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New and Noteworthy Venezuelan Glanapterygine Catfishes (Siluriformes, Trichomycteridae), with Discussion of Their Biogeography and Psammophily PDF

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Preview New and Noteworthy Venezuelan Glanapterygine Catfishes (Siluriformes, Trichomycteridae), with Discussion of Their Biogeography and Psammophily

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3496, 27 pp., 10 figures, 2 tables November 29, 2005 New and Noteworthy Venezuelan Glanapterygine Catfishes (Siluriformes, Trichomycteridae), with Discussion of Their Biogeography and Psammophily SCOTT A. SCHAEFER,1 FRANCISCO PROVENZANO,2 MARIO DE PINNA,3 AND JONATHAN N. BASKIN4 CONTENTS Abstract ....................................................................... 2 Introduction .................................................................... 2 Methods ....................................................................... 4 Institutional Abbreviations ..................................................... 5 Comparative Material Examined ................................................ 5 Results ........................................................................ 5 Key to the Species of Pygidianops Myers, 1944 .................................. 5 Pygidianops cuao, new species ................................................. 5 Pygidianops magoi, new species ............................................... 10 Key to the Species of Typhlobelus Myers, 1944 ................................. 14 Typhlobelus guacamaya, new species .......................................... 14 1DivisionofVertebrateZoology (Ichthyology),AmericanMuseumofNaturalHistory([email protected]). 2Instituto de Zoolog´ıa Tropical, Facultad de Ciencias and Curator of Fish Collection, Museo de Biolog´ıa de la Universidad Central de Venezuela, Apartado de Correos 47058, Caracas 1041-A, Venezuela ([email protected]. ucv.ve). 3Division of Vertebrate Zoology (Ichthyology), American Museum of Natural History; Associate Professor and Scientific Director, Museu de Zoologia da Universidade de Sa˜o Paulo, Av. Nazare´ 481, Sa˜o Paulo-SP 04263-000, Brasil([email protected]). 4Biological Sciences Department, California State Polytechnic University Pomona, Pomona, California 91768; ResearchAssociate,SectionofFishes,NaturalHistoryMuseumofLosAngelesCounty([email protected]). CopyrightqAmericanMuseumofNaturalHistory2005 ISSN0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3496 Typhlobelus lundbergi, new species ............................................ 17 Discussion .................................................................... 19 Generic Diagnoses ........................................................... 19 Interspecific Comparisons ..................................................... 20 Biogeography and Psammophily ............................................... 22 Acknowledgments ............................................................. 25 References .................................................................... 25 ABSTRACT Four new species of the trichomycterid subfamily Glanapteryginae are described from the R´ıo Orinoco basin of Venezuela. Two new species each in Pygidianops Myers 1944 and Typhlobelus Myers 1944 represent the first documented occurrence of these genera in Vene- zuela, and for Pygidianops the first occurrence outside the R´ıo Negro basin. The new species werecapturedfromsand-bottomhabitatsintwodisparatelocationsintheOrinocoRiverbasin and display a remarkable suite of reductive features, such as loss of eyes, fins, and pigment, and reductions or absence of laterosensory canals and odontodes. Pygidianops cuao, n.sp. from the R´ıo Cuao, a clear-water tributary of the upper Orinoco River, is diagnosed from its congeners by the presence