Metadata taiytad ASAE Aaa, where R ahd peta tchat MA AMAA cAiyeah pie Fact at a AS Bae Ota Ay ba anes sa 08 SCEPC! “i PONe RRP ‘ is “ Ere) mets AMAL Ogi set Te Beas ayraeg ces Dita liaete Ley yer acy ycu ane eseetore tr ta~ara poe med er yee ys ‘i PROCEEDINGS of the Biological Society of Washington VOLUME 117 2004 Vol. 117(1) published 1 June 2004 Vol. 117(3) published 7 December 2004 Vol. 117(2) published 4 August 2004 Vol. 117(4) published 20 December 2004 WASHINGTON PRINTED FOR THE SOCIETY Supplement to the Proceedings of the Biological Society of Washington EDITOR RICHARD V. STERNBERG RICHARD C. BANKS ASSOCIATE EDITORS Classical Languages Invertebrates FREDERICK M. BAYER STEPHEN L. GARDINER CHRISTOPHER B. BOYKO JANET W. REID Plants Vertebrates CAROL HOTTON GARY R. GRAVES CAROLE C. BALDWIN EDWARD O. Murpy Insects Invertebrate Paleontology WAYNE N. MaTuis GALE A. BISHOP All correspondence should be addressed to the Biological Society of Washington, National Museum of Natural History Washington, D.C. 20013 Printed by ALLEN PREss INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 2004—2005 OFFICERS President ROY W. McDIARMID President-Elect W. RONALD HEYER Secretary CAROLE C. BALDWIN Treasurer T. CHAD WALTER COUNCIL Elected Members MICHAEL D. CARLETON F. CHRISTIAN THOMPSON W. DUANE HOPE JEFFREY T. WILLIAMS MARILYN SCHOTTE NEAL WOODMAN ‘a wits , pam alae mab Tae te tae pat TABLE OF CONTENTS Volume 117 Alvarenga, Herculano M. F. and Storrs L. Olson, A new genus of tiny condor from the Pleistocene of Brazil (Av essavill tii dae) eee merece ae asters eee ee ahehrus wgeeectsues oteees ade cosbece es, Goes Graves, Gary R., Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 13. An undescribed intra- generic combination Heliodoxa imperatrix xX Heliodoxa jacula ...............++-+++++++00-- Gill, Anthony C. and Hiroyuki Tanaka, Pholidochromis cerasina, a new species of pseudochromine dotty back fish from the west Pacific (Perciformes: Pseudochromidae) ....................... Kawai, Tadashi and J. F. Fitzpatrick, Jr. Redescription of Cambaroides japonicus (De Haan, 1841) (Crustacea: Decapoda: Cambaridae) withallocation of a type locality and month of collection of (NOES “ao oes Sie'o's-o-o.ple: Gay Geontn to Mua ce: OE n eC Ceo: Creer Re HIE RPS aN er Sree a ee eee Campos, Martha R. and Diego M. Valencia, Two new species of freshwater crabs of the genus Chaceus Pretzmann, 1965 from the Serrania de Perija of Colombia (Crustacea: Decapoda: Escudo thelphusiG ae) mesg tone ere eres Ta sesso ey leone eI ay eueeade send Ges eper suits. haapoaiparnaaas McLaughlin, Patsy A. and Akira Asakura, Reevaluation of the hermit crab genus Parapagurodes McLaughlin & Haig, 1973 (Decapoda: Anomura: Paguroidea: Paguridae) and a new genus for Parapagurodes doederleini (Doflein, 1902) ...........-. 0.0 c ee eee Asakura, Akira and Takeharu Kosuge, Pseudopaguristes bicolor, a new species of hermit crab (Crustacea: Decapoda: Diogenidae) from Japan, the third species of the genus ................. Felder, Darryl L. and Brian Kensley, A new species of axiid shrimp from chemosynthetic communities of the Louisiana continental slope, Gulf of Mexico (Crustacea: Decapoda: Thalassinidea) ........ Moore, Wendy, Description of a new Synidotea species (Crustacea: Isopoda: Valvifera: Idoteidae) from LAW Aller ee er uate arent oie tore ke Aes See ae mse we ee wee ees aisen eos ES Peta ese ee Schotte, Marilyn and Richard Heard, A new species of Synidotea (Crustacea: Isopoda: Valvifera) from themorcherns Gulia i NICxdC Ot enc ate sens erie eee Se hehe eR PGi tae weeeG Bee oan Sem Oe Ho, Ju-shey and Il-Hoi Kim, A new genus of the Clausidiidae (Copepoda: Poecilostomatoida) associ- ated with a polychaete from Korea, with discussion of the taxonomic status of Hersiliodes Canu, IS 5S eereae eee eee cate ates eee a area mr ata nh iene atielvoy eo atie Voeeaien se sinsadi dec ees a ah bce cl Sia Man Se ah eye Shae Dreyer, Jennifer, Tomoyuki Miura, and Cindy Lee Van Dover, Vesicomyicola trifurcatus, a new genus and species of commensal polychaete (Annelida: Polychaeta: Nautiliniellidae) found in deep-sea clamsiromithe BlakesRidserc oldiscc piesa eae een ene eee nna ae ees israel Cairns, Stephen D. and Frederick M. Bayer, Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 4: The genus Paracalyptrophora Kinoshita, 1908 .....................---..-- Wasshausen, D. C. and J. R. I. Wood, Notes on the genus Dicliptera (Acanthaceae) in Bolivia ..... AVOSvAnnualiNicetin geViiMUteS aires eerie sac eee Scan Stay Aedes Shiites eee aoe ae Asakura, Akira, Pseudopaguristes shidarai, a new species of hermit crab (Crustacea: Decapoda: Diogenidae) from Japan, the fourth species of the genus ...............---.----+--+--+--+--- L6pez-Mejia, Marilu, Fernando Alvarez, and Luis M. Mejia-Ortiz, A new species of Procambarus (Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico ..............-.---++-+++-eeeeee: Lewis, Julian J., Brackenridgia ashleyi, a new species of terrestrial isopod from Tumbling Creek Cave, Missouri (Isopoda: Oniscidea: Trichoniscidae) .............--222-- eee eee eee eee eee Markham, John C., New species and records of Bopyridae (Crustacea: Isopoda) infesting species of the genus Upogebia (Crustacea: Decapoda: Upogebiidae): the genera Orthione Markham, 1988, and Gyre ComaliasSAbanceni aS Olle. care ies hoe eee ee Oe Oe eR a Hee eee Duplessis, Kirk and Henry M. Reiswig, Three new species and a new genus of Farreidae (Porifera: Mexatinellida-gHexaGtim@si cal paeresws rns ten cs een ree ose eis AHS) Si eee, ctroncheira en eaeren She es eos er eue har ewetreus Meyer, Stephen C., The origin of biological information and the higher taxonomic categories ....... Dickerman, Robert W., A review of the North American subspecies of the Great Blue Heron (Ardea [QROGHOS) 23 4-0-6 9.0.4 8:80 BS LES OS ELS AS Ol RT TT ee Re as eee ae are ee ae eee Goodman, Steven M. and Voahangy Soarimalala, A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Forét des Mikea of southwestern Madagascar ..................-..-.- Woodman, Neal, Designation of the type species of Musaraneus Pomel, 1848 (Mammalia: SOHCOMOLPNAgS OLICIG ae) benaey wei el ruereee a eee uss Slay Sls, Sie yea yee enum asl Geena subieashereeteerenane. Esselstyn, Jacob A., Peter Widmann, and Lawrence R. Heaney, The mammals of Palawan Island, LPAONUUY DY OVUNSS aad. OB @, alo Bb eoatyog: Cumgte auctor beacG- cra Ota ee CN Ce Ee RSET TS eS ee Kraus, Fred and Allen Allison, A new species of Tropidonophis (Serpentes: Colubridae: Natricinae) from the D’Entrecasteaux Islands, Papua New Guinea ...........------ 22-2 eee eee eee 10 23 35 42 57 68 76 88 95 106 114 140 150 153 169 176 186 199 213 242 Dail 266 271 303 6 McCranie, James R. and Franklin E. Castafieda, A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras .... . Malabarba, Luiz R., Flavio C. T. Lima, and Stanley H. Weitzman, A new species of Kolpotocheirodon (Teleostei: Characidae: Cheirodontinae: Compsurini) from Bahia, northeastern Brazil, with a new diagnosisiof the Genus) oa cye te cesussicue.s celtic te easesee snsveae tele Co Seclseeeban: siaaspetee cackenc ie) tare eee ee Castro, Ricardo M. C. and Richard P. Vari, Astyanax biotae, a new species of stream fish from the Rio Paranapanema basin, upper Rio Parana system, southeastern Brazil (Ostariophysi: Characiformes: (Oi nkctes(cis Clo) ere a ce era ic culos doc od ce 6 ocd Soo oucds 66s e500 Benine, Ricardo C., Gabriela Zanon Peligao, and Richard P. Vari, Tetragonopterus lemniscatus (Characiformes: Characidae), a new species from the Corantijn River basin in Suriname ......... Worsaae, Katrine, Wolfgang Sterrer, and Thomas M. Iliffe, Longipalpa saltatrix, a new genus and species of the meiofaunal family Nerillidae (Annelida: Polychaeta) from an anchihaline cave in | 5X¢) 101 (6 - ene nae eee ere ergs eee ete erect etme Sint in oA ci ciniS h Gidis sa cg hele Glos cio 5 0 0-0 Campos, Martha R., Neostrengeria lemaitrei, a new species of freshwater crab from Colombia (Crustacea: Decapoda: Pseudothelphusidae), and the vertical distribution of the genus........... Alvarez, Fernando, José Luis Villalobos, and Thomas M.. Iliffe, A new species of Agostocaris (Caridea: Agostocarididae) from Acklins Island, Bahamas .........................----2.---2.----- Wicksten, Mary K. and Joel W. Martin, A new species of caridean shrimp of the family Stylodactylidae from the eastern Pacific'OCeam 4). in bestia lee oe 3 eine ee ope eee eee eee wt Buhl-Mortensen, L. and W. A. Newman, A new pedunculate barnacle (Cirripedia: Heteralepadidae) fromthe Northwest Atlamtic= cise cce ccd c iy ar eae ep ene ail 6 enn eee ae Kornicker, Louis S. and J. A. Rudjakov, Two new species of seven-spined Bathyconchoecia from the North Atlantic and Indian oceans (Crustacea: Ostracoda: Halocypridae)...................... Yanagi, Kensuke and Marymegan Daly, The hermaphroditic sea anemone Anthopleura atodai n. sp. (Anthozoa: Actiniaria: Actiniidae) from Japan, with a redescription of A. hermaphroditica ....... Robinson, Harold and Abigail J. Moore, New species and new combinations in Rhysolepis (Helvantheaés, Asteraceae) five) a0 sean eee tee asec ee uy artnet) Casi oan uo ee oc Re Cairns, Stephen D. and Frederick M. Bayer, Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 5. The genera Plumarella Gray, 1870; Acanthoprimnoa, n. gen.; and Candidella Bayer V994) ss. cece gists aie Riana muattti wera in, sheen See are aeons arene cee ot Si oe Se aS ate ee ee Ardelean, Adorian and Daphne Gail Fautin, A new species of the sea anemone Megalactis (Cnidaria: Anthozoa: Actiniaria: Actinodendridae) from Taiwan and designation of a neotype for the type Species OL thezSenUs’ ax avs. esau oy wee SAAS eI TE aA RON GEC oT eee cet ee Vazquez-Bader, Ana Rosa and Adolfo Gracia, A new genus and new species of crab of the family Xanthidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the southwestern Gulf of Mexico Sternberg, Richard v. and Marilyn Schotte, A new anchialine shrimp of the genus Procaris (Crustacea: Decapoda: Procarididae) from the Yucatan Peninsula ............-...--+---+-+--++---+-++:> Phone, Hla and Hiroshi Suzuki, Macrobrachium patheinense, a new species of freshwater prawn (Crustacea: Decapoda: Palaemonidae) from Myanmar .................-2---00 ee eee eee eees Gomez, Samuel, A new species of Enhydrosoma Boeck, 1872 (Copepoda: Harpacticoida: Cletodidae) fromthe Eastern Tropical! Pacific: 2: 3 4eccsie condenses cle Aen che Sasi oa SO eee Borrero-Pérez, Giomar Helena and Milena Benavides-Serrato, New record of Ophiosyzygus disacan- thus Clark, 1911 (Echinodermata: Ophiuroidea: Ophiomyxidae) in the Caribbean Sea ........... Knapp, Leslie W. and Hisashi Imamura, Sunagocia sainsburyi, a new flathead fish (Scorpaeniformes: Platycephalidae)itroninorthwestemeAus tralia eee ie nein niente enna eee Vari, Richard P. and Carl J. Ferraris, Jr., A new species of Nannocharax (Characiformes: Distichodontidae) from Cameroon, with the description of contact organs and breeding tubercles in Li SYR e{=) 1b eee names eee nee eRe etn enema ey CSIC OOD OG A.S OU oe oo soe 516 dt DoNascimiento, Carlos, Francisco Provenzano, and John G. Lundberg, Rhamdia guasarensis (Siluriformes: Heptapteridae), a new species of cave catfish from the Sierra de Perijé, northwestern WEMEZUCL A eccd st Semen dilatere ican ie ses ses wins sareecaachni sion cease ee al Se CA poses TALE oe AR Sem Olson, Storrs L., Taxonomic review of the fossil Procellariidae (Aves: Procellariiformes) described from Bermuda iby Re We ShULeldt =< gic cnc wre cos erenep aes ne ures ey cre ee crist er nee ae ee oS Espinasa, Luis and Bethany Burnham, Revision of the genus Squamigera (Insecta: Zygentoma: Nicoletiidae) with descriptions of two new species ................ 