SNS fan ste ba A 0 eV tat eae pees He hese an bre? 1a cWichatebegy fig 26 Ree aT Te Aa tha} VeVi are treyianh Rusti Peet 3 Bh i i \ \ . Agar eh Menthe VWenryial yey CPL Ia) Alita nah Aa Lea One vty oe Viiane ed Saye UPPED EEN Sy the ther hel via Bravaees iy Sey niga re “8 Tyee a peaceeio} abit ee Neaply yb ate aim ea Erie tangie te Matt ritpecetts Mibbieites if DED PUN ‘ Er } es TaN ined 0 assy! wtih i 4 D i i i ny re: aD Fegty 4 Se ry y DORSAL Rel WO Lane tal ; ; ‘ Se Saa tetanic abe EOE VO iP AE AUPE eS i 1 Bo gp esa ty Bote ii ‘ Be ‘i } a ti LE NA LE ie AE Tas i rf “ id Re Ren EOL Ue itb sg Pere eee eh Rg ae art tien: ne stn ay f Y isi Z CLP eae et g hat ? ie ta sbte MOF Ehren yo i mae? ¥ Pi ata , tle Aiea Ara ee ; Haber j figgana ea is see Oe PH IPESE PD BERT B ig ge eaty fe encuning gy ahs tot itn, ¢ baa H Sgt tyes wily Sn i ed cieute aa gry ante yee? et as Teataoon spel tetgenceg? hee iiss nen ag RecA Vga he bey ota 5 woe erg ate a) iinet de ' $ Tele Be faery lpr deen etait vanes f f Vance V3 ‘ a / Ba eee ( i ye sage ces eeiNy ta Meaty =H i : yc nae th cus Hae u Snaepay sos eed a ies i } win ; (iia : AOA ihe Stay ere 4 d \ 4 ate # heat { Wi og : ; } , nis a 4 ih i yi "| LAE Hit He Heh hha Bas PROCEEDINGS of the Biological Society of Washington VOLUME 111 1998 Vol. 111(1) published 6 April 1998 Vol. 111(3) published 18 September 1998 Vol. 111(2) published 24 June 1998 Vol. 111(4) published 23 December 1998 WASHINGTON PRINTED FOR THE SOCIETY EDITOR C. BRIAN ROBBINS ASSOCIATE EDITORS Classical Languages Invertebrates FREDERICK M. BAYER STEPHEN L. GARDINER FRANK D. FERRARI RAFAEL LEMAITRE Plants Vertebrates DAviD B. LELLINGER Gary R. GRAVES Insects WAYNE N. MATHIS All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN PRESS INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1998-1999 OFFICERS President RICHARD P. VARI President-Elect BRIAN F. KENSLEY Secretary CAROLE C. BALDWIN Treasurer T. CHAD WALTER COUNCIL Elected Members MICHAEL D. CARLETON RAFAEL LEMAITEE W. DUANE HOPE ROY W. MCDIARMID SUSAN L. JEWETT JAMES N. NORRIS i) ne onil ] WS, eo x TABLE OF CONTENTS Volume 111 Abe, Wataru, Kazuo Utsugi, and Masatsune Takeda. Pseudechiniscus asper, a new TYardigrada (Heterotardigrada: Echiniscidae) from Hokkaido, northern Japan .......................0. 0. sees eee Ahyong, Shane T. and Raymond B. Manning. Two new species of Erogusquilla from the Indo- West Pacific (Crustacea: Stomatopoda: Squillidae) ................ 00. c eee cece eect eee eee e teen eee Ahyong, Shane T., Tin-Yam Chan, and Y. J. Laio. A new stomatopod (Crustacea: Malacostraca) of the genus Harpiosquilla Holthuis, 1964 from Taiwan and Australia ............................ Blow, Warren C. and Raymond B. Manning. Eohalimede sandersi, the correct name for the species described as Eohalimede saundersi Blow & Manning, 1997 (Crustacea: Decapoda: Xan- HEGRE) 5 ncoood code GAMO AAC AGATE COT OBORAT GORGE SECC tC ER ICICI IPLAC CIES ACRE REAL Arto on rae Bueno-Soria, Joaquin and Ralph Holzenthal. Studies in aquatic insects XIV: Description of eight new species of Ochrotrichia Mosley (Trichoptera: Hydroptilidae), from Costa Rica ............. Campos, Ernesto and Alma Rosa de Campos. Taxonomy and distribution of the parasitic isopod Progebiophilus bruscae Salazar-Vallejo & Leija-Tristan, 1990 (Crustacea: Bopyridae) ......... Campos, Emesto and Ramond B. Manning. Pinnotheres malaguena Garth, 1948, a new member of the genus Fabia Dana, 1851 (Crustacea: Brachyura: Pinnotheridae) .................-.-+---00+ Campos, Ernesto, Victoria Diaz, and J. A. Gamboa-Contreras. Notes on distribution and taxonomy of five poorly known species of pinnotherid crabs from the eastern Pacific (Crustacea: Brachyura: LETHRADCUMODES)) ea scageSonec Gee ae AAP ES ORE SCO A NOES orate eISEI aes Te SAREE TEs roe eR aD Campos, Martha R. A new species of freshwater crab of the genus Phallangothelphusa Pretzmann, 1965 from Colombia (Crustacea: Decapoda: Pseudothelphusidae) ..................22.c0ee cece eeeee Campos, Martha R. and Rafael Lemaitre. A new freshwater crab of the genus Neostrengeria Pretzmann, 1965, from Colombia (Crustacea: Decapoda: Brachyura; Pseudothelphusidae), with 2% [Rey (© Whe SISEIES WHINE ENG: Baoseqonccndcoseddacanausaaordsadeaspaatdecsdadanacapon abe scee conn adoue Cooper, John E. A new species of crayfish of the genus Procambarus, subgenus Ortmannicus (Decapoda: Cambaridae), from the Waccamaw River basin, North and South Carolina ......... da Silva, Maria Nazareth F. Four new species of spiny rats of the genus Proechimys (Rodentia: Echimyidac)) trom the western Amazon of Brazil) 22322222: ..seeeeee secs oee eee eae eceeee soon Dalkey, Ann. A new species of amphipod (Crustacea: Amphipoda: Lysianassoidea) from the Pacific (CORE OF IN@iin ATIC cusp caguaseenatsnoonsd caowauned Siebue daceiaoaaacr CEA CnenSnnntand ree i ren an eeee de Pina, Gloria M. Alonso. Metatiron bonaerensis, a new species (Crustacea: Amphipoda; Syno- pidae)momathersouthwest Atlantic: 7csh-s sc eee eeece esc e eee denecseeae net uhneie eae teue seers emesme sec de Pinna, Mario C. C. A new species of the catfish genus Glanapteryx (Siluriformes: Tricho- EEMCIORECEES) cocdecagnadboanddnne Heed pseerEdeadH Eade 6 oaaO RUA GOST TO CHORUS CRE cn Oconee TBE GHEE neceta tare ee Dean, Harlan K. A new species of Hesionidae, Glyphohesione nicoyensis (Annelida: Polychaeta), PEOMMiNCsGULiROl@NiCoyas © Osta Rica® iacsseessechee ances shen seh ones clda neue satu acecetenesnat seve! Elias-Gutiérrez, M. and E. Sudrez-Morales. Redescription of Microdiaptomus cokeri (Crustacea: Copepoda: Diaptomidae) from caves in central Mexico, with the description of a new diaptomid subfamily Felder, Darryl L. and Raymond B. Manning. A new ghost shrimp of the genus Lepidophthalmus from the Pacific coast of Colombia (Decapoda: Thalassinidea: Callianassidae) ................... Ferrari, Frank D. and Adam Benforado. Setation and setal groups on antenna 1 of Ridgewayia kKlausruetzleri, Plearomamma xiphias, and Pseudocalanus elongatus (Crustacea: Copepoda: Cal- anoida) during the copepodid phase of their development ..................00000cceceeeeeeeeeeneees Fitzpatrick, Jr., J. F. and Mary K. Wicksten. A new crayfish of the genus Procambarus (Crustacea: DecaApodas Gani bandae) nomacenttalishexasy weeas-f eset te cece een oe ies meee cere ee nnGiGe cteneeaes Forest, Jacques and Patsy A. McLaughlin. Descriptions of two new Japanese hermit crabs (De- CASOGAE PACUING EIS IDYOY SITES) 9 jocdecrosoc sno droecaeese rca dco se SeBEeEURGGeae 5 Gd GeEEM nee lect ea ane Formas, J. Ramon, César Cuevas, and José Nufiez. A new species of Alsodes (Amphibia: Anura: Pe MOdAchyindac) piLOmMmsouthenyChiley mean -anies te cen here emcee oo eeese eee a ace oe sec isitine tates Fraser, Thomas H. A new species of cardinalfish (Apogonidae) from the Philippines, with com- ments on species of Apogon with six first dorsal spines Gerken, Sarah and Les Watling. Diastylis tongoyensis, a new diastylid (Crustacea: Cumacea) from the northern central coast of Chile, with an amendment to the description of Diastylis crenellata Watling & McCann, 1997 Gomez, Patricia. First record and new species of Gastrophanella (Porifera: Demospongiae: Lithis- tida) from the central East Pacific 843-848 653-662 929-935 409 604-612 288-294 912-915 372-381 92-96 899-907 81-91 436-471 621-626 627-633 35-42 257-262 199-208 398—408 209-221 146-152 188-198 521-530 986-99 1 857-874 774-780 vi Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 5. Probable hybrid origin OF Aimeriiia GUSTS W/CUMORO C2 IATEOS scccoosacooaesoeqoansccoqgs0 ned scaaaesenedaadcacsnoavdueRsenee Graves, Gary R. Taxonomic notes on hummingbirds (Aves: Trochilidae). 1. Eriocnemis dyselius Elliot, 1872 is a melanistic specimen of Eriocnemis cupreoventris (Fraser, 1840) ............... Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 6. An intergeneric hybrid, Aglaiocercus kingi X Metallura tyrianthiana, form Venezuela ......................se ee eee eee eee Guidetti, Roberto. Two new species of Macrobiotidae (Tardigrada: Eutardigrada) from the United States of America, and some taxonomic considerations of the genus Murrayon ................-- Hajdu, Eduardo and Klaus Riitzler. Sponges, genus Mycale (Poecilosclerida: Demospongiae: Porif- era), from a Caribbean mangrove and comments on subgeneric classification ...................- Hanger, Rex Alan and Ellen E. Strong. Helicoprion nevadensis (Wheeler, 1939) from the Penn- sylvanian—Permian Antler Peak Limestone, Lander County, Nevada (Pisces: Selachii: Heli- GOprionidae)) .cacae das sree ce sree a ak are cee neue SATO ict aes cae als a ete eee ee R Ee eee Cee EC LER EEE Hanger, Rex Alan and Ellen E. Strong. Acteonina permiana, a new species from the Permian Coyote Butte Formation, central Oregon (Mollusca: Gastropoda: Actaeonidae) .................. Harold, Antony S., James H. Wessel, III, and Robert K. Johnson. A new species of Polyipnus (Teleostei: Stomiiformes) from the western Indian Ocean, with comments on sternoptychid SCOLOBY ai ssnierseis oerceciemiecyasn abe aerate minceienis wsaaasb maa sisteyeis telus laaeis neleineic Okt Baste spear roe ese ERECT Heard, Richard W. and Raymond B. Manning. A new genus and species of ghost shrimp (Crus- tacea: Decapoda: Callianassidae) from the Atlantic Ocean .................. cesses eee eee eee nent ne ees Hendrickx, Michel E. A new genus and species of “‘goneplacid-like’’ brachyuran crab (Crustacea: Decapoda) from the Gulf of California, Mexico and a proposal for the use of the family Pseu- dorhombiidae: Alcock 1990) 2.55.2) eiiaaaasca ister e cession te yan «ise otsepatodesleeietererye estes eres Hendrickx, Michel E. and Ma. del Carmen Espinosa-Pérez. A new species of Cassidinidea Hansen (Isopoda: Sphaeromatidae) and first record of the genus from the eastern tropical Pacific ...... Hendrickx, Michel E. and Ma. del Carmen Espinosa-Pérez. A new species of Excorallana Stebbing (Crustacea: Isopoda: Corallanidae) from the Pacific coast of Mexico, and additional records for Ei DrUusCat Delaney, rts) ai cienra tease nie Sales eR SI oes Od LEE EEE CSE eRe Hernandez, F. and S. Jiménez. Cave chaetognaths in the Canary Islands (Atlantic Ocean) ........ Hernandez-Alcantara, Pablo and Vivianne Solis-Weiss. Capitellids (Polychaeta: Capitellidae) from the continental shelf of the Gulf of California, México, with the description of a new species, NOLOMGSTUS GNBELICGE 5 ajar ep hace spa eee eas STR eds oie ohe VEIN EY OTR EOS Hutchings, Pat, Patrick Frouin, and Christian Hily. Two new species of Spionidae (Polychaeta) from Tahiti,.