of diminutive eyes and a triangular skin flap at the corner of the mouth.Pygidianopsmagoi,n.sp.,knownfromnearthedeltaoftheOrinocoRiver,isdiagnosed from its congeners by the absence of pectoral and anal fins, presence of four laterosensory pores, and nine or ten caudal-fin rays. Typhlobelus guacamaya, n.sp. from the R´ıo Cuao is diagnosedrelativetoitscongenersbythepresenceofthreebranchiostegalrays,posteriornaris absent, lack of pleural ribs, and is further distinguished from both T. ternetzi and T. macro- mycterus by the absence of eyes and from T. lundbergiby the presenceofthreelaterosensory canalpores.Typhlobeluslundbergi,n.sp.fromthelowerOrinocoisdiagnosedbythepresence of four laterosensory canal pores and further distinguished from T. ternetzi andT.macromyc- terus by the absence of eyes. We review the characters useful in diagnoses of Pygidianops and Typhlobelus among trichomycterid catfishes and discuss morphological patterns in the diversification of the Glanapteryginae. Species of Pygidianops and Typhlobelus are known only from the rivers draining the Guyana and Brazilian shields, yet within these areas they occupy all major water types. Such broad ecological range suggests that the geographic dis- tribution of species of these two genera are not limited by water type. That observation, plus their common occurrence in the ubiquitous shallow sand-bottom habitats of the larger rivers of the shield regions of northern South America, indicate that species of Pygidianops and TyphlobelusmaybeexpectedtooccurthroughouttheentireAmazonandOrinocobasins.The evolution of habitat preference in glanapterygines seems to follow a trend toward increased specialization for interstitial environments. The degree of psammophilic adaptation in species of Pygidianops and Typhlobelus is remarkable, without parallel in siluriforms and perhaps in any other freshwater fishes. We describe the physical characteristics of the sand and review the suite of morphological specializations for life in interstitial sand that are shared by these species, such as loss or reduction of certain structures and presence in these species of paired metapleural keels along the ventral edges of the abdomen formed by a long ridge of stiffened integument, underlain by well-differentiated medial infracarinalis muscles, that are superfi- cially similar to the metapleural folds of sand-dwelling cephalochordates and other interstitial organisms. INTRODUCTION phic adaptations known within any single The siluriform family Trichomycteridae is catfish family assemblage. Apart from the widely distributed throughout the Neotropics typical free-living, generalized predators of and includes perhaps the widest rangeoftro- small invertebrates, the rather atypical tro- 2005 SCHAEFER ET AL.: VENEZUELAN CATFISHES 3 phic modes represented amongtrichomycter- of glanapterygines is supported by five syn- id species include the parasitic hematopha- apomorphiesinvolvingreductionsinthefins, gous ‘‘candirus’’ of the subfamily Vandeli- caudal skeleton, and laterosensory system inae and lepidophagous and muciphagous (de Pinna, 1989). Glanapteryx was proposed species of Stegophilinae (Machado and Sa- by de Pinna (1989) as the sister group to a zima, 1983), necrophagous species of the clade composed of Pygidianops and Typhlo- subfamily Pareiodontinae (Goulding, 1979, belus, with Listrura the sister group to those 1980),andtherecentlydiscoveredandpartly three. algivorous species of the subfamily Copio- The genera Pygidianops and Typhlobelus dontinae (de Pinna, 1992). The rangeofboth are the most highly modified glanapterygi- ecological habitats and elevations occupied nes, and species of both genera share ex- by members of the family rivals the rangeof treme reduction of pigmentation, loss of dor- trophic specializations represented, and in- sal fin, loss or