Copeia, 2002(3), pp.

739–747

New Species of Trichomycterus from the Andes of Argentina with a

Redescription of Trichomycterus alterus (Siluriformes: Trichomycteridae)
LUIS FERNÁNDEZ AND RICHARD P. VARI

Trichomycterus belensis, new species, is described from a tributary to Laguna Blan-

ca, a high-elevation drainage in the Departamento Belén, Provincia de Catamarca,
Argentina. The new species is distinguished from other members of the apparently
nonmonophyletic genus Trichomycterus by a combination of features involving the
premaxilla, dentition, barbel form, pigmentation, and morphometrics. Trichomycte-
rus alterus, whose original description was based on a limited number of specimens,
is redescribed based on numerous specimens from localities in the central and
northwestern portions of Argentina.

Trichomycterus belensis, nueva especie, es descripta de un tributario de Laguna

Blanca de elevada altura en el departamento Belén, Provincia de Catamarca, Argen-
tina. La nueva especie se distingue de otros miembros del género Trichomycterus,
aparentemente no monofilético, por la combinación de rasgos del premaxilar, den-
tición, forma de la barbilla, pigmentación y morfométricos. Trichomycterus alterus,
cuya descripción original estaba basada en un número limitado de ejemplares, es
redescripta basada en numerosos especimenes de localidades de la parte central y
noroeste de Argentina.

T HE speciose, but apparently nonmonophy-

letic, catfish genus Trichomycterus has more
than 100 recognized species and numerous yet-
to-be described forms (de Pinna, 1989, 1992,
1998). Trichomycterus occurs across broad reach-
es of Central and South America and is highly
diverse along the lengthy Andean Cordilleras, a
region where it is one of the few fish taxa that
inhabit higher elevations (Ortega, 1992). Col-
lecting efforts by the first author and colleagues
at high-elevation localities in northwestern Ar-
gentina yielded several undescribed Trichomyc-
terus species. Two of these species were recently
described by Fernández and Vari (2000) and
Fernández (2000a) with the third formally de-
scribed herein. Collecting activities across cen-
tral and northwestern Argentina also yielded
numerous specimens of Trichomycterus alterus, a
species originally described on the basis of four
specimens by Marini et al. (1933). Subsequently,
Ringuelet et al. (1967) summarized the meristic
and morphometric features of this species based
on five additional specimens. We redescribe this
poorly known species in detail based on an ex-
tensive series of specimens. The assignment of
the species treated herein to Trichomycterus fol-
lows the concept of that genus used by recent
authors (e.g., Arratia and Menu Marque, 1984).
MATERIAL AND METHODS
binocular microscope following the methods
outlined by Tchernavin (1944) and de Pinna
(1992). Cleared-and-counterstained specimens
for osteological study were prepared following
the procedure of Taylor and Van Dyke (1985)
and osteological nomenclature follows Baskin
(1973) and de Pinna (1989, 1998). Counts of
dorsal and anal fin rays follow the method pro-
posed by de Pinna (1992). As proposed by de
Pinna (1992), the vertebral counts do not in-
clude the vertebrae in the Weberian apparatus,
and the compound caudal centrum is counted
as one element. Museum abbreviations are as
listed in Leviton et al (1985) with the addition
IADIZA, Instituto Argentino de Investigación de
Zonas Aridas, Mendoza. All elevations are given
in meters above sea level (asl).
Trichomycterus belensis, n. sp.
Figure 1
Holotype.—FML 2530, 63.5 mm SL; Argentina,
Provincia de Catamarca, Departamento Belén,
stream tributary to Laguna Blanca, 11 km
northeast from Belén on provincial road 43
along route from Belén to Antofagasta de la Si-
erra, near Los Nacimientos de San Antonio (ap-
proximately 268309S, 678039W), elevation 3500
m, collected by L. Fernández, G. Scrocchi, and
E. Lavilla, 8 September 1993.

