q 2004 by the American Society of Ichthyologists and Herpetologists
Copeia, 2004(4), pp. 876?882
New Species of Trichomycterus from Midelevation Localities of
Northwestern Argentina (Siluriformes: Trichomycteridae)
LUIS FERNA? NDEZ AND RICHARD P. VARI
Trichomycterus pseudosilvinichthys, new species, is described from midelevation
drainages in the Provincia de La Rioja, Argentina. The new species is distinguished
from other members of the apparently nonmonophyletic genus Trichomycterus by
having the insertion of the ?rst proximal dorsal-?n pterygiophore at, or posterior
to, vertebra 20 to 22; the presence of the pelvic ?n and girdle; a fronto-lachrymal
tendon bone with a lateral expansion but an incomplete laterosensory canal seg-
ment; the absence of a portion of the supraorbital laterosensory canal running be-
tween the frontal and nasal bones with the resultant loss of pore 3; the possession
of 17?19 ribs; a ?rst pectoral-?n ray that is not prolonged as a short ?lament; and
the tip of the pelvic ?n falling short of the anus.
Trichomycterus pseudosilvinichthys, nueva especie, es descripta para una cuenca de
mediana altura de la Provincia de La Rioja, Argentina. La nueva especie se diferen-
cia de los otros miembros del aparentemente ge?nero no mono?le?tico Trichomycterus
por tener la insercio?n del primer pterigio?foro proximal de la aleta dorsal en o
posterior a las ve?rtebras 20 a 22; la presencia de la aleta y cintura pe?lvica; el hueso
fronto-lachrimal con una expansio?n lateral y un segmento del canal incompleto, la
ausencia de la porcio?n del canal laterosensorial supraorbital entre los huesos frontal
y nasal con la pe?rdida resultante del poro 3; la posesio?n de 17 a 19 costillas; el
primer radio de la aleta pectoral que no esta? prolongado como un corto ?lamento
y el extremo de la aleta pe?lvica pro?ximo al ano.
THE nearly 90 species of the cat?sh genusTrichomycterus inhabit a diversity of habitats
ranging from Amazonian lowlands to Andean
uplands and occur across broad reaches of
South and southern Central America (de Pinna
and Wosiacki, 2003). Ichthyological collecting
during recent years have yielded a number of
species of Trichomycterus previously unknown to
science from various mid- to high-elevation lo-
calities in western Argentina (Ferna?ndez, 2000,
2001; Ferna?ndez and Vari, 2002; and references
therein). In those upland regions the species of
Trichomycterus are among the few, or sometimes
only, ?shes occupying water bodies at middle to
higher elevations. Collecting efforts in Provincia
de La Rioja, western Argentina in midelevation
river systems that are severely adversely affected
by erosion resulting from mineral ore extrac-
tion projects yielded a species of Trichomycterus
previously unknown to science that we herein
formally describe.
MATERIALS AND METHODS
Measurements were taken from the left side
of the specimens with digital calipers under a
binocular microscope following the methods
outlined by Tchernavin (1944) and de Pinna
(1992). Cleared-and-counterstained specimens
(C&S) for osteological study were prepared fol-
lowing the procedure of Taylor and Van Dyke
(1985). Osteological nomenclature follows Bas-
kin (1973) and de Pinna (1989, 1998), and the
numbering system and terminology for latero-
sensory pores of the head follows Arratia and
Huaquin (1995) and Arratia (1998). Counts of
dorsal- and anal-?n rays follow the methods pro-
posed by de Pinna (1992). As proposed by de
Pinna (1992), the vertebral counts exclude the
vertebrae in the Weberian apparatus and the
compound caudal centrum is counted as one
element. Counts of caudal vertebrae follow Fer-
na?ndez and Schaefer (2003). Institutional ab-
breviations are as listed in Leviton et al. (1985).
Elevations are given in meters above sea level
(asl).
Trichomycterus pseudosilvinichthys, n. sp.
Figures 1?2, Table 1
Holotype.?FML 2588, 61.1 mm SL; Argentina,
Provincia de La Rioja, Departamento Chilecito,
R??o Amarillo at Famatina, a small assemblage of
houses near Fundicio?n de Oro Santa Florentina
(288559S, 678319W), on east slope of Sierra de
Famatina, collected by L. Ferna?ndez, R. Mon-
tero, and J. Scroci, 8 September 1994.