0.000 ee eee eee eee eee eee Minutesiof the: 20047AnnualiMeeting saaeeisem camo ccn ci ccic ae cctcie sc cteer om ely Stele renee ConstitutionandiBy laws: sess ee eis a eas ats tinal oe Ren in eee 311 317 330 339 346 551 564 575 582 594 597 SMITHSONIAN INSTITUTION LIBRARIES WWIII 140 0934 ISSN 0006-324X | PROCEEDINGS oF THE BIOLOGICAL SOCIETY or WASHINGTON 24 JUNE 2004 VOLUME 117 NUMBER 1 .THE BIOLOGICAL SOCIETY OF WASHINGTON 2003-2004 Officers President: Roy W. McDiarmid Secretary: Carole C. Baldwin President-elect: W. Ronald Heyer Treasurer: T. Chad Walter Elected Council Michael D. Carleton G. David Johnson Clyde Roper Michael Vecchione Marilyn Schotte Don Wilson Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: Richard v. Sternberg Associate Editors Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner Plants: Carol Hotton Christopher B. Boyko Insects: Wayne N. Mathis Janet W. Reid Vertebrates: Gary R. Graves Invertebrate Paleontology: Gale A. Bishop Ed Murdy Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Pro- ceedings of the Biological Society of Washington. Annual dues are payable on or before January 1 of each year. Renewals received after January 1 must include a penalty charge of $5.00 for reinstatement. 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OF ZOOLOGY MRC-116 NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C. 20013-7012 Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Periodicals postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). SNITHSON 4g JUL 1 6 2004 ( a condor from the Pleistocene of Brazil (Aves: Vulturidae) Herculano M. EF Alvarenga and Storrs L. Olson (HA) Museu de Historia Natural de Taubaté, Rua Colombia 99, Jardim das Nagoes, Taubaté SP, CEP 12030-520, Brazil (SLO) Division of Birds, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—A new genus and species of Vulturidae (Cathartidae auct.), Win- gegyps cartellei, is described from Pleistocene cave deposits in the states of Bahia and Minas Gerais, Brazil. This species is closely related to condors Gymnogyps and Vultur, particularly the former, as opposed to the smaller ca- thartid vultures, but is much smaller, being slightly smaller than the smallest living member of the family, the Lesser Yellow-headed Vulture Cathartes bur- rovianus. The Vulturidae appears to consist of two basic divisions (condors vs. other vultures) that differ profoundly in the morphology of the skull. Each appears to have been more diverse in the past and to contain larger or smaller species than survived to the present. Resumo.—Um novo género e nova espécie de Vulturidae (Cathartidae auct.), Wingegyps cartellei, é descrito dos depositos pleistocénicos de cavernas da Bahia e Minas Gerais, Brasil. Este é mais relacionado aos condores Gymnogyps e Vultur, principalmente com o primeiro, do que com os verdadeiros urubus, embora seja de tamanho reduzido, menor ainda que Cathartes burrovianus, 0 menor membro vivente da familia. Os Vulturidae se constituem de dois grupos basais, condores e urubus, que diferem entre si basicamente pela morfologia do cranio (o tamanho nao é fundamental), sendo que ambos parecem ter sido OCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):1—9. 2004. bastante diversificados no passado. Peter Wilhelm Lund was a Danish natu- ralist who resided in Brazil from 1832 until his death in 1880. Between 1835 and 1849 he shipped masses of Quaternary fossils from the state of Minais Gerais back to Denmark for study (Voss and Myers 1991). The great majority of these fossils were of mammals, including extinct megafauna, but also rodents and bats (Voss and Myers 1991, Czaplewski and Cartelle 1998, Car- telle 1999). The mammals were originally studied by Herluf Winge, who published his excep- tionally perceptive findings in a series of volumes entitled E Museo Lundii from 1887 to 1915. The study of fossil birds from these deposits fell to his brother Oluf Winge (1888) who produced a list of some 126 species. Only one of these, the anatid Chenalopex (now Neochen) pugil, was named as new, and many were referred to modern taxa. Others could not be assigned either for lack of comparative material or because Winge considered them probably to represent unknown species that he left unnamed. Among the last was a vulture that Winge (1888: 33) regarded as probably belonging to a new genus and species (“‘G. sp. indet. magnitudine Catharistae atrati” [= gyps atratus|). This was represented by the distal end of a humerus and an ulna lacking the distal end. He described these speci- mens in considerable detail and illustrated Cora- 7, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the humerus in comparison with a fossil of the Black Vulture Coragyps atratus. Noth- ing further was ever made of this discovery during the succeeding 115 years. In identifying fossil bird remains from a cave in the state of Bahia, we puzzled over a peculiar ovoid cranium that defied place- ment to family until we happened to notice that fossil crania of Gymnogyps from Ran- cho La Brea, California, seemed to be sim- ilar in shape, although larger. We identified two humeri from Bahia as probably belong- ing to the same species as the cranium, and a well-preserved distal fragment appeared to be identical to that illustrated by Winge as his unidentified new genus. We were able to borrow Winge’s original material and confirmed that he was quite correct that a new genus and species is indicated. This, however, turns out not to be closely related to the smaller genera of Vulturi- dae,Cathartes or Coragyps, but to the much larger condors, especially Gymnogyps. Comparative material examined.—Pre- liminary comparisons were made with al- most all families of non-passerine birds and all species of South American vultures in Museu de Histéria Natural de Taubaté. The original material of Vulturidae collected by Lund in Minas Gerais was borrowed from the Zoological Museum University of Co- penhagen (ZMUC) and restudied and com- pared. Modern skeletons examined in the Division of Birds, National Museum of Natural History, Smithsonian Institution (USNM) included: Gymnogyps californi- anus 3369, 492447; Vultur gryphus 345384, 429839; Sarcoramphus papa 345434, 559318; Coragyps atratus 613353; Cathartes aura 490864, 612254; C. melam- brotus 621939, C. burrovianus 431336, 622341. Systematics Class Aves Family Vulturidae Within the family, there is marked oste- ological distinction, particularly in the neu- rocranium, between the two living genera of condors (Vultur and Gymnogyps) on one hand, and all of the other genera (hereafter ‘‘vultures’’) on the other. The more salient of these were first noted by Miller and Howard (1938) and were further docu- mented by Fisher (1944). The extinct genus and species Breagyps clarki was also shown to belong with the condors based on cranial characters (Miller and Howard 1938). Cranial differences were detailed and extended to additional fossil specimens by Emslie (1988). In the following com- parisons, “‘condors” includes the new ge- nus. Neurocranium.—In dorsal view the neu- rocranium of condors is relatively longer and narrower, appearing almost ovoid in shape; the tranverse nuchal crest is visible because the attachments of the cervical musculature extend much farther dorsally than in the vultures, and the cerebellar prominence is much larger and more dis- tinct. In posterior view, the last two features are equally distinct. The foramen magnum is much larger and more elliptical in con- dors, as opposed to nearly circular in the vultures. The occipital condyle is more dis- tinctly stalked (better seen in ventral view) and is rounded, lacking the notch in the dorso-posterior surface seen in vultures. In condors, the dorsolateral margins of the fo- ramen magnum give rise to distinct crests that angle ventro-laterally to the extremely well developed paroccipital processes (su- praoccipital processes of Suarez and Emslie 2003; opisthotic processes of Fisher 1944; postauditory processes of Miller & Howard 1938) that parallel the similarly well devel- oped processes that angle out from the oc- cipital condyle (occipital processes of Sua- rez and Emslie 2003; medial basitemporal processes of Bock 1960; exoccipital pro- cesses of Fisher 1944). In the vultures these processes were always much smaller and differently shaped. In lateral view, the temporal fossa is much larger in condors, so that the postor- VOLUME 117, NUMBER 1 bital and zygomatic processes are farther apart, and the orbit is relatively smaller than in the vulture. Humerus.—Differs from Cathartes or Coragyps in having the distal end more ex- panded and the ectepicondylar prominence situated more distally on the shaft. Sarco- ramphus differs in having a large pneumat- ic Opening in the depression between the entepicondylar prominence and the ulnar condyle. The entepicondylar prominence is less developed than in Vultur. In almost ev- ery respect, down to the slightest detail of pneumatization, the distal end of the hu- merus of the new genus is a perfect dupli- cate in miniature of that of Gymnogyps. The complete humerus of the new genus is so worn as to preserve few useful characters, but it does have on the palmar surface a slightly pneumatized depression just distal to the head as in Gymnogyps. This depres- sion is absent in Cathartes, only slightly in- dicated in Vultur, and bears a very large pneumatic foramen in Coragyps and Sar- coramphus. Wingegyps, new genus Type species.—Wingegyps cartellei, new species. Diagnosis.—A tiny condor most similar to Gymnogyps in the narrowness and elon- gation of the neurocranium, but even nar- rower, with the braincase in dorsal view be- ing of nearly uniform width, rather than ex- panding posteriorly. Muscle scars on either side of the cerebellar prominence are cor- respondingly narrower. The foramen mag- num opens directly posteriorly rather than partly ventrally. The paroccipital processes and their associated crests are angled more ventrally than in Gymnogyps or Vultur. Dif- fers from other condors in having the entire occipital condyle and its stalk visible in lat- eral view (not visible in Breagyps, only par- tially visible in Gymnogyps and Vultur). The ulna is like that of condors and Sar- coramphus in having very little pneumati- 3 zation in the brachial depression (well de- veloped in Cathartes and Coragyps) but differs from all but Vultur in having the olecranon more distinctly set off from the margin of the internal cotyla. The olecranon is narrower, however, than in Vultur. Etymology.—Winge + Greek gyps, vul- ture, in commemoration of the perspicacity of Oluf Winge for recognizing the distinc- tiveness of this remarkable new genus. Wingegyps cartellei, new species Figs. 1-4 Holotype.—Neurocranium lacking the parasphenoid rostrum and ethmoid region, with damage to the anterior margin of the frontals and left otic area, MCL CLA782 (Fig. 1B, 2B). Type-locality.—Brazil, Bahia State, Municipio de Morro do Chapeu, Gruta dos Brejoes (11°00’30"S, 41°26'07’W), eleva- tion ca. 600 m. Horizon and age.—Probably late Pleis- tocene or early Holocene. A radiocarbon date of 12,200 + 120 radiocarbon years be- fore present was obtained from a coprolite of a ground sloth from the type-locality (Czaplewski and Cartelle 1998). Associated mammals from caves in Bahia and Minas Gerais are considered to be of Pleistocene age (Cartelle 1999). Measurements (mm) of holotype.