extreme reduction of pectoral cludes species restricted to elevations above fins, reduced laterosensory canal, and com- 4000 m in the Andes (Arratia and Menu- plete loss of eyes in some species. Bothgen- Marque, 1984; Ferna´ndez and Vari, 2000; era are markedly miniaturized, yet retain a Ferna´ndez and Schaefer, 2003), Andean relatively well-ossified skeleton comparable lakes, off-shore coastal islands (Ferna´ndez in both bone differentiation and degree of and Schaefer, 2005), lowland species known calcificationtothatobservedinlargertricho- only from large rapids (Myers, 1944), leaf- mycterids. Along with Glanapteryxanguilla, litter puddles, and the bottom of torrential the type species of both genera were collect- rivers (Arratia, 1998). ed at the same time and place, namely, Feb- Representatives of the poorly known sub- ruary 1, 1925, at the rocky pools below the family Glanapteryginae, a group currently Sa˜o Gabriel Rapids on the upper Rio Negro, comprising four genera and nine species, in- Amazonas State, Brazil. Their smallsizeand clude someof themostbizarremorphologies occurrence in atypical habitats are probably and extreme reduction of features thus far most responsible for the fact that glanapter- known among members of the Siluriformes. ygines are rarely collected and currentlyrep- The subfamily was described by Myers resented in collections by few specimens. (1944) to include two new genera and spe- However,recentnewcollectionsindicatethat cies, Pygidianops eigenmanni and Typhlo- glanapterygines are more widely distributed belus ternetzi, plus the previously described than previously thought. For example, Nico Glanapteryx anguilla Myers, 1927, all cap- and de Pinna (1996), following upon the re- tured from a single locality in the upper Rio port of de Pinna (1989), reported on addi- NegroofBrazil.Nonewglanapterygineshad tional populations of Glanapteryx in the R´ıo been discovered for more than four decades, Orinoco basin of Venezuela. Recent collec- until de Pinna (1988)describedanewgenus, tions made in Guyana by J. Armbruster, M. Listrura, with two species from southeastern Sabaj, and colleagues have revealed addi- Brazil, representing the first record of glan- tional populations of Typhlobelus and Pygi- apterygine occurrence outside the Amazon dianops, perhaps representing additional un- River basin. Costa and Bockmann (1994) described species. based their description of Typhlobelus ma- For similar reasons, a more widespread cromycterus on a single specimen from the distribution for both Pygidianops and Ty- Rio Tocantins of Brazil. A new species of phlobelus has been suspected for some time, Glanapteryx, G. niobium, was described by yet unconfirmed. We report herein the exis- dePinna(1998a)fromtheRioNegroofBra- tence of four new glanapterygine species zil. Landim and Costa (2002) described Lis- from the R´ıo Orinoco basin of Venezuela, trura tetraradiata, a glanapterygine with nu- twonewspecieseachinPygidianopsandTy- merous plesiomorphic features, from coastal phlobelus, from two geographically distant streams of Rio de Janeiro State, Brazil.Most locations. One new species each of Pygidi- recently, de Pinna and Wosiacki (2002) re- anops and Typhlobelus was collected by one ported on still another species of Listrura,L. of us (JNB et al.) during the 1979 trawling boticario, from southern Brazil. Monophyly expedition of the R/V Eastward in whitewa- 4 AMERICAN MUSEUM NOVITATES NO. 3496 ter of the Orinoco River delta. After a 0.5 m a set of measurements that are relevant in plankton net, which was being towed behind taxonomic descriptions of those fishes. Most the boat, inadvertently struck the bottom of measurements involve distances that are too theriver,specimensofbothspeciesweredis- small to be obtained with calipers and were covered among the coarse sand and gravel instead