PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

117(4):564-574. 2004.

Rhamdia guasarensis (Siluriformes: Heptapteridae), a new species of

cave catfish from the Sierra de Perija, northwestern Venezuela

Carlos DoNascimiento, Francisco Provenzano, and John G. Lundberg

(CDN) Seccion de Ictiologia, Museo de Historia Natural La Salle, Fundacion La Salle de

Ciencias Naturales, Apdo. 1930 Caracas 1010- A, Venezuela,
Carlos. donasciiniento@fundacionlasalle. org. ve;
(FP) Laboratorio de Biosistemadca de Feces, Institute de Zoologia Tropical, Universidad Central
de Venezuela, Apdo. 47058 Caracas 1041-A, Venezuela, fprovenz@strix.ciens.ucv.ve;
(JGL) Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin
Parkway, Philadelphia, Pennsylvania 19103, USA, lundberg@acnatsci.org

Abstract. —
Rhamdia guasarensis n. sp. is described from subterranean waters
in the Rio Guasare drainage of northwestern Venezuela. The new species is
distinguished from congeners by its concave head profile; medially sutured
frontal bones; small, circular vestige of the anterior cranial fontanelle; and
troglomorphic characters such as absence of eyes and pigmentation, wide ce-
phalic laterosensory pores, and wide fossae of preoperculomandibular sensory
canal in preopercle and dentary. Cave catfish diversity in the Sierra de Perija
region of Venezuela is reviewed and compared to cave catfish diversity else-
where in South America.
Resumen. —Se describe Rhamdia guasarensis sp. n. proveniente de aguas
subterraneas de la cuenca del Rio Guasare en el noroccidente de Venezuela.
La nueva especie se diferencia de las restantes especies que conforman el
genero por su perfil dorsal de la cabeza concavo; huesos frontales suturados
medialmente; fontanela craneal anterior reducida a un pequeno foramen cir-
cular; y caracteres troglomorficos tales como ausencia de ojos y pigmentacion,
poros cefalicos latero sensoriales anchos, fosas ensanchadas del canal sensorial
preoperculomandibular en el preoperculo y dentario. La diversidad de bagres
cavernicolas de la Sierra de Perija es revisada y comparada con la diversidad
de bagres cavernicolas de otras regiones de Suramerica.

The family Heptapteridae has invaded Trajano, Reis & Bichuette, 2004 is a re-
and adapted to hypogean waters multiple cently described without
troglophile
times. Among Neotropical catfish families, marked specializations for hypogean life,
heptapterids have the greatest diversity of Taxonomic practice has shifted away from
truly Phreatobius cister-
troglobitic taxa: assigning supra-specific rank to cave-dwell-
narum, Pimelodella kronei, Rhamdia lalu- ing fishes solely on account of their trog-
chensis, Rhamdia laticauda typhia, Rham- lobitic adaptations. Among Heptapteridae,
dia macuspanensis, Rhamdia quelen urichi, the nominalmonotypic genera Caecorham-
Rhamdia redelli, and Rhamdia zongolicen- dia, Caecorhamdella, and Typhlobagrus
sis. Trajano & Bockmann (2000) described have long been treated as synonyms of
the ecology and behavior of Taunayia sp., Rhamdia and Pimelodella respectively. Fur-
a troglobitic catfish, inhabiting caves of thermore, Silfvergrip (1996) synonymized
northeastern Brazil, but the species has not all cave populations of Rhamdia described
been formally named. Pimelodella spelea as separate species with R. quelen or R.
 —

VOLUME 1 17, NUMBER 4

laticauda, both wide-ranging epigean spe- scription of Diplomystes and Nematogenys