877FERNA? NDEZ AND VARI?ARGENTINIAN TRICHOMYCTERUS
Fig. 1. Trichomycterus pseudosilvinichthys, new species, holotype, FML 2588, 61.1 mm SL, left lateral view.
TABLE 1. MORPHOMETRIC DATA FOR HOLOTYPE AND 29 PARATYPES OF Trichomycterus pseudosilvinichthys, NEW SPE-
CIES. Standard length is expressed in millimeters; measurements 1?11 are percentages of standard length; 12
to 17 are percentages of head length.
Holotypes
Paratypes
Range Mean 6 SD
Standard length
1. Body depth
2. Caudal peduncle length
3. Caudal peduncle depth
4. Predorsal length
61.06
17.29
20.60
9.89
66.46
29.4?66.9
10.4?18.0
17.9?23.4
7.5?10.5
60.1?68.0
49.3 6 10.1
15.5 6 1.9
20.7 6 1.3
9.3 6 0.7
65.9 6 1.5
5. Preanal length
6. Prepelvic length
7. Dorsal-?n base length
8. Anal-?n base length
9. Head length
70.55
56.93
11.04
7.63
17.56
67.1?81.8
53.5?59.3
9.9?12.4
6.8?10.0
15.8?20.2
71.6 6 5.6
56.5 6 1.4
10.7 6 0.8
8.7 6 0.7
17.9 6 1.1
10. Head width
11. Head depth
12. Interorbital width
13. Snout length
14. Nasal barbel length
15.72
9.14
26.12
45.52
45.34
14.4?18.4
7.1?11.2
21.8?33.1
41.2?49.1
35.6?67.2
16.5 6 1.1
9.5 6 0.9
25.1 6 2.2
45.5 6 2.0
51.6 6 9.7
15. Maxillary barbel length
16. Submaxillary barbel length
17. Mouth width
74.07
52.24
48.88
44.7?90.9
34.0?57.7
33.8?47.6
68.6 6 12.1
45.1 6 6.3
40.6 6 3.8
Paratypes.?Twenty-nine specimens, 29.4?66.9
mm SL. All from Argentina, Provincia de La
Rioja. FML 2589, 9, 54.5?62.2 mm SL (1, 62.9
mm SL C&S); USNM 374759, 4, 43.5?52.9 mm
SL (1, 61.3 mm SL C&S), collected with holo-
type. FML 2558, 6, 29.4?42.6 mm SL (2, 35.6?
41.0 mm SL C&S), Departamento General La-
madrid, near Puerto Alegre (298269S, 678579W),
collected by L. Ferna?ndez and R. Montero, Sep-
tember 1994. FML 2595, 2, 36.9?48.8 mm SL;
CAS 218432, 1, 49.6 mm SL; FMNH 112974, 1,
52.8 mm SL; Departamento General Lamadrid,
near Puerto Alegre (298269S, 678579S), collected
by L. Ferna?ndez and R. Montero, September
1994. AMNH 233620, 3, 45.1?54.8 mm SL; De-
partamento Chilecito, Cuesta de Miranda
(298209270S, 678479310W) 1,850 m, collected by
M. Archangelsky and F. Cruz, 3 October 1998.
MLP 7370, 3, 57.2?66.9 mm SL, Departamento
Chilecito, Valle Guanchin (298119S, 678399W),
collected February 1962.
Nontype specimens.?Eighty-seven specimens,
28.9?53.1 mm SL. All from Argentina, Provincia
de La Rioja. FML 2103, 34, 39.5?53.1 mm SL,
collected with holotype. FML 2116, 53, 28.9?
38.8 mm SL, Departamento General Lamadrid,
near Puerto Alegre (298269S, 678579W), collect-
ed with FML 2558, part of paratype series.