taken using an ocular micrometerfit- that was recovered. An additional new spe- ted to a stereomicroscope. Thus, they are cies of each genus was collected by three of measured as projections rather than straight- us (FPR, SAS, JNB) during a joint expedi- line distances. Standard length (SL) was tion in 2001 to the R´ıo Cuao, a clear water measured with the micrometer for purposes tributary of the R´ıo Sipapo, itself a western of proportional values, but with digital cali- Guyana Shield tributary of the upper Ori- pers for absolute values in mm given in lists noco River of Amazonas State, Venezuela. of material. During measurement, specimens These collections represent the first docu- were positioned in ethanol under a glass mi- mented occurrence of Pygidianops and Ty- croscope slide, which served tomaintainori- phlobelus from Venezuela and, for Pygidi- entation, straighten and compress the speci- anops, the first occurrence of the genus out- mens onto the viewing plane, and thereby side the Rio Negro basin. All previously minimize measurement error. described Typhlobelus species possess di- In addition to standard length, the follow- minutive eyes, whereas the two new species ing measurements were obtained: caudal pe- lackeyesentirely.Conversely,thepreviously duncle length—from posterior margin of described Pygidianops eigenmanni, along anustomiddleofposteriormarginofhypural with one of the new Orinocoan species, lack plate; body depth—maximum vertical dis- eyes, while the new species from the R´ıo tance through anterior margin of anus, ex- Cuao has eyes present, although diminutive. cluding fleshy dorsal-fin fold, when present; The possibility that these new speciesareju- head length—from rostrum tip to posterior veniles of other recognized species has been margin opercular flap at pectoral-fin origin; considered and rejected based on their well- head width—maximum distance perpendic- ossified skeletons and the apparent presence ular to longitudinal axis through posteriortip of mature gonads. of opercle; prenasal length—from rostrum Inthispaper,wediagnoseanddescribethe tipatmidlinetoanteriorbaseofnasalbarbel; fournewspeciesofPygidianopsandTyphlo- rostrum length—from snout tip to anterior belusfromtheR´ıoOrinocosystem.Keysare mouth margin; rostrum width—horizontal provided as an aid to the identification of all maximum through mouth corners; internar- species of the two genera. We conclude with ial width—between anteriormarginsofnasal a discussion of the biogeographic implica- barbel bases; preanal length—from rostrum tions and their psammophilic ecology. tip at midline to base of first anal-fin ray; preproctal length—from rostrum tip to an- METHODS terior margin of anus. Data for rostrum length and prenasal length wereexcludedfor Due to the minute size of the taxa treated specimens having a damaged or distorted herein, obtaining morphometric data was rostrum. problematic and made more difficult by the Because all fin rays in Pygidianops and absenceofvariousfinsandeyes,therebypre- Typhlobelus species are unbranched and un- cluding the use of certain traditional land- segmented (except as noted below), counts mark reference points. The lack of integu- forthecaudalfinincludedthoseraysdirectly ment pigmentation also makes the visuali- associated with the hypural plate. Anal-fin zation of superficial traits, such as the nares, ray counts included all rays with a corre- extremely difficult under opticalmicroscopy. sponding pterygiophore, and the posterior In view of those particulars inherent to the closely set rays were counted separately.La- morphology of the species, and in order to terosensory canal pores were countedonwet provide a standardized future reference for specimens in air; visualization was aided by obtaining morphometric data fromspeciesof directing a jet of compressed air