Measurements were taken from the left side Paratypes.—Twenty-eight specimens, 25.5–58.7
of the specimens with digital calipers under a mm SL: FML 2531, 26 specimens, 25.5–58.7 mm
q 2002 by the American Society of Ichthyologists and Herpetologists
740 COPEIA, 2002, NO. 3
Fig. 1. Trichomycterus belensis, new species, holotype, FML 2530, 63.5 mm SL, left lateral view; Argentina,
Provincia de Catamarca, Departamento Belén, stream tributary to Laguna Blanca.
SL (five cleared and counterstained), and
USNM 364371, two specimens, 37.5–48.5 mm
SL; collected with holotype.
Nontype specimens.—Four hundred eighty-six
specimens, 9.8–55.5 mm SL, collected with ho-
lotype. FML 2532, 71 specimens, 9.8–42.0 mm
SL; FML 2533, five specimens, 9.4–35.9 mm SL;
FML 2534, 405 specimens, 27.1–55.5 mm SL;
USNM 364372, three specimens, 40.8–44.9 mm
SL.
Diagnosis.—The combination of the possession
of an oval premaxilla smaller than maxilla, the
presence of the pelvic fins and the associated
pelvic girdle, seven or eight branched dorsal fin
rays, 15 to 17 ribs, a compressed caudal pedun-
cle, the presence of an unpigmented region on
the body anteroventral to the origin of the dor-
sal fin, the marmoration on the dorsal fin, the
lack of a very thick, rugose layer of fatty tissue
on the head and body, the presence of a portion
of the laterosensory canal system within the
sphenotic, the lack of an extensive perforation
of the skin surface by ampullary organs, and a
maximum overall body size of 63.5 mm SL in
Trichomycterus belensis distinguishes that species
from all other known members of the subfamily
Trichomycterinae with the exception of T. alte-
rus, Trichomycterus areolatus, Trichomycterus boylei,
and Trichomycterus ramosus. Trichomycterus belensis
differs from T. alterus in caudal peduncle depth
(59.5–64.1% of body depth, vs 48.1–55.8%, re-
spectively), body coloration (darkly marmorat-
ed, vs spotted, respectively), and degree of de-
velopment of the first pectoral fin ray (usually
extends beyond fin margin, but does not form
filament, vs always in form of distinct filament,
respectively); from T. areolatus in the number of
premaxillary tooth rows (2 or 3 vs 5, respective-
ly), number of caudal vertebrae (29 to 32 vs 35,
respectively), and number of ribs (15 to 17 vs
13, respectively); from T. boylei in the number
of rows of teeth on the premaxilla (2 or 3 vs 4
to 7, respectively), tooth form (teeth with
straight vertical margins, vs teeth spatulate and
distally expanded, respectively), the degree of
development of the first pectoral fin ray (with
distinct extension beyond the fin margin, vs
without extension beyond fin margin, respec-
tively), and the form of the papillae-like struc-
tures on the body (minute and slightly sharp-
pointed, vs large and blunt or conic, respective-
ly); and from T. ramosus in the form of the bar-
bels (unbranched, vs some or all barbels
branched one or more times, respectively) and
body pigmentation (darkly marmorated, vs uni-
formly brown, respectively).
Description.—Morphometric data for the holo-
type and paratypes presented in Table 1. Body
elongate; roughly circular in cross-section and
only slightly compressed transversely in trunk
region, gradually becoming progressively more
compressed transversely posteriorly. Dorsal and
ventral profiles of trunk region ranging from
slightly convex to slightly concave. Dorsal and
ventral profiles of caudal peduncle straight to
slightly convex and smoothly continuous with
profile of trunk region. Small, slightly sharp-
pointed, papillae-like structures present on
body. Anterior most portion of laterosensory ca-
nal system on body surrounded by one small,
tubular ossicle. Very thin layer of fatty tissue
overlies much of head and body.
Head approximately triangular overall when
examined from dorsal view; relatively wide pos-
teriorly, with anterior portion narrower. Snout
with broadly rounded margin from dorsal view.
Head depressed, with eye located on dorsal sur-
face. Eye ovoid and slightly anteroposteriorly
elongate. Skin covering eye thin, transparent,
and separate from surface of eye, with eye read-
ily visible. Orbital rim not free. Regions lateral,
and particularly posterolateral, of eye somewhat
expanded as consequence of well-developed jaw
muscles.
Anterior nostril slightly smaller than posterior
nostril and surrounded by fleshy flap of integ-
ument medially and by nasal barbel laterally.
Posterior nostril bordered anteriorly by flap of
skin.
Mouth distinctly subterminal, with rictus di-
rected posteriorly. Premaxilla oval, smaller than
 FERNÁNDEZ AND VARI—ARGENTINIAN TRICHOMYCTERUS 741

TABLE 1. MORPHOMETRIC DATA FOR HOLOTYPE AND 23 PARATYPES OF Trichomycterus belensis, NEW SPECIES. STAN-
DARD AND HEAD LENGTHS ARE EXPRESSED IN MM; MEASUREMENTS 1–11 ARE PERCENTAGES OF STANDARD LENGTH;
12–17 ARE PERCENTAGES OF HEAD LENGTH.