—Total length as preserved 48.6; width at level of postorbital processes 27.5; width at level of base of zygomatic processes 29.4; greatest depth at midline 27.2; width and depth of foramen magnum; 8.9 X 8.1; width of oc- cipital condyle 4.1 Paratypes.—Topotypes: Complete but very worn left humerus MCL CLA670 (Fig. 4C); distal third of left humerus MCL CLA1678 (Fig. 3B). Lapa do Tit, Minais Gerais, Brazil: distal half of right humerus ZMUC 1116 (Fig. 3A); right ulna lacking distal end ZMUC 1118 (Fig. 4A). Measurements (mm) of paratypes.—Hu- meri (in the same sequence as above): total PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ioe Ie B, Wingegyps cartellei, new species, holotype MCL CLA782; C, Cathartes aura USNM 612254. Seale = 2 cm. length 129.5, —, —; length from head to distal end of pectoral crest 54.4, —, —; shaft width and depth at midpoint 10.0 x 8.5, —, 9.7 X 7.6; distal width —, 24.8, 23.2; greatest dimension of brachial depres- sion 13.4; 11.7, 13.0; greatest dimension of radial condyle —, 10.8 11.1. Ulna: proxi- mal width 12.5; proximal depth 15.8; length of brachial depression 23.4. Etymology. Dedicated to paleotologist Castor Cartelle of the Universidade Federal de Minas Gerais in recognition of his ex- Neurocrania in dorsal (top) and posterior (bottom) views: A, Gymnogyps californianus USNM 3369; cavations at Gruta dos Brejoes (Cartelle 1983) and his contributions to the paleon- tology of Brazil. Diagnosis—Much smaller than any known condor; slightly smaller than the smallest living cathartid vulture (Lesser Yellow-headed Vulture Cathartes burrovi- anus). Discussion.—Wingegyps is indisputably a condor based on the very distinct features of the neurocranium and on similarities of the distal end of the humerus. Its extraor- VOLUME 117, NUMBER 1 5 Fig. 2. Neurocrania in lateral view: A, Gymnogyps californianus USNM 3369; B, Wingegyps cartellei, new species, holotype MCL CLA782; C, Cathartes aura USNM 612254. Scale = 2 cm. 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. dinarily small size is quite unanticipated, being somewhat smaller than the smallest living species of the family (Carthartes burrovianus). The humerus is only slightly shorter than in females of the Black Vulture Coragyps atratus from the tropics, which are smaller than individuals at the temperate ends of the species’ range (Brodkorb 1944). But the humerus is proportionately much shorter in Coragyps atratus than in Cathar- tes, so that this species is otherwise much larger than Cathartes burrovianus (1875 g in a female Coragyps from Panama vs 960 g ina male C. burrovianus form Guyana). Wingegyps shows that condors were much more diverse in size in the past. The family Vulturidae may be viewed as being divisible into two basic groups: the condors (Vultur, Gymnogyps, Breagyps, Winge- Humeri in anconal view: A, Wingegyps cartellei, new species, paratype ZMUC 1116; B, Wingegyps cartellei, new species, paratype MCL CLA1678; C, Gymnogyps californianus USNM 492447, reduced to the same size as B; D, same natural size. Scale = 2 cm except for C. gyps), which appear to be derived (Emslie 1988) and the remaining living genera (Ca- thartes, Coragyps, Sarcoramphus), which may be paraphyletic. Both may have been more diverse at one time and perhaps some of the larger fossil taxa (Geronogyps, Plio- gyps, “Sarcoramphus”’ kernense—see Em- slie 1988), for which cranial material is un- known, may prove to be more closely re- lated to the assemblage of smaller vultures than to condors. Known only from a rather limited area in eastern Brazil, Wingegyps doubtless had a greater range than indicated at present, pos- sibly much greater. If it has been collected in fossil deposits elsewhere the material might easily be overlooked as belonging to Cathartes or Coragyps because of its small size. VOLUME 117, NUMBER 1 Fig. 4. paratype ZMUC 1118; B, Cathartes burrovianus USNM 431336; C, Wingegyps cartellei, new species, paratype MCL CLA670; D, Cathartes burrovianus USNM 431336. Scale = 2 cm. What sort of feeding niche might such a tiny condor have occupied? The habits of living species of the family are briefly sum- marized from Olson et al. (1967), Sick (1993), and Hertel (1994). The living con- dors Vultur and Gymnogyps forage by sight and prefer soft viscera from large carcasses. Sarcoramphus and Coragyps forage by sight and are very aggressive at carcasses. Coragyps takes food in small bits, tearing even small carcasses such as a frog or mouse to pieces before eating. The species of Cathartes are very different in finding food with their keen sense of smell. Thus, they specialize in finding caracasses of Right ulnae (A, B) and left humeri (C, D) in palmar view: A, Wingegyps cartellei, new species, small animals either before they are located by sight foragers or detecting food that can- not be seen at all from above. They are also very docile and not at all competitive with other vultures at carcasses. The small size of Wingegyps would have placed severe limitations on its ability to process the majority of carcasses or to com- pete at carcasses with other species of vul- tures. If we assume that it was like its clos- est relatives in lacking the olfactory capa- bilities of Cathartes, Wingegyps would have had little success competing with any of the species of Cathartes for small car- casses. There does seem to be a potential 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON niche in the New World, however, that is not as fully exploited as it is in the Old World, viz. palm fruits. In Africa, the Palm-nut Vulture (Gypho- hierax, Accipitridae) feeds mainly on the soft mesocarp of the African oil palm Eleis guineensis. This palm has been introduced to Brazil and Sick (1993:149) describes Turkey Vultures Cathartes aura as being a “nuisance” in palm plantations in Amazon- ia, where they consume the fruits. He also records them as feeding on the native palm Acrocomia sclerocarpa (= A. aculeata), a very widespread species occurring through the West Indies and from Mexico south to southern Brazil and Paraguay (Henderson et al. 1995), and overlapping the small known range of Wingegyps. Although Wingegyps may possibly have been the New World ecological equivalent of the unrelated Old World Palm-nut Vulture, its habits might also have been like that of the Egyptian Vulture Neophron percnopterus in subsist- ing on scraps thrown off of carcasses by larger vultures. Such habits might better ex- plain the extinction of Wingegyps, as many of the larger avian scavengers in the New World also went extinct at the time of dis- appearance of much of the mammalian megafauna (Steadman and Martin 1984). Acknowledgments Travel by SLO to Brazil was made pos- sible by the Alexander Wetmore Endow- ment Fund, National Museum of Natural History, Smithsonian Institution. We are grateful to Castor Cartelle of the Museu de Ciéncias Naturais (MCL) of the Pontificia Universidade Catdlica de Minas Gerais, Belo Horizonte, Brazil, for making the fos- sil material from Gruta dos Brejoes avail- able for study, to Jon Fjeldsa and Kim Aar- is-Sgrensen, of the Zoological Museum University of Copenhagen (ZMUC), Den- mark, for lending the material studied by Oluf Winge. Frederick V. Grady, Smithson- ian Department of Paleobiology, cleaned and repaired fossil specimens. Kevin Sey- mour, Royal Ontario Museum, suggested a reference. Photographs are by John Steiner, Smithsonian Center for Scientific Imaging and Photography, and the figures were ar- ranged by Brian Schmidt, Division of Birds, Smithsonian Institution. Literature Cited Bock, W. J. 1960. Secondary articulation of the avian mandible.—Auk 77:19-55. Brodkorb, P. 1944. Geographical variation in the Black Vulture.—Papers of the Michigan Academy of Science, Arts, and Letters 29:115-121. Cartelle, C. 1983. Tesouro féssil no sertao baiano.— Ciéncia Hoje 1(5):35-43. . 1999. Pleistocene mammals of the cerrado and caatinga of Brazil. Pp. 27-46 in J. EK Ei- senberg and K. H. Redford, eds., Mammals of the Neotropics, vol. 3. The Central Neotropics. Ecuador, Peru, Bolivia, Brazil. Chicago, Uni- versity of Chicago Press, 609 pp. Czaplewski, N. J., & C. Cartelle. 1998. Pleistocene bats from cave deposits in Bahia, Brazil—Jour- nal of Mammalogy 79:784-803. Emslie, S. D. 1988. The fossil history and phyloge- netic relationships of condors (Ciconiiformes: Vulturidae) in the New World.—Journal of Ver- tebrate Paleontology 8:212-228. Fisher, H. I. 1944. The skulls of the cathartid vul- tures.—Condor 46:272-296. Henderson, A., G. Galeano, & R. Bernal. 1995. Palms of the Americas. Princeton, New Jersey, Prince- ton University Press, 352 pp. Hertel, E 1944. Diversity in body size and feeding morphology within past and present vulture as- semblages.—Ecology 75:1074-1084. Miller, L. H., & H. Howard. 1938. The status of the extinct condor-like birds of the Rancho La Brea Pleistocene.—Publications of the University of California at Los Angeles in Biological Scienc- es 1:169-176. Olson, S. L., H. Loftin, & J. Wiese. 1967. Observa- tions on the behavior of Black and Turkey Vul- tures at traps and in captivity.—Bird-Banding 38:75-76. Sick, H. 1993. Birds in Brazil. Princeton, New Jersey, Princeton University Press, 703 pp. Steadman, D. W., & P. S. Martin. Extinction of birds in the Late Pleistocene of North America. Pp. 768-780 in P. S. Martin and R. G. Klein, eds., Quaternary Extinctions. A Prehistoric Revolu- tion. Tucson, University of Arizona Press, 892 Pp. Suarez, W., & S. D. Emslie. 2003. New fossil material with a redescription of the extinct condor Gym- nogyps varonai (Arredondo, 1971) from the VOLUME 117, NUMBER 1 Quaternary of Cuba (Aves: Vulturidae).—Pro- ceedings of the Biological Society of Washing- ton 116:29-37. Winge, O. 1888. Fugle fra Knuglehuler i Brasilien. E Museo Lundii 1 (2): pp. 54 + 5 + 1 plate. Voss, R. S., & P. Myers. 1991. Pseudoryzomys simplex (Rodentia: Muridae) and the significance of Lund’s collections from the caves of Lagoa San- ta, Brazil—Bulletin of the American Museum of Natural History 206:414-432. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):10-16. 2004. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 13. An undescribed intrageneric combination, Heliodoxa imperatrix x Heliodoxa jacula Gary R. Graves Department of Zoology, MRC-116, National Museum of Natural History, Smithsonian Institution, PO. Box 37012, Washington, D.C. 20013-7012, U.S.A. Abstract.—An enigmatic specimen collected by Perry O. Simons, presum- ably on the Pacific slope of the Ecuadorian Andes, is demonstrated to be a hybrid between Heliodoxa imperatrix and Heliodoxa jacula jamesoni. This represents the only known instance of intrageneric hybridization in Heliodoxa. External measurements of the hybrid are consistent with the proposed parental hypothesis. At the monthly meeting of the British Or- nithologists’ Club on the 17 January i900, Ernst Hartert (1900:39) exhibited a speci- men of hummingbird, “obtained in Ecuador by Mr. Simons, combin[ing] in a striking way the shape and colours of Eugenia [He- liodoxa] imperatrix and Heliodoxa jacula jamesoni . . . to be described in detail in the ‘Novitates Zoologicae’.”’ Although Hartert never published a description or reported a museum registration number, this brief ex- hibition notice has been cited in catalogs of avian hybrids (Gray 1956, Panov 1989). Here I provide a taxonomic assessment of the specimen employing the methods and assumptions outlined in Graves (1990) and Graves & Zusi (1990), as modified by the findings of Graves (1998, 1999). Methods The specimen, now deposited in the Nat- ural History Museum (registration number, 1902.3.13.2211), bears two labels, one from the British Museum marked “P. O. Si- mons,’ and an older one from the Roths- child Museum. Perry O. Simons collected mammals and birds for Oldfield Thomas (British Museum) from 1898 until his mur- der near Cuervas, Argentina, in 1901 (Allen 1903, Chubb 1919). Both specimen labels are marked with Hartert’s taxonomic deter- mination. Curiously, neither the specimen labels nor the Natural History Museum cat- alog indicate when or where the specimen was collected. I compared the specimen (Figs. 1, 2) with all species in the subfamily Trochilinae, the typical hummingbirds (Zusi & Bentz 1982, Sibley & Monroe 1990, Bleweiss et al. 1997), deposited in the Natural History Mu- seum, Tring, and the National Museum of Natural History, Smithsonian Institution. The specimen appears to be a male in de- finitive plumage as judged by the absence of striations on the maxillary ramphotheca and the presence of a well-defined, strongly iridescent gorget and coronal stripe. De- scriptions in this paper refer to definitive male plumage. Simons’ specimen is clearly assignable to the genus Heliodoxa in pos- sessing a unique combination of characters: (a) robust, moderately long (22.7 mm), nearly straight bill (Fig. 1); (b) feathers ex- tend forward on the bill obscuring the nos- trils; (c) unmodified regimes; (d) tarsal feathers extend to the base of toes; (e) mod- erately forked tail (fork depth = 35.3 mm; Fig. 2), (f) unspotted rectrices; (g) small brilliant gorget; and (h) brilliant coronal stripe. VOLUME 117, NUMBER 1 11 Fig. 1. According to Chubb (1919), Stephens & Traylor (1983), and Paynter (1993), Si- mons’ collecting itinerary overlapped the known range of the genus Heliodoxa on the Pacific slope of the Ecuadorian Andes (prov. Azuay, Cafar, Chimborazo, Guayas, and El Oro), and on the Amazonian slope of the Andes in Peru (depto. Junin and Puno) and Bolivia (depto. La Paz). For the purposes of the hybrid diagnosis, I restrict- ed the pool of potential parental species (Graves 1990, Graves & Zusi 1990) to He- liodoxa aurescens, H. rubinoides, H. lead- beateri, H. schreibersii, H. branickii, H. im- peratrix, and H. jacula jamesoni (taxonomy A probable hybrid, Heliodoxa imperatrix X H. jacula jamesoni (BMNH 1902.3.13.2211). of Schuchmann 1999). I measured selected specimens with digital calipers (rounded to the nearest 0.1 mm): wing chord; bill length (from anterior extension of feathers); and rectrix length (from point to insertion of the central rectrices to the tip of each rectrix) (Table 1). Pairs of rectrices are numbered from the innermost (R1) to the outermost (R5). I evaluated the color of the breast at the ventral midline and of the medial vane of the dorsal surface of R4 (7 mm from tip) with a calibrated colorimeter (CR-221 Chroma Meter, Minolta Corporation) equipped with a 3.0 mm aperture. The mea- Table 1.—Ranges (mean + standard deviation) of measurements (mm) of adult males of Heliodoxa imperatrix, H. jacula jamesoni, and a probable hybrid, Heliodoxa imperatrix X H. jacula jamersoni (BMNH 1902.3.13.2211). Heliodoxa imperatrix (n = 10) Wing chord 70.4—75.8 (WBA = 1.7) Bill length 23.3—24.9 (23.4 + 0.8) Rectrix 1 22.6-26.6 (4S) = 12) Rectrix 2 27.9-32.9 (29.7 = 1.4) Rectrix 3 38.7-46.5 (41.3 + 2.3) Rectrix 4 51.0-60.3 (54.8 += 2.8) Rectrix 5 62.2-76.2 (68.4 + 5.7) Heliodoxa jacula jamesoni BMNH (n = 12) 1902.3.13.2211 73.3-79.0 75.5 (75.9 + 1.8) 21.8-24.4 Del @32 = 0.7) 32.7—35.5 DISeu (3-7 25 10) 36.7-40.0 34.3 (38.5 + 1.0) 42.3-46.5 43.7 (44.4 + 1.1) 47.7-51.3 55.6 (CY7/ = 1123) 48.5-54.3 64.0 @ilo7/ 2 21) 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON suring head of the CR-221 uses 45° circum- ferential illumination. Light from the pulsed xenon arc lamp is projected onto the spec- imen surface by optical fibers arranged in a circle around the measurement axis to pro- vide diffuse, even lighting over the mea- suring area. Only light reflected perpendic- ular to the specimen surface is collected for color analysis. Colorimetric data from iri- descent feathers are acutely dependent on the angle of measurement, the curvature of plumage surfaces in museum skins, and the degree of pressure applied to the plumage surface by the Chroma Meter aperture. In order to reduce measurement variation, I held the aperture flush with the surface of the breast plumage or rectrix without de- pressing it. The default setting for the CR- 221 Chroma Meter displays mean values derived from three sequential, in situ mea- surements. I repeated this procedure twice, removing the aperture between trials. Thus, each datum summarized in Table 2 repre- sents the mean of six independent colori- metric measurements. Colorimetric characters were described in terms of opponent-color coordinates (L, a, b) (Hunter & Harold 1987). This system is based on the hypothesis that signals from the cone receptors in the human eye are coded by the brain as light-dark (L), red- green (a), and yellow-blue (b). The ratio- nale is that a color cannot be perceived as red and green or yellow and blue at the same time. Therefore “redness” and ““sreenness” can be expressed as a single value a, which is coded as positive if the color is red and negative if the color is green. Likewise, “‘yellowness” or “blue- ness”’ is expressed by b for yellows and -b for blues. The third coordinate, L, ranging from 0 to 100, describes the “lightness” of color; low values are dark, high values are light. The more light reflected from the plumage, the higher the L value will be. Vi- sual systems in hummingbirds (e.g., Gold- smith & Goldsmith 1979) differ signifi- cantly from those of humans and the rele- vance of opponent color coordinates to col- ors perceived by hummingbirds is unknown. Results and Discussion I considered hypotheses that the speci- men represents (7) an undescribed geo- graphic variant or genetic color morph of one of the aforementioned species of Helio- doxa; (ii) a hybrid; or (iii) an undescribed species of Heliodoxa. Simons’ specimen does not appear to represent an unknown color morph or geographic variant of any described species because of its unique tail morphology (Table 1). As noted by Hartert (1900), the specimen combines characters of Heliodoxa imperatrix and Heliodoxa ja- cula (Figs. 1-3; Tables 1, 2). The hybrid diagnosis focuses on the identification of apomorphic character states of possible parental species in puta- tive hybrids (Graves 1990). Complete dom- inance and polygenic inheritance of plum- age characters, however, may preclude or obscure the expression of parental apomor- phies in hybrids. When parental apomor- phies are not identifiable, the parentage of a hybrid may be indicated, although less conclusively, by the presence or absence of a suite of plesiomorphic characters. The pool of potential parental species may first be narrowed by focusing on the absence of rufous or buff pigmentation in the hybrid’s plumage. Because brown and reddish-brown pigments appear to exhibit consistent penetrance in hummingbird hy- brids (Banks & Johnson 1961, Graves & Newfield 1996), Heliodoxa rubinoides (1u- fous on inner vanes of secondaries and pri- maries; cinnamon-buff margins of breast and abdomenal feathers), H. aurescens (ru- fous pectoral band), and H. branickii (ru- fous inner vanes of rectrices) can be elim- inated from further consideration as paren- tal species. In a similar fashion, H. schrei- bersi (black throat, breast, and abdomen) and H. leadbeateri (brilliant violet coronal stripe; coppery-bronze hindcrown and neck) are exceedingly unlikely to be paren- VOLUME 117, NUMBER 1 Fig. 2. tal species because they possess characters not observed in the hybrid. Based on plum- age characters, the hybrid is most likely the product of the species, H. imperatrix X H. jJacula jamesoni. Below, I present a synop- sis of the essential evidence. The visual display of iridescence in He- liodoxa imperatrix and H. jacula has evolved to be viewed head-on. Both paren- tal species possess brilliant gorgets and cor- onal stripes that exhibit metallic irides- cence. In H. imperatrix, the green coronal stripe is bluntly triangular in shape, extend- ing from the base of the bill and narrowing to a point along the midline of the crown (even with the anterior edge of the eye). The bluish-green coronal stripe in H. jacula extends from the bill to the hindcrown forming a coronal stripe. The coronal stripe of the hybrid is intermediate in appearance between those of H. imperatrix and H. ja- cula. Heliodoxa imperatrix possesses a small purplish-pink gorget that appears to be surrounded by a field of dimly glowing, greenish-black plumage when viewed head- on. The blue gorget of H. jacula is sur- rounded by a field of green plumage, which is spangled with glowing iridescence when viewed head-on. In the hybrid, the color Dorsal surface of the rectrices of a probable hybrid, Heliodoxa imperatrix * H. jacula jamesoni (BMNH 1902.3.13.2211). and quality of iridescence exhibited by the gorget (purple exhibiting pinkish tones at certain angles) and the surrounding plum- age are intermediate in appearance between those of H. imperatrix and H. jacula. The ventral plumage of Heliodoxa im- peratrix exhibits brilliant golden-green iri- descence on the lower breast, flanks, and abdomen when viewed head-on. The breast and abdominal plumage is significantly darker in H. jacula and exhibits far less ir- idescence than in H. imperatrix. The color and quality of iridescence in the hybrid is intermediate between those of the postulat- ed parental species (Table 2). The rectrices of H. imperatrix are dark bronzy-olive be- coming progressively darker from R1 to RS5, whereas those of H. jacula are bluish- black (the lateral webs of R1 are tinted with olive in some individuals). Rectrix color in the hybrid is roughly intermediate between that of the postulated parental species (Ta- ble 2): As a second step, the parental hypothesis was tested with an analysis of size and ex- ternal proportions (Table 1, Fig. 3). Mea- surements of avian hybrids fall within the mensural ranges exhibited by their parental species as a consequence of a polygenic PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 14 (=) (j=) ce) Ww G xljOey YyjHue7 (oe) © py Xioey Yyjbue7 Xo) Ko) (e) Ko) 91 (OLS) 8 0- U1 97 (9'0¥) Cl ET Cll (L1=) eel VSI Col (Ves) sat 6 0c 6 01— (6'€=) 6 e1— LO VVC (S'1+) € 6C CEE LI@c el € C06 (as +) urs “xP HNWa& (ZI = 4) 1uosaulvl pnovl “Fy € xloey yjbue7 Som 0) € Ol Ss Ol T0c— c9C “Ul (OT) (C1) (TH) (LT#) (O'E=) (Cex) (as +) ice) (9) 6L 60 LSI Sel SCI— VIC urs N oD N Z Xlqjoey Yyybue7] (or = 4) ell EC © 6l 881 9 3— 8 8C “XR xipiadu pxopoyay Length Rectrix 1 Bivariate plots of measurements (see Table Fig. 3. 