onto the PygidianopsandTyphlobelus,weheredefine structures. Vertebral counts and the vertebral 2005 SCHAEFER ET AL.: VENEZUELAN CATFISHES 5 association of the first anal-finpterygiophore M. de Pinna, 10 Nov. 1996. Typhlobelus ter- were taken fromradiographsandclearedand netzi SU 36558 paratype (25.3 mm SL) Bra- stainedspecimenpreparationsandincludeall zil, Amazonas, Rio Negro, rock pools below freevertebra(thoseincorporatedintotheWe- rapids at Sa˜o Gabriel, C. Ternetz, 1 Feb. berian complex not counted) plus the ural 1925. T. macromycterus MNRJ 12129 ho- complex counted as a single element. lotype (21.9 mm SL) Brazil, Para´, Rio To- Osteological preparations (denoted ‘‘c&s’’) cantins near Tucuru´ı, L. C. Alvarenga, Sept. were made following Taylor and Van Dyke 1984. Typhlobelus sp. INPA 12929 (5) Bra- (1985). One specimen of each species was zil, Rio Xingu at Arroz Cru, near Senador critical point dried, gold coated, and exam- Jose´ Porf´ırio, J. Zuanon, 7 Oct. 1996. ined using a Hitachi S-4700 scanning elec- tron microscope. Histological preparations RESULTS were made from 4 mm serial sections that were imbedded in Paraffin and stained with KEY TO THE SPECIES OF PYGIDIANOPS Mason’s trichome. Drawings were prepared Myers, 1944 using a camera lucida attached to a stereo- 1A. Anal and pectoral fins absent; caudal fin microscope. In the listing of material exam- with nine or ten rays associated with hy- ined from the R/V Eastward collections,dis- puralplate;fourlaterosensorypores;low- tance from the buoy marking the ocean/river er jaw pointed in ventral view, two man- mouth are indicated in nautical miles for dibular rami forming acute angle at mid- each collections station. Size fractions for line; distance between middle of lower R´ıoCuaosedimentweredeterminedbypass- jawandanteriormarginofrostrumlarger ing a wet sample through a series of sieves thanmouthwidth;rostrumlesswidethan of progressively smaller pore openings, and head width .......... P. magoi, n.sp. the material collected on each sieve was 1B. Anal and pectoral fins present (the latter as quantified by volume displacement, de- ashortone-rayedflap);caudalfinwith12 to 14 rays associated with hypural plate; scribed, and measured using an ocular mi- six laterosensory pores; lower jaw form- crometer fitted to a stereomicroscope. ingcontinuouscurveinventralview;dis- tance between middle of lower jaw and INSTITUTIONAL ABBREVIATIONS anterior margin of rostrum smaller than mouthwidth;rostrumaswideas,orwid- AMNH American Museum of Natural History, er than, head width ............... 2 New York 2A. Eyes present; triangular skin flap at mouth CAS California Academy of Sciences, San corner; maxillary and rictal barbels ex- Francisco tending posteriorly beyond vertical INPA Instituto Nacional de Pesquisas da through base of pectoral fin; nasal barbel Amazoˆnia, Manaus, Brazil extending posteriorly to posterior half of MBUCV Museo de Biologia, Universidad Cen- pectoral fin or longer ... P. cuao, n.sp. tral de Venezuela, Caracas 2B. Eyes absent; no triangular skin flap at MNRJ MuseuNacional,RiodeJaneiro,Brazil mouth corner; maxillary and rictal bar- MZUSP Museo de Zoologia, Universidade da bels posteriorly not reaching vertical Sa˜o Paulo, Sa˜o Paulo, Brazil through base of pectoral fin; nasal barbel SU former Stanford University collections, reaching posteriorly to base of pectoral now housed at CAS fin or shorter ..................... .......... P. eigenmanni Myers,1944 COMPARATIVE MATERIAL EXAMINED Pygidianops eigenmanni SU 36557 para- Pygidianops cuao, new species types (3:12.3–18.4 mm SL), CAS 11121 Figures 1, 2; Table 1 paratypes(7:10.5–13.6mmSL)Brazil,Ama- zonas, Rio Negro, rock pools below rapids HOLOTYPE: MBUCV-V-30917 (18.70 mm at Sa˜o Gabriel, C. Ternetz, 1 Feb. 1925. Py- SL) Venezuela, Estado Amazonas, R´ıo Cuao gidianops sp. INPA 