cies. (1995). However, our numbering of sensory
In this paper we describe a new troglob- pores in Rhamdia reflects anteroposterior or
itic Rham-
heptapterid species in the genus mesiolateral pore order, and does not imply
dia. Our placement of the new species is individual homologies of pores among cat-
more a matter of convenience than firm fishes. All measurements were made on the
phylogenetic resolution. Rhamdia is taxo- left side of the specimens using a Mitutoyo
nomically complex. In the latest revision of digital, needlepoint caliper at a precision of
the genus, Silfvergrip (1996) consolidated 0.1 mm. For osteological observation one
its approximately 100 nominal species into paratype (101.1 mm
SL) was cleared and
eight and he described three new species. stained using the method of Taylor Van &
In 1998, Weber & Wilkens described the Dyke (1985). A second paratype (93.4 mm
blind species R. macuspanensis, and in SL) was radiographed. Only these two
2003, Weber et al. described Rhamdia lal- specimens were used for counts of verte-
uchensis, another troglobitic species from brae, branchiostegal rays, ribs, and ptery-
Mexico. In the most thorough phylogenetic giophores. The vertebral count includes the
study of Heptapteridae to date, Bockmann first five vertebrae incorporated into the
(1998) concluded that Rhamdia is non- Weberian apparatus whereas the compound
monophyletic but he did not attempt to re- caudal centrum is counted as one. Institu-
solve the genus into phylogenetically di- tional abbreviations follow Leviton et al.

agnosable units. As it stands, Rhamdia is a (1985). Other abbreviations are: —

SL stan-
non-monophyletic assemblage of common dard length, HL—head length, —
CS cleared
fishes with an immense geographic distri- and stained skeletal preparation, ale
bution in South and Middle America from whole specimen preserved in alcohol.
the lower Parana Basin in Argentina to cen-
tral Mexico.
Rhamdia guasarensis, new species
The new species, from a cave in the Si- Figs. 1-4
erra de Perija region of northwestern Ve-
nezuela, is distinct both in its typical trog- //o/o/jpe.— MBUCV-V-29604: 106.8
lobitic specializations and other apomorph- mm SL; Surgencia del Tigre at 2.5 km W
ic features, but overall it is most similar to of Cerro Yolanda, Rio Guasare basin. Sierra
other Rhamdia. Discovering the relation- de Perija, Estado Venezuela
Zulia,
ships of the new species and, more gener- (10°52'53"N, 72°30'03"W). Elevation 200
ally, resolving the relationships of Rhamdia m asl; collected by J. Lagarde, 3 April
species are major problems quite beyond 1999.
our present scope. Our immediate concern Paratypes. —All collected with the ho-
is to name and describe this previously un- lotype: MBUCV-V-29622, two specimens,
seen species that has a highly restricted dis- 87.2-101.1 mm SL, the second cleared and
tribution in a marginal and potentially frag- stained; ANSP 179878, one specimen, 93.4
ile habitat. We comment also on the sub- mm SL.
terranean catfish fauna of the Perija region. Diagnosis. —Rhamdia guasarensis is dis-
tinguished from congeneric species by two
Material and Methods characters: dorsal profile of head concave
(Fig. 1, vs. convex or straight); and frontal
Morphometric measurements follow the bones broadly sutured to each other anterior
criteria set out by Lundberg & McDade to small, circular remnant foramen of an-
(1986) and Bockmann (1994). Terminology terior cranial fontanelle that is anteriorly ad-
of cephalic laterosensory canals and jacent to epiphyseal bar (Fig. 2, vs. frontals
branches follows Arratia & Huaquin's de- separated by anterior fontanelle widely
 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Rhamdia giiasarensis. Holotype MBUCV-V-29604, 106.8 i ; B, dorsal view

of head; C, ventral view of head.

open from mesethmoid to epiphyseal bar). eight or nine soft rays in other species, data
Rhamdia guasarensis differs from all epi- from Silfvergrip 1996); both lobes of the
gean Rhamdia by the following troglo- caudal fin pointed (vs. at least one lobe
morphic characters: absence of eyes, com- rounded); caudal skeleton with three hy-
plete depigmentation, widened cutaneous pural plates, PH; 1 + 2; 3 + 4 + 5 (vs.
pores of the cephalic laterosensory system, modally four PH; 1 + 2; 3 + 4; 5 in the
preoperculomandibular sensory canal form- other species with the exception of R. laii-

ing wide fossae in the dentary and preo- kidi and R. jequitinhonha, see Silfvergrip
percle (Fig. 3, vs. narrow pores and canals). 1996).
In addition to these characteristics, R. Description. — Morphometric data are
guasarensis can be distinguished from other presented in Table 1 . Body elongate, strong-
species of the genus by the following com- ly depressed anteriorly and gradually more
bination of characters: pectoral fins with a compressed from origin of pectoral fins to
spine and ten branched rays (vs. modally caudal peduncle. Shape approximately tri-
VOLUME 1 17, NUMBER 4