Diagnosis.?The combination of the presence of
the pelvic ?ns and a pelvic girdle, the possession
of a rectangular premaxilla, seven branched
dorsal-?n rays, 17?19 ribs, the form of the body
and caudal ?n, the lack of a very thick, rugose
layer of fatty tissue on the body and head, the
extension of a portion of the laterosensory ca-
nal system through the sphenotic, the reduction
of the segment of the portion of the laterosen-
sory canal within the frontal with a loss of a seg-
ment of the canal between pores 2 and 6 with
the resultant absence of intervening pore, the
lack of an extensive perforation of the skin sur-
878 COPEIA, 2004, NO. 4
face by ampullary organs, the termination of
the ?rst pectoral-?n ray at the margin of the ?n,
details of the pigmentation on the body and
?ns, and the overall body size of T. pseudosilvi-
nichthys separates that species from all other
known members of the subfamily Trichomycte-
rinae. Trichomycterus pseudosilvinichthys further
differs from all congeners from southern South
America, with the exception of Trichomycterus
areolatus, Trichomycterus catamarcensis, Trichomyc-
terus chiltoni, Trichomycterus vittatus, and Tri-
chomycterus yuska, in having the insertion of the
?rst proximal dorsal-?n pterygiophore located
at, or posterior to, the neural spine of vertebrae
20 to 22 (vs inserting posterior to vertebra 15
to 19). Trichomycterus pseudosilvinichthys can be
distinguished from T. areolatus by the presence
of a fronto-lachrymal tendon bone (the supra-
orbital of Arratia, 1998) with a lateral expansion
and an incomplete supraorbital laterosensory
canal segment in that bone with the consequent
loss of pore 3 (vs a fronto-lachrymal tendon
bone without a lateral expansion but incorpo-
rating a complete laterosensory canal segment),
from T. catamarcensis by the presence of the pel-
vic ?n and girdle (versus the absence of both
structures), from T. chiltoni by the presence of
a lateral expansion on the fronto-lachrymal ten-
don bone and the possession of 17?19 ribs (vs
the absence of a lateral expansion and the pres-
ence of 10?15 ribs), from T. vittatus in having
the ?rst pectoral-?n ray terminating at the mar-
gin of the ?n and in having the tip of the pelvic
?n falling short of the anus (vs having the ?rst
pectoral ?n ray prolonged as a short distal ?la-
ment and having the tip of the pelvic ?n reach-
ing to the anus), and from T. yuska in having
the ?rst pectoral-?n ray terminating at the mar-
gin of the ?n (versus having the ?rst pectoral
?n ray prolonged as a short distal ?lament). In
addition to the internal features listed above, T.
pseudosilvinichthys further differs from the synto-
pic T. corduvensis in having the pelvic ?n falling
short of the anal opening (vs reaching the rear
of the opening) and in the possession of 1 or 2
pores on the anterior most portion of the lat-
eral line (vs 3 or 4 pores).
Description.?Descriptive morphometric features
for holotype and paratypes of T. pseudosilvi-
nichthys provided in Table 1. Body elongate, ap-
proximately cylindrical but slightly compressed
transversely in trunk region, gradually and pro-
gressively becoming more compressed trans-
versely toward caudal ?n. Dorsal and ventral
pro?les of trunk region ranging from nearly
straight to slightly convex or concave. Caudal
peduncle smoothly continuous with dorsal and
ventral pro?les of trunk. Small papilla-like struc-
tures present on body. Specimens originating in
R??o Amarillo at Puerto Alegre and Cuesta Mi-
randa with distinctly larger amounts of subcu-
taneous fat on head and body than present in
other populations samples of species. Urogeni-
tal opening closer to anal-?n origin than to pel-
vic-?n insertion.
Head dorsoventrally ?attened and obtusely
pointed in pro?le. Head pro?le in dorsal view
rectangular. Eyes located on dorsal surface of
head but visible in both lateral and dorsal views.
Eyes ovoid and slightly anteroposteriorly elon-
gate. Skin covering eye thin, transparent, and
separate from surface of eyeball, with eyes read-
ily visible.
Anterior nostril slightly smaller than posterior
nostril and surrounded by ?eshy integument
?ap medially and by base of nasal barbel later-
ally. Anterior margin of posterior nostril bor-
dered by ?ap of thin skin.
Infraorbital canal incomplete (Fig. 2). Ante-
rior segment absent with loss of infraorbital
pores 1 and 3. Posterior portion of infraorbital
canal situated posterior to rim of orbit and aris-
ing from temporal canal running within sphe-
notic. Posterior portion of infraorbital canal
with infraorbital pores 10 and 11. Supraorbital
laterosensory canal segment incomplete with
short anterior nasal canal separated from re-
duced portion of canal located in posterior por-
tion of frontal by distance approximately equal
to horizontal length of lateral ethmoid, and
with only several pores (pore 3 absent). Epiph-
yseal branch of supraorbital canal discontinuous
with pair of median pores posterior to eye.