12427 (5) Brazil, Ama- at Raudal Guacamaya, 8.1 miles upstream zonas, Igarape´ do Acara´ (tributaryoftheRio from Raudal El Danto, 05807.719N, Taruma˜, Rio Negro basin), A. Kirovsky and 67831.539W, SAS01–03, F. Provenzano, S.A. 6 AMERICAN MUSEUM NOVITATES NO. 3496 Fig.1. Pygidianopscuao,holotype,MBUCVV-30917,21.06mmSL,Venezuela,EstadoAmazonas, R´ıo Cuao at Raudal Guacamaya, 8.1 miles upstream from Raudal El Danto. Scale bar is 1 mm. 2 0 0 5 TABLE1 S Counts and Measurements of Holotype (HT) and Paratypes (PT) of New Venezuelan Pygidianops and Typhlobelus Species C H Measurements given as percent head length, unless noted otherwise; mean (median for counts) and standard deviation in parentheses. A E F Pygidianops Typhlobelus E R cuao magoi guacamaya lundbergi E T HT PT(N518) HT PT(N55) HT PT(N59) HT PT(N54) A L Standardlength(SL) 18.7 17.0 (2.8) 12.6 12.6 (.81) 22.2 22.3 (3.1) 29.0 25.9 (2.6) .: Headlength(%SL) 16.8 16.1 (1.8) 18.5 17.4 (1.0) 12.2 11.7 (1.0) 11.0 9.7 (0.6) V Body depth (%SL) 17.5 16.1 (1.4) 14.1 14.0 (2.9) 5.1 5.0 (0.4) 4.6 4.2 (0.7) E N Headwidth 63.5 69.0 (4.0) 54.9 56.9 (4.7) 39.3 39.3 (2.7) 44.8 46.2 (3.4) E Z Preanallength(%SL) 54.8 58.5 (1.7) — — 70.4 70.1 (2.2) 73.1 70.7 (1.3) U Preproctallength(%SL) 51.5 56.3 (1.6) 63.8 61.7 (3.0) 69.7 68.5 (1.9) 71.8 68.9 (1.7) E L Caudal pedunclelength(%SL) 44.8 44.0 (1.9) — — 29.5 29.4 (0.9) 24.8 30.8 (1.4) A Rostrumlength 12.7 11.4 (1.7) 17.2 16.4 (3.8) 33.3 35.0 (2.5) 24.5 22.5 (1.8) N Rostrumwidth 26.3 31.0 (3.7) 18.5 20.7 (2.9) 13.7 15.0 (2.6) 17.2 15.4 (1.8) C A Prenasallength 23.2 27.6 (4.3) 24.9 21.2 (2.0) 32.2 34.9 (2.6) 31.3 28.0 (1.3) T Internarialwidth 26.3 30.4 (5.0) 23.6 23.5 (3.7) 17.4 15.3 (2.9) 18.8 13.7 (4.4) F I S Caudal finrays 13 13 (.24) 9 9.5 (.58) 9 9 (0) 10 9 (0) H Analfinrays 4 4 (.51) — — 5 5 (0) 5 5 (0) E S 7 8 AMERICAN MUSEUM NOVITATES NO. 3496 Schaefer, J.N. Baskin, A. Rojas (hereafter DESCRIPTION: General appearance com- ‘‘PSBR’’), 3 March 2001. pact, oblate; head slightly depressed and PARATYPES (all Venezuela, Estado Ama- body compressed, trunk neither cylindrical zonas):AMNH232970(23:16.59–24.00mm nor ribbonlike. Dorsal and ventral body pro- SL), MBUCV-V-29905 (22: 8.13–21.55 mm files gently convex. Dorsal profile of head SL), MZUSP 82103 (5: 13.4–19.0 mm SL) slightly concave at vertical through opercle, same dataasholotype;MBUCV-V-29843(1: gently convex across occipital region. Out- 20.00 mm SL) R´ıo Cuao, Can˜o Ceje, 5 min- lineofheadrectangularindorsalview,great- utesupstreambyboatfromRaudalElDanto, est width (75% HL) at opercle. Anteriorros- 05805.869N, 67831.379W, SAS01–02, PSBR, trum margin blunt, nearly straight between 3 March 2001; AMNH 232978 (11: 12.3– maxillary barbel bases, rounded in lateral 23.15 mm SL), MBUCV-V-29922 (10: profile. Ventrum between pectoral and anal 12.41–20.30 mm SL), MZUSP 82104 (3: fins forming shallow sulcus with margin of 10.6–18.4mmSL)R´ıoCuaoatIsladeCielo, laterally divergent medial infracarinalis 21.3 kilometers upstream from Raudal El (Winterbottom, 1974) fiber bundles (fig. 3). Danto, 05811.019N, 67831.119W, SAS01–04, Three pairs of barbels; all similar in general PSBR, 5 March 2001; AMNH 232995 (4: appearance, robust at base. Maxillary barbel 13.95–19.32 mm SL), MBUCV-V-29933 (4: continuous with anterior lip margin; its base 14.46–18.22mmSL)R´ıoCuaoatRaudalPia slightly depressed. Rictal barbel compressed Poco, 21.5 kilometers upstream from Raudal at base, merging with maxillary barbel base El Danto, 05810.959N, 67830.819W, SAS01– at mouth corner. Nasal barbel distinctly sep- 05, PSBR, 6 March 2001; MBUCV-V-29977 arate,extendingposteriorlywellbeyondpec- (3: 8.12–12.15 mm SL) R´ıo Cuao, approxi- toral fin origin. Mouth ventral, upper lip a mately20minutesbymotorboatdownstream broad fleshy pad, anterior margin of lower from base camp at SAS01–4, 