\FrenchePolymesia\cccus serosa cscnecie cise cisaecnon eee eee eae eee eee eee CEE EEE Ivanenko, V. N. Laperocheres koorius, a new genus and species (Copepoda: Siphonostomatoida: Asterocheridae) associated with the sponge Amphimedon in Australia .................0.22eee+e0+: Karawawa, Hiroaki. Typilobus kishimotoi, a new leucosiid crab (Crustacea: Decapoda: Brachyura) fromutheyViiocenesKatsutak Groups apalleeere eer ee EE CEe ree eee ee eee EEE ECE EEE Eee e ee EEE CEE EEE ner eeerre Kensley, Brian and Kerry Clark. A new isopod species from Key Largo, Florida (Crustacea: Iso- poda: Hologmathidae)) .peadssjscenkasaeroceisrg sees yt teens SIs EE CEE RTE eee Kim, Won. Chelomalpheus koreanus, a new genus and species of snapping shrimp from Korea (GmustaceaDecapodavAllpheidac) aaeppeerer etna eee EEE EE CE EEE ee CE REL eee EERE EEE CCE E Ee cE rere Komai, Tomoyuki, Tin- Yam Chan, and Ding-An Lee. The discovery of Glyphocrangon stenolepis Chace (Decapoda: Caridea: Glyphocrangonidae) from Taiwan and Japan, with notes on individ- Wall VarbathOmd a ose cic catielanarssteyeceretaieve ayessiereve ets asia apslefeis scaltuseeesiel ie Geter Oe AICS ES SITES Ee LT eee leer eee Lambert, Philip. Pentamera rigida and P. pediparva, two new species of sea cucumber from the west coast of North America (Echinodermata: Holothuroidea) .................0..0.ceceeee eee ee eee Lechapt, Jean-Paul and David W. Kirtley. New species of bathyal and abyssal Sabellariidae (An- nelida: Polychaeta) from near New Caledonia (southwest Pacific Ocean) ....................2+0-+ Leon Gonzalez, Jesis Angel de and Vivianne Solis-Weiss. The genus Perinereis (Polychaeta: Nereididae) from Mexican littoral waters, including the description of three new species and the nEdeSCrIpP MONSTOLeL Nanderssoniands > nClenaGdSOGCMPEEEEEEEE EEE Ee TEE EEE EEE ELEC EC EERE EE Eee Eeeee Le6n-Gonzalez, J. A. de and V. Diaz-Castafieda. Two new species of Nereis (Polychaeta: Nerei- didae) from Todos Santos Bay, Ensenada, Baja California, México .......................00.e eee Lin, Ching-Long and Ju-shey Ho. Two new species of ergasilid copepods parasitic on fishes cul- tured.im brackish) water: muPaiwamn -j.2sace.osntesan ene aradae enc ce se meee ee eee oE eee nee cee Lu, Hua and Kristian Fauchald. Description of Eunice weintraubi and E. wui, two new species of eunicidipoly chaetesstrommorthems" GultsoieViecxicOmer eer EeEEEEEEEEEEEE eee EEE eee eee EEE eee eeere Lu, Hua and Kristian Fauchald. Marphysa belli (Polychaeta: Eunicidae) and two related species, Marphusa oculata and M. totospinata, a new species, with notes on size-dependent features 28-34 420-424 511-520 663-673 737-173 531-535 795-7198 942-953 883-888 634-644 295-302 303-313 916-920 708-719 799-806 263-271 97-101 314-319 140-145 921-928 535-550 807-822 674-693 823-828 15-27 230-240 829-842 Lucas, Spencer G., Robert J. Emry, and Scott E. Foss. Taxonomy and distribution of Daeodon, an Oligocene-Miocene entelodont (Mammalia: Artiodactyla) from North America .............-.... Lucas, Spencer G., Robert J. Emry, and Pyruza A. Tleuberdina. Franconictis (Mammalia: Carmiv- Brenton tie tate Oloocene of ecastemm Kazakstan 2222 208. -donseen oe ood cewek oa se aes ne tee e sce Ludwig, Craig A. Type locality and taxonomic status of Saltator plumbiceps ~“Baird. MS.-° Law- HEC MENG CAVES: Passeriornes: Cardimahidae) =. 3205.2. 322. 252 fo hc ook nce eee ko nhc cto ccc en nee Manning, Raymond B. and Akio Tamaki. A new genus of ghost shrimp from Japan (Crustacea: 2 BSS TG Te (Sa TEE SS GEIS) SE ee ae ea a ee ee ee Manoleli, Dan G., Donald J. Klemm, and Serban M. Sarbu. Haemopsis caeca (Annelida: Hirudinea: Arhynchobdellida: Haemopidae), a new species of troglobitic leech from a chemoautotrophically PESEL TER SL TT a oe Marshall, Harold G. and Lubomira Burchardt. Phytoplankton composition within the tidal fresh- NOPE? GEOG GE Lie WETS a fee TT ee 2 ee ee McLaughlin, Patsy A. and Michéle de Saint Laurent. A new genus for four species of hermit crabs formerly assigned to the genus Pagurus Fabricius (Decapoda: Anomura: Paguridae) ............ McLelland, Jerry A. and Gabriele H. Meyer. Ekleptostylis heardi (Diastylidae), a new cumacean SPSMES HOUT SON gs B10 ee Medellin, Rodrigo A., Alfred L. Gardner, and J. Marcelo Aranda. The taxonomic status of the Yucatan brown brocket, Mazama pandora (Mammalia: Cervidae) ...............-...2220222222+++- Murano, Masaaki and Manuel Rafael Bravo. Parapetalophthalmus suluensis, a new genus and species (Crustacea: Mysidacea: Petalophthalmidae) from the Sulu Sea ............................ Murano, Masaaki and Anton McLachlan. A new species of the genus Gastrosaccus (Crustacea: Pesiel tec ee NOUS OAC) SFO OHNAD 6 assess o’ Colombia (n = 8), Venezuela (n = 8). series of Amazilia and Hylocharis in the American Museum of Natural History, New York. Color descriptions were made under Ex- amolites (MacBeth). Measurements of wing chord, bill length (from anterior extension of feathers), and rectrix length (from point of insertion of the central rectrices to the tip of central and outermost rectrices) were taken with digital calipers and rounded to the nearest 0.1 mm (Table 1). Measure- ments and least squares regression lines were projected on bivariate plots to illus- trate size differences (Wilkinson 1989). The hybrid diagnosis was approached in a hierarchical manner. The presumed paren- tal species of A. distans were hypothesized through the comparative analysis of plum- age pattern and color, feather shape, and bill color. As a second step, the restrictive hy- pothesis was tested with the quantitative analysis of size and external proportions. Concordance of results is regarded as strong support for the hypothesis (Graves 1990, 1993a, 1993b, 1996a; Graves & Zusi 1990). Atavism or hybrid luxuriance has not been demonstrated in hybrid humming- birds (Banks & Johnson 1961, Graves 1990). For brevity, A. distans will be re- 29 ferred to as a hybrid in the remainder of this paper. Results and Discussion Several characters of the hybrid permit its parental species to be identified: (a) bill red tipped with black in life; (b) base of bill conspicuously swollen, nasal flanges un- feathered and exposed; (c) crown glittering bluish-green; (d) throat glittering bluish- green, chin and upper throat streaked with white; (e) indistinct white pectoral spot; (f) abdomen gray along midline; and (g) rec- trices black, innermost and outermost about the same length (Appendix 2; Fig. 1, 2; Ta- ble 1). None of the potential parental spe- cies considered one at a time exhibits this suite of character states in definitive or sub- definitive plumage. The red bill of the hybrid appears to be the most useful character for initially nar- rowing the field of potential parental spe- cies. Adult males of several species in Ap- pendix 1 have pink or red mandibular (low- er jaw) ramphothecae (Lophornis delattrei, L. stictolophus, Chrysuronia oenone, Hy- locharis cyanus, Amazilia versicolor, A. fimbriata, and A. viridigaster), and some specimens of A. fimbriata have pinkish- brown maxillary ramphothecae (upper jaw). However, bright red maxillary ramphothe- cae are found in only three species, L. de- lattrei, L. stictolophus, and Hylocharis cy- anus. Lophornis can be eliminated as pos- sible parents of A. distans because they possess elongated rufous crests and pre- dominately rufous rectrices, which would almost certainly be expressed in a hybrid. Hylocharis cyanus is thus identified as the parental contributor of the red maxillary ramphotheca of the hybrid. Identifying the second parental species is most easily accomplished by focusing on the plumage characters of the hybrid that are lacking in Hylocharis cyanus. Plumage of the head, chin, throat and upper breast of H. cyanus is glittering purple. The in- heritance of iridescence in hybrid hum- 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Lateral and ventral views of male Amazilia fimbriata (top), Hylocharis cyanus (bottom), and their putative hybrid, A. distans Wetmore & Phelps (USNM 461695). VOLUME 111, NUMBER 1 Fig. 2. mingbirds is poorly understood (Graves 1990, Graves & Zusi 1990). In this case, however, I assume that hybridization be- tween two purple-crowned species would not result in offspring with a bluish-green crown. The glittering bluish-green crown and throat of the hybrid suggest that the second parental species has iridescent green plumage in these areas, ruling out Klais guimeti (purple chin and upper throat) as a parental species. In another example, the rectrices of Chrysuronia oenone are shining coppery-gold on both dorsal and ventral surfaces, whereas the rectrices of the hybrid are black, similar to those of H. cyanus (bluish-black). Hybridization of C. oenone and H. cyanus would likely produce off- spring with bronze-colored or dark brown rectrices that are significantly paler, less melanized, than those of the hybrid. In a similar fashion, Campylopterus falcatus (chestnut rectrices, thickened primary ra- chises), Colibri thalassinus and C. corus- cans (purple auricular tufts, banded rectri- ces), Chlorostilbon poortmani (shining golden-green tail), Chlorestes notatus (bril- liant bluish-green plumage from breast to 31 Lateral view of head and bill of the type of Amazilia distans Wetmore & Phelps (USNM 461695). undertail coverts), Chalybura buffonii (lengthened silky-white undertail coverts), Heliomaster longirostris (tail spots, brilliant magenta gorget), Thalurania furcata (pur- ple lower breast, deeply forked tail), Helio- doxa leadbeateri (violet crown patch), Ster- noclyta cyanopectus (violet breast patch, white-tipped rectrices, heavy curved bill), Coeligena coeligena (brown plumage), Ocreatus underwoodii (racket-tipped rectri- ces, tibial ‘“‘puffs’’), Aglaiocercus kingi (greatly elongated rectrices with metallic bluish-green dorsal surfaces), and Chaeto- cercus jourdanii (rufous shafts of rectrices, rose throat), can be removed from the list of potential parental species because they exhibit plumage characters not expressed in the hybrid. By the process of elimination, the second parental species appears to be one of three species of Amazilia that are sympatric with Hylocharis cyanus in Tach- ira, Venezuela (Appendix 1). Wetmore & Phelps (1956:4) noted that the type of A. distans had the general ap- pearance of Amazilia fimbriata, differing from that species “in the glittering blue foreneck and upper breast, and in possess- 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 60 55 G = 2 @® = D & = 50 45 24 26 28 30 32 34 Length Rectrix 5 Fig. 3. 24 22 20 Bill Length 24 26 28 30 32 34 Length Rectrix 5 Bivariate plots of selected measurements (see Table 1) of male Hylocharis cyanus (diamonds), Ama- zilia fimbriata (triangles), and their putative hybrid (filled circle), A. distans Wetmore & Phelps (USNM 461695). Least squares regression lines are illustrated for comparison. ing a crown spot differing in color from the rest of the head . . . . the appearance of the specimen is so distinct from that of other species of the genus [Amazilia] that we have no hesitance in describing it as rep- resenting a new species.”’ I concur with Wetmore and Phelps that A. fimbriata bears more than a fleeting resemblance to Ama- zilia distans in plumage pattern. In fact, the hybrid is nearly intermediate in appearance between Hylocharis cyanus and Amazilia fimbriata. Significantly, the hybrid lacks plumage traits that characterize A. versicol- or (e.g., dark subterminal band on the out- ermost rectrices) and A. viridigaster (e.g., brown or buff undertail coverts). In conclu- sion, evidence gleaned from bill and plum- age characters suggest that A. distans rep- resents a hybrid of Hylocharis cyanus and Amazilia fimbriata. External measurements.