HOLOTYPE PARATYPES MEAN STANDARD DEVIATION

STANDARD LENGTH 63.5 25.5–58.7 48.1

HEAD LENGTH 10.9 5.2–10.6 8.7
1. BODY DEPTH 14.8 11.5–17.9 15.5 2.2
2. CAUDAL PEDUNCLE LENGTH 23.8 20.2–26.7 23.7 1.3
3. CAUDAL PEDUNCLE DEPTH 10.7 9.3–12.5 10.4 0.9
4. PREDORSAL LENGTH 63.8 62.1–67.8 64.6 1.6
5. PREANAL LENGTH 71.2 67.0–72.0 69.4 1.3
6. PREPELVIC LENGTH 55.6 42.7–58.8 54.0 3.3
7. DORSAL FIN BASE LENGTH 9.3 8.8–11.4 10.0 0.6
8. ANAL FIN BASE LENGTH 6.4 5.9–10.0 7.8 0.8
9. HEAD LENGTH 17.2 17.1–20.4 18.1 0.8
10. HEAD WIDTH 16.2 16.3–20.1 18.8 1.0
11. HEAD DEPTH 10.5 9.6–12.9 10.9 0.8
12. NASAL BARBEL LENGTH 45.9 29.6–61.2 41.7 8.1
13. MAXILLARY BARBEL LENGTH 62.4 34.6–82.7 60.5 12.3
14. SUBMAXILLARY BARBEL LENGTH 45.9 26.3–48.0 38.9 5.9
15. MOUTH WIDTH 38.5 32.7–44.8 40.4 3.0
16. INTERORBITAL WIDTH 34.9 25.0–34.3 29.7 2.8
17. SNOUT LENGTH 40.4 38.9–47.9 42.4 2.3