1) of males in definitive plumage: Heliodoxa impera- trix (@), H. jacula jamesoni (A), and a hybrid (x), Heliodoxa imperatrix X H. jacula jamesoni (BMNH 1902.3.13.2211). SV Cm ell 601 931— /LII “UN mode of inheritance (see Buckley 1982). Measurements of H. imperatrix and H. ja- NY sgn ss cula overlap for four of seven characters. The percent difference in character means (larger species divided by smaller) varies va Jsvoig (LIZZ ELE ZO6L HNING) osawol vjnovl “YX X14josadun pxopoyay ‘piiqdky e pue ‘uosaiupl pjnovl FY ‘x14josadu DxOpoyay JO aseunyd dAIUYOp Ul soyeUr IO} (py) p XMMIO0I pue jsvoIq Jo (q ‘D “7) Sa}JeUIPIOOS IO[O9 jUsUOddoO Jo (UOTJeIAOpP pIepuL}s |) suUvOW pu ‘eUTTUTW ‘eUTxeAJ—'Z 2GR VOLUME 117, NUMBER 1 from negligible to moderate: wing chord (3.7%), bill length (0.9%), R1 (37.6%), R2 (29.6%), R3 (7.5%), R4 (10.3%), and R5 (32.3%). Measurements of the hybrid fall within the cumulative range of parental measurements for all seven characters and within the parental means for five charac- ters (wing chord, R1, R2, R3, R5). In sum- mary, evidence obtained from plumage col- or and pattern, as well as from external size and shape, is consistent with the hypothesis that Simons’ specimen is an intrageneric hybrid between Heliodoxa imperatrix and H.. jacula jamesoni. Simons collected avian specimens on the Pacific slope of the Ecuadorian Andes in the provinces of Azuay, Chimborazo, Guayas, El Oro, and Pichincha from 1 No- vember 1898 to 12 July 1899 (Chubb 1919, Paynter 1993). His northernmost collecting locality, Guaillabamba, Pichincha (0°04’S, 78°21'W), lies in a semi-arid intermontane valley some 30 km southeast of the zone of sympatry for Heliodoxa imperatrix and H. Jacula jamesoni in humid cloud forest on the Pacific slope (see Ridgely & Greenfield 2001). This suggests one of three possibil- ities: (1) Simons collected the specimen along the Quito-Guaillabamba-Gualea road, but on the Pacific slope; (2) he purchased the specimen from a third party, possibly a native collector; or (3) he obtained the spec- imen at an unknown area of sympatry be- tween the parental species on the Pacific slope in west-central or southwestern Ec- uador. Whatever the source, Simons’ spec- imen represents the only known instance of intrageneric hybridization in Heliodoxa. Acknowledgments I am grateful to Robert Prys-Jones, Mi- chael Walters, Mark Adams, Don Smith, and the Schliisselmeister, Frank Steinhei- mer, of The Natural History Museum, Tring, for permission to study Simons’ specimen and for loaning it for long-term study. I thank Richard C. Banks and Rich- ard L. Zusi for comments on the manu- 15 script. Travel was supported by the Re- search Opportunities Fund, the Alexander Wetmore Fund, and the Department of Ver- tebrate Zoology, Smithsonian Institution. Literature Cited [Allen, J. A.] 1903. [Perry O. Simons, widely known as an energetic and careful collector. . .]|—Auk 20:94-96. Banks, R. C., & N. K. Johnson, 1961. A review of North American hybrid hummingbirds.—Con- dor 63:3-28. Bleweiss, R., J. A. W. Kirsch, & J. C. Matheus. 1997. DNA hybridization evidence for the principal lineages of hummingbirds (Aves: Trochili- dae).—Molecular Biology and Evolution 14: 325-343. 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. Chubb, C. 1919. Notes on collections in the British Museum, from Ecuador, Peru, Bolivia, and Ar- gentina, part 1. Tinamidae-Rallidae.—Ibis (11th series):1:1-55. Goldsmith, T. H., & K. M. Goldsmith. 1979. 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Proceedings of the Biolog- ical Society of Washington 109:755-763. , & R. L. Zusi. 1990. An intergeneric hybrid hummingbird (Heliodoxa leadbeateri X Helian- gelus amethysticollis) from northern Colom- bia——Condor 92:754-760. Gray, A. P. 1958. Bird hyrids. Commonwealth Agri- cultural Bureaux, Bucks, England, 390 pp. Hartert, E. 1900. [Mr. Ernst Hartert exhibited two hy- brids of hummingbirds].—Bulletin of the Brit- ish Ornithologists’ Club 10:39-40 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hunter, R. S., & R. W. Harold. 1987. The measurement of appearance, 2nd edition. Wiley, New York, All pp. Panov, E. N. 1989. Natural hybridisation and etholog- ical isolation in birds. Nauka, Moscow, 510 pp. Paynter, R. A., Jr. 1993. Ornithological gazetteer of Ecuador, 2nd edition. Museum of Comparative Zoology, Harvard University, Cambridge, Mas- sachusetts, 247 pp. Ridgeley, R. S., & P. J. Greenfield. 2001. The birds of Ecuador. Volume 1: status, distribution, and tax- onomy. Cornell University Press, Ithaca, New York, 848 pp. Schuchmann, K. L. 1999. Family Trochilidae. Pp. 468-680 in Handbook of the Birds of the World, vol. 5. Barn-owls to Hummingbirds (J. del Hoyo, A. Elliott, & J. Sargatal, Eds.). Lynx Edicions, Barcelona, 759 pp. Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Uni- versity Press, New Haven, Connecticut, 1111 PPp- Stephens, L., & M. A. Traylor, Jr. 1983. Ornithological gazetteer of Peru. Museum of Comparative Zo- ology, Harvard University, Cambridge, Massa- chusetts, 271 pp. Zusi, R. L., & G. D. Bentz. 1982. Variation of a muscle in hummingbirds and swifts and its systematic implications—Proceedings of the Biological Society of Washington 95:412—420. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):17—22. 2004. Pholidochromis cerasina, a new species of pseudochromine dottyback fish from the west Pacific (Perciformes: Pseudochromidae) Anthony C. Gill and Hiroyuki Tanaka (ACG) Fish Research Group, Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.:; (HT) Jinguh Clinic, 2-2-79 Jinguh, Miyazaki, Miyazaki 880, Japan. Abstract.—Pholidochromis cerasina is described from the 43.9-mm SL ho- lotype from Talisei Island, off the northern tip of Sulawesi, Indonesia. It is distinguished from its congener P. marginata (Lubbock) from Papua New Guinea and the northern Solomon Islands in lacking both dark submarginal markings on the median fins and prominent dark grey to black spots surround- ing sensory pores on the head. Fishes of the Indo-Pacific subfamily Pseudochrominae were recently revised by Gill (2003), who recognised 80 species in 10 genera, four of which were newly de- scribed. One of the newly described genera, Pholidochromis, was erected to accommo- date a single species, Pseudochromis mar- ginatus Lubbock, 1980, and distinguished from other pseudochromine genera in hav- ing the following combination of external characters: lower lip complete (uninterrupt- ed at symphysis); dorsal-fin rays III, 22; anal-fin rays III, 13; scales in lateral series 28—32; dorsal and anal fins with well-de- veloped scale sheaths; and predorsal scales extending anteriorly to or forward of pos- terior nostrils. It is also unique among pseu- dochromid genera in having the following combination of osteological characters: three equal-sized supraneural bones; first dorsal pterygiophore posterior lamina run- ning most of the length of the bone; and 11—12 consecutive dorsal pterygiophores inserting in a 1:1 relationship with inter- neural spaces directly behind neural spine 4. Gill (2003) recorded Pholidochromis marginata from the east coast of Papua New Guinea, Bougainville Island, and off the northern tip of Sulawesi, Indonesia. The latter record was based on a single speci- men (USNM 136954) collected at Talisei Island in 1909 by the United States Bureau of Fisheries Steamer Albatross; it bears a silk tag with the number “‘2038.”’ The spec- imen differs from other examined speci- mens (all of which were collected 50 or more years after the Sulawesi specimen) in lacking conspicuous dark spots on the head and dark submarginal stripes on the median fins. Although no comments on the condi- tion of these markings were made in his revision, the first author attributed their ab- sence to the age of the specimen, with the assumption that it was badly faded. In 1995, the first author received a colour illustration of an aquarium individual of a pseudochromid from W. E. Burgess (for- merly of Tropical Fish Hobbyist Publica- tions Inc.). It was a pale pink, deep-bodied fish with orange to red spots on the body and median fins, and a yellow ring around the eye. The first author was unable to iden- tify 1t confidently with any known species, but suggested that it was perhaps an unusu- ally coloured individual of either Pseudoch- romis fuscus Miller & Troschel, 1849 (which is often yellow with blue to grey spots and a similar body shape) or a poor illustration of P. marshallensis Schultz, 1953 (which, though usually more slender 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with a darker ground coloration, has yellow to orange or red spots on the body). In May 2000, the second author sent the first author a photograph of a pseudochrom- id from a recent article in the Japanese aquarium journal Aqualife, as well as ad- ditional aquarium photographs of the spec- imen. The fish depicted was very similar in coloration and shape to the one in Burgess’s illustration, thus rekindling interest in its identity. A search of the first author’s col- lection of pseudochromid photographs re- vealed an illustration of a similar specimen collected on the Albatross expedition (orig- inal housed in the National Museum of Nat- ural History, Smithsonian Institution). The number “2038” was written in pencil next to the illustrated fish. As Fowler (1931) had reported on pseu- dochromids collected by the Albatross, his paper was searched in attempt to locate a reference to the number “2038.” No such reference was found, but a colour descrip- tion closely matching the illustration was found for a specimen numbered “22731” from Talisse Island, which Fowler had iden- tified as Pseudochromis xanthochir Bleeker, 1855 (a junior subjective synonym of P. fuscus). The number “22731” refers to a linen tag attached to a 45.0-mm-SL speci- men of the pseudoplesiopine Pseudople- siops typus Bleeker, 1858 (now registered USNM 146624). However, the Albatross il- lustration (and Fowler’s description) is ob- viously not based on the specimen of P. typus. Although P. typus may be pale pink to pale grey with a ring around the eye (which is red to black in life), it does not possess red spots on the body. Moreover, the illustration depicts a fish with relatively short, broad pelvic fins, whereas they are long and slender in specimens P. typus (n- cluding the specimen in USNM 146624). The other illustration and photographs of aquarium specimens are also not referable to P. typus. However, the P. typus specimen was col- lected from Talisse (=Talisei) Island on the same date as the Albatross Pholidochromis specimen (presumably from the same sta- tion), and this, coupled with the close sim- ilarity in body form and pelvic-fin shape, led us to question whether the illustration was of the Pholidochromis specimen, and whether the “2038” may refer to the silk tag number on that specimen. We therefore asked S. L. Jewett and J. T. Williams of the National Museum of Natural History to check whether there were further details that might corroborate this. Jewett consult- ed the original illustration and responded (pers. comm., 4 Aug 2000): ‘It not only says 2038 in pencil, but Leonard Schultz {former Curator of Fishes at USNM, and author of a paper on the pseudochromid ge- nus Labracinus, based mostly on Albatross specimens] wrote a note in the margin that says “see USNM 136954.’” She also noted that there is a small tag in the jar containing USNM 136954 indicating that the specimen was drawn. We therefore conclude that the illustra- tion is based on the Pholidochromis speci- men. Clearly, then, the absence of dark markings in the specimen are not due to fading, as such markings are not indicated in the Albatross illustration, nor are they evident in the illustrations or photographs of live aquarium specimens. Thus, we con- clude that the specimens represent a species distinct from P. marginata, and therefore describe it as new. Materials and Methods Methods of counting, measuring and pre- sentation follow Gill (2003). Institutional codes follow Leviton et al. (1985). Pholidochromis cerasina, new species Cherry Dottyback Fig. 1 Pseudochromis xanthochir [non Bleeker, 1855]; Fowler, 1931: 32 (color descrip- tion). Pholidochromis marginata [non Pseudoch- romis marginatus Lubbock, 1980]; Gill, VOLUME 117, NUMBER 1 Fig. 1. (Photo by P. Hurst.) 