—Measurements of avian hybrids fall within the mensural ranges exhibited by their parental species as a consequence of a polygenic mode of in- heritance (see Buckley 1982). External measurements of adult male Hylocharis cy- anus and Amazilia fimbriata overlap and the difference in character means (larger species divided by smaller) is modest: wing chord (9.8%) bill length (17.2%); rectrix 1 (11.2%); and rectrix 5 (14.6%). Consistent with the hypothesis derived from plumage color and pattern, measurements of the hy- brid fall between the character means of the parental species (Table 1, Fig. 3). Had the hybrid’s measurements fallen outside the range of those of Hylocharis cyanus and Amazilia fimbriata, this particular hybrid hypothesis would have been rejected. In summary, both plumage and morpho- logical data are consistent with the hypoth- esis that Amazilia distans represents a hy- brid between Hylocharis cyanus and Ama- zilia fimbriata. These species overlap ex- tensively in Amazonia. For taxonomic pur- poses the Amazilia distans Wetmore & Phelps is available only for the purpose of homonymy. Berlioz (1929) described a supposed hy- brid specimen, Hylocharis cyanus X Ama- zilia fimbriata, prepared in the “Bahia” style and presumably collected in Brazil. Unfortunately, he failed to report the spec- imen’s registration number or in what mu- seum the specimen was deposited. Later, he (Berlioz 1951:287 equivocated in his iden- tification, suggesting that the specimen might represent Hylocharis pyropygia (Sal- vin & Godman 1881), poorly-known and VOLUME 111, NUMBER 1 somewhat doubtful species from Bahia, Brazil (see Sibley & Monroe 1990): “D/ailleurs, faute de connaitre alors des Hyl. pyr- opygia authentiques, j’avais primitivement décrit ce specimen . . . comme étant probablement un hy- bride: Agyrtrina [Amazilia] fimbriata nigricauda X Hylocharis cyanus. Sans rejeter définitivement cette hypothése, trés justifiable par l’apparence de 1’ Oiseau, il me semble pourtant plausible, mainten- ant que l identification, comme espéce distincte, d’Hyl. pyropygia s'est affirmée par l’existence de plusieurs spécimens identiques, de considérer dub- itativement cet Oiseau comme référable aussi a cette derniére espéce.” To further complicate matters, Berlioz (1938) had proposed in earlier paper that Hylocharis pyropygia was actually a hybrid between Chlorostilbon aureoventris and Hylocharis cyanus. In any case, there ap- pears to be no previous verified examples of the hybrid combination reported here (Aylocharis cyanus X Amazilia fimbriata). Sight records.—Sight records of “‘Ama- zilia distans” in northwestern Venezuela and adjacent Colombia (see Hilty & Brown 1986, Collar et al. 1992) are problematic, and, to my knowledge, none is supported by diagnostic photographs. Although these sightings may refer to Hylocharis X Ama- zilia hybrids, they more likely represent the manifestation of imaginations fertilized by the possibility of observing a narrowly dis- tributed endemic. Identification of hum- mingbird hybrids under field conditions is virtually impossible (Graves 1996b). Acknowledgments I thank Nigel Collar, Steve Hilty, Miguel Lentino, and David Wege for clarifying the status of Amazilia distans in the Coleccién Ornitol6gica Phelps. The manuscript was improved by the reviews of Richard Banks, Douglas Stotz, and Richard Zusi. I thank Leo Joseph and David Agro (Academy of Natural Sciences of Philadelphia) for loan- ing comparative material, and the curators and staff of the American Museum of Nat- ural History, New York, for permitting me to work in their collections. Finally, I thank 33 Carl Hansen (Smithsonian photographic services) for providing good photographs. Literature Cited Banks, R. C., & N. K. Johnson. 1961. A revision of North American hybrid hummingbirds.—Con- dor 63:3-28. Berlioz, J. 1929. Un cas nouveau d’hybridité chez les Trochilidés.—L Oiseau 10:340—343. . 1938. Notes critiques sur des Trochilidés.— L Oiseau (new series) 8:3—-19. . 1951. Etude systématique de quelques espe- ces litigieuses de Trochilidés——L Oiseau 21: 278-288. Buckley, P. A. 1982. Avian genetics. Pp. 21-110 in M. Petrak, ed., Diseases of cage and aviary birds. 2nd ed. Lea and Febiger, Philadelphia, 680 pp. Collar, N. J., L. PR Gonzaga, N. Krabbe, A. Madrofio Nieto, L. G. Naranjo, T. A. Parker, II, & D. C. Wege. 1992. Threatened birds of the Americas: The ICBP/IUCN Red Data Book, 3rd edition, part 2. International Council for Bird Preser- vation, Cambridge, U.K., 1150 pp. Graves, G. R. 1990. Systematics of the “‘green-throat- ed sunangels”’ (Aves: Trochilidae): valid taxa or hybrids?—Proceedings of the Biological Soci- ety of Washington 103:6—25. . 1993a. Relic of a lost world: a new species of sunangel (Trochilidae: Heliangelus) from “Bogota.” —Auk 110:1-8. . 1993b. A new hybrid manakin (Dixiphia pi- pra X Pipra filicauda) (Aves: Pipridae) from the Andean foothills of eastern Ecuador.—Pro- ceedings of the Biological Society of Washing- ton 106:436—441. 1996a. Hybrid wood warblers, Dendroica striata X Dendroica castanea (Aves: Fringilli- dae: Tribe Parulini) and the diagnostic predict- ability of avian hybrid phenotypes.—Proceed- ings of the Biological Society of Washington 109:373-390. 1996b. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 1. Characterization of Ca- lypte anna X Stellula calliope and the possible effects of egg volume on hybridization poten- tial—Proceedings of the Biological Society of Washington 109:755-763. , & R. L. Zusi. 1990. An intergeneric hybrid hummingbird (Heliodoxa leadbeateri * Helian- gelus amethysticollis) from northern Colom- bia.—Condor 92:754-760. Hilty, S. L., & W. L. Brown. 1986. A guide to the birds of Colombia. Princeton University Press, Princeton, New Jersey, 836 pp. Meyer de Schauensee, R., & W. H. Phelps, Jr. 1978. 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A guide to the birds of Venezuela. Princeton University Press, 424 pp. Morony, J. J., Jr, W. J. Bock, & J. Farrand, Jr. 1975. Reference list of the birds of the world. Amer- ican Museum of Natural History, New York, 207 pp. Ortiz-Crespo, FE I. 1972. A new method to separate immature and adult hummingbirds.—Auk 89: 851-857. Phelps, W. H., & W. H. Phelps, Jr. 1958. Lista de las aves de Venezuela con su distribucion. Tomo 2, Parte 1. Editorial Sucre, Caracas, 317 pp. Salvin, O., & EF D. Godman. 1881. On some new and little-known species of Trochilidae.—Ibis (se- ries 4) 5:595—-597. Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Uni- versity Press, New Haven, Connecticut, 1111 pp. Wetmore, A., & W. H. Phelps, Jr. 1956. Further ad- ditions to the list of birds of Venezuela.—Pro- ceedings of the Biological Society of Washing- ton 69:1—10. Wilkinson, L. 1989. SYSTAT: the system for statis- tics. SYSTAT, Inc., Evanston, Illinois, 822 pp. Appendix 1 Species of hummingbirds that occur regularly below 1000 m elevation in southwestern Estado Tachira and extreme western Estado Apure, Venezuela: Campylop- terus falcatus, Colibri thalassinus, C. coruscans, Klais guimeti, Lophornis delattrei, L. stictolophus, Chlores- tes notatus, Chlorostilbon mellisugus, C. poortmani, Thalurania furcata, Hylocharis cyanus, Chrysuronia oenone, Amaczilia versicolor, A. fimbriata, A. viridi- gaster, Chalybura buffonii, Heliodoxa leadbeateri, Sternoclyta cyanopectus, Coeligena coeligena, Ocrea- tus underwoodii, Aglaiocercus kingi, Heliomaster lon- girostris, Chaetocercus jourdanit. Appendix 2 General comparative description of definitive plum- ages of male Hylocharis cyanus, Amazilia fimbriata, and the hybrid, H. cyanus X A. fimbriata (=Amazilia distans Wetmore & Phelps, 1956; USNM 461695). Descriptions of structural colors are unusually subjec- tive, as color seen by the observer varies according to the angle of inspection and direction of light. For this reason I use general color descriptions. The forecrown and crown (to a line drawn across the crown at the rear of the orbits) of cyanus are glit- tering purple, bordered posteriorly by dark bluish- green on the hindcrown. The hindneck, upper back, and scapulars are dark green, gradually turning to bronzy green and then coppery on the lower back and rump, respectively; the upper-tail coverts are purplish black. In fimbriata the dorsal plumage (capital and spi- nal tracts) is primarily dark green, with bronze reflec- tions on the crown and upper-tail coverts. The dorsum of the hybrid is intermediate in appearance between cyanus and fimbriata, but more closely resembling the latter species. The forecrown is glittering greenish-blue and the upper-tail coverts are dark bronzy green. The sides of the head, throat, and upper breast of cyanus are deep glittering purple; exposed white feath- er bases on the chin impart a spotted or mottled ap- pearance. Feathers of the lower breast, sides, and flanks are dark brownish-gray tipped with a dark green disc; greenish feather tips are less apparent near the midline. Vent feathers are white and the under-tail co- verts are dull brownish-black (blue reflections in bright light). Feathers of the chin, throat, and upper breast of fimbriata have glittering green discs (when viewed head-on); feathers are white basally, narrowly fringed with white. White feather margins and a few complete- ly white feathers form an indistinct spot near the center of the lower breast. The belly, sides, and flanks are green with an indistinct grayish-white stripe along the midline. Vent feathers are white; under-tail coverts are dark gray (with greenish reflections) moderately mar- gined with white or pale grayish-white. The venter of the hybrid more closely resembles that of fimbriata. Feather discs of the chin, throat, and upper breast are bluish-green, a few are distinctly purple. Traces of the white pectoral spot of fimbriata are present (one com- pletely white feather); under tail coverts are dark slate gray margined with dull white. The tail of cyanus is bluish-black. In fimbriata, the outer rectrices (2—5) are dull bluish-black; the outer margins of rectrices 2—4 are glossed with dark green. The central rectrices (1) are dark green, becoming dull bluish-black distally. The tail of the hybrid is similar in color to that of cyanus, but the outer margins of rectrices 2—4 are faintly glossed with bronzy-green; the basal two-thirds of the central rectrices (1) are glossed with bronzy-green. The maxillary ramphotheca is red, tipped with black in cyanus, and moderately to heavily melanized (pink- ish-brown to black in life) in fimbriata. Ramphotheca of the hybrid exhibits an intermediate amount of mel- anin; the specimen tag notes that the bill was red with a black tip in life. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(1):35—42. 1998. A new species of the catfish genus Glanapteryx (Siluriformes: Trichomycteridae) Mario C. C. de Pinna Department of Zoology, Universidade de Sao Paulo, Caixa Postal 11461 Sao Paulo-SP 05422-970 Brazil Abstract.—A new species of the glanapterygine trichomycterid genus Glan- apteryx is described from the upper Rio Negro in Brazil, State of Amazonas. The new species, currently represented by a single specimen, is the only fish so far known to occur in the remote “Morro dos Seis Lagos’”’ lake complex, a region with high levels of natural radioactivity. Glanapteryx niobium, new species, is distinguished from its only congener, G. anguilla, by a white collar immediately posterior to head, the dark pigmentation on the ventral surface of the head, the longer pectoral-fin remnant, the lanceolate caudal fin, and the narrow union of branchial membranes to the isthmus. Resumo.— Uma nova espécie do género Glanapteryx (Trichomycteridae, Glanapteryginae) é descrita do alto Rio Negro, Estado do Amazonas, Brazil. A nova espécie, atualmente conhecida por um Unico exemplar, é 0 Unico peixe encontrado até 0 momento no remoto complexo de lagos chamado Morro dos Seis Lagos, uma regiao com altos niveis de radioatividade natural. Glanapteryx niobium, espécie nova, distingue-se da Unica outra espécie do género, G. an- guilla, pelo colar branco logo apés a cabecga, a pigmenta¢ao escura na face ventral da cabega, 0 maior comprimento da nadadeira peitoral, a forma lan- ceolada da nadadeira caudal e a uniao estreita das membranas branquiais ao istmo. The remarkable catfish genus Glanapte- ryx was established by Myers (1927) to in- clude a single eel-like species, G. anguilla, from the upper Rio Negro in Brazil. The species was subsequently illustrated in My- ers (1944), along with descriptions of other members of the subfamily Glanapteryginae, which was established in the same paper. Glanapteryx anguilla was redescribed by de Pinna (1989) on the basis of additional ma- terial mostly from the rio Negro in Brazil. That publication also included osteological data which formed the basis for a phylo- genetic diagnosis of the genus and species, and a hypothesis of phylogenetic relation- ships among glanapterygines. The occur- rence of G. anguilla in the Orinoco basin was suggested by de Pinna (1989) and later confirmed by Nico & de Pinna (1996). Glanapteryx has to date been restricted to its type species, and although represen- tatives of the genus are now being collected regularly, nothing is known of their biolo- gy. The present paper reports on a distinc- tive second species of the genus. The new species is remarkable mainly in its occur- rence in a remote region in the Amazon called Morro dos Seis Lagos (meaning “hill of the six lakes’), a large and poorly- known complex of relatively high-elevation lakes in the upper Rio Negro. The region is known for its high level of natural radio- activity. The new Glanapteryx seems to be the only fish species occurring in these lakes. Material and methods.