05810.139N, jawconvex.Smalltriangularflapofskinpro- 67831.769W, SAS01–7, PSBR, 7 March jects from corner of mouth (figs. 1, 2) be- 2001; AMNH 233036 (6: 12.25–19.27 mm tween lower lip margin and rictal barbel SL), MBUCV-V-29997 (7: 11.05–17.49 mm base.Posterodorsalcornerofbranchialopen- SL)R´ıoCuaoatRaudalPauj´ı,approximately ing located immediately anterior to pectoral 10 minutes by motorboat downstream from fin base, gill membrane continuing antero- base camp at SAS01–4, 05808.929N, ventrally, its posterior margin straight to 67832.189W, SAS01–8, PSBR, 8 March slightly concave, not united across isthmus; 2001; MBUCV-V-30105 (4: 8.80–21.65 mm fivebranchiostegalrays.Eyespresent,butre- SL) R´ıo Cuao, approx. 10 minutes by mo- duced, appearing as compact spherical mass torboat below Raudal El Danto, 05802.399N, of darkly pigmented cells below slightly 67833.359W, SAS01–13, PSBR, 10 March raised transparent skin surface. Spherical 2001. pigment mass with dorsal indentation filled DIAGNOSIS: Distinguished from congeners with mass of opaque cells,yieldingrudimen- by the presence of diminutive eyes (vs. eyes taryeyeballandlensconfiguration.Naresnot absent), posterior naris absent (vs. nares bi- juxtaposed or connected, but rather wellsep- laterallypaired),presenceofatriangularskin arated; anterior naris at dorsomesial base of flap at mouth corner (vs. skin flap absent), nasal barbel, posterior naris larger, located dorsal and ventral fin fold deep and extend- slightly mesial to and one eye diameter an- ing forward on the head to a verticalthrough terior to eye, its opening directed laterocau- tip of pectoral fin (vs. fin fold shallow, ex- dad. Laterosensory system reduced to short, tending forward on head to point approxi- wide trunk canal segment posterior to supra- mately two times head length from rostrum cleithrum and dorsal to pectoral fin, canal tip). Further distinguished from P. magoi by continuing onto head through supracleith- presence of six laterosensory pores on head rum, pterotic, sphenotic, and frontal to ter- (vs. four pores), presence of pectoral and minate just posterior to eye. Six pores total anal fins (vs. fins absent), presence of 12–13 (fig. 2 top, middle), first (anteriormost) im- caudal fin rays associated with the caudal mediately behind eye and homologous with plate (vs. 9–10). thesupraorbitalepiphysealpore(‘‘i6’’ofAr- 2005 SCHAEFER ET AL.: VENEZUELAN CATFISHES 9 Fig. 3. Histological cross section through the caudal region of P. cuao showing dorsal fin fold and ventrally projecting paired metapleural keels (arrows). Scale bar is 1 mm. homologous with pterotic branch (Schaefer andAquino,2000;‘‘L1’’ofArratiaandHua- Fig. 2. Pygidianops cuao, paratype, AMNH quin, 1995), fifth at canal entrance to supra- 232995, 19.2 mm SL, scanning electron micro- cleithrum and homologous with first lateral- graphs of head. Scale bar is 1 mm. line pore, sixth (posteriormost) on trunk at vertical through middle of pectoral fin and ratia and Huaquin, 1995), second at frontal/ representingposteriorterminusofshorttrunk sphenotic junction and homologous with in- canal. fraorbital canal branch (canal segment ab- Dorsal fin absent, dorsum of body with sent),thirdatpteroticnearitsanteriormargin thin, semitransparent fin fold (fig. 3) from and homologous with preoperculomandibu- head to dorsal margin caudal fin; its greatest lar canal branch (canal segment absent), depth at vertical through anus, contained fourthatpterotic/supracleithrumjunctionand three times in body depth. Anal fin present, 10 AMERICAN MUSEUM NOVITATES NO. 3496 4–5 (mostly 4) rays, all unsegmented, un- ETYMOLOGY: The specific name is for the branched; first ray rudimentary, short, about R´ıo Cuao, a clearwater tributary of the R´ıo 0.25 length second ray; first pterygiophore Orinoco and the region of first discovery of associated with