Preoperculo-mandibular canal represented by
reduced preopercular canal with one pore on
surface of skin. Postotic canal with pterotic
branch (sensu Schaefer and Aquino, 2000) pre-
sent at junction of pterotic and posttemporal-
supracleithrum. Laterosensory canal along mid-
lateral portion of trunk reduced, without ossi-
?ed tubules and limited to pores 11 and 12 of
anterior most portion of lateral line.
Mouth distinctly subterminal, with rictus di-
rected posteriorly. Premaxilla rectangular with
width of bone greater than maximum dimen-
sion of palatine. Premaxilla with three or four
rows of distally narrowing, incisiform teeth; with
seven to 10 teeth in outer tooth row. Dentary
with one to four rows of distally narrowing, in-
cisiform teeth; with 10?13 teeth in outer tooth
row. Lower lip with prominent ?eshy lobes
along lateral limits; lobes situated medial to
base of rictal barbels. Lower lip ?eshy anteriorly,
with anterior, and to lesser degree, anteroven-
879FERNA? NDEZ AND VARI?ARGENTINIAN TRICHOMYCTERUS
Fig. 2. Dorsal view of the neurocranium and associated structures of Trichomycterus pseudosilvinichthys show-
ing laterosensory canal system on left side of head. Abbreviations: ep 5 epioccopital; f 5 frontal; ff 5 frontal
fontanelle; i10, 11 5 pores 10 and 11 of infraorbital sensory canal; lat. l 5 lateral line; le 5 lateral ethmoid;
me 5 mesethmoid; pr 5 preopercular pore; ptts 5 posttemporal-supracleithrum; pt 5 pterotic; pt. br 5
pterotic branch of postotic sensory canal; s1, 2, 6 5 pores 1, 2, and 6 of supraorbital sensory canal; s 1 pro
1 ptr 5 sphenotic 1 prootic 1 pterosphenoid; so 5 supraorbital; su 5 supraoccipital; and w 5 weberian
complex and capsule.
880 COPEIA, 2004, NO. 4
tral surfaces covered with papillae. Upper lip
?eshy and bearing numerous papillae.
Barbels relatively short and tapering distally
but not threadlike or with distal branching pre-
sent in some congeners. Tips of maxillary bar-
bels extending posteriorly to middle of patch of
interopercular odontodes but falling short of
pectoral-?n origin. Nasal barbels reaching pos-
teriorly to distinctly beyond posterior border of
eye. Submaxillary barbels shorter than maxil-
lary barbels.
Branchiostegal rays seven. Interopercular
odontode patch anteroposteriorly elongate;
with odontodes imbedded in ?eshy covering of
interopercle; 32?42 odontodes present on patch
in four C&S specimens. Opercular patch of
odontodes small, rounded, and slightly antero-
posteriorly elongate, with odontodes imbedded
in ?eshy covering of opercle, 7?11 odontodes
present on patch in four C&S specimens.
Dorsal-?n rays 12 [12] or 13 [4], with seven
branched and ?ve [8] or six [8] unbranched
rays. Distal margin of dorsal ?n semicircular
when ?n expanded. Dorsal ?n ?eshy basally.
Dorsal-?n origin located slightly anterior to ver-
tical through anterior limit of vent. First proxi-
mal dorsal-?n pterygiophore inserting posterior
to vertebra 20 [6], 21 [6], or 22 [3]. Anal-?n
rays 11 [7] or 12 [8], with six [7] or seven [8]
branched and ?ve unbranched rays. Anal ?n
relatively elongate, equal in size to, or slightly
smaller than, dorsal ?n, with distal margin of
?n slightly rounded. Anal-?n origin approxi-
mately at vertical through middle of base of dor-
sal ?n. Pectoral-?n rays eight, with lateral most
ray unbranched. Distal margin of pectoral ?n
slightly convex. First pectoral-?n ray terminat-
ing at ?n margin without forming distal ?la-
ment. Pelvic-?n rays ?ve, plus small splint an-
terior to ?rst unbranched ray; second and third
rays longest. Tip of pelvic ?n falling distinctly
short of anal opening. First proximal anal-?n
pterygiophore inserting posterior to vertebra
23. Distal margin of caudal ?n nearly straight
but with outer ?n rays slightly shorter than mid-
dle rays. Principal caudal-?n rays 6 1 6 or 6 1
7. Dorsal procurrent caudal-?n rays 12 [1], 13
[1], 14 [1], or 15 [1] and ventral procurrent
caudal-?n rays 12 [2] or 13 [2] in four C&S
specimens.