—All measure- ments are straight-line, taken according to the protocol described in de Pinna (1989). 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Abbreviations are: CAS (Califomia Acad- emy of Sciences, San Francisco); INPA (In- stituto Nacional de Pesquisas da Amazonia, Manaus); MZUSP (Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo); ex (number of specimens); C&S (material cleared and stained); SL (standard length); HL (head length); an (anus); pf (pelvic fin); and up (urogenital papilla). Comparative glanapterygine material ex- amined-Glanapteryx anguilla: CAS 56048 (holotype), MZUSP 36530 (21 ex, 2 C&S); Listrura nematopteryx: MZUSP 36974 (ho- lotype), MZUSP 36975 (12 paratypes), MZUSP uncat. (Sex C&S); Listrura cam- posit: MZUSP uncat. (lex); Pygidianops ei- genmanni: CAS 11121 (@ paratypes, 1 C&S); Pygidianops sp.: INPA 8080 (3ex); Typhlobelus ternetzi: CAS 56201 (2 para- types, 1C&S); Typhlobelus sp: INPA 12929 (10 ex, 2 C&S). Comparative material of other trichomycterids is listed in de Pinna (1992). Glanapteryx niobium, new species Fig. 1 Holotype.—INPA 12421, 55.3 mm SL; Brazil, State of Amazonas, Pico da Neblina National Park, Morro dos Seis Lagos (ap- prox. 0°17’N, 66°41'W), Lago Esperanga. Collected by Ulysses C. Barbosa and Victor Py-Daniel, 11 Sep 1990. Diagnosis.—Distinguished from its only congener, G. anguilla, by the following characteristics: 1—presence of a well-de- fined wide white collar-like band shortly posterior to head (Fig. 1; coloration uniform in G. anguilla); 2—ventral part of head darkly-pigmented (Fig. 1B; dark pigmen- tation on ventral surface of head scarce or absent in G. anguilla); 3—caudal fin lan- ceolate, its middle portion projecting be- yond rest of fin (Fig. 2; caudal fin round in G. anguilla); 4—branchial membranes nar- rowly united to isthmus (membranes more broadly united to isthmus in G. anguilla); 5—pectoral fin long (40% of HL), and of even width, not narrowing towards tip (Fig. 3; fin 9—25% of HL and narrowing gradu- ally to tip in G. anguilla). Characters 1 to 3 are considered autapomorphic for the spe- cies (see Discussion). Description.—Morphometric data are provided in Table 1. Holotype preserved in strongly contorted position, SL and propor- tions thereof may be inexact. Overall form of body similar to that of G. anguilla (see de Pinna, 1989, fig. 1). Body eel-like, head continuous with trunk. Body round in cross section for most of its length, gradually more compressed posterior to anal opening. Caudal peduncle gently tapering to caudal fin. Dorsal and ventral profiles of body nearly straight. Anterior portion of caudal peduncle slightly deeper than remainder of body. Dorsal and ventral profiles of caudal peduncle converging gradually to caudal fin. Head small (HL approx. 7% of SL), less deep than body, its dorsal surface flat. Bran- chial membranes narrowly united to isth- mus, gill openings wide, not constricted. Eyes very small but well formed, with dis- tinct lenses and covered by thin transparent integument. Eyes located on nearly vertical surface of head, facing almost laterally. Posterior nares slightly anterior to eyes, very close to mesial margin of eyeball, but separated from it by narrow strip of integ- ument (eye and naris not adpressed as in G. anguilla). Anterior nares opening located on short tube of integument, continuous posterolaterally with nasal barbel. Three large sensory pores situated serially on dor- solateral surface of head and anterior por- tion of trunk. Posterior pore located slightly posterior to vertical through origin of pec- toral fin. Middle pore dorsal to uppermost point of branchial membrane. Anterior pore smallest and positioned dorsal and slightly anterior to middle pore. Mouth subterminal, upper jaw slightly longer than lower, cor- ners not markedly externded posteriorly. Lips poorly differentiated, continuous with remainder of head and with covering of sensory papillae comparable to those over rest of head. All barbels large and robust, VOLUME 111, NUMBER 1 Fig. 1. Glanapteryx niobium, holotype, INPA 12421. Views of head. A—dorsal; B—ventral; Scale bars = 1 mm. 37 C—Jateral. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. A—Glanapteryx anguilla, MZUSP 36530; B—G. ni- obium, holotype, INPA 12421. Scale bar = 1 mm. Caudal fin and part of caudal peduncle. with visible internal cores and similar to each other in general aspect. Maxillary and rictal barbels asymmetric in single known specimen. Right maxillary barbel reaching tip of extended pectoral fin, left barbel reaching only to midlength of same fin. Right rictal barbel extending slightly pos- terior to pectoral-fin base, barbel on left reaching beyond fin tip. Maxillary barbel longer than rictal on right, situation re- versed on left side. Nasal barbels both ex- tending to anterior margin of white collar. Pectoral fin vestigial, reduced to small flap on side of body, originating immedi- ately posterior to dorsal-most point of bran- chial membrane. Width of pectoral fin near- ly even from base to tip, about as wide as rictal barbel at base, its tip round. Three pectoral-fin rays present, but visible only with strong transmitted light, apparently un- branched and unsegmented. First ray longer and thicker than other two, extending to tip of fin, third ray shortest. Pelvic fin vestigial, reduced to two roughly triangular flaps an- terior to anal opening (Fig. 4). Dorsal, anal, and adipose fins absent. Caudal fin small and inconspicuous, continuous with re- mainder of caudal peduncle and body, lan- ceolate in shape, with middle rays longest. Caudal-fin rays 2 + 2 (ie., only two B Fig. 3. Profile of pectoral fin. A—Glanapteryx an- guilla, MZUSP 36530; B—G. niobium, holotype, INPA 12421. branched rays). Branched rays with incipi- ent segmentation proximally. Procurrent caudal-fin rays numerous, at least 25 dor- sally and ventrally. Exact number difficult to determine in alcoholic specimen. Pigmentation in preservative.—Overall coloration uniform dark tan, lighter on ven- tral half of head and body. Wide white area (about 0.5 HL) located on body shortly pos- terior to head, forming well-defined collar encircling whole circumference of body. Collar formed by abrupt disappearance of dark chromatophores distributed on rest of Table 1—Morphometric data for holotype of Glan- apteryx niobium, INPA 12421 (in mm or as proportion of SL, TL, or HL, as indicated in parentheses). head length 4.0 (mm) head width 0.75 (HL) head depth 0.52 (HL) mouth width 0.48 (HL) interorbital 0.41 (HL) eye diameter 0.06 (HL) anterior internarial width 0.33 (HL) posterior internarial width 0.24 (HL) collar width (lateral view) 0.40 (HL) pectoral-fin length 0.40 (HL) standard length 55.3 (mm) total length 1.05 (SL) preanal length 0.65 (TL) body depth 0.05 (TL) caudal peduncle length 0.28 (TL) caudal peduncle depth (max.) 0.05 (TL) VOLUME 111, NUMBER 1 39 Fig. 4. Pelvic fins and surrounding structures in Glanapteryx niobium, holotype, INPA 12421, ventral view. Scale bar = 1 mm. body. Caudal peduncle with more frag- mented covering of dark pigmentation. Ventral surface of body, except for collar, with uniform scattering of dark chromato- phores, less dense than on dorsal surface but still conspicuous. Head with slightly denser covering of dark chromatophores than those on body. Ventral surface of head and anterior portion of trunk with dense covering of melanophores, only slightly sparser than on dorsal surface. Cephalic sensory-canal pores with narrow white rim. All barbels with fields of dark chromato- phores at base, abruptly fading shortly dis- tal to that point. Narrow white ring around eyes, widest at posteroventral margin. Dis- tal portion of branchial membranes lacking melanophores. Pectoral fin almost totally white, only few dark chromatophores lo- cated near base. Base of caudal fin with same pigmentation as caudal peduncle, its distal portion white. Distal portion of pro- current-ray dorsal and ventral areas without dark chromatophores. Pelvic fin remnant lacking dark pigmentation. Etymology.—tThe specific epithet, niobi- um, a noun in apposition, refers to niobium, the chemical element chiefly responsible for the high background radiation of the Morro dos Seis Lagos, the known fish fauna of which is so far limited to the new Glan- apteryx species. Discussion.—Although as yet represent- ed by a single specimen, there is little doubt that G. niobium is distinct from G. anguilla. The latter is currently known by tens of specimens, a sample which allows a satis- factory estimate of the degree of intraspe- cific variation expected for the genus (cf. de Pinna 1989). The differential characters displayed by G. niobium, summarized in the diagnosis above, have not been seen in any examined specimen of G. anguilla. The new species is readily distinguisha- ble from its only congener mainly by the white collar formed by a well-defined area lacking dark chromatophores. This area forms a white ring around the whole cir- cumference of the body, and is striking against the dark pigmentation of the re- 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mainder of the fish. In G. anguilla, the dark pigmentation is even along the whole length of the body, with no fading in the collar region. The white collar of G. nio- bium is unique among the Trichomycteri- dae, and can be considered autapomorphic for the species. The lack of pigmentation in the collar region is particularly striking be- cause the remainder of the body and head surface in G. niobium is more heavily pig- mented than in G. anguilla, which contrib- utes to a marked contrast. The white collar is evident even in ventral aspect, because in G. niobium the ventral surface of the head and anterior portion of the abdomen is more heavily pigmented than what is usual in other glanapterygines and trichomycterids in general. The heavy dark pigmentation on the ventral side of G. niobium is itself au- tapomorphic. The lanceolate shape of the caudal-fin in G. niobium is also unique to that species among trichomycterids. The caudal fin in the family is commonly round, furcate or emarginated, and the unique lanceolate shape in G. niobium is considered an aut- apomorphy for the species. The peculiar caudal shape is a result of the prolongation of the five middle caudal-fin rays, which are far longer than the others. The pectoral-fin length readily distin- guishes G. niobium and G. anguilla. In G. anguilla the length of the fin is at most 25% of HL, never reaching 40% of HL as seen in G. niobium. This characteristic, however, is not autapomorphic, because the relative length of the fin varies