maxilla or width of palatine. Premaxilla with 2
or 3 rows of teeth. Outer premaxillary tooth row
regularly arranged, with 6 to 8, relatively elon-
gate, incisiform teeth with parallel margins.
Teeth on inner rows of premaxilla somewhat
shorter than those of outer row, with teeth
small, conical, and somewhat irregularly ar-
ranged. Maxilla unusually prolonged relative to
form in most Trichomycterus species and lacking
distinct process on lateral portion common to
all examined nominal congeners. Dentary with
3 tooth rows medially and 2 rows laterally. Outer
dentary tooth row with 8 relatively elongate, in-
cisiform teeth with parallel margins. Teeth of
inner tooth rows on dentary of form similar to
that of teeth of outer row but relatively smaller
and irregularly arranged. Lower lip with prom-
inent fleshy lobe along lateral limit; lobe situ-
ated internal to base of rictal barbel. Anterior
portion of lower lip fleshy, with anterior, and to
lesser degree anteroventral, surfaces covered
with papillae. Upper lip fleshy; bearing numer-
ous papillae.
Barbels relatively short and tapering distally,
but not threadlike or with distal branches pre-
sent on some congeners. Maxillary barbel ex-
tending posteriorly to interopercular batch of
odontodes but falling short of pectoral fin ori-
gin. Nasal barbel reaching to posterior border
of eye. Origin of nasal barbel on posterolateral
portion of skin flap along margin of anterior
naris. Submaxillary barbel shorter than maxil-
lary barbel.
Branchiostegal membranes thick, with bran-
chiostegal rays barely visible through skin. Bran-
chiostegal rays 5 to 7, most commonly 6.
Distal margin of pectoral fin rounded. First
pectoral fin ray extending as short filament
slightly beyond fin margin in holotype and most
paratypes, and not present in other examined
specimens. Pectoral fin rays 7 or 8, with lateral
most ray unbranched. Distal margin of dorsal
fin rounded; semicircular when fin distended.
Total dorsal fin rays 11 to 13, usually 12, with 7
or 8, usually 8, branched rays and 4 or 5 un-
branched rays. Externally visible dorsal fin rays
8 or 9, with remaining fin rays hidden by thick
integument that overlies anterobasal portion of
fin. Dorsal fin fleshy basally. Dorsal fin origin
located slightly anterior to vertical through an-
terior limit of anal aperture. Total anal fin rays
9 or 10, usually 10, with 5 branched rays and 4
or 5, usually 5, unbranched rays. Externally vis-
ible anal fin rays 6 or 7 in holotype and paraty-
pes, with remaining fin rays covered by thick
integument that overlies anterobasal portion of
fin. Vertical through anal fin origin extends
though posterior portion of dorsal fin base.
Anal fin slightly smaller than dorsal fin but with
most rays proportionally elongate and with dis-
tal margin slightly rounded. Pelvic fin rays 5,
first ray unbranched and second and third rays
742 COPEIA, 2002, NO. 3
longest. Tip of adpressed pelvic fin reaching to,
or extending slightly beyond, anus but falling
short of anal fin origin. Caudal fin margin ei-
ther nearly straight or slightly emarginate. Cau-
dal fin with 616 principal rays. Dorsal procur-
rent rays 10 to 13. Ventral procurrent rays 10 to
13.
Interopercular patch of odontodes antero-
posteriorly elongate; odontodes embedded in
thick integument that covers interopercle.
Opercular patch of odontodes rounded, with
odontodes embedded in thick integument that
covers opercle. Opercle with 8 to 13 odontodes
and interopercle with 18 to 40 odontodes in 5
cleared-and-stained specimens.
Vertebrae 36 to 39, with 5 to 9 precaudal and
29 to 32 caudal vertebrae. Ribs on each side 15
to 17.
Color in alcohol.—Dorsal and dorsolateral por-
tions of head with pattern of dark marmoration.
All barbels, other than submaxillary barbels,
darkly pigmented with diffuse pattern of small,
dark chromatophores. Interopercle with small
patch of dark pigmentation. Opercular patch of
odontodes with weblike pattern of dark pigmen-
tation around base of odontodes. Jaw teeth and
opercular and interopercular odontodes unpig-
mented even in overall darker individuals.
Body with dark marmoration on all areas oth-
er than for abdominal region and rounded area
anteroventral to dorsal fin origin. Abdominal
region largely lacking dark coloration, other
than for scattered small, dark chromatophores
in smaller specimens. Rounded area anteroven-
tral to dorsal fin origin either unpigmented or
with scattered dark pigmentation.
Dorsal fin irregularly marmorated. Caudal fin
dusky, with rays outlined by small dark chro-
matophores. Dark pigmentation on basal por-
tions of caudal fin rays more pronounced, form-
ing slightly anteroventrally angled vertical bar.
Anal fin hyaline or with scattered irregular
patches of dark pigmentation. Lateral most pec-
toral fin ray with irregular dark pigmentation.
Dorsal surface of pectoral fin with scattered, ir-
regular patches of dark pigmentation. Pelvic fin
hyaline or with scattered irregular patches of
dark pigmentation.
Ecology.—Only two other aquatic vertebrates
were collected at the type locality of Trichomyc-
terus belensis: T. catamarcensis, and tadpoles and
adults of an undescribed Telmatobius species
(Anura: Leptodactylidae). The only other Tri-
chomycterus species known from proximate water
bodies is T. ramosus, which was collected in a
river system approximately 1 km from Laguna
Blanca, but without surface connections with it.
The type locality for T. belensis, which is at ap-
proximately 3500 m, is a small, clear water
stream, approximately 0.5 m deep, running
over a sandy bottom and with marginal vegeta-
tion limited to grasses. The stream disappears
into the substrate at its lower terminus with an
apparently subterranean flow into the saline La-
guna Blanca. As noted by Fernández and Vari
(2000) few other Neotropical fishes occur at
such elevations, with only four other Trichomyc-
terus species reported to inhabit higher-eleva-