Total vertebrae 37 [1], 38 [1], 39 [1], or 40
[1], with 9 [1], 10 [2], or 11 [1] precaudal and
26 [1], 27 [1], 28 [1], or 29 [1] caudal verte-
brae, and 17 [1], 18 [2], or 19 [1] ribs on each
side in four C&S specimens.
No external obvious sexual dimorphism ob-
served in available population samples. Popu-
lation samples from R??o Amarillo with cysts on
body and ?ns, perhaps as a consequence of pol-
lution resulting from upstream mining opera-
tions.
Color in alcohol.?Trichomycterus pseudosilvi-
nichthys demonstrates considerable variation in
intensity of dark pigmentation on head and
body. Holotype (Fig. 1) and most of paratypes
with pattern of distinct, albeit faint, marmora-
tion formed by patches of small, dark chro-
matophores. Several more lightly pigmented pa-
ratypes lack distinct marmoration on head and
body. Darkly pigmented specimens with mar-
morated, dark pigmentation present on dorsal
and dorsolateral surfaces of head and trunk and
all but ventral most portion of caudal peduncle.
Ventral surface of head ranging from lightly pig-
mented to hyaline. All barbels pigmented at
least to some degree; less so in overall less dark-
ly pigmented individuals. Dorsal, anal, and pec-
toral ?ns with irregular, dark pigmentation pre-
sent on rays and membranes. Caudal ?n with
membranes irregularly darker than membranes
of other unpaired ?ns. Pelvic ?n ranging from
lightly pigmented to hyaline. Opercular and in-
teropercular odontodes and oral dentition un-
pigmented even in overall more darkly pig-
mented individuals.
Relationships.?The derived reduction in the de-
gree of development of the supraorbital canal
within the frontal of T. pseudosilvinichthys is
unique to that species and T. yuska, another Ar-
gentinean species, among congeners that have
been examined osteologically. The common
possession of this feature may indicate that
these are sister species, but con?rmation of that
hypothesis requires a broader comparative anal-
ysis incorporating information from multiple
character systems.
Ecology.?The type locality of T. pseudosilvi-
nichthys was a small, clear water stream with a
rock and pebble bottom at an elevation of ap-
proximately 1800 m. The only other species of
?sh collected at that site was T. corduvensis.
Distribution.?Trichomycterus pseudosilvinichthys is
known from various localities in Departamento
Chilecito and Departamento General Lama-
drid, Provincia de La Rioja, western Argentina.
Etymology.?The speci?c name, pseudosilvi-
nichthys, is in reference to the similarity in the
external appearance of the new species and the
trichomycterid genus Silvinichthys.
881FERNA? NDEZ AND VARI?ARGENTINIAN TRICHOMYCTERUS
Remarks.?Trichomycterus pseudosilvinichthys is
very similar in terms of external features to the
trichomycterid Silvinichthys mendozensis, a species
that also occurs in western Argentina (Arratia
et al. 1978). The two species differ most tren-
chantly in the presence of a laterosensory canal
segment in the sphenotic of T. pseudosilvinichthys
in contrast to the absence of that canal segment
in Silvinichthys mendozensis.
Discussion.?Berg (1897) described Trichomycte-
rus riojanus based on one specimen from an in-
de?nite locality described as an ``arroyo of the
Cordillera in [Provincia de] La Rioja'' (our
translation; addition in square brackets ours).
The original description of T. riojanus is, unfor-
tunately, uninformative as to a number of tax-
onomically important morphological features.
Furthermore, the holotype (MACN 5175) is
now dried and in very poor condition making
it impossible to either con?rm much of the data
presented in the original description of that
species, or unequivocally compare the meristic
and morphometric features of the holotype of
T. riojanus with those of T. pseudosilvinichthys. Al-
though all known specimens of both T. riojanus
and T. pseudosilvinichthys originated in the Pro-
vincia de La Rioja, the two nominal species
clearly differ in the proportional length of the
pelvic ?n (tip of ?n reaching posteriorly to the
anus in the holotype of T. riojanus versus dis-
tinctly falling short of that landmark in T. pseu-
dosilvinichthys) and in the degree of develop-
ment of the ?rst ray of the pectoral ?n (ray ex-
tending beyond the margin of the ?n as a short
distal ?lament versus terminating at the margin
of the ?n, respectively). We consequently rec-
ognize them as distinct species.