widely in glanapter- ygines, making a polarity assessment of the condition in G. niobium uncertain. The same uncertainty applies to the shape of the fin, another character that apparently distin- guishes the two species but which cannot unabiguously be determined as autapo- morphic for either. An additional difference that may be ob- served between the holotype of G. niobium and most specimens of G. anguilla is the number of sensory pores on the posterior part of head (3 versus 4, respectively). These pores trace the highly reduced latero- sensory canal system of glanapterygines. The two anterior pores are openings of the postotic canal (running through pterotic, see Arratia & Huaquin 1995) and the two pos- terior ones represent the lateral-line canal (running through supracleithrum and end- ing shortly posterior to it). The posterior branch of the lateral line in some specimens splits further and opens into a tiny addi- tional posterior pore (not considered in the discussion below). On the basis of topo- graphical correspondence, the pore lost in G. niobium seems to be the last one (pos- terior lateral-line pore) present in four- pored G. anguilla. However, the number of pores is intraspecifically variable in G. an- guilla. While most specimens examined of the species indeed have four pores (cf. fig. 2C in de Pinna 1989), a few specimens have only three pores, like in the only known specimen of G. niobium. Therefore, the difference in sensory-pore number can- not be used to separate the two Glanapteryx species confidently. The infraorbital canal in G. anguilla (ancomplete as in all other trichomycterids except Trichogeninae and Copionodontinae) is bifurcated distally, and opens through two minute pores posterior to the eye (cf. de Pinna 1989, fig. 5A). These pores could not be located in G. ni- obium, but they are also invisible externally in some specimens of G. anguilla, and can- not be confirmed as absent in G. niobium until more specimens are available for an- atomical studies. Not all characters diagnostic for Glan- apteryx given by de Pinna (1989: 363) can be checked in G. niobium, which is known only from the holotype. Of the eight char- acters proposed by de Pinna, numbers 2 (triangle-shaped premaxilla), 3 (simplified pelvic bone, when present), and 5 (pro- nounced interdigitations between frontals, sphenotics and supraoccipital in fully- grown individuals) are internal traits pres- ently unobservable in the new species. Glanapteryx niobium, however, demon- strates all of the remaining characters. VOLUME 111, NUMBER 1 Those are (numbering of de Pinna 1989): 1—absence of the anal fin; 4—reduced, diphycercal caudal fin; 6-eighty-eight or eighty-nine vertebrae (not directly observed in G. niobium, but likely in view of its body elongated to a degree similar to that of G. anguilla); 7—posterior naris mesial to eye, adjoined to mesial margin of eyeball; 8 combination of three-rayed and short pec- toral fin. Not all of those characters are un- ambiguous evidence of monophyly. The anal fin is also lacking in some recently dis- covered, as of yet undescribed glanaptery- gine species, seemingly more closely relat- ed to genera other than Glanapteryx. The same applies to the diphycercal and reduced caudal fin. Character 8 is a combination character, and although appropriate for identification, does not hold as evidence for monophyly (Pygidianops has a short but one-rayed pectoral fin, while Listrura cam- posi has a three-rayed but long fin). Remaining characters seem to provide relatively reliable but still circumstantial evidence of relationships. The position of the posterior nares is indeed unique, but other glanapterygines (Pygidianops and Ty- phlobelus) have eyes greatly reduced or lost, making a comprehensive comparison impossible. Still, allocation of G. niobium to the genus Glanapteryx is the best course of action based on the combination of all the evidence above. Some internal characters illustrated and described by de Pinna (1989) seem unique to G. anguilla among the trichomycterids so far examined, although not yet explicitly proposed as synapomorphic. These are the posteriorly tripartite palatine; the enlarged head of the vomer; the anterior canal-bear- ing part of the sphenotic separated from the frontal; and the cartilage plug on the pos- terolateral margin of the premaxilla. All those characteristics are probably either au- tapomorphies for G. anguilla or synapo- morphies for Glanapteryx. Their exact level of generality will have to await examination of the internal anatomy of G. niobium. Habitat notes.—Glanapteryx niobium is 41 the only fish species known to date from the Morro dos Seis Lagos lake complex. The collecting effort in the area intensively sampled various microhabitats with differ- ent fishing gear. Still, only the single spec- imen of G. niobium was found in the Lago Esperanga (a second specimen of seemingly the same species was captured in that lake, but was subsequently lost). Similar collect- ing efforts were undertaken in all the other five lakes, but failed to secure any fish. The Morro dos Seis Lagos is located in- side the Yanomami indigenous preserve, it- self part of the Pico da Neblina National Park. It is located approximately 60 km northeast of Sao Gabriel da Cachoeira. The Morro dos Seis Lagos is an isolated round outcrop 6 km in diameter, about 40 km away from the nearest elevated areas (Serra do Padre, to the north). It is covered by thick laterite crust, reddish brown in color. Morro dos Seis Lagos includes six major lakes at an altitude of 300 m, plus a number of smaller water bodies. Those are the only true lakes in the Brazilian Amazon, and are permanently isolated from other water courses. The lake beds were a consequence of the collapse of underlying rocky blocks. The level of radiation in the region is ex- tremely high, because of the concentration of radioactive minerals naturally in the soil, mainly niobium, thorium and cerium. Ra- diation detectors worn by expedition mem- bers in the field recorded daily radiation ex- posures equivalent to the maximum consid- ered tolerable for a whole week according to international standards. In several places the measured emission was well over 25 milliroentgens. One of the creeks around the lake, Igarapé Ya-Mirim, is so radioac- tive as to cause itching in bathers after re- peated exposure, and is called “itching creek”’ by indians, who avoid settling in the region. There is also a thermal spring in the area. The spot where G. niobium was collected with a hand seine was about 1 m deep, and had a thick layer of leaf litter on the bottom, amidst which the fish was hiding. The water 42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON was transparent, green at distance, still, and acidic (pH 4.0). The invertebrate fauna was reported as rich by the collectors. Acknowledgments I thank Victor Py-Daniel and Ulysses Barbosa for collecting and bringing to my attention the single known specimen of G. niobium. Py-Daniel also provided the hab- itat data. The expedition to Morro dos Seis Lagos was part of the First Brazilian Mul- tidisciplinary Expedition to Pico da Nebli- na, organized and funded by INPA in 1990. A visit to Manaus during which I was able to examine the INPA ichthyological collec- tions was sponsored by the Graduate Pro- gram in Entomology of that institution, through José A. Rafael. I am also grateful to Cristina Cox-Fernandes, Paulo Petry, Ef- rem Ferreira, and Labish N. Chao for their help during my visit. The manuscript ben- efitted from reviews by Richard Vari and Scott Schaefer. Research funding is provid- ed by CNPq and FAPESP. Literature Cited Arratia, G., & L. Huaquin. 1995. Morphology of the lateral line system and of the skin of diplomys- tid and certain primitive loricarioid catfishes and systematic and ecological considerations.— Bonner Zoologische Monographien 36:1—110. Myers, G. S. 1927. Descriptions of new South Amer- ican fresh-water fishes collected by Dr. Carl Ternetz.—Bulletin of the Museum of Compar- ative Zoology 68:107—135. . 1944. Two extraordinary new blind nematog- nath fishes from the Rio Negro, representing a new subfamily of Pygidiidae, with a rearrange- ment of the genera of the family and illustra- tions of some previously described genera and species from Venezuela and Brazil.—Proceed- ings of the California Academy of Sciences 23: 591-602. Nico, L. G., & M. C. C. de Pinna. 1996. Confirmation of Glanapteryx anguilla (Siluriformes, Tricho- mycteridae) in the Orinoco river basin, with notes on the distribution and habitats of the Glanapteryginae.—Ichthyological Exploration of Freshwaters 7(1):27—32. de Pinna, M. C. C. 1989. Redescription of Glanap- teryx anguilla, with notes on the phylogeny of Glanapteryginae (Siluriformes, Trichomycteri- dae).—Proceedings of the Academy of Natural Sciences of Philadelphia 141:361—-374. . 1992. A new subfamily of Trichomycteridae, lower loricarioid relationships, and a discussion on the impact of additional taxa for phyloge- netic analysis (Teleostei, Siluriformes).—Zoo- logical Journal of the Linnean Society 106:175— 229. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(1):43-51. 1998. A new species of Nannosquilla (Crustacea: Stomatopoda: Nannosquillidae) from the eastern Pacific and new records of species of Neogonodactylus (Gonodactylidae) from the Pacific coast of Mexico José Salgado-Barragan and Michel E. Hendrickx Estacion Mazatlan UNAM, Apartado Postal 811, Mazatlan, Sinaloa 82000, Mexico Abstract.—A large series of specimens of stomatopods was collected from intertidal and shallow subtidal habitats along the Pacific coast of Mexico. A new species of Nannosquilla Manning is described from the southeastern part of the Baja California Peninsula, and new records are provided for species of Neogonodactylus Manning, including the first record for Mexico of N. laliber- tadensis (Schmitt), previously known from Ecuador and Panama. A total of 50 species and 22 genera of stomatopod crustaceans are known from the eastern Pacific. They belong to three of five currently recognized superfamilies of Sto- matopoda: Gonodactyloidea, Lysiosquillo- idea and Squilloidea (see Hendrickx & Sal- gado-Barragan 1991). Among these, 29 species are known from the Pacific coast of Mexico, eight species belong to the genus Neogonodactylus Manning, and seven to the genus Nannosquilla Manning. Species of the family Nannosquillidae and Gonodactylidae have been somewhat rarely reported in literature dealing with the eastern Pacific. This is mostly due to three factors: with a few exceptions, they are of small size, they often are burrowing species or live among coral or rubble and they gen- erally feature a depth range too shallow to be sampled by research vessels, yet often too deep to be visited by non-scuba divers. Recent collection of specimens of crus- taceans in intertidal and shallow subtidal habitats along the Pacific coast of Mexico yielded some interesting specimens of sto- matopods, including an undescribed species of Nannosquilla. Additional material de- posited in the crustacean collection of UNAM at Mazatlan and belonging to the recently described genus Neogonodactylus Manning (Manning 1995: 80) allows us to report an additional species for the Pacific coast of Mexico and add records for two otherwise scarcely cited species. Abreviations and acronyms used are: coll., collector; TL, total length Gin milli- meters); EMU, Estacion Mazatlan UNAM, Invertebrates Reference Collection; USNM, National Museum of Natural History, Smithsonian Institution, Washington D.C.; UABCS, Invertebrates Collection, Depar- tamento de Biologia, Universidad Auténo- ma de Baja California Sur. Latitude and longitude were obtained in the field with a ICOM global positioning system (GPS). Family Nannosquillidae Manning, 1980 Nannosquilla raymanningi, new species Figs. 1-3 Material examined.