Fig. 3. Rhamdia guasarensis. Enlarged preoper-

culomandibular laterosensory canal and associated fo-
ramina. MBUCV-V-29622, lOI.l mm SL. Abbrevia-
tions: ANG, anguloarticular; D, dentary; HYO, hy-
omandibula; MPT, metapterygoid; OP, opercle; POP,
preopercle; Q, quadrate.

lower jaw similar to premaxillary teeth, in

six irregular tooth rows. Palatine and vomer
edentulous. Maxillary barbels long, extend-
ing beyond base of pelvic fins. Mental bar-
bels relatively short, inner mentals scarcely
reaching posterior border of branchial
membrane; outer mentals surpass pectoral
fin bases. Inner mental barbel bases inserted

slightly in advance of outer mental barbel

bases. Anterior nares tubular, near border of
Fig. 2. Rhamdia guasarensis. Skull roof illustrat- snout. Posterior nares with elongated orific-
ing reduction of the anterior cranial fontanelle and
es, bounded anterolaterally by membrane of
midline contact of frontal bones. MBUCV-V-29622,
fine skin. Internarial length less than width
101.1 mm SL. Abbreviations: ACF, anterior cranial
fontanelle; FR, bone contact on midline; LET,
frontal between posterior nares. Eyes completely
lateral ethmoid; MES, mesethmoid; PCF, posterior cra- absent. Branchial membranes overlapping
nial fontanelle; PT, pterotic; SPH, sphenotic. medially; united to isthmus only anteriorly.
Cephalic lateralis sensory system with
paired supraorbital (SO), infraorbital (lO),
angular in transverse section at dorsal-fin preopercular (POP), mandibular (MA), otic
origin. Dorsal profile sinusoidal anterior to (OT), and post-otic (POT) canals, without
dorsal fin, then approximately straight to tubular commissure connecting supraorbital
middle of adipose fin, then slightly concave canals. Sensory pores simple, not branched
along caudal peduncle. Ventral profile near- and multiple. SO canal with six pores:
ly straight to anal-fin origin, then slightly S01-S03 associated with nasal bone, SOI
concave posteriorly. medially adjacent to anterior naris, wide
Head depressed, its dorsal profile con- and delimited by membrane of fine skin,
cave, its lateral and ventral profiles nearly S02 between anterior and posterior nares,
straight. Mouth terminal, upper jaw slightly slightly closer to first,S03 posteromedially
in advance of lower jaw. Rictal folds little near posterior naris. S04 near dorsal mid-
developed. Upper and lower lips with weak line at end of short medial tube and separate
sulci, slightly evident in holotype, forming from its counterpart of opposite side. SOS
single labial fold. Premaxillaries with single lateral to its canal midway between S04
band of diminutive teeth, arranged in ten and union of SO and lO canals. S06 medial
irregular tooth rows, the posterolateral cor- to its canal a little posterior to union of SO
ners rounded, not produced. Dentition of and lO canals.
 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1
.
—Measurement data for the type series of Rhamdia guasarensis. Measurement 1 expressed in mm.
Proportional measurements expressed as thousandths of standard length (2-19; 26-28) or head length (20-25).

MBUCV-V-29622 ANSP 179878

. Standard length 106.8 87.2 101.1 93.

. Total length 1057 1093 1083
. Body depth 162 178 167
. Body width 171 180 168 173
. Predorsal length 351 367 352 359
. Preanal length 645 653 659 656
. Prepelvic length 462 458 490
. Preadipose length 555 545 530
. Caudal peduncle length 211 210 214 224
. Caudal peduncle depth
. Dorsal fin spine length
. Length of first branched dorsal fin ray
. Dorsal fin base
. Adipose fin length
. Dorsal fin to adipose fin