2003:000, fig. 5 (description and distri- bution in part). Holotype.—USNM 136954, 43.9 mm SL, Indonesia, Sulawesi, Talisei (=Talisse) Island, R/V Albatross, 9 November 1909. Diagnosis.—Pholidochromis cerasinus is distinguished from other pseudochromi- nes in having the following combination of characters: dorsal-fin rays III, 22; anal-fin rays III, 13; scales in lateral series 29-30; dorsal and anal fins with well-developed scale sheaths; predorsal scales extending anteriorly to just behind anterior nostrils; and no prominent dark grey to black spots surrounding sensory pores on head. Description.—Dorsal-fin rays IJ, 22, at least last 18 segmented rays branched (ray preceding first apparent branched ray dam- aged); anal-fin rays III, 13, at least last 12 segmented rays branched (anteriormost ray damaged); pectoral-fin rays 19/19; upper procurrent caudal-fin rays 6; lower procur- rent caudal-fin rays 5; total caudal-fin rays 28; scales in lateral series 29/30; anterior lateral-line scales 23/24; anterior lateral line terminating beneath segmented dorsal-fin Pholidochromis cerasina, USNM 136954, 43.9 mm SL, holotype, Talisei Island, Sulawesi, Indonesia. ray 17/17; posterior lateral-line scales 10 + 0/9 + O; scales between lateral lines 3/3; horizontal scale rows above anal-fin origin 12 + 1 + 3/13 + 1 + 3; circumpeduncular scales 16; predorsal scales 21; scales behind eye 3; scales to preopercular angle 4; gill rakers 5 + 10; pseudobranch filaments 9; circumorbital pores 17/18; preopercular pores 9/8; dentary pores 4/4; posterior in- terorbital pores 0. Lower lip complete; dorsal and anal fins with well-developed scale sheaths; predor- sal scales extending anteriorly to just be- hind anterior nostrils; posterior margin of opercle with 4 inconspicuous serrations; outer gill rakers of ceratobranchial-1 with teeth mostly confined to raker tips; anterior dorsal-fin pterygiophore formula S/S/S + 3/ 1 + WI/WI/II/I/I/I/I/1/1/1 + 1; dorsal- fin spines pungent and moderately slender; anterior anal-fin pterygiophore formula 3/1/ 1 + 1/I/1/1 + 1; anal-fin spines pungent and moderately slender to stout, second spine stouter than third; pelvic-fin spine slender, tip weakly pungent; second seg- mented pelvic-fin ray longest; caudal fin 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rounded (inferred for holotype from Alba- tross illustration); vertebrae 10 + 16; epi- neurals 13; epurals 3. Upper jaw with 2 pairs of curved, en- larged caniniform teeth anteriorly, medial pair smallest, and about 6 (at symphysis) to 2-3 (on sides of jaw) irregular rows of small conical teeth, outermost of rows of teeth much larger and more curved than those of inner rows; lower jaw with 2 pairs of curved, enlarged caniniform teeth ante- riorly, medial pair smallest, and about 5 (at symphysis) to | (on sides of jaw) inner rows of small conical to caniniform teeth, those on middle of jaw large and canini- form; vomer with 1—2 rows of small conical teeth arranged in chevron; palatine with 2— 3 rows of small conical teeth arranged in elongate ovoid patch, anterior part of tooth patch more-or-less contiguous with postero- lateral arm of vomerine tooth patch; ectop- terygoid edentate; tongue moderately point- ed and edentate. As percentage of SL: head length 28.0; orbit diameter 9.1; snout length 7.7; fleshy interorbital width 6.2; bony interorbital width 3.9; body width 13.9; snout tip to posterior tip of retroarticular bone 15.9; predorsal length 35.3; prepelvic length 35.3; posterior tip of retroarticular bone to pelvic-fin origin 21.0; dorsal-fin origin to pelvic-fin origin 34.9; dorsal-fin origin to middle dorsal-fin ray 34.4; dorsal-fin origin to anal-fin origin 46.2; pelvic-fin origin to anal-fin origin 27.6; middle dorsal-fin ray to dorsal-fin termination 26.2; middle dor- sal-fin ray to anal-fin origin 33.5; anal-fin origin to dorsal-fin termination 39.0; anal- fin base length 29.6; dorsal-fin termination to anal-fin termination 17.1; dorsal-fin ter- mination to caudal peduncle dorsal edge 10.9; dorsal-fin termination to caudal pe- duncle ventral edge 19.6; anal-fin termina- tion to caudal peduncle dorsal edge 19.8; anal-fin termination to caudal peduncle ven- tral edge 10.7; first dorsal-fin spine 2.7; sec- ond dorsal-fin spine 5.2; third dorsal-fin spine 7.1; first segmented dorsal-fin ray 13.7; fourth from last segmented dorsal-fin ray broken; first anal-fin spine 3.0; second anal-fin spine 5.7; third anal-fin spine 7.1; first segmented anal-fin ray broken; fourth from last segmented anal-fin ray broken; third pectoral-fin ray broken (both sides); pelvic-fin spine 9.8; second segmented pel- vic-fin ray 22.6; caudal-fin length not de- termined (ray tips broken). Live coloration (based on a color illus- tration of holotype and photographs and an illustration of aquarium specimens).—Head and body pale pinkish grey to pinkish olive dorsally, becoming pale pink to pale yellow or white ventrally; posttemporal pore in dusky grey spot (not apparent in illustra- tions); orbital rim yellow to bright orange or bright red; pale blue to mauve stripe ex- tending from anteroventral edge of eye to middle of upper lip (not apparent on illus- tration of holotype); iris silvery white, blue dorsally, with grey to blue suboval ring around pupil; body with small (about half pupil size) pale orange to bright orange or bright red spots, these best developed dor- sally and posteriorly, and more or less ar- ranged along horizontal scale rows; dorsal and anal fins pale pink to pale blue with blue distal margin, and 2—S horizontal rows of pale orange to bright orange or crimson spots (crimson spots encircled with pale pink in photographed individuals); caudal fin pale pink to pale blue with blue distal margin and bright red to crimson spots (en- circled with pale pink in photographed in- dividuals), these irregularly arranged on basal part of fin, becoming arranged in con- vex columns on remainder of fin; pectoral fins hyaline with pinkish to yellowish hue; pelvic fins pale pink to pale blue. Preserved coloration.—Head and body pale brown, paler ventrally; posttemporal pore in dusky grey spot; fins whitish hya- line to plain hyaline. Habitat and Distribution.—No_ habitat data are known for the holotype. We also lack precise locality or habitat information for aquarium individuals of the species; however, K. Endoh (pers. comm.) informed VOLUME 117, NUMBER 1 Fig. 2. Province, Papua New Guinea. (Photo by P. Crabb; after Gill, 2003:fig. 20.) us that they were collected in the Philippine Islands. Comparisons.—Pholidochromis cerasina agrees closely with its congener, P. margin- ata (Fig. 2), in most details, but differs in lacking conspicuous dark spots around the sensory pores on the head (only the post- temporal pore of P. cerasina has an incon- spicuous dusky grey spot whereas P. mar- ginata has conspicuous dark grey to black spots on at least the posterior suborbital, upper preopercular, anterior interorbital, posttemporal and parietal pores) and in lacking dark submarginal markings on the median fins (present as dark grey to black convex marking on the caudal fin, and short dark grey to black stripe on the posterior part of the dorsal and anal fins in P. mar- ginata). Values for 13 morphometric characters of the holotype of P. cerasina lay at the ex- treme or outside ranges observed in P. mar- ginata (15 specimens, 27.2—45.6 mm SL). Although more specimens are needed to de- termine whether these are truly diagnostic, they are at least suggestive. We also docu- ment these in order to correct Gill’s (2003) Pholidochromis marginata, CAS 65783, 32.4 mm SL, southern side of Nagada Harbour, Madang description of P. marginata, as this includ- ed data from the holotype of P. cerasina. The characters are as follows (values ex- pressed as % SL, and given first for P. cer- asina, followed by P. marginata): fleshy in- terorbital width (6.2; 5.0—6.1); predorsal length (35.3; 36.0—39.0); middle dorsal-fin ray to dorsal-fin termination (26.2; 20.5— 25.2); anal-fin termination to caudal pedun- cle ventral edge (10.7; 10.8—12.4); first dor- sal-fin spine (2.7; 2.7—5.1); second dorsal- fin spine (5.2; 5.1—7.9); third dorsal-fin spine (7.1; 7.0—10.3); first segmented dor- sal-fin ray (13.7; 11.0—13.9); first anal-fin spine (3.0; 3.7—5.7); second anal-fin spine (5.7; 6.8-9.6); third anal-fin spine (7.1; 7.0—11.1); pelvic-fin spine (9.8; 9.6—13.2); and second segmented pelvic-fin ray (22.6; 22.6—26.3). Pholidochromis cerasina might also be confused with Pseudochromis fowleri Her- re, 1934, from Sabah and the Philippine Is- lands, and Pseudochromis fuscus, from throughout the West Pacific, which it re- sembles in general body shape. These spe- cies differ from Pholidochromis cerisina in having an incomplete lower lip (interrupted 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON at symphysis) and more segmented dorsal- fin rays (23-25, usually 24 in fowleri and 25-29 in fuscus versus 22 in cerasina). Remarks.—The live coloration of P. marginata is unknown, but, accepting the dark pigmentation on the head and median fins, is likely to be similar to P. cerasina. Moreover, as noted by Gill (2003), some specimens of P. marginata have pale spots on the body and median fins, and these pos- sibly correspond with the red to orange spots shown by P. cerasina. Etymology.—tThe specific epithet is from the Latin cerasinus, meaning “‘of cherry.” It alludes to the cherry-like bright orange to red spots on the body and median fins. Material examined.—See above. Acknowledgments We thank S. L. Jewett and J. T. Williams for lending the holotype for study, and for their help in checking the history of the AI/- batross specimen and its illustration. We thank W. E. Burgess and K. Endoh for sending an illustration and photographs, re- spectively, of the species. S. E. Reader as- sisted with radiographing the holotype. Literature Cited Bleeker, P. 1855. Zevende bijdrage tot de kennis der ichthyologische fauna van Celebes.—Naturrk- undig Tijdschrift Nederlandsch Indié 8:435— 444. . 1858. Bijdrage tot de kennis der vischfauna van den Goram Archipel.—Naturrkundig Tijdschrift Nederlandsch Indié 15:197—218. Fowler, H. W. 1931. Contributions to the biology of the Philippine Archipelago and adjacent re- gions. The fishes of the families Pseudochrom- idae, Lobotidae, Pempheridae, Priacanthidae, Lutjanidae, Pomadasyidae, and Teraponidae, collected by the United States Bureau of Fish- eries Steamer “Albatross’’, chiefly in Philippine seas and adjacent waters.—United States Na- tional Museum Bulletin 100(11):1—388. Gill, A. C. 2003. Revision of the Indo-Pacific dotty- back fish subfamily Pseudochrominae (Percifor- mes: Pseudochromidae).—Smithiana Mono- graph 1:1—213, 12 pls. Herre, A. 1934. Notes on fishes in the Zoological Mu- seum of Stanford University. 1. The fishes of the Herre Philippine Expedition of 1931. The Newspaper Enterprise, Hong Kong, 106 pp. Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. Dawson. 1985. Standards in herpetology and ichthyology: Part 1. Standard symbolic codes for institutional resource collections in herpe- tology and ichthyology.—Copeia 1985(3):802— 832. Lubbock, R. 1980. Five new basslets of the genus Pseudochromis (Teleostei: Pseudochromidae) from the Indo-Australian Archipelago.—Revue Suisse de Zoologie 87(3):821—834. Miiller, J.. & E H. Troschel. 