hemal spine of vertebra 17. the species. Treated as a noun in apposition Ventrum with thin fin fold between base last to the generic name. anal-fin ray to ventral margin caudal fin; its greatest depth about 0.5 times depth dorsal Pygidianops magoi, new species fin fold. Anus and urogenital papilla located Figures 5, 6; Table 1 immediately anterior to first anal-fin ray. Urogenital papilla length approximately HOLOTYPE: MBUCV-V-31035 (20.0 mm equal to anus diameter, papilla connected to SL) Venezuela, Estado Delta Amacuro, R´ıo anal-fin base by median ridge of translucent Orinoco at Puerto Cabrian, 8834.89N, tissue. Caudal fin with 12–13 raysassociated 62815.99W, R/V Eastward station 36957, mi with the caudal plate, all unbranched, all but 151, field number T-35-79, 10 ft bottom thedorsalandventralmostrayswith2–4seg- trawl, capture depth 25 m, 13:55 hrs., John ments per ray; 4–5 dorsal and 5–7 ventral G. Lundberg et al., 10 Nov. 1979. procurrent rays; spacing between dorsal pro- PARATYPES (all from Venezuela): AMNH current rays compacted posteriorly, yielding 233706 (11.71 mm SL), Estado Bol´ıvar, R´ıo slightly upturned appearance to caudal fin Orinoco between Ciudad Bol´ıvar and Puerto relative to longitudinal body axis; posterior Ordaz, 8820.169N, 62857.819W, R/V East- fin-margin rounded. Pectoral fin present, mi- ward station 36901, mi 203, field number T- nute, its length about 0.3 times head length; 12-79, 10 ft bottom trawl, capture depth 18 one unbranched unsegmented ray, fin mem- m, 16:30 hrs., F. Mago-Leccia et al., 7 Nov. brane expanded at base, compressed; distal 1979;AMNH233707(12.69mmSL)Estado tip pointed. Pelvic fin and girdle absent. Delta Amacuro, R´ıo Orinoco just down- Jaw teeth unicuspid, blunt; 7–9 mandibu- stream from Isla Portuguesa, 88389N, lar and 6–8 premaxillary teeth per jaw ele- 618499W, field numberT-59-79, mi116.5,10 ment. Odontodes absent. Trunk myomeres ft bottom trawl, capture depth 30–40 m, 15: 39–42 (holotype 42), total vertebrae 41. 07 hrs., J.G. Lundberg et al., 14 Nov. 1979; COLORATION: Pale tan to grey in life. In AMNH 233708 (11.96 mm SL), MZUSP ethanol,creamwhiteoverall,withdensecon- 84304 (1c&s: 11.4 mm SL) Estado Bol´ıvar, centration of dark melanophores on occipital R´ıoOrinoco,8818.39N,62856.19W,R/VEast- region, extending ventral to posteriormargin ward station 36895, field number P-2-79, mi swimbladder capsule; deeper lying melano- 200.5, 1 m plankton net, capturedepth26m, phores along mesethmoidlateralmarginpos- 12:55 hrs., H. Lopez-Rojas et al., 7 Nov. terior to nasal barbel base and at opercle an- 1979; MBUCV-V-31036 (14.5 mm SL) Es- terior margin. Trunk with diffuse band of tado Delta Amacuro, Rı´o Orinoco, just melanophores along dorsal fin-fold base, downstream from Isla Portuguesa, 88389N, scattered diffuse line along ventral fin fold, 618499W, field numberT-61-79, mi116.5,10 midlateral line with fewsmall,irregularlyar- ft bottom trawl, capture depth 25 m, J.G. ranged, discrete clusters of melanophores, Lundberg et al., 14 Nov. 1979; MBUCV-V- ventrum of head and trunk otherwise unpig- 31037 (13.4 mm SL) Estado Bol´ıvar, R´ıo mented. Orinoco, Isla Isabella, between PuertoOrdaz DISTRIBUTION: Known only from six col- and Ciudad Bol´ıvar, 8819.79N, 62856.89W,R/ lection localities within the R´ıo Cuao drain- V Eastward station 36924, field number T- age basin (fig. 4). 18-79, mi 203, 10 ft bottom trawl, capture fi Fig. 4. Geographic distribution of the Venezuelan Glanapteryginae. Drainage map of Venezuela (center);rectanglesdepictregionsofinterestexpandedinAandB.A,lowerR´ıoOrinoco;B,R´ıoCuao. Symbols designate collection localities; some symbols represent more than one lot. Open symbols des- ignate holotype; grey star symbol denotes shared type locality for both P. cuao and T. guacamaya.

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