tion water bodies. This total has been increased
by the subsequent description of T. ramosus
from a location at 3680 m (Fernández 2000).
The stomachs of three cleared-and-stained T.
belensis specimens contained dipteran larvae
(Chironomidae and Simuliidae), coleopterans
(Elmidae), trichopterans, and plecopterans
(Perlidae). Thus, the species apparently shares
the diet of autochthonous benthic macroinver-
tebrates common to many congeners (Casatti
and Castro, 1998; Ferriz, 1998; Fernández,
2000b).
Distribution.—Trichomycterus belensis is known
only from the type locality.
Etymology.—The specific name, belensis, is in ref-
erence to Departamento Belén, Provincia de
Catamarca, Argentina, where the type locality is
located.
Remarks.—As noted in the introduction, the ge-
nus Trichomycterus is likely nonmonophyletic
and no encompassing studies have been carried
out on the phylogenetic relationships within the
genus. Nonetheless, it is noteworthy that T. be-
lensis shares with T. alterus, T. areolatus, T. boylei,
and T. ramosus an ovoid premaxilla that is small-
er than the maxilla. Such a shape of the pre-
maxilla distinguishes these five species from all
other Trichomycterus species, all of which have a
rectangular premaxilla. A rectangular premax-
illa also occurs in Hatcheria and Silvinichthys, the
proximate outgroups to Trichomycterus. An oval
premaxilla smaller than the maxilla may, thus,
delimit a monophyletic group within Trichomyc-
terus. Alternatively, T. belensis and some conge-
ners share a distinctive number of principal cau-
dal fin rays with Scleronema and Bullockia that
may, in turn, represent a derived condition. A
more detailed phylogenetic analysis, which lies
beyond the scope of this study, is necessary to
determine the significance of these similarities.
 FERNÁNDEZ AND VARI—ARGENTINIAN TRICHOMYCTERUS
Fig. 2. Trichomycterus alterus, FML 2085, 49.2 mm SL, left lateral view; Argentina, Provincia de La Rioja,
Departamento San Blás, Andalucas.
Trichomycterus alterus (Marini, Nichols, and
La Monte 1933)
Figure 2
Pygidium alterum Marini et al., 1933:2 [type lo-
cality: Rı́o Los Sauces, La Rioja, Argentina];
Pozzi 1945:261 [distribution]; Ringuelet and
Arámburu, 1962:47 [citation]; Ringuelet et al.
1967:352 [in key, brief redescription. Argen-
tina: Jujuy Province, Quebrada de Humahua-
ca; La Rioja Province, Rı́o Los Sauces and Va-
lle Guanchı́n]; Miquelarena and Moly, 1974:
159–161 [papillae]; Eschmeyer, 1998:78 [ci-
tation].
Pygidium alternum, Gosline, 1946:56 [citation;
species name misspelled]; Burgess, 1989:321
[citation].
Trichomycterus alterum, Arratia et al., 1978:176–
178 [comparison]; Arratia et al., 1983:48–107
[Argentina, La Rioja Province and Jujuy Prov-
ince, Quebrada de Humahuaca, Rı́o Grande
above 2940 m]; Arratia and Menu-Marque,
1984:514–518 [comparison]; Fernández,
1994:12, 23, 25 [citation]; López et al., 1987:
28 [citation]; Miquelarena et al., 1990:273
[Argentina, Rı́os Calera, Artaza, Vipo]; Fer-
nández, 2000b:32 [comparisons].
Material examined.—Argentina. Provincia de La
Rioja: AMNH 12241, 1, 27.7 mm SL (holotype);
AMNH 12242, 3 (paratypes), 22.9–27.2 mm SL.
FML 2118, 3 specimens, 39.1–45.3 mm SL, De-
partamento Sanagasta, Rı́o de los Sauces. FML
2085, 199 (1 cleared and counterstained), 18.9–
49.2 mm SL, Departamento San Blás, Andalu-
cas. FML 2086, 1, 37.9 mm SL, Departamento
San Blas, Gualco or Hualco. IADIZA 121, 3, De-
partamento San Blas, Rı́o Los Sauces. IADIZA
125, 3, Departamento San Blas, Alpasinche. IA-
DIZA 140, 3, Departamento Arauco, Rı́o Colo-
rado or Rı́o Salado. Provincia de Catamarca:
FML 2269, 6, 10.5–44.2 mm SL, Departamento
Belén, Dique Rı́o Belén. FML 2089, 19, 33.0–
46.5 mm SL, Departamento Belén, Rı́o Belén.
FML 2088, 7 (2 cleared and counterstained),
22.9–41.9 mm SL, IADIZA 136, 8; Departamen-
743
to Paclı́n, Rı́o Paclı́n. FML 2114, 59, 56.5–60.0
mm SL, Departamento Belén, Rı́o Agua Clara.
FML 2115, 17, 12.5–49.6 mm SL, USNM
364373, 3, 36.6–39.5 mm SL, FML 2087, 46 (3
cleared and counterstained), Departamento Be-
lén, Los Nacimientos. FML 2585, 20, 18.8–44.3
mm SL, Departamento Belén, Villa Vil. Provin-
cia de Córdoba: FML 2586, 1, 49.1 mm SL, De-
partamento Cruz del Eje, Arroyo Barriales. FML
2587, 2, 27.8–50.9 mm SL, Departamento Cruz
del Eje, Rı́o San Marcos.
Diagnosis.—The combination of the possession
of an oval premaxilla, the presence of pelvic fins
and the associated pelvic girdle, 6 to 9 branched
dorsal fin rays, 11 to 14 ribs, a compressed cau-
dal peduncle, the rounded, unpigmented re-
gion on the body anteroventral to the origin of
the dorsal fin, the lack of a prominent pattern
of dark pigmentation on the fins, the lack of a
very thick, rugose layer of fatty tissue on the
body and head, the presence of a portion of the
laterosensory canal system within the sphenotic,
the lack of an extensive perforation of the skin
surface by ampullary organs, the lack of a dor-
sally directed spine on the scapulo-coracoid pro-
cess, and the maximum body size of 60 mm SL
in T. alterus differentiates that species from all
other known members of the subfamily Tri-
chomycterinae with the exception of T. boylei,
which also has an oval premaxillae and the lack
of a spine on the scapulo-coracoid process. Tri-
chomycterus alterus differs from T. boylei in the
number of precaudal vertebrae (3 to 5 vs 7 or
8, respectively), the number of principal caudal
fin rays (12 vs 13, respectively), the form of the
first pectoral fin ray (a distinct filament, vs de-
veloped slightly beyond fin margin, but not in
form of distinct filament, respectively), the form
of the maxillary barbel (distinctly expanded at
the base into a bulblike region, vs not expanded
basally, respectively), and the form of the teeth
on the outer row of the premaxilla (straight sid-
ed incisiform, vs spatulated and distinctly distal-
ly expanded, respectively).
744 COPEIA, 2002, NO. 3