1849. Horae Ichthyolo- gicae. Beschreibung und Abbildung neuer Fis- che. 3. Verlag von Kleit and Comp., Berlin, 28 pp. 5 pls. Schultz, L. P 1953. Family Pseudochromidae, pp. 380-411, pl. 33a in L. P. Schultz, E. S. Herald, E. A. Lachner, A. D. Welander, & L. P. Woods, Fishes of the Marshall and Marianas Islands; vol. 1. Families from Asymmetrontidae through Siganidae.—United States National Museum Bulletin 202(1):1—685. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):23-—34. 2004. Redescription of Cambaroides japonicus (De Haan, 1841) (Crustacea: Decapoda: Cambaridae) with allocation of a type locality and month of collection of types Tadashi Kawai and J. E Fitzpatrick, Jr.* (TK) Hokkaido Nuclear Energy Environmental Research Center, 261-1 Miyaoka, Kyowa, Hokkaido 045-0123, Japan, e-mail: kawaita@fishexp.pref.hokkaido.jp; (JFF) Museum of Natural History, Tulane University, Belle Chasse, Louisiana 70037, U.S.A. Abstract.—The Japanese crayfish, Cambaroides japonicus (De Haan), is re- described and illustrated, and details of its distribution and morphological var- iation are provided. Notable character differences between the populations of Honshu Island and Hokkaido Island indicate that gene flow between them is precluded. Analysis of geographical variation demonstrates that the undesig- nated type locality of the species is in central-western Aomori Prefecture, Hon- shu. The analysis of the gastrolith weights of the lectotype and possible top- otypes indicates that the lectotype was collected in June. The German medical doctor, Philip Franz von Siebold, was the first to introduce the natural history of Japan to European aca- demics (Siebold 1897). He also taught Eu- ropean medicine to traditional Japanese practitioners, and on 23 February 1826, at Shimonoseki City, Yamaguchi Prefecture, received specimens of a crayfish used as a Japanese drug from his student, Kosai Ya- maguchi (Siebold 1897). These were sent to the Netherlands and were described as Cambaroides japonicus by De Haan (1841). The brief description of the species included no locality or other collection data. Heretofore, taxonomic studies of C. japon- icus have been limited to examining cyclic dimorphism (Kawai & Saito 1999), and the genus Cambaroides has yet to be the sub- ject of modern morphological studies. This paper provides a redescription of C. japonicus, allocates a type locality based on an analysis of geographic variation, and suggests a probable month of collection of * Deceased 11 July 2002. the types based on an analysis of monthly changes in gastrolith weight. Abbreviations used in the text are: GVM, geographical variation in morphology; POCL, postorbital carapace length; RMNH, Nationaal Natuurhistorisch Museum, Lei- den; and TCL, total carapace length. Calculation of GVM: The geographical variation of each specimen was divided into three different levels (see Fig. 1), and the mean of the levels among specimens was calculated in each collection (Appendix I). The mean in each collection was classified into three degrees; 1.0—1.6, 1.7—2.3, 2.4— 3.0. Cambaroides japonicus (De Haan, 1841) Fig. 2, Table 1 Diagnosis.—Body pigmented; eyes well developed, pigmented. Carapace subcy- lindrical, dorsal and lateral surfaces with large punctations, without tubercles; cervi- cal spines absent. Rostrum acuminate, broadest at base; margins thickened, strong- ly convergent, lacking spines or tubercles; median carina present, often very weak; 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5 A, rostrum DREMEL RK ote tere es, B, telson C, sternal plates Fig. 1. Z, Definition of morphological variations. A-1, median carina present on rostrum; A-2, carina inter- mediate between A-1 and A-3; A-3, carina almost absent. B-1, caudomedian excavation present on telson; B-2, excavation intermediate between B-1 and B-3; B-3, excavation absent; C-1, sternal plates closed; C-2, sternal plates intermediate between C-1 and C-3; C-3, sternal plates open. acumen comprising 27.5—-59.5% (X = 47.8%, SD = 4.6, n = 200) of rostrum length, latter consisting 14.7—26.7% OX = 17.3%, SD = 2.5, n = 200) of TCL. Areola 112.3 0% = 1.9%, SD = O2, 1m = BOO) times as long as broad, constituting 26.3— AN 1% 0X = 25%, SD = 3.1, 2 = ZOO) oF TCL and 30.9-46.9% (X = 35.5%, SD = 3.3, n = 200) of POCL. Antennal scale 1.6— 2.8 (X = 2.2%, SD' = 0.3, n = 100) times as long as broad, widest at midlength, lat- eral margin thickened, terminating in large, stout spine. Pleura of somites 2 and 3 with rounded to subtruncate ventral margins. Palm of chela of cheliped with scattered large punctations on dorsal, lateral, and ventral surfaces, without setae; palm inflat- ed, width 1.2-1.6 (X = 1.4%, SD = O01, n = 100) times length of mesial margin. Large punctations on dorsal, lateral and ventral surface of fixed finger and dactyl. Hooks present on ischia of second and third pereiopods in males, hooks simple and not reaching basioischial articulation. In situ gonopods (first pleopods) of adult male symmetrical, bases not contiguous. In me- sial aspect (Fig. 2A), apex directed cephal- odistally nearly 45° to axis of shaft, with strong endopodite and protopodite; apex (Fig. 2B) sclerotized, at least distally, ce- VOLUME 117, NUMBER 1 25 Nese Fig. 2. Cambaroides japonicus (De Haan, 1841), all figures from lectotype (RMNH 5602, RMNH 5603), except B (redrawn from Hart 1953), I from paralectotype female (RMNH 2912), and j from paralectotype male#1: A, mesial view of first pleopod; B, mesial view of distal portion of first pleopod; C, lateral view of mandible; D, ventral view of ischum of third maxilliped; E, lateral view of first three abdominal segments; F dorsal view of carapace; G, epistome; H, dorsal view of distal podomeres of right cheliped; I, annulus ventralis; J, proximal podomeres of pereiopods; K, dorsal view of telson and uropods. Line = 2 mm. 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Measurements of type series of Cambaroides japonicus. Lectotype Carapace Total length 31.6 Postorbital length 25.4 Width 16.4 Height 11.1 Areola Length 98 Width 4.9 Rostrum Length 6.6 Width 7.3 Chela Length of mesial margin of palm 10.9 Width of palm 11.4 Length of lateral margin 24.5 Length of dactyl 14.2 Abdomen width 14.7 phalolaterally swollen into straight, sub- acute, stout, cephalodistally directed mesial process, cephalic process, and central pro- jection with blade-like caudal process; 3 subequal spines near mid-width of apex, length about one-tenth of width of apex. Proximal part of gonopod subcylindrical in cross section, becoming subtriangular dis- tally. Sperm groove along caudomesial face of gonopod shallow, open between mesial process and central projection, ending in relatively blunt tip. Adult male gonopod with “juvenile suture’’. Annulus ventralis (Fig. 21) immovable, symmetrical, rounded in outline, about 1.2 times as long as wide. Preannular plate transversely subdivided into 2 subtriangular plates, cephalic margin of anterior part broadly attached to preceding sternite, mid- dle section of posterior part with shallow depression as fossa without sinus. Postan- nular sclerite subcircular, about 1.7 times as broad as long, and 0.5 times as wide as an- nular plate. Measurements of type specimens provid- ed in Table 1. Description of lectotype.—Cephalotho- Paralectotype Paralectotype Paralectotype male #1 male #2 female WM 21.7 30.4 18.6 17.6 25.1 11.1 11.7 15.1 8.7 9.4 11.1 3 7.0 9.4 3.7 2.6 4.2 4.7 43 Voll 4.9 4.5 V3 6.9 5.6 8.9 7.6 733) 8.9 16.4 15.8 20.4 8.3 8.0 10.8 10.6 10.2 17.3 rax (Fig. 2F) subcylindrical, slightly com- pressed laterally; dorsoventrally depressed (greatest width of thoracic section 1.5 times depth); POCL 80.4% of TCL. Areola 2.0 times longer than wide, dense punctate, length 31.0% of TCL (38.6% of POCL). Rostrum acuminate, tip barely reaching dis- tal margin of antennal scale and midlength of ultimate podomere of antennal pedencle; acumen comprising 48.4% of rostrum length, latter consisting 20.9% of TCL; floor (dorsal surface) of rostrum dense punctate; median carina nearly absent. Postorbital ridges poorly defined. Sub- orbital angle obtuse, without tubercle or spine. Antennal scale with strong distolat- eral spine, tip reaching tip of rostrum and midlength of ultimate podomere of anten- nular peduncle. Greatest width of abdomen 89.6% great- est width of carapace. Proximal podomere of uropod lacking spine or tubercle on lat- eral lobe, mesial lobe broadly rounded; me- sial ramus of uropoda with caudolateral spine, and submedian dorsal ridge termi- nating in small caudomedian spine, tip of which not reaching caudal margin; lateral VOLUME 117, NUMBER 1 ramus with stout caudolateral spine; lateral ramus divided into cephalic and caudal sec- tions, separated by transverse flexure bear- ing spines. Telson divided into cephalic and caudal sections, each caudolateral corner with pair of stout, fixed spines. Caudal mar- gin of telson with deep median excavation. Epistome (Fig. 2G) with subovate ce- phalic lobe bearing prominent cephalome- dian projection, margins of lobe markedly elevated; fovea of epistome scarcely visi- ble; central portion of epistome with pair of transverse grooves, and deep transverse grooves along cephalic margin of weakly arched zygoma. Third maxilliped (Fig. 2D) with mesial margin bearing 21 denticles; mesial half of ischium with row of clusters of long, stiff setae. Incisor ridge of right margin (Fig. 2C) with 5 corneous denticles. Palm of right chela (Fig. 2H) subovate in cross section, moderately depressed dorso- ventrally. Total chela length 77.5% of TCL. Palm 2.1 times as long as wide, length of mesial margin 44.5% of total chela length; dorsal surface with deep, widely scattered punctations, which become scarce laterally and caudolaterally; ventral surface less punctate. Dorsal surface of both fingers with poorly defined, longitudinal submedi- an ridge, and rows of deep punctations; tip of fingers corneous, subacute. Opposable surface of fixed finger with row of 9 tuber- cles, third from base largest. Opposable sur- face of dactyl with row of 5 tubercles; length of dactyl 1.3 times length of mesial margin of palm. Carpus (Fig. 2H) longer than broad, dorsal surface with prominent longitudinal furrow, lateral and mesial sur- face with large, crowded punctations; me- sial surface with large, blunt subdistal spine, lateral surface with proximal spine; ventral surface with oblique furrow, short longitudinal furrow, and deep punctations. Merus with row of prominent tubercles on ventromesial margin, punctuate dorsally and ventrally. Gonopods as described in “‘Diagnosis’’. In addition, tips of gonopods extending be- 27 yond cephalic margin of coxae of fourth pe- reiopods. Description of paralectotype male#].— Differing from lectotype as follows: great- est width of thoracic section 1.3 times depth. POCL 84.2% of TCL. Areola length 33.0% of TCL (39.2% of POCL). Acumen comprising 46.1% of rostrum length, latter consisting 21.3% of TCL. Greatest width of abdomen 95.5% greatest width of carapace. Total chela length 74.2% of TCL; palm of right chela 2.2 times as long as wide; length of mesial margin 42.1% of total chela length. Dactyl length 1.2 times length of mesial margin of palm; opposable surface of fixed finger with 10 tubercles; opposable surface of dactyl with 6 tubercles. Description of paralectotype—female.— Differing from lectotype, except in second- ary sexual characteristics, as follows: great- est width of thoracic section 1.4 times depth. POCL 82.6% of TCL. Areola 2.2 times as long as broad. Areola length 30.9% of TCL (37.5% of POCL). Acumen com- prising 50.0% of rostrum length, latter con- sisting 23.4% of TCL. Opposable surface of fixed finger with 6 tubercles, proximal largest; opposable surface of dactyl with 6 tubercles, length of finger 1.2 times length of mesial margin of palm. Disposition of types.