—Holotype, 1 female (TL 22.2 mm), Ensenada Grande, Isla Par- tida, Baja California Sur, Mexico (24°33'N, 110°24’W), 17 Aug 1994, 13.7 m, sandy bottom, scuba diving, collected by hand us- ing quinaldina (coll. C. Sanchez Ortiz) (EMU-4620); paratypes 1 male (TL 20.3 mm) and 1 female (TL 22 mm), same lo- cality (EMU-4621); paratypes 1 male (TL 20.7 mm) and 1 female (TL 23.6 mm), same locality (USNM 285512); paratypes 2 females (TL 20.3 and 21.0 mm), same lo- 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Nannosquilla raymanningi, new species, female holotype. a, anterior part of body; b, right raptorial claw, inner face; c, right uropod, ventral view; d, right uropod, dorsal view (EMU-4620). cality (EMU-4622); 1 male (TL 20.3 mm) and 1 female (TL 22.8 mm) (UABCS). Description.—Eye small, short, not over- reaching antennular peduncle. Cornea subglobular, not bilobed, set obliquely on stalk and slightly expanded laterally. Ocular scales fused at base and most of length, but distinctly separated in distal portion; apex acute or rounded. Upper antennular flagel- lum with 11 free articles; lower longer, with VOLUME 111, NUMBER 1 a 45 Fig. 2. Nannosquilla raymanningi, new species, female holotype. a, sixth abdominal somite and telson, dorsal view; b, telson, posterior view; c, telson, ventral view (EMU-4620). 8 to 10 (holotype with 9-9), lower shorter with 4 free articles; antennular process vis- ible laterally, projecting beyond sides of rostral plate and overreaching anterolateral corners of rostral plate. Antennal peduncles short, not overreaching eyes, flagella with 11 articles; antennal scale not extending be- yond midpoint of last segment of antennal peduncle. Rostral plate subrectangular, wid- er than long, covering only proximal mar- gin of ocular peduncles; lateral margins subparallel, slightly convex, anterolateral comers rounded, anterior margins slightly concave, apex angled, not ending in spine. Mandibular palp absent, four epipods pres- ent. Propodus of raptorial claw with four movable teeth on proximal inner margin; dactylus with two proximal notches (in most specimens) and 9 to 11 teeth (9-10 in holotype), including terminal one. Sixth ab- dominal somite with posterolateral corners acute but not produced as spines. Telson short in dorsal view, aproximately 1.3 times wider than long. False eave trilobed, the median distinctly wider, rounded; lateral lobes obtuse in dorsal view; dorsum of tel- son with short, shallow sulcus on each side of median projection of false eave, con- verging to level of median projection; me- dian projection of false eave low, ventrally curved posteriorly, flanked on each side by narrow, deeply concave depressions; sub- median projections rounded. False eave merges with true margin at about level of last fixed lateral tooth. Marginal armature on each side of midline consisting of 9—12 denticles (holotype with 9-10), entire row forming an inverted (widely open) ““V” in posterior view; 1 movable submedian tooth, originating anteriorly, and seven fixed lat- eral teeth and denticles; third denticle small and inserted at a lower (ventrally) level than second denticle; outermost denticle located on false eave. Basal segment of uropod 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. dorsal view of female paratype (EMU-4622). Nannosquilla raymanningi, new species, with ventral, proximal tubercle. Dorsal spine of basal segment of uropod short, not extending to midpoint of endopod. Outer spine of basal prolongation curved, much longer than inner one which is much nar- rower and almost straight. Proximal seg- ment of exopod with 1—4 stiff setae (holo- type with 2 on each side) on inner distal corner, and 5 to 6 spatulate spines on outer distal margin (holotype with 5-5). Color.—Specimens in ethanol are pale, with few, variable, stellate chromatophores distributed over the eyes, rostral plate and dorsum. Carapace, thoracic and abdominal segments 1—4 with sparse chromatophores; a pair of chromatophores lateral to the dor- sal midline. Fifth abdominal somite with posterolateral regions bearing a symmetri- cal dark area (Fig. 3). Remarks.—Nannosquilla raymanningi 1s so far the only species of Nannosquilla from the eastern Pacific with the external spine of the basal prolongation of the uro- pod considerably longer than the inner spine. Five out of the seven species of Nan- nosquilla previously reported from this re- gion feature a longer inner spine: N. decem- spinosa (Rathbun, 1910); N. similis Man- ning, 1972b; N. galapagensis Manning, 1972b; N. canica Manning & Reaka, 1979; N. anomala Manning, 1967. The other two species feature subequal spines. The new species, however, does not show the 10—12 projections seen on the false eave of N. cal- iforniensis (Manning, 1961a); it also pos- sesses 9-12 submedian fixed denticles on the telson instead of eight denticles, and rounded anterolateral angles of the rostral plate contrary to acute angles in N. califor- niensis. It is also easily distinguished from N. chilensis (Dahl, 1954), a species with rostral plate with a broadly rounded antero- lateral angle, and the dactylus of the rap- torial claw with 17 teeth (9-11 in N. ray- manningi). The Atlantic species of Nannos- quilla with the external spine of the basal prolongation of the uropod either longer or similar in size to the inner spine, have few (4-7) submedian denticles and, at most, nine teeth on the dactylus of the raptorial claw; these species are Nannosquilla adkin- soni Camp & Manning, 1982; N. antillensis (Manning, 1961b); N. carolinensis Man- VOLUME 111, NUMBER 1 ning, 1970; N. disca Camp & Manning, 1986; N. heardi Camp & Manning, 1982; N. tobagoensis Schotte & Manning, 1993; N. virginalis Camp & Manning, 1986; N. yucatanica Camp & Manning, 1986. The specimens were collected in an area inhabited by the gregarious “‘garden eel’’, Taenioconger digueti Pellegrin, on sandy bottom, near reefs. Although the specimens of N. raymanningi and T. digueti were ob- tained in the same sample, there is no evi- dence the stomatopods use the burrows of eels as a shelter. Etymology.—We are pleased to name this new species in honor of Raymond B. Man- ning, worldwide expert in stomatopod tax- onomy and ecology, for his astounding con- tribution to the knowledge of this group of crustaceans. Family Gonodactylidae Giesbrecht, 1910 Neogonodactylus lalibertadensis (Schmitt, 1940) Fig. 4 Gonodactylus’ festae lalibertadensis Schmitt, 1940:223, fig. 33. Gonodactylus lalibertadensis.—Manning, 1974:102.—Reaka & Manning, 1980: 5.—Hendrickx & Salgado-Barragén, 1991:29 (table 6). Neogonodactylus lalibertadensis.—Man- ning, 1995:80. Material examined.—Sayulita Bay, Jal- isco (20°52.3'N, 105°28.0’W), 10 Apr 1996, 1 male (TL 20.6 mm) and 1 female (TL 16.7 mm), rocks and algae, 2 m (EMU- 4623).—Chilenos Bay, Baja California Sur (22°56.0'N, 109°48.0’'W), 1 female (TL 13.1 mm), coral and algae, 4 m, 20 Jul 1996 (EMU-4624). Previous records.—Known only from La Libertad, Santa Elena Bay, Ecuador (type locality), and Taboga, Panama (Manning 1974). Color.—The three specimens examined were preserved in ethanol. They have a very similar pattern of chromatophores and dark spots, with only slight variations. 47 Propodus and dactylus of first maxilliped darkened; a dark spot close to the colored spot of the merus of the raptorial claw. There are two submedian spots in the pos- terior dorsal half of the carapace. The fe- male specimen from Chilenos features two additional marginal spots anteriorly. The male sixth thoracic somite bears two sub- median spots, two intermediate spots, and a lateral spot; females show a similar pattern. Abdominal segments 1—5 bear a pair of large submedian boomerang-shaped spots and a pair of intermediate spots; spots are more difuse in the female specimens. The fifth abdominal segment has an additional central spot on all three examined speci- mens. There are two anterior submedian spots on the telson of the Sayulita male and Chilenos female (Fig. 4). Remarks.—The specimens examined agree well with the original description of N. lalibertadensis, including the laterally projected ocular scales and the anterolateral angle of rostral plate which is apically blunt [not spiny or sharp, as in N. festae (Schmitt, 1940) or N. bahiahondensis (Schmitt, 1940)]. Number and location of spines and spinules also correspond to the original de- scription of N. lalibertadensis, except for a higher number of spines on the intermediate accesory carinae of examined females (Ta- ble 1). The only male specimen collected presents an inflated telson with a reduced number of dorsal spines. The material cited under G. lalibertad- ensis by Manning (1972a) was later rec- ognized by Manning & Reaka (1979) as an undescribed species, which they described and named G. costaricensis. Neogonodactylus stanschi (Schmitt, 1940) Gonodactylus stanschi Schmitt, 1940:215, Fig. 30.—Steinbeck & Ricketts, 1941: 429.—Manning, 1972a:110.—Reaka & Manning, 1980:8.—Hendrickx & Salga- do-Barragan, 1989:244 (table 6); 1991: 36, fig. 17. 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Neogonodactylus lalibertadensis (Schmitt, 1940). a, body of male in dorsal view; b, telson of female, dorsal view (EMU-4623). Neogonodactylus 1995:80. Material examined.—San Juan de Alima, Michoacan (18°36.1’N, 103°42.1’W), 5 Nov 1996, 1 male (TL 9.8 mm), intertidal, rocks and algae (EMU-4233).—Punta San- tiago, Manzanillo, Colima, (19°06.5'N, 104°21.0’W), 6 Nov 1996, 1 male (TL 30.5 stanschi.—Manning, mm) and | juvenile (TL 7.2 mm), 2—3 m, rocks and algae (EMU-4234).—El Tama- rindo Beach, Tenacatita Bay, Jalisco (19°15.9’N, 104°47.9'W), 4 Nov 1996, 2 males (TL 13.2 and 18.8 mm), 1 female (TL 11.8 mm), and 2 juveniles (TL 7.4 and 7.5 mm), 1—2 m, rocks and algae (EMU- 4235).—Ensenada de Litigu, Nayarit VOLUME 111, NUMBER 1 49 Table 1.—Pattern of dorsal spination of telson in several Neogonodactylus from the eastern Pacific region. Modified from Manning (1972a) and Schmitt (1940). bahiahondensis festae Median Carina 1 0-1 Accessory Median 0-1 0-4 Anchor absent 34 Knob 2 4-5 Anterior Submedian 1 + 1-2 1 + 0-3 Submedian 1-3 4-6 Accessory Intermediate in | row in 2 rows Lateral Denticle — + (20°47.4’N, 105°31.9'W), 9 Apr 1996, 1 fe- male (TL 29.4 mm), 1 m, dead coral (EMU- 4236).—Chilenos Bay, Baja California Sur (22°56.0'N, 109°48.0’W), 20 Jul 1996, 2 fe- males (TL 29.1 and 30.5 mm), 3—5 m, coral and rocks (EMU-4237).—Playa Calerita, La Paz, Baja California Sur (24°21.0'N, 110°16.0’W), 18 Jul 1996, 1 male (TL 23.7 mm), | juvenile (TL 13 mm), 2—3 m, rocks and algae (EMU-4238).—Los Algodones Bay, Guaymas, Sonora (27°58.6’N, 111°07.7'W), 25 Mar 1997, 1 female (TL 18.3 mm), 1.5—2.5 m, rocks and algae (EMU 4239). Previous records.—E]| Datil, Espiritu Santo, Angel de La Guarda, Isabel and Tres Marias Islands, Gulf of California; San Car- los Bay, Sonora, Tangola-Tangola Bay and Puerto Huatulco, Oaxaca; Punta Mita, Nay- arit; Guaymas, Sonora; Teacapan, Sinaloa; Barra de Navidad, Jalisco; Zihuatanejo, Guerrero; Puerto Lobos and Punta Mar- quez, Baja California Sur; Chamela, Jalis- co, Mexico. Salera and del Cafio Islands, Costa Rica (Hendrickx & Salgado-Barragan 1991). Remarks.—Present records are the first available for the coast of Colima and Mi- choacan, Mexico. Neogonodactylus zacae (Manning, 1972a) Gonodactylus oerstedii Schmitt, 1940:221 (part) figs. 27-28 (not fig. 26 = G. oer- stedii Hansen, 1895; not fig. 29 G. lalibertadensis Female from laliberta- Male from Female from Chilenos costaricensis — stanschi densis Sayulita Sayulita Bay 1 1 1 1 1 1 1 1 2-3 2 2 3 absent absent absent absent absent absent 4 — 24 2-3? 