. Anal fin base

. Pectoral fin spine length
. Length of first branched pectoral fin ray
. Pelvic fin length
. Head length 262
. Head width 645 633
. Head depth 522 528
. Internarial length
. Anterior internarial width
. Posterior internarial width
. Maxillary barbel length 610 557 578
. Outer mental barbel length 262 258 262
. Inner mental barbel length

lO canal with four pores; I01-3 wide POT canal with two pores, POTl over
like SOI. lOl posterior to anterior nostril; pterotic dorsal to gill opening; POT2 dorsal
I02 emerges dorsal to groove for maxillary to supracleithrum and above main lateralis
barbel, posterior to base of barbel; I03 near canal at level of first pore. First pore of la-
point where lO canal curves dorsally; I04 teralis canal at end of ventral branch dorsal
at tip of short posterior tube near union with to postcleithral process. Several following
SO Holotype and one paratype (87.2
canal. pores also at tips of short postero-ventral
mm SL) have different single supernumer- branches. Lateral line canal complete to
ary lO pores; extra pore of holotype from base of middle upper-lobe caudal rays.
left canal between the I02 and I03; extra Dorsal fin with a spinelet, spine and six
pore of paratype from right lO canal be- branched rays; its margin rounded. Dorsal
tween I03 and I04. spine weakly developed, only its basal part
POP canal with four pores; MA canal rigid and unsegmented; dentations diminu-
with seven pores; all except and MAI tive and scarcely visible, limited to basal
POP4 originate from much enlarged cavi- part of anterior margin. The distal two-
ties in dentary and preopercular bones. thirds of dorsal spine flexible and obliquely
MAI in mental position near to midventral segmented. Adipose fin long and low, its
line at tip of its branch from lower jaw sym- origin near tip of depressed dorsal fin, and
physis. extending posteriorly to approximately 80%
VOLUME 117, NUMBER 4

Fig. Rhamdia giiasarensis. Pectoral spine in

dorsal \ w. Holotype MBUCV-V-29604, 106.8 mm

of caudal peduncle length; posterior end of

adipose fin adnate to caudal peduncle with-
out a free fleshly tab. Caudal fin deeply
forked; both caudal lobes pointed; upper
lobe slightly longer than lower; membrane
uniting innermost caudal rays complete.
Principal caudal rays i,7-8,i, except i,7-7,i
in one paratype. Anal fin with 12 rays, an-
teriormost two or three rays simple, others
branched; margin rounded.
its

Pectoral fins with a spine and ten

branched rays. Pectoral spine (Fig. 4) with Fig. 5. The Lago de Maracaibo —
Sierra de Perija
region, showing type locality (star) of
Venezuela,
weak dentations proximally on anterior Rhamdia Map based on shaded relief im-
giiasarensis.
margin; distal half of spine flexible and age PIA03388, Shuttle Radar Topography Mission,
obliquely segmented. First branched pec- National Aeronautics and Space Administration
toral-fin ray longest, posterior branched (NASA).
rays diminishing in length. Postcleithral
process short, sharp. Pelvic fins with one
simple ray and five branched rays, its origin giophores preceded by small supraneural;
posterior to end of dorsal fin. first dorsal-fin pterygiophore inserted be-
Skull roof (Fig. 2) with anterior cranial tween rami of neural spine of fourth ver-
fontanelle extremely reduced to a small cir- tebra. Eleven anal-fin pterygiophores, first
cular foramen located in front of epiphyseal inserting posterior to hemal spine of verte-
bar; mesethmoid posteriorly lacking con- bra 21. Caudal skeleton with three hypural
cave notch of fontanelle, and frontals meet- plates: rectangular parhypural; triangular
ing medially along most of their length. hypurals 1-1-2; triangular hypurals 3 + 4
Posterior cranial fontanelle reduced to oval + 5.