Comparative material.?Includes material cited in
Ferna?ndez (2000) and Ferna?ndez and Vari
(2000) with the addition of the following: Tri-
chomycterus alterus, FML 2087, 10 (3 C&S); FML
2088, 7 (2 C&S). Trichomycterus atochae, CAS
64576, holotype. Trichomycterus banneaui, FMNH
56025, holotype; FMNH 70014, 2. Trichomycterus
belensis, FML 2530, holotype, FML 2533, 5
(C&S). Trichomycterus boylei, KU 20188, 1. Tri-
chomycterus caliense, FMNH 56029, holotype;
FMNH 70338, 1. Trichomycterus catamarcensis,
FML 2507, holotype; FML 2509, 10 (2 C&S).
Trichomycterus chaberti, ANSP 140068, 1 paratype.
Trichomycterus chapmani, CAS 58128, 1 paratype.
Trichomycterus chungaraensis, KU 19218, 2; KU
19392, 2 (C&S). Trichomycterus dispar, ANSP
21174, 4. Trichomycterus duellmai, KU 20194, 4 (2
C&S). Trichomycterus eigenmanni, MCZ 8301, ho-
lotype (radiograph). Trichomycterus fassli, USNM
302757, 1 (1 C&S). Trichomycterus gabrieli, CAS
64583, 1 syntype; SU 36556, 1 syntype. Trichomyc-
terus immaculatus, MCZ 8300, 1 syntype (radio-
graph). Trichomycterus laucaensis, KU 19403, 2
(C&S), KU 19404, 1 (C&S). Trichomycterus ma-
racaiboensis, AMNH 91133, 1 (C&S). Trichomyc-
terus nigromaculatus, UMMZ 187674, 2. Tri-
chomycterus punctulatus, FMNH 58672, 1 paraty-
pe. Trichomycterus ramosus, FML 2070, holotype;
FML 2071, 5 paratypes (2 C&S). T. romeroi,
ANSP 69331, holotype; ANSP 69332, 2 paraty-
pes. Trichomycterus schmidti, MACN 5176, 1
syntype; MACN 5174, 2; MACN 4595, 1. Tri-
chomycterus tiraquae, ANSP 69126, holotype;
UMMZ 204202, 4; AMNH 39740, 2 (C&S). Tri-
chomycterus vittatus, ANSP 149683, 3 (1 C&S).
Trichomycterus weyrauchi, ANSP 71639, holotype.
Trichomycterus yuska, FML, 2535, holotype; FML
1132, 12 paratypes (2 C&S). Trichomycterus zo-
natus, UMMZ 231757, 4.
ACKNOWLEDGMENTS
The expedition that collected the type series
of Trichomycterus pseudosilvinichthys was funded
by the Fundacion Miguel Lillo and Consejo Na-
cional Investigacion Cienti?ca y Tecnica. The
latter and the Neotropical Lowland Research
Program funded the research that was the basis
of study. For the loan of specimens and other
assistance, we thank S. A. Schaefer, B. Brown,
M. Sabaj, D. Catania, M. A. Rogers, A. Bentley,
O. T. Oyakawa, and D. Nelson. Access to the
holotype of Trichomycterus riojanus was provided
by L. Braga. S. J. Raredon prepared Figure 1
and radiographs. This paper bene?tted from
the comments and suggestions of S. H. Weitz-
man and T. A Munroe.
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(LF) FUNDACION MIGUEL LILLO, DEPARTAMENTO
ZOOLOG??A, DIVISION ICTIOLOG??A, MIGUEL LIL-
LO 251, 4000 TUCUMA? N, ARGENTINA; AND
(RPV) DIVISION OF FISHES, SMITHSONIAN IN-
STITUTION, P.O. BOX 37012, NATIONAL MUSE-
UM OF NATURAL HISTORY, WG-14, MRC-159,
WASHINGTON, DC 20013-7012. E-mail: (RPV)
vari.richard@nmnh.si.edu. Send reprint re-
quests to RPV. Submitted: 25 March 2004. Ac-
cepted: 11 Aug. 2004. Section editor: J. W.
Armbruster.