3 4 1+ 1 1 2-5 Da 2 Saad Bar 3 2 0-1 2-5 D) y) 2 in 1 row — 2-3 isp il Ghee 4 3 db Al pumilus Manning, 1970).—Steinbeck & Ricketts, 1941:428 (not G. oerstedii Han- sen, 1895). Gonodactylus zacae Manning, 1972a:107, hea OVA N03 Se hie. pel 976223: Reaka & Manning, 1980:8.—Brusca, 1980:244, figs. 13-10.—Hernandez- Aguilera et al., 1986:190.—Hendrickx & Salgado-Barragan, 1991:39, fig. 19. Neogonodactylus zacae.—Manning, 1995: 80. Material examined.—El Tesoro Beach, La Paz, Baja California Sur (24°18.0/N, 110°19.0'W), 17 Jul 1996, 5 males (TL 25.2—33.5 mm) and 5 females (TL 25.2— 35.8 mm), intertidal to 1 m, rocks and algae (EMU-4240).—Calerita Beach, La Paz, Baja California Sur (24°21.0’N, 110°16.0’W), 18 Jul 1996, 1 male (TL 23.7 mm), 1 female (TL 13 mm), 1.5—2.5 m, rocks, algae and sponges (EMU-4625). San Juan de la Costa, La Paz, Baja Cali- fornia Sur (24°27.0'N, 110°42.0’W), 19 Jul 1996, 4 males (TL 11.5—27.7 mm), 3 fe- males (TL 10.8—32.9 mm), and 10 juveniles (TL 5.8—9.3 mm), rocks, algae and sponges, 1-2 m (EMU-4626).—Sendero Viejo Bay, Guaymas, Sonora (27°52.0'N, 110°52.4’W), 27 Mar 1997, 1 male (TL 13 mm), 1-3 m, rocks and algae (EMU-4627). Previous records.—Puerto Huatulco, Oa- xaca; Santa Inés Bay, Concepcion Bay, Arena Bank, Gorda bank and Puerto Es- condido, Baja California Sur; Revillagigedo 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Tres Marias Islands, Mexico. Puerto Parker and Isla del Coco, Costa Rica. Hon- da Bay, San José and Perlas Islands, Pana- ma. La Plata and Galapagos Islands, Ec- uador (Camp & Kuck 1990, Hendrickx & Salgado-Barragan 1991). Remarks.—Material from Guaymas, So- nora, corresponds to the first continental record on the east coast of the Gulf of Cal- ifornia. It also represents a slight extension of the northernmost distribution limit of UN. zacae. Acknowledgments Authors are grateful to C. Sanchez Ortiz (Departamento de Biologia, Universidad Autonoma de Baja California Sur) who col- lected the specimens of N. raymanningi. Those specimens were collected under the auspices of the Reef Fauna Program, UABCS/Brich Aquarium, Scripps Collec- tions. The collections of the gonodactylids reported here were partially financed by the Comisi6n Nacional para el Conocimiento y Uso de la Biodiversidad (Project H-017). We thank the staff of the Laboratorio de Invertebrados Bentonicos, Estacion Maza- tlan, UNAM for their help in the collections of specimens. We acknowledge R. B. Man- ning for his help in the confirmation of the identity of N. lalibertadensis. Drawings were made by G. Valenzuela. Literature Cited Brusca, R. C. 1980. Common intertidal invertebrates of the Gulf of California. Revised and expanded 2™4 edition, The University of Arizona Press, 513 p. Camp, D. K., & R. B. Manning. 1982. Five new spe- cies of Nannosquilla from the northwestern At- lantic (Crustacea:Stomatopoda).—Smithsonian Contributions to Zoology 368:1-15. , & 1986. Observations on Nannos- quilla with descriptions of three new species from the northwestern Atlantic (Crustacea:Sto- matopoda).—Smithsonian Contributions to Zo- ology 444:1-17. , & H. G. Kuck 1990. Additional records of stomatopod crustaceans from Isla del Coco and Golfo de Papagayo, east Pacific ocean.—Pro- ceedings of the Biological Society of Washing- ton 103:847-853. Dahl, E. 1954. Stomatopoda. Report of the Lund Uni- versity Chile Expedition. 1948—49, 15.—Lunds Universitets Arsskrift, new series, avdelning 2, 49(17):1-12. Hansen, H. J. 1895. Isopoden, Cumaceen und Sto- matopoden der Planktonexpedition—Ergebnis- se der Plankton-expedition der Humboldt-Stif- tung 2:1—105. Hendrickx, M. E., & J. Salgado-Barragan. 1989. Ecol- ogy and fishery of stomatopods in the Gulf of California. Pp. 241-249 in E.A. Ferrero, ed., (R.B. Manning, M.L. Reaka, & W. Wales, co- eds), Biology of stomatopods, Collana UZI: Se- lected Symposia and Monographs, Mucchi Ed- itore, Modena (Italy). » & . 1991. Los estomatdpodos (Crus- tacea:Hoplocarida) del Pacifico mexicano.— Publicaciones Especiales, Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de México 10:1—200. Hernandez-Aguilera, J. L., Y. Lopez-Salgado, & P. Sosa-Hernandez. 1986. Fauna carcinologica Insular de México. I. Crustaceos estomatopodos y decapodos de Isla Clari6n.—Investigaciones oceanograficas Secretaria de Marina, Direccion General de Oceanografia Naval. B 3 (1):183— 250. Manning, R. B. 196la. A new Lysiosquilla (Crusta- cea:Stomatopoda) from the Gulf of California, with a redescription of L. decemspinosa Rath- bun.—Proceedings of the Biological Society of Washington 74:29—36. 1961b. Stomatopod Crustacea from the At- lantic coast of northern South America——Allan Hancock Atlantic Expedition Report 9:1—46. . 1967. Nannosquilla anomala, a new stomat- opod crustacean from California.—Proceedings of the Biological Society of Washington 88: 147-150. . 1970. Nine new American stomatopod crus- taceans.—Proceedings of the Biological Society of Washington 84:225—230. . 1972a. Stomatopoda: eastern Pacific expedi- tion of the New York Zoological Society.— Zoologica 56 [1971]:95—-113. . 1972b. Three new stomatopod crustaceans of the family Lysiosquillidae from the Eastern Pa- cific Region.—Proceedings of the Biological Society of Washington 85:271—279. . 1974. Stomatopods collected by Th. Morten- sen in the Eastern Pacific Region (Crustacea: Stomatopoda).—Steenstrupia 3:101—109. 1976. Notes on some Eastern Pacific sto- matopod Crustacea, with descriptions of a new genus and two new species of Lysiosquilli- VOLUME 111, NUMBER 1 dae.—Proceedings of the Biological Society of Washington 89:221—231. . 1980. The superfamilies, families and genera of Recent stomatopod Crustacea with diagnoses of six new families.—Proceedings of the Bio- logical Society of Washington 93:362-372. . 1995. Stomatopod Crustacea from Vietnam: the legacy of Raoul Seréne. Crustacean Re- search, Special Number 4. The Carcinological Society of Japan, Tokyo, 339 pp. , & M. L. Reaka. 1979. Three new stomatopod crustaceans from the Pacific coast of Costa Rica.—Proceedings of the Biological Society of Washington 92:634—639. Rathbun, M. J. 1910. The stalk-eyed Crustacea of Peru and the adjacent coast——Proceedings of 51 the United States National Museum 38:531— 620, pls. 36—56. Reaka, M. L., & R. B. Manning. 1980. The distribu- tional ecology and zoogeographical relation- ships of shallow water stomatopod Crustacea from Pacific Costa Rica——Smithsonian Contri- butions to the Marine Sciences 7:1—29. Schmitt, W. L. 1940. The stomatopods of the west coast of America based on collections made by the Allan Hancock Expeditions, 1933—38.—AI- lan Hancock Pacific Expeditions 5:129—225. Schotte, M., & R. B. Manning. 1993. Stomatopod Crustacea from Tobago, West Indies.—Proceed- ings of the Biological Society of Washington 106:566-—581. Steinbeck, J., & E. FE Ricketts. 1941. Sea of Cortez. New York: Viking Press, 598 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(1):52—80. 1998. A revision of the southeast Asian freshwater crabs of the genus Isolapotamon Bott, 1968 (Crustacea: Decapoda: Brachyura: Potamidae) Peter K. L. Ng and S. H. Tan School of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 119260, Republic of Singapore Abstract.—The taxonomy of the potamid freshwater crab genus /solapota- mon Bott, 1968, is revised. Fifteen Bornean species (including two new ones) and three Philippine species are recognized. The fifteen Bornean species are: I. anomalum (Chace, 1938) (type species), I. bauense Ng, 1987, I. beeliae Ng, 1986, I. borneense, new species, I. collinsi Holthuis, 1979, I. consobrinum (De Man 1899), I. doriae (Nobili 1900), 7. griswoldi (Chace, 1938), I. grusophallus Ng & Yang, 1986, I. ingeri, new species, I. kinabaluense (Rathbun, 1904), J. mahakkamense (De Man, 1899), I. naiadis Ng, 1986, I. nimboni Ng, 1987, and I. stuebingi Ng, 1995. The three Philippine species are: J. mindanaoense (Rath- bun, 1904), 7. sinuatifrons (H. Milne Edwards, 1853) and J. spatha Ng & Takeda, 1992. These species are distinguished mainly by means of their gon- opodal and carapace features. The family Potamidae is represented by three genera in the island of Borneo (Bru- nei, Malaysian Sarawak and Sabah, and In- donesian Kalimantan), i.e., [solapotamon Bott, 1968, Cerberusa Holthuis, 1979, and Ibanum Ng, 1995. Cerberusa contains only two troglobitic species, known only from the caves in northern Gunung Mulu Sara- wak (Holthuis 1979). Ibanum contains two non-troglobitic species, one from central Kalimantan and the other from Sarawak (Ng 1995). Isolapotamon is the largest ge- nus, with 13 described species (Bott 1970b, Ng 1986, 1987, 1995). Three other species are known from the island of Mindanao in the southern Philippines (Bott 1970b, Ng & Takeda 1992). Bott (1970b) included /solapotamon, Malayopotamon Bott, 1968, and Nanhai- potamon Bott, 1968, in a new family, Iso- lapotamidae, which he defined as having stout or elongate male first pleopods, and whose members presumably occurred mainly in Sumatra, Java, Borneo and the Philippines, with some species present in Taiwan, southern China and Peninsular Ma- laysia. The Isolapotamidae, however, lacks sufficient distinguishing characters from typical potamids to warrant its recognition as a separate family (Ng 1986, 1987, 1988a, 1988b; Ng & Yang 1985, 1986). None of the three genera in the Isolapotamidae (Js- olapotamon, Malayopotamon and Nanhai- potamon) possess characters unique to themselves. Substantial changes have also occured with the discovery of additional Sundaic species of, for example Cerberusa and Jbanum and clarification of various genera such as Nanhaipotamon (see Ng & Dudgeon 1992, Ng & Takeda 1992, Dai & Ng 1994, Dai 1997), which casts serious doubts on the validity of the Isolapotami- dae. In this paper, the genus Jsolapotamon is revised and placed in the family Potam- idae Ortmann, 1896 (sensu Ng 1988a). All measurements, in millimeters, are in- cluded as carapace widths by lengths. The terminology used follows that used by Ng (1988a). Specimens are deposited in the Museum of Comparative Zoology, Harvard VOLUME 111, NUMBER 1 University (MCZ); National Museum of Natural History (USNM), Washington, D.C: National Natuurhistorische Museum (former Rijksmuseum van Natuurlijke His- torie), Leiden (NNM); The National His- tory Museum, London (NHM); Museum Civico di Storia Naturale, Genoa (MGE); Museo ed Istituto di Zoologia Sistematica della Universita di Torino, Turin (MUT); National Science Museum, Tokyo (NSMT); Zoological Reference Collection, School of Biological Sciences, National University of Singapore (ZRC); Museum of Zoology, Cambridge University, (CMZ); Muséum National d’Histoire naturelle, Paris (MNHN); Sarawak Museum, Kuching (SM); and the Museum Zoologicum Bo- goriense, Bogor (MZB). The abbreviations G1 and G2 are used for the male first and second pleopods re- spectively; Mt. and Sg., for Mount and Sun- gai respectively, the latter term meaning river in Malay. Altitudes above sea level are indicated in meters (m). Localities of col- lecting sites have been derived from Anon (1971) or from the field collector’s data. Taxonomy Family Potamidae Ortmann, 1896 Genus Isolapotamon Bott, 1968 Isolapotamon (Isolapotamon) Bott, 1968: 119. Tsolapotamon.—Bott, 1970a:333; Bott, 1970b:190. Diagnosis.