foramen near center of supraoccipital. Su- Color in alcohol. —

Body and fins com-
praoccipital process short, its length about pletely depigmented; most of skin, rayed-
equal to length of supraoccipital body. and adipose-fin membranes hyaline and
Anus and urogenital papillae separated, translucent; musculature appearing yellow-
anus located equidistant between medial ish, particularly jaw adductors and dorsal
edge of pelvic-fin base and urogenital pa- trunk myomeres; parts of head and fin bases
pilla, approximately at midlength along pel- whitish.
vic fins; urogenital papilla conspicuous and Distribution and habitat. — R. guasaren-
elongated, located closer to base of anal fin sis is known only from the Surgencia del
than to base of pelvic fin. Tigre (Zu. 23), in the middle basin of the
Total vertebrae 40-42; neural spines of Rio Guasare, north of the Sierra de Perija
vertebrae 6-10 bifid; hemal arch closed in in northwestern Venezuela (Fig. 5). The
vertebra 12 or 13, first hemal spine on ver- cave is near the margin of Rio Guasare and
tebra 14, 15 or 16; eight pairs of ribs borne is the source of a spring during seasonal
on vertebrae 6-13. Seven dorsal-fin ptery- rains (Sociedad Venezolana de Espeleologia
 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1991). The cave's lower conduit has a 280
m course, 2-3 m wide and 1-2
rowly communicating with the access gal-
lery. The underground river is permanently
fed by a spring about 60 m
into the lower
gallery. At the time the cave was surveyed,
the average depth of this water course was
1.5 m with deeper pools along its course
where the catfishes were observed (Socie-
dad Venezolana de Espeleologia 1991).
Etymology. —
The name is based on Rio
Guasare, parent stream of the subterranean
waters in which this endemic catfish species
lives.
higher level synapomorphies, and others,
m high, neu-- some of uncertain polarity, have wider and
among heptapterids.
variable distributions
Thus, placement of this new species in
Rhamdia must be considered provisional
because the genus has not been supported
as monophyletic. Bockmann's (1998) phy-
logenetic analysis of Heptapteridae placed
one representative species, R. laticauda,
sister to Pimelodella but a second species,
R. quelen, is deeper in his cladograms. In
this context R. guasarensis has one derived,
although non-unique, feature listed by
Bockmann as diagnostic for R. quelen. This
is the highly reduced posterior cranial fon-

Discussion tanelle, long used as one of the diagnostic

characters of Rhamdia. Indeed, we find the
Rhamdia guasarensis is placed in Hep- posterior fontanelle closed or reduced to a
tapteridae by its possession of four syna- small foramen in the supraoccipital in most
pomorphies identified for the family (Lund- other Rhamdia examined: R. laukidi, R. ni-
berg & McDade 1986, Ferraris 1988, Bock- caraguensis, R. quelen (including speci-
mann & Guazelli 2003): posterior limb of mens originally identified as R. guatema-
fourth transverse process expanded and lensis, R. hilarii, R.vilsoni, R. wagneri) and
notched; posterodorsal corner of hyoman- some R. laticauda. Silfvergrip (1996) re-
dibula greatly expanded for attachment of ported the fontanelle to be variably open or
levator operculi muscle; dorsal margin of reduced in R. laticauda, and our sample
quadrate free, not sutured to hyomandibula also shows such variability among speci-
and metapterygoid; ventrolateral corner of mens. We find that R. muelleri has an open
mesethmoid anteriorly recurved. However, posterior fontanelle. The fontanelle is also
the new species lacks a fifth synapomorphy closed in the heptapterids Brachyglanis,
of heptapterids: a straight-edged vertical Brachyrhamdia, Leptorhamdia, and Myog-
bony lamina on the Weberian complex cen- lanis (Bockmann 1998, pers. obs.). Fur-
trum. Instead, the vertical lamina has a con- thermore, R. guasarensis has an uncinate
cave margin in R. guasarensis. process on hypobranchial 1, unlike R. que-
Except for the obvious lack of a free or- len that lacks the process (listed as a second
bital rim, R. guasarensis possesses the non-unique derived feature of R. quelen by
character combination presented as diag- Bockmann 1998). Accordingly, we do not
nostic oi Rhamdia by Silfvergrip (1996:74). take the foregoing as evidence for a partic-
This includes: three pairs of barbels, double ularly close phylogenetic relationship be-
lip fold, vomer without teeth, transverse tween R. guasarensis and R. quelen. The
processes of fourth vertebra expanded midline union of frontal bones (Fig. 2) and
branched distally, supraoccipital process concomitant extreme reduction of the an-
not contacting anterior nuchal plate, adi- terior fontanelle are a distinctive apomorph-
pose fin with free posterior margin, poste- ic character of R. guasarensis. Although not
and postcleithral pro-
rior fontanelle closed all species have been examined for this fea-
cess well developed. However, none of ture, we have not observed it in any Rham-
these are unambiguous synapomoiphies of dia nor has it been previously reported, and
the group of species comprising Rhamdia. in his description of the genus, Silvergrip
Instead, some characters are heptapterid or (1996:74) reported the anterior fontanelle to
VOLUME 1 17, NUMBER 4 571

