mm SL, Guatemala, Pete´n,
near Sayaxche, Rı´o Usumacinta basin; AUM 19407, 5, 112–
159 mm SL, Guatemala, Izabal, Lago Izabal, shore at beach
off Finca Carolina; AUM 32198, 3, 161–166 mm SL,
Guatemala, Pete´ n, mouths of ´o San Juan and Rı´o Pucte´,
´o La Pasio´n, Rı´o Usumacinta basin; ECO-SC 4268, 1,
217 mm SL, Mexico, Chiapas, Montes Azules, Selva
Lacandona, Rı´o Usumacinta basin; ECO-SC 4270, 1,
198 mm SL, Mexico, Chiapas, Montes Azules, Selva
Lacandona, Rı´o Usumacinta basin; UF 35320, 4, 51–
119 mm SL, Guatemala, Izabal, Lago Izabal, off Finca
Carolina; UMMZ 188018, 1, 276 mm SL, Guatemala, Pete´n,
Arroyo Tamarindo 0.5 km above mouth in Laguna Petexba-
tum, above jct with Arroyo Aquateca, 16u249150N,
90u119200W, Rı´o Usumacinta basin; USNM 134330, 1,
holotype Cathorops kailolae, 181 mm SL, Guatemala, Lago
Izabal, embayment about 3 mi. W of El Estor; USNM 385739,
5, 145–214 mm SL, collected with holotype.
Cathorops mapale: ICN-MHN 8244, 1, 166 mm SL, Colombia,
Magdalena, Cie´ naga Grande de Santa Marta, 10u599N,
74u179W; ICN-MHN 8245, 2, 164–196 mm SL, Colombia,
Magdalena, Golfo de Salamanca, 11u69N, 74u189W; ICN-MHN
8246, 2, 165–179 mm SL, Colombia, Co´rdoba, mouth of ´o
Sinu´, 9u269N, 75u559W; ICN-MHN 8247, 1, 143 mm SL,
Colombia, Antioquia, Golfo de Uraba´, Rı´o Atrato, mouth El
Roto, 8u079N, 76u589W; INVEMAR-PEC 277, 1, 256 mm SL,
Colombia, Magdalena, Tasajera, 11u09N, 74u209W; INVEMAR-
PEC 1584, 1, 166 mm SL, Colombia, Magdalena, Costa Verde,
Cie´naga Grande de Santa Marta, 11u029N, 74u159W; INVE-
MAR-PEC 3654, 1, 180 mm SL, Colombia, Co´rdoba, Cie´naga
de Soledad, 9u209N, 75u529W; INVEMAR-PEC 5197, 1, 156 mm
SL, Colombia, Co´rdoba, mouth of tı´o Tinajones, 9u259N,
75u489W; INVEMAR-PEC 5333, 1, holotype of Cathorops
mapale, 183.5 mm SL, Colombia, Magdalena, Cie´ naga Grande
de Santa Marta, 10u599N, 74u179W; INVEMAR-PEC 5348, 1,
118 mm SL, Colombia, Antioquia, Golfo de Uraba´, Rı´o Atrato,
mouth El Roto, 8u079N, 76u589W; INVEMAR-PEC 5501, 2,
151–152 mm SL, Colombia, Antioquia, Golfo de Uraba´,
8u299N, 77u139W; INVEMAR-PEC 5730, 1, 157 mm SL,
Colombia, Magdalena, Golfo de Salamanca, 11u69N,
74u189W; INVEMAR-PEC 5731, 2, 139–172 mm SL, Colombia,
Antioquia, Golfo de Uraba´, 8u179N, 76u489W; INVEMAR-PEC
5732, 1, 157 mm SL, Colombia, Antioquia, Golfo de Uraba´,
7u569N, 76u539W; INVEMAR-PEC 5733, 26 CS, 110–144 mm
SL, Colombia, Magdalena, Golfo de Salamanca, 11u69N,
74u189W; USNM 286396, 2, 178–180 mm SL, Colombia,
Cienaga Grande de Santa Marta; USNM 286398, 4, 175–
186 mm SL, Colombia, Cienaga La Virgen.
Cathorops melanopus: AMNH 35241, 2, 115.0–143.0 mm SL,
Guatemala, Izabal, Rı´o Motagua at Finca Hopi; BMNH
1865.4.29. 51–53, 1, syntype of Arius melanopus, 172 mm
SL, Guatemala, Rı´o Motagua; UMMZ 197336, 3, 126–
188 mm SL, Guatemala, Izabal, Rı´o Motagua at Finca Hopi.
ACKNOWLEDGMENTS
We are indebted to S. Raredon (USNM) for obtaining x-ray
and data, and to P. Be´ arez, J. Gregorio, C. Ferrara, P. Pruvost,
M. Hautecoeur, R. Causse, S. Raredon (USNM), for providing
logistic support at the MNHN. C. Ferraris Jr., N. Menezes,
and F. de Lima critically reviewed the manuscript. This work
was supported by the Fundac¸a˜o de Amparo a` Pesquisa do
estado de Sa˜o Paulo (FAPESP, Proc. 03/04509-3 to APM), the
Conselho Nacional de Desenvolvimento Cientı´fico e Tecno-
lo´ gico of the Brazilian Federal Government (CNPq, Proc.
152782/2007-9 to APM), and the All Catfish Species
Inventory, supported by the National Science Foundation
(NSF DEB-0315963). This is contribution 355 of Centro de
Estudios en Ciencias del Mar, CECIMAR, Nacional de
Colombia sede Caribe.
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... In the laboratory the species were identified using the descriptions and identification keys proposed by Acero (2003), Marceniuk (2007) and Marceniuk et al. (2012). Analysis included specimens preserved in the Fish Collection of the Federal University of Maranhão (Coleção de Peixes da Universidade Federal do Maranhão-CPUFMA). ...
... In this section "material deposited" indicates specimens collected in field in the present study and "material examined" indicates specimens analyzed during our visit to CPUFMA. The diagnosis of species where based on the descriptions and diagnosis of the papers of Miles (1945), Acero & Betancur-R (2006), Marceniuk (2007), Marceniuk et al. (2012) and Marceniuk et al. (2017). ...
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During 1994, the catfish "chivo mapalé" (Cathorops spixii) was the second most abundant fishing resource in the "Ciénaga Grande de Santa Marta" and the "Complejo de Pajarales", the main lacustrine-estuarine system of Colombia. Using commercial catch statistics and fishing explorations, the population parameters were determined reproductive cycle, growth, mortality, yield and biomass per recruit. Mature individuals were present all year around with peaks of ripeness and spawning from June to August during periods of little rain and dry, respectively. The mean maturity size was estimated at 23 cm. The growth parameters of the Von Bertalanffy function were: Loo = 32.5 cm and K = 0.38 year1, with a growth performance index of 2.6. Total (Z), natural (M) and fishing (F) mortality rates were estimated as 2.83, 0.96 and 1.87 year1, respectively. The exploitation rate E = 0.66 indicates that the resource is overfished. Therefore it is recomended a fishery regulation by decreasing the mortality rate to F = 1.0 year1 and increasing the cath mean size to 23 cm.
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