—Carapace broader than long; epigastric and postorbital cristae well de- veloped, separate; anterolateral margin con- vex, serrated; external orbital angle usually broadly triangular, outer margin distinctly longer than inner margin; epibranchial tooth present. Exopod of third maxilliped very broad, especially proximally, outer margin distinctly convex outwards; distal tip reach- es or slightly exceeds half length of merus; flagellum distinct, longer than width of me- rus. Male abdomen distinctly triangular, lat- eral margins of last segment convex or al- most straight; telson triangular. G1 slender, 53 elongated; terminal segment subequal to or longer than subterminal segment, slender, tip often dilated. G2 slender, elongated; dis- tal segment distinct, subequal to half length of basal segment. Type species.—Potamon anomalus Chace, 1938, by original designation. Remarks.—Bott (1968) erected Jsolapo- tamon with three subgenera, 1.e. [solapota- mon, Malayopotamon Bott, 1968, and Nan- haipotamon Bott, 1968. Bott (1970a) sub- sequently raised all three to distinct genera and placed them in a new family, the Iso- lapotamidae (Bott 1970b), recognising eight species in the genus /solapotamon. Bott (1970b) also excluded the Philippine spe- cies “Telphusa artifrons Burger, 1894,” which he had earlier (1968) placed in Jso- lapotamon (Isolapotamon), and referred it to the genus Tiwaripotamon Bott, 1970. Ng (1992) reviewed the problems associated with Bott’s genus Tiwaripotamon, and sub- sequently Ng & Takeda (1992) established a new genus, Ovitamon, for Biirger’s spe- cies as well as two new species from the Philippines. Bott (1970b) placed Potamon (Potamis- cus) chaseni Roux, 1934, in the genus Js- olapotamon, noting that it was the only member of the genus from Peninsular Ma- laysia. Isolapotamon chaseni differs from typical Jsolapotamon species in at least one very major character, the form of the exo- pod of the third maxilliped. In the Bornean and Philippine /solapotamon, the exopod of the third maxilliped is very broad, especial- ly proximally, and the outer margin is dis- tinctly convex outwards. When the exopod is appressed against the endopod, the outer margin of the third maxilliped has a round- ed appearance. This type of third maxilliped exopod is known in only one other sundaic taxon, Allopotamon tambelanense (Rath- bun, 1905), from the Tambelan Islands west of Borneo (Ng 1988b). In J. chaseni, the exopod is proportionately more narrow, the outer margin being almost straight or only slightly convex, giving the third maxilliped a more quadrate appearance (see Ng 1988). 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The structure of the G1 of I. chaseni, and the absence of a flagellum on the third max- illiped exopod, suggests that this species should be excluded from the genus J/sola- potamon. Ng (1988b) referred this species to the Malayan genus Stoliczia Bott, 1966. Seven Isolapotamon species have been described from China, viz. J. sinense Tai & Sung, 1975, I. papilionaceum Dai, Song, He, Cao, Xu & Zhong, 1975, I. physalisum Dai, Song, Li, Chen, Wang & Hu, 1984, J. aflagellum Dai, Song, Li & Liang, 1980, J. nasicum Dai, Chen, Song, Fan, Lin & Zeng, 1979, I. sheni Dai, Chen, Liu, Luo, Yi, Liu, Gu & Liu, 1990, and J. obtortum Dai, Song, Li, Chen, Wang & Hu, 1984. All these Chi- nese Isolapotamon species, however, have the exopod of the third maxilliped slender (not broad) and the outer margin being straight (not convex outwards). In addition, the G1 terminal segment of the Chinese species is proportionately much longer than the subterminal segment, where as it is equal to or more than half the length of the subterminal segment in [solapotamon. In a recent reappraisal of these species using the above features as well as the structures of the external orbital angle, exopod of the third maxilliped, male abdomen and Gl, Dai & Tiirkay (1997) referred the Chinese species to four new genera, viz. Neilupo- tamon, Yarepotamon, Minpotamon, Vado- sapotamon and Latopotamon. As a result of this, [solapotamon s. str. has a wholly Bor- nean and Philippine distribution. The form of the exopod of the third max- illiped exopod in Malayopotamon and Nan- haipotamon is very different from that of Isolapotamon. Examination of Malayopo- tamon and Nanhaipotamon specimens shows that, like other Southeast Asian po- tamids, their exopods are slender and the outer margins slightly concave. This obser- vation, together with the fact that the Gls of the three genera are markedly different, indicates that the three genera are not close- ly related, contrary to Bott’s (1970b) belief. The generic diagnostic characters defined by Bott (1968) for [solapotamon are gen- erally valid and are used here in a modified form. Ng (1986, 1987) and Ng & Yang (1986) partially revised the genus, adding five new species to the Bornean fauna and resurrecting another species which Bott (1970b) had synonymised with J. mahak- kamense (De Man, 1899). A total of 15 Bornean species are now recognized, two of which are here described as new: J. an- omalum (Chace, 1938) (type species), J. bauense Ng, 1987, I. beeliae Ng, 1986, I. borneense, new species, I. collinsi Holthuis, 1979, I. consobrinum (De Man, 1899), J. doriae (Nobili, 1900), J. griswoldi (Chace, 1938), I. grusophallus Ng & Yang, 1986, I. ingeri, new species, I. kinabaluense (Rath- bun, 1904), I. mahakkamense (De Man, 1899), I. naiadis Ng, 1986, I. nimboni Ng, 1987, and J. stuebingi Ng, 1995. Although the present paper deals essen- tially with Bornean potamids, three poorly known Philippines species, 7. mindan- aoense (Rathbun, 1904), J. sinuatifrons (H. Milne Edwards, 1853) and I. spatha Ng & Takeda, 1992, are also discussed for com- pleteness. Key to the Species of Jsolapotamon la. Carapace relatively deep, carapace height to width ratio 0.4—0.5; G1 ter- minal segment about equal to subter- minal segment, terminal segment sin- uous, tip dilated, forming a nght-an- gled triangular projection (Fig. LE—H) (Satawak)icz nk (yee oats I. bauense 1b. Carapace normal or flat, carapace height to width ratio distinctly less than 0.4; G1 terminal segment equal or subequal to subterminal segment, terminal segment sinuous or straight, tip dilated, laterally flattened, rounded, or with a secondary projection 2a. Dorsal surface of carapace evenly flat; epibranchial tooth distinct, well devel- oped, separated from the anterolateral margin by distinct triangular cleft... 3 2b. Dorsal surface of carapace inflated or only inflated in 1 or 2 regions; epi- branchial tooth indistinct or distinct VOLUME 111, NUMBER 1 3a. 3b. 4a. 4b. Sa. but separated from anterolateral mar- gin by shallow cleft Carapace wider than long to almost squarish (Fig. 3), dorsal surfaces smooth; anterolateral regions striated; intestinal region smooth; epibranchial tooth well developed, tip sharp, sepa- rated from anterolateral margin by deep, distinctly triangular cleft; epi- gastric cristae sharp; G1 terminal seg- ment tip dilated to form “‘knob-like”’ structure or appearing vaguely trian- gular (Figs. SE-H) (Sarawak, north- western Kalimantan) .... J. consobrinum Carapace wider than long, dorsal sur- faces rugose, especially anterolateral and posterolateral regions; intestinal regions granulose; epibranchial tooth prominent but tip not sharp, separated from anterolateral margin by relative- ly shallower triangular cleft; epigastric cristae rugose; Gl terminal segment tip not dilated or rounded (Fig. 12E— H) (Sarawak, northwestern Kaliman- (ZN) -<. oo A 3 Caen cae ek a I. stuebingi Dorsal surface of carapace granulated, especially on anterolateral, posterolat- eral, epigastric, postorbital, branchial and intestinal regions (Fig. 9); epigas- tric cristae rugose but low; posterolat- eral margins distinctly convex; outer margin of external orbital angle about 4 times longer than inner margin; cer- vical groove shallow; no groove dis- cernible between cardiac and intesti- nal regions; Gl not known (central Kalimantan) I. mahakkamense Dorsal surface of carapace granulated or striated only on anterolateral and/or posterolateral regions; epigastric cris- tae not rugose, distinct and usually prominent; posterolateral margins straight to concave; external orbital angle about 2 or 3 times longer than internal orbital angle; cervical groove shallow or deep, groove between car- diac and intestinal region present or absent; G1 straight to or sinuous ... 5 Epigastric cristae distinct, sloping from centre and merging gradually with postorbital cristae; dactylus of last ambulatory leg very short; G1 sin- uous, terminal segment subequal in Sb. 6a. 6b. Ta. 7b. 8a. 8b. 9a. 9b. 10a. 10b. lla. 11b. 55 length to subterminal segment, distal part bifurcated, subdistal process sub- equal in length to distal process (Fig. SM-—P) (Sabah) ............ I. griswoldi Epigastric cristae indistinct, distinctly separated from postorbital cristae by wide, disjunct notch; dactylus of last ambulatory leg long or medium leng- thed; Gl sinuous to almost straight, terminal segment of varying lengths, distal part of various forms G1 slightly sinuous to almost straight, not distinctly sinuous, tip foming head-like structure or bifurcated .... 7 G1 very slender, distinctly sinuous, tip rounded, forming “‘knob-like”’ struc- (UR ie a eee Re relma ces cat Sucecm Me ie tan erate 14 Distal part of Gl terminal segment forming a “‘head-like” structure .... 8 Distal part of G1 terminal segment bi- MUNCALCOR easiest ars catty ck 8 sen Higgs 16 Tip of distal part of G1 terminal seg- ment distinctly rounded, outer margin of dilated part gradually curving to meet cylindrical part of terminal seg- ment, inner margin of distal part curv- ing gradually without distinct hump Tip of distal part of G1 terminal seg- ment not rounded, usually trapezoidal or triangular, outer margin of dilated part gradually curving to meet cylin- drical part of terminal segment, inner margin of distal part curving gradually with or without distinct hump ...... 11 Distal part of G1 terminal segment at right angles to the G1, forming an in- verted “‘boot-like”’ structure (Figs. 10M-—P) (Mindanao) ..... I. sinuatifrons Distal part of G1 terminal segment not at right angles, usually directed at 45° to the perpendicular, forming a ““knob- like”’ structure G1 tip dilated, twice as long as broad (Fig. 1OA—D) (Mindanao) ......... ET GiielsLe Mes a2 Goats on) erases I. mindanaoense G1 tip dilated, as long as broad (Fig. 12A—D) (Mindanao) ......... I. spatha G1 gently sinuous, especially terminal segment, tip resembling a “chicken head” (Figs. 1M—P) (Sarawak 7?) eG Ash. SAR OA the Pen eater! 8 I. borneense G1 almost straight, terminal segment 56 12a. 12b. 13a. 13b. 14a. 14b. 15a. 15b. 16a. 16b. 17a. 17b. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tip dilated, trapezoidal or triangular, not resembling a “‘chicken head”... 12 G1 subterminal segment with distinct notch at proximal end, terminal seg- ment tip dilated, rounded, forming a “club-like”’ structure (Fig. 8) (Sabah) ea Eee eR = Be eer I. kinabaluense G1 subterminal segment without dis- tinct notch at proximal end, terminal segment tip dilated, flattened, forming a “horse head-like”’ structure ..... 13 Distal side of G1 tip longer than prox- imal side (Fig. 1A—D) (Sabah) EAN Ga ee eee oe I. anomalum Distal side of G1 tip shorter than prox- imal side (Figs. 6E—H) (Sabah) .. J. ingeri G1 terminal segment distinctly longer than subterminal segment, tip evenly rounded (Fig. 5I-L) (Sarawak) . . I. doriae G1 terminal segment subequal to or slightly longer than subterminal seg- ment, distal part unevenly dilated, up- per margins gently convex to almost Sirarveht se hve ean ameter eee 15 Outer part of dilated G1 terminal seg- ment tip projecting slightly outwards, upper margin gently convex, curving gently downwards about 90° to form dilation (Fig. 1OE-—H) (southeastern I