be invariably open. This is at least a diag- surface waters of Rio Guasare, thus R.
nostic autapomorphy of the species, al- guasarensis is not a cave-dwelling ecotype
though these features are potentially infor- of a proximate epigean species. Two Rham-
mative about relationships. Bockmann dia species have been reported from north-
(1998) illustrated a variety of conditions of western Venezuela. From the Lago de Ma-
anterior fontanelle narrowing and closure in racaibo Basin, Schultz (1944) published on
other heptapterids including Myoglanis, specimens now identified as R. quelen (Sil-
Taunayia, Imparfinis pristos, and an unde- fvergrip 1996). Fernandez- Yepez & Martin
scribed species, but all of these are struc- (1953) reported R. wagneri based on spec-
turally different from that in R. guasarensis. imens collected in the Rio Negro in the
Another peculiar character of the new southern part of the Perija range. One of us
species is the concave dorsal profile of the (CDN) has reidentified these specimens at
head. In general, Rhamdia species, includ- the Museo de Historia Natural La Salle as
ing most cave populations, have convexly R. quelen. As noted above, there is no ev-
rounded heads. The cave species R. macus- idence for a uniquely close relationship be-
panensis recently described from Mexico tween R. guasarensis and R. quelen.
(Weber & Wilkens 1998) has a straight dor- The fauna of troglobitic catfishes of the
sal head profile, somewhat more similar to Sierra de Perija region includes: Ancistrus
that of R. guasarensis than to other con- galani Perez & Viloria, 1994, Trichomyc-
geners. Rhamdia macuspanensis is readily terus spelaeus DoNascimiento, Villarreal &
distinguished from R. guasarensis by its Provenzano, 2001, and Rhamdia guasar-
strong development of pectoral spine den- ensis. There is another hypogean population
tations and rounded tips of the caudal lobes. of Trichomycterus, possibly an undescribed
Rhamdia guasarensis possesses typical species, living in a cave drained by the Rio
reductive characteristics in common with Yasa (Rio Negro system) in the southern
other cave-dwelling species and popula- part of the Sierra de Perija (DoNascimiento,
tions of the genus. Furthermore, the greater in prep.).
relative length of the head and elevated The diversity of three cave catfishes of
number of pectoral-fin rays are also shared the Rio Guasare system is among the high-
by other troglobitic species of the genus est of any Neotropical karst region, al-

(Weber 1996). It has been suggested that though the species are not found syntopi-
larger head size is related to an increase in cally in the same cave. By contrast Bichu-
the development of the cephalic laterosen- ette & Trajano (2003) list five troglobitic
sory system (Langecker & Longley 1993, species in caves of the Sao Domingos karst
Weber 1996), and the greater number of area, Goias, Brazil. Ancistrus cryptophthal-
pectoral-fin rays is possibly correlated with mus and the trichomycterid Ituglanis pas-
the increased mass of the anterior part of sensis coexist in the Sao Vicente cave, To-
the body, compensating this increase with a cantins Basin, Goias, Brazil (Trajano &
greater fin area for hydrodynamic lift (We- Souza 1994, Fernandez & Bichuette 2002).
ber 1996). If there is a functional relation- Also, the inundated caves of the Formosin-
ship between head size and pectoral-fin area ho karst region of Bodoquena, Mato Grosso
in cave dwelling Rhamdia, it does not ex- do Sul, southeastern Brazil, are co-inhabit-
tend to the heptapterid genus Pimelodella, ed by Ancistrus formoso and an unde-
wherein the large-headed P. kronei has scribed troglomorphic population of Tri-
eight or nine pectoral-fin rays (Trajano chomycterus (Sabino & Trajano 1997).
1997, Trajano & 1992) and the
Britski Cave-dwelling and specialized troglobitic
small-headed P. spelaea has ten pectoral- neotropical catfishes belong to the families
fin rays (Trajano et al. 2004). Astroblepidae, Heptapteridae, Loricariidae,
No Rhamdia species are known from the and Trichomycteridae. Within the last three
572 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

of these families the genera Rhamdia, An- —

Comparative material. Rhamdia lati-
cistrus, and Trichomycterus are most com- cauda:ANSP 104034, one specimen. X-ray
monly represented in cave faunas. Their and ale, 86 mm SL, Panama, Code;
prevalence suggests possession of morpho- UMMZ 197078, two dry skeletons, Bl-
logical, physiological, behavioral, and eco- ue mm SL, Honduras. R. laukidi: ANSP
logical features (preadaptations) that facili- 139184, one of three specimens. X-ray and
tate existence in cave waters (Eigenmann ale, 127 mm
SL, Colombia, Meta. R. muel-
1919, Norman 1926, Hubbs 1936). leri: ANSP 162521, two of four specimens.