TABLE 2. MORPHOMETRIC DATA FOR HOLOTYPE, THREE PARATYPES (A), AND 20 NONTYPE SPECIMENS (B) OF Tri-
chomycterus alterus FROM OTHER LOCALITIES. Standard and head lengths are expressed in millimeter; measure-
ments 1–11 are percentages of standard length; 12–17 are percentages of head length.

Paratypes Nontypes
Holotype A B Mean SD

Standard length 27.7 22.9–27.2 26.5–51.0 42.7

Head length 4.9 4.8–5.7 5.1–9.3 8.1
1. Body depth 14.8 14.7–16.3 15.3–22.0 19.2 1.7
2. Caudal peduncle length 25.9 25.1–28.6 24.7–32.0 28.0 1.8
3. Caudal peduncle depth 9.0 8.9–10.8 8.3–12.9 10.4 1.1
4. Predorsal length 62.4 61.6–62.9 58.6–65.0 62.3 1.7
5. Preanal length 66.8 69.1–69.4 62.0–70.5 65.8 2.0
6. Prepelvic length 53.1 53.7–55.6 47.5–54.1 51.5 1.9
7. Dorsal fin base length 10.1 10.6–12.0 8.4–13.2 11.4 1.4
8. Anal fin base length 7.9 6.9–9.3 6.0–8.7 7.6 0.6
9. Head length 17.7 18.5–21.1 16.9–21.1 19.1 1.1
10. Head width 19.1 18.1–21.6 17.9–22.9 20.4 1.4
11. Head depth 11.9 11.1-13.1 9.7–13.3 11.8 0.9
12. Nasal barbel length 53.1 32.5–56.2 23.7–51.1 37.5 8.0
13. Maxillary barbel length 67.3 53.8–72.9 36.8–81.5 56.8 12.7
14. Submaxillary barbel length 44.9 36.7–42.1 23.4–45.6 33.9 5.2
15. Mouth width 36.7 33.5–37.5 33.3–46.2 38.8 3.3
16. Interorbital width 38.8 34.3–36.0 26.8–35.1 31.3 2.4
17. Snout length 51.0 45.0–50.0 35.9–48.7 43.3 2.5

Description.—Morphometric data for the holo- rected posteriorly. Premaxilla approximately

type, paratypes, and nontype specimens from
which counts and measurements were taken are
presented in Table 2. Body elongate; slightly
compressed transversely in trunk region and
gradually becoming distinctly compressed trans-
versely in region of caudal peduncle. Dorsal and
ventral profiles of trunk region ranging from
nearly straight to slightly convex or concave.
Dorsal and ventral profiles of caudal peduncle
straight to slightly concave and smoothly contin-
uous with profile of trunk region. Small papil-
lae-like structures present on body. Anterior
most portion of laterosensory canal system on
body surrounded by one small, tubular ossicle.
Head approximately triangular overall in dor-
sal view; much wider posteriorly than anteriorly.
Snout with broadly rounded margin in dorsal
view. Head depressed, with eye located on dor-
sal surface. Eye ovoid, slightly anteroposteriorly
elongate. Skin covering eye thin and transpar-
ent, separate from surface of eye, with eye read-
ily visible. Orbital rim not free. Regions lateral,
and particularly posterolateral, of eye expanded
to varying degrees as consequence of well de-
veloped jaw muscles.
Anterior nostril slightly smaller than posterior
nostril and surrounded by fleshy flap of integ-
ument medially and by nasal barbel laterally.
Posterior nostril partially bordered anteriorly by
flap of skin.
Mouth distinctly subterminal, with rictus di-
oval and equal in size to, or smaller than, max-
illa or width of palatine. Premaxilla with 2 or 3
rows of teeth. Outer tooth row on premaxilla
regularly arranged, with 7 or 8 distally narrow-
ing, incisiform teeth. Inner rows of teeth on
premaxilla somewhat shorter than those of out-
er row and regularly arranged, with dentition of
similar shape to that of outer row. Dentary teeth
of shape similar to those of the premaxilla, with
3 rows of teeth. Outer dentary tooth row with 7
teeth. Inner tooth rows on dentary as long as,
to somewhat shorter than, outer row, with teeth
shorter than those of outer row. Lower lip with
prominent fleshy lobes along lateral limit; lobe
situated internal to base of rictal barbel. Ante-
rior portion of lower lip fleshy, with anterior,
and to lesser degree anteroventral, surfaces cov-
ered with papillae. Upper lip fleshy, bearing nu-
merous papillae.
Barbels relatively short and tapering distally,
but not threadlike or with distal branches pre-
sent in some congeners. Maxillary barbel ex-
tending posteriorly to interopercular patch of
odontodes but falling short of pectoral fin ori-
gin. Maxillary barbel distinctly widened and
bulblike basally, tapering distally. Nasal barbel
reaching posterior margin of eye. Origin of na-
sal barbel on posterolateral portion of skin flap
along margin of anterior nostril. Submaxillary
barbel shorter than maxillary barbel. Branchios-
tegal rays 6 or 7, most commonly 6.
 FERNÁNDEZ AND VARI—ARGENTINIAN TRICHOMYCTERUS
Distal margin of pectoral fin broadly round-
ed. First pectoral fin ray prolonged as distinct
filament in adults but proportionally much
shorter in juvenile of 16.4 mm SL. Pectoral fin
rays 7 or 8, with lateral most ray unbranched.
Distal margin of dorsal fin rounded, semicircu-
lar when fin distended. Total dorsal fin rays 10
to 13, usually 11, with 6 to 8, rarely 6, branched
rays, and 4 or 5 unbranched rays. Externally vis-
ible dorsal fin rays 8 or 9, with remaining fin
rays hidden under thick integument that over-
lies anterobasal portion of fin. Dorsal fin origin
located distinctly posterior to vertical through
anterior limit of urogenital aperture. Distinct
median fin fold present in juveniles between
posterior limit of dorsal fin base and posterior
limit of caudal peduncle. Fold continuous with
caudal fin in very small individuals. Total anal
fin rays 9 or 10, with 5 branched rays and 4 or
5 unbranched rays. Externally visible anal fin
rays 6 to 8, usually 7, with other anal fin rays
covered with thick integument that overlies an-
terobasal portion of fin. Vertical through anal
fin origin extending through posterior portion
of dorsal fin base. Anal fin slightly smaller than
dorsal fin; with most rays proportionally elon-
gate and distal margin slightly rounded. Distinct
median fin fold present between posterior limit
of anal fin base and posterior limit of caudal
peduncle in juveniles. Fold continuous with cau-
dal fin in very small individuals. Pelvic fin rays
5, first ray unbranched and second and third
rays longest. Tip of adpressed pelvic fin extend-
ing posteriorly beyond anus but usually falling
short of anal fin origin. Caudal fin margin
straight or slightly emarginate. Caudal fin with
616 principal rays. Procurrent caudal fin rays
inconspicuous and relatively few in number.
Dorsal procurrent rays 7 to 11 and ventral pro-
current rays 6 to 12.
Interopercular patch of odontodes antero-
posteriorly elongate; odontodes embedded in
thick integument that covers interopercle.
Opercular patch of odontodes rounded and
slightly anteroposteriorly elongate, with odon-
todes embedded in thick integument that cov-
ers opercle. Interopercle with 29 to 36 odonto-
des and opercle with 8 to 14 odontodes in 6
cleared and stained specimens.
Vertebrae 36 to 39, usually 37, with 3 to 5
precaudal vertebrae and 31 to 34 caudal verte-
brae. Ribs on each side 11 to 14, usually 13.
Color in alcohol.—Trichomycterus alterus demon-
strates considerable range in intensity of head
and body pigmentation, with holotype and pa-
ratypes being nearly unpigmented, perhaps as
consequence of original preservation or subse-
745
quent conservation. Small, more recently col-
lected, individuals unpigmented other than for
scattered, dark chromatophores on dorsal and
dorsolateral surfaces of head and body. Larger,
recently collected, specimens typically with mar-
morated dark pigmentation covering dorsal and
dorsolateral surfaces of head. Scattered pigmen-
tation present on remainder of dorsal and lat-
eral portions of head other than for interoper-
cle. Interopercle with small patch of dark pig-