Wilkens (1986) proposed a correlation X-ray and ale, 109-1 10 mm

between degree of morphological reduction
and time of subterranean evolution based
on a neutral mutation model for the regres-
sive evolution of eyes and pigmentation in
cave fishes and crustaceans. The subterra-
nean catfishes of the Sierra de Perija, es-
pecially Trichomycterus spelaeus and R.
guasarensis, are highly advanced in their
troglobitic features, suggesting that they are
not new arrivals in their subterranean en-
SL, Venezuela,
Amazonas. R. nicaraguensis: ANSP 8444,
one specimen, ale, 135 mm SL, Nicaragua,
Lago Nicaragua. Rhamdia quelen: ANSP
141578, two of five specimens, X-ray and
ale, 100-105 mm SL, Venezuela, Bolivar;
ANSP 45365 (original identification R.
guatemalensis), one specimen, X-ray and
ale, 120 mm SL, Panama, Canal Zone;
ANSP 172138 (original identification R. hi-
larii), two of 37 specimens, X-ray and ale,

vironment. Ocular and pigmentation reduc- 107-110 mm SL, Brazil, Minas Gerais;
tion of R. guasarensis and T. spelaeus are ANSP 16020 (original identification R. vil-
complete. These species exhibit additional soni), one specimen, ale, 200 mm SL, Trin-
autapomorphies such as extremely elongate idad; ANSP 71621 (original identification
barbels in Trichomycterus and much en- R. wagneri), one specimen. X-ray and ale,
larged head laterosensory organs in Rham- 125 mm SL, Colombia, Magdalena; DU-
dia. These characters, too, may indicate a F1021, one dry skeleton, 202 mm SL;
long period of hypogean evolution. Indirect MBUCV-CT-561, eight specimens, CS, 23-
evidence suggests the availability of an am- 57 mm SL, Venezuela, Zulia; MHNLS-
ple period of time for the evolution of the 1645, two specimens, ale, 59-223 mm SL,
Perija cave fishes. Paleogeographic recon- MHNLS-1734, three specimens, ale, 107-
structions of northwestern Venezuela sug- 228 mm SL, Venezuela, Zulia.
gest that uplift of the Sierra began in the
early Cenozoic (Gonzalez de Juana et al.
Acknowledgments
1980). It is reasonable to assume that these
fishes originated in situ after subterranean We are indebted to the members of So-
waters carved out their habitat within Cre- ciedad Venezolana de Espeleologia, espe-
taceous limestones of the Sierra de Perija. cially O. Villarreal who both brought us the
On the fish side of the equation, the only specimens described here and assisted with
fossil record of Rhamdia are fin spines of illustration of the skull. K. Luckenbill ably
relatively young Pleistocene age (Clone prepared the final figures. H. H. Ng gener-
1982). However, based on much older Mio- ously provided us with character data on
cene fossils of phylogenetically related pi- comparative skeletal specimens at UMMZ.
melodid and pseudopimelodid catfishes, the We are indebted to two anonymous and
heptapterids are expected to have originated careful reviewers for many useful com-
and diversified long before the late Pleis- ments on the manuscript. N. Milani de Ar-
tocene (Lundberg 1998). Thus it is possible nal photographed the holotype, and M. Litt-
that subterranean aquatic habitats and cave mann radiographed the ANSP paratype.
fishes have been present in this region for Partial support of publication costs was pro-
tens of millions of years. vided by the All Catfish Species Inventory
 —

VOLUME 117. NUMBER 4

(NSF DEB-03 15963) and an NSF research ical catfish genus Nemuroglanis, with a descrip-

award to JGL (DEB-0089612). tion of a new species (Osteichthyes: Silurifor-

mes: Pimelodidae). —Proceedings of the Bio-
logical Society of Washington 101(3):509-516.
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