mentation. Opercular patch of odontodes with
weblike pattern of dark pigmentation around
base of odontodes. Premaxillary teeth, opercu-
lar, and interopercular odontodes unpigmented
even in overall darker individuals.
Body slightly to distinctly dark overall laterally
and dorsally other than for largely unpigment-
ed ventral most portion of caudal peduncle and
rotund patch situated immediately anteroven-
tral to origin of dorsal fin. Some specimens with
horizontal stripe extending anteriorly from an-
teroventral margin of less darkly pigmented
patch located anteroventral of origin of dorsal
fin. Stripe lighter than surrounding regions and
gradually narrowing anteriorly.
Caudal fin rays variably outlined by small dark
chromatophores. Most examined specimens
with dark pigmentation on basal portions of
caudal fin rays more pronounced and forming
slightly anteroventrally angled vertical bar. Dor-
sal fin irregularly marmorated, darker basally.
Anal fin hyaline. Lateral portion of pectoral fin
dark; irregular patches of dark pigmentation
scattered over dorsal surface of pectoral fin
base. Pelvic fin hyaline.
Ecology.—Trichomycterus alterus was collected in
mountainous rivers and streams that were gen-
erally 0.5–4.0 m wide and 0.2–1.0 m deep, with
clear waters running over sandy and rock-peb-
ble substrates, at elevations of approximately
500–2500 m. The stomachs of two cleared-and-
stained specimens of T. alterus contained dipter-
an larvae (Chironomidae and Ceratopogoni-
dae), coleopterans (Elmidae), and ostracod
crustaceans. The diet of autochthonous benthic
macroinvertebrates in T. alterus is common to T.
belensis and many congeners (Casatti and Cas-
tro, 1998; Ferriz, 1998; Fernández and Vari,
2000).
Distribution.—Trichomycterus alterus has been col-
lected at localities in La Rioja, Jujuy, Catamarca,
and Córdoba Provinces of central and north-
western Argentina. The samples from Córdoba
are the first records of the species for that prov-
ince.
746 COPEIA, 2002, NO. 3
Remarks.—We examined the holotype of Pygidi-
um alterum (5 Trichomycterus alterus), three pa-
ratypes collected with it in the Rı́o Los Sauces,
La Rioja Province, and more recently collected
material of the species from a number of local-
ities across the central and northwestern por-
tions of the Argentina. The more recently col-
lected specimens cover a much greater size
range (10.5–49.6 mm SL) than the four speci-
mens (22.9–27.2 mm SL) available to Marini et
al. (1933). Comparisons of details of meristics,
morphometrics, dentition, and other features
indicate that the only difference between the
specimens reported on by Marini et al. (1933)
and the more recently collected specimens in-
volves the degree of development of dark col-
oration on the head and body. The holotype
and paratypes of T. alterus, although well pre-
served, lack a definite pigmentation pattern and
were cited by Marini et al. (1933:3) as being
‘‘pale, unmarked.’’ As noted, smaller individuals
of T. alterus also lack the dark pigmentation that
is present in most examined specimens of the
species. In the absence of any other difference
between the type series and the more of T. al-
terus recently collected samples of that species
it is most reasonable to assume that the samples
are conspecific.
Although Marini et al. (1933) reported lower
numbers of interopercular and opercular odon-
todes in T. alterus than observed herein, their
counts were based on the examination of whole
specimens in which it is difficult to determine
the number of such elements. The data pre-
sented herein were taken from cleared-and-
counterstained specimens from which it is pos-
sible to determine the exact numbers of odon-
todes.
Ringuelet et al. (1967:352) considered T. al-
terus to be a member of the subunit of Tri-
chomycterus characterized by the possession of
papillae. Arratia and Menu-Marque (1984), in
contrast, did not find papillae on the skin of the
holotype or paratypes of T. alterus. They also
noted that Marini et al. (1933) did not mention
the presence of such papillae in their descrip-
tion of the species. Examination of the holotype
of T. alterus, in light of the discovery of papillae
in larger nontype specimens of this species, has
revealed that papillae are present in that speci-
men, albeit very small and, thus, difficult to see.
An ontogenetic increase in the proportional
size of the papillae similarly occurs in nontype
material of that species. A similar ontogenetic
increase in the proportional size of the papillae
occurs in T. boylei, T. corduvensis, and T. spegaz-
zinii in which small-sized specimens have rela-
tively small papillae.
COMPARATIVE MATERIAL EXAMINED
Specimens which served as the basis for the
comparative observations in this paper were list-
ed in Fernández and Vari (2000) with the ad-
dition of Trichomycterus ramosus, FML 2070, 1,
63.1 mm SL (holotype), FML 2071, 11 58.9–66.8
mm SL (paratypes) and FML 2106, 40, 22.3–
42.6 mm SL; Argentina, Catamarca, Departa-
mento Belén.
ACKNOWLEDGMENTS
Research associated with this project was sup-
ported by funds from the Neotropical Lowland
Research Program of the Smithsonian Institu-
tion (RPV) and a fellowship from the Consejo
Nacional Investigacion Cientifica y Tecnica
(LAF). The expedition which collected the type
series of Trichomycterus belensis was funded by the
Fundacion Miguel Lillo. Figures 1 and 2 were
prepared by T. B. Griswold. This paper benefit-
ted from the comments and suggestions of C. J.
Ferraris Jr., T. A. Munroe, R. E. Reis, and S. H.
Weitzman.
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(LF) CONICET-FUNDACIÓN MIGUEL LILLO, DI-
VISION ICTIOLOGı́A, MIGUEL LILLO 251, 4000
TUCUMÁN, ARGENTINA; AND (RPV) DEPART-
MENT OF SYSTEMATIC BIOLOGY (FISHES), NA-
TIONAL MUSEUM OF NATURAL HISTORY, SMITH-
SONIAN INSTITUTION, WASHINGTON, DC
20560–0159. E-mail: (LF) majose@unt.edu.ar;
and (RPV) vari.richard@nmnh.si.edu. Send
reprint requests to RPV. Submitted: 5 Oct.
2001. Accepted: 8 March 2002. Section editor:
S. A. Schaefer.