—All dry and lack- ing most appendages. Lectotype: 1 male, RMNH 5602, RMNH 5603. Paralectotypes: 2 males and | female, RMNH 2912. The lectotype has the mark ““@”’ written on the areola, but is a male. A milky-white gastro- lith is included with the lectotype. Its dry weight (dried at 80°, 48 hr) is 0.0305 g, and its shape is semi-globular, with the greatest diameter 4.5 mm, the least diameter 4.2 mm, and the greatest height 1.8 mm. Type locality.—No type locality for C. japonicus has ever been designated. In or- der to establish a type locality, we exam- ined geographic variation in morphology (GVM) to identify any unique characters that might be displayed by the type speci- mens. Earlier, Fitzpatrck (1995) detected possible geographically defined races based PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 145°E 4sN “oly A Sakhalin Hokkaido ie ae ( 140 “67 : = “\ Hokkaido 4 : ‘ a re 10 an ; ——_——= 28 z 40°N- 21 * 11 : ‘é Z L » we 30 29 16 4 YA ~— Korean \ es ie °° Pen. { Honshu / 343 me : sh be ~ i S20 eo ie ob , 130 140°E é is ray a ES Uae Aomori Pref. “e330 6064 7\ 40° = ee 77 (ee AkitaePref. ( Iwate Pref. i | = oe, fog Fig. 3. sampling sites listed in Appendix I. on a distinct rostral carina and pleural mar- gin shape of the abdominal segment. Sam- ples, however, were too small and too wide- ly scattered for definite conclusions. In our larger series from far more localities, we examined the presence or absence of a me- dian carina on the rostrum, median exca- vation in the caudal margin of the telson, and open or closed sternal plates. The GVM was Classified into three levels (Fig. 1), and mean GVM in each collection was sum- marized according to three categories by previously mentioned calculation (Fig. 4). Three GVM characters were found to be common in all the type series specimens, and similar to characters found only in specimens from central-western Aomori Prefecture, Honshu (Figs. 1—4, Appendix I, 55—62). This strongly suggests that the type Known geographical range of Cambaroides japonicus (De Haan, 1841). Numbers correspond to specimens were collected in that area. The central western Aomori Prefecture was de- signed as a probable locality of the type se- ries. Kurimi (1811) and Ohtsuki (1817), in a paper published at the time Siebold re- ceived specimens of C. japonicus, remarked that the species commonly inhabited cen- tral-western Aomori Prefecture. This lends support to our assumption about the type locality. Date of type collection.—Monthly sam- plings of C. japonicus were made from April to November 1989 in Iwaki City (Fig. 3, Appendix I, 59), in the central-western part of Aomori. For each sampling, gastro- liths were removed from the stomachs of 30 individuals, ranging in size from 17.7 mm to 25.4 mm POCL, which corresponds to size range of the types. The result indi- VOLUME 117, NUMBER 1 Aomori @ Pref. ree B, caudomedian : AkitaePref. excavation of telson Iwate Pref. C, sternal plates Fig. 4. Geographical variation of (A) carina on rostrum, (B) caudomedian excavation of telson, and (C) sternal plates. Solid circle, 1.0—1.6 level of mean GVM in collection; semi-open circle, 1.7—2.3; open circle, 2.4—3.0. A, Sapporo City; A’, Kazuno City. cates that the dry weight (dried at 80°, 48 hr) of the gastrolith from lectotype (0.0305 g) is similar to that of the June sample (Fig. 5). Thus, it is believed the type series was likely collected in June. Range and specimens examined.—We examined a total of 405 specimens from Hokkaido Islands and four of its larger nearby islands (Rebun, Rishiri, Teuri, and Yagishiri), as well as the northern part of Honshu Island (major parts of Aomori Pre- fecture, and northern part of Akita and Iwa- te Prefecture). Information on sampling sites is provided in Fig. 3 and Appendix I. As far as known, C. japonicus is endemic to the entire Japanese Archipelago. How- 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON S ‘2 ° =) 5 ui = ° v9) Weight of gastrolith (g) i=) ° 0 —— A M J Fig. 5. J A S O N Monthly change of dry gastrolith weight in Cambaroides japonicus (De Haan, 1841) from Iwaki City, Aomori Prefecture (see Fig. 3, 59). Vertical bar indicates SD. Dotted horizontal line shows the dry gastrolith weight of the lectotype (0.0305 g). ever, Okada (1933:155—156) mentioned that “Mr. T. Urita, a director of the Girl’s High School at Maoka in south part of Sakhalin, U.S.S.R., informed me that C. japonicus seems not to occur in the stream and rivers, and that if it is found in anywhere in Sa- khalin, it is very rare; however, I examined Outomari of Sakhalin specimens in the col- lection of Professor Iijima of Tokyo Impe- rial University, these are preserved in the Zoological Institute, Faculty of Sciences, Tokyo Imperial University”. And Urita (1942:39) said “I consequently spent con- siderable time and labour in search of this species, especially in Outomari and its neighbourhood, but unfortunately, without success; presumably, this species does not exist here in south Sakhalin, not, at any rate, at present”. On 11 November 2001, all specimens in the Tokyo Imperial Uni- versity were transferred to the University Museum, the University of Tokyo, but no specimen from the Sakhalin could be found there. Size.—The largest lake specimen is a male from Lake Akan, Hokkaido, measur- ing 39.2 mm POCL; the largest brook spec- imen is a male from Hamamasu with a POCL of 34.3 mm. The smallest ovigerous female is 17.6 mm POCL. Variation.—Most variations were noted in the number and comparative sizes of tu- bercular ornamentation, particularly on the cheliped. The caudolateral corner of the ce- phalic section of the telson bears one to three fixed spines. The lateral margin of the telson of most specimens gently tapers to a rounded caudal margin, but in some the lat- eral margins are subparallel and the caudal margin is flat. Some specimens have bosses between the sternal plates (Fig. 21), but in most specimens these bosses are nearly ab- sent. Color.—Dorsal and lateral surfaces of cephalothorax, abdomen, chelae, and tail fan dark brown to chocolate, ventral surface light brown. Ventral surface of chela dark orange. Tips of pereiopods dark orange. Caudal process and three spines of distal adult male gonopods amber. Background colors translucent to light brown in freshly molted individuals. The whitish-blue col- orations or “blue color phase” (Fitzpatrick 1987) on the dorsal and lateral surfaces of thoracic carapace, abdomen, chelae, and tail fan, was found in specimens from Abashiri, VOLUME 117, NUMBER 1 Obihiro, Iwamisawa, and Hamamasu, Hok- kaido Prefecture, and in Shichinohe, Ao- mori Prefecture (see Appendix I). Crayfish associates and conservation sta- tus.—During the past decade, local extinc- tions of C. japonicus have been reported from throughout its range. In eastern Hok- kaido its numbers have been declining rap- idly, while population numbers of the intro- duced crayfish, Pacifastacus leniusculus (Dana, 1852) in the same area have been increasing (Kawai et al. 2002). Also, Kawai et al. (2002) demonstrated that following the introduction of P. leniusculus into Lake Kussharo and Lake Shikaribetsu, C. japon- icus disappeared. Pacifastacus leniusculus is known to be a vector of crayfish plague fungus, Aphanomyces astaci (Schikora), to which it is resistant, but to which C. japon- icus 1s highly susceptible (Unestam 1969). It is possible that Aphanomyces may be a factor affecting displacement of C. japoni- cus at some localities, but there is as yet no investigation of infection to the natural pop- ulations in Hokkaido. The mechanisms un- derlying the negative impacts of P. lenius- culus on C. japonicus required further in- vestigation. Cambaroides japonicus was designated an endangered species by the Japanese Fisheries Agency in 1995 and by the Jap- anese Environmental Agency in 2000. Ecological notes.—Cambaroides japon- icus appears to be restricted to lentic habi- tats, either lakes or small brooks in which current velocity is less than 10.0 cm/sec. In brooks, the species is found beneath boul- ders, or burrows in the banks. It appears to be a secondary burrower, and retreats un- derground to remain below the frost line in winter. Females enter burrows prior to ovu- lation, and remain in them to lay eggs. Most burrows are Y- or T-shaped, with two open- ings slightly above or below the water sur- face. Reproduction.—Mating in C. japonicus is unique (Kawai & Saito 2001). The male moves beneath the female to deposit its spermatophore, and does not grasp the fe- 31 male with its chelae. In Hokkaido, mating pairs were encountered only in September and October, and ovigerous females during the subsequent May. Spermatozoa obvious- ly are stored in the annulus ventralis for a six-month period during winter. Number of ova ranges from 50 to 100, and egg diam- eter is 2.3—2.7 mm. Name in Japanese.—In Japan, it is usual for organisms to have one or more local names. To prevent possible ambiguity in this pragmatic system, and make it easier to incorporate taxonomic and distributional in- formation, the common, Japanese name Zarigani, is proposed. This name, which re- fers to an animal that moves backward (Ohtsuki 1817), is also mentioned in older papers (e.g., Kurimi 1811). The names “‘Sa- rugani’’ which is the local name in Aomori Prefecture, and “‘Sarukani,” the local name in Akita Prefecture, means “the backward creeping crab.”’ Two local names are on the label attached to the specimens of C. ja- ponicus at Saito Ho-Onkai Museum, Sen- dai, Japan (Nos. 1039, 1369). Also, the Ainu people, former occupants of Hokkaido and northern Honshu, know C. japonicus as ‘““Tekinpekorupe,” alluding to an armed knight (Ohtsuki 1817). Discussion Cambaroides japonicus occurs in north- ern parts of Honshu and Hokkaido Islands (Fig. 3). It is likely that populations of the species inhabiting certain areas of Honshu were introduced from Hokkaido, but differ- ences in the GVM (Fig. 4, Appendix I) in- dicate that the majority of populations on Honshu are native. An exception is seen in the GVM of specimens from Kazuno City (A’), Akita Prefecture, Honshu, which agrees with that of Sapporo City (A), Hok- kaido. In 1943, a locality report in a small stream in Kazuno City originated from in- troduction of a population in Sapporo City (Mr. T. Komoriya, Japanese regional report 1978). The distribution of Asian branchiobdel- 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lidans, which are symbionts on crayfish, in- cluding C. japonicus, may shed some ad- ditional light on this issue, since there is a high degree of endemism in the various species. Cirrodrilus aomorensis and C. tsu- garensis occur in Honshu (Gelder & Oht- aka 2000), while C. inukaii and C. uchidai, and others occur only in Hokkaido (Ya- maguchi 1934). There is no overlap in the natural distributions of these two species. However, C. inukaii and C. uchidai have both been found in Kazuno City, Akita Pre- fecture, northern Honshu, an occurrence that might be explained by an introduction of C. japonicus from Hokkaido (Gelder & Ohtaka 2000). Acknowledgments We thank A. Ohtaka, J. E. Cooper, and Y. Hanamura, who offered many useful suggestions concerning the present study. Thanks are extended to C. H. J. M. Fransen, S. E Mawatari, M. Takeda, K. Sakamoto, T. Urano, Y. Yabumoto, G. Scholtz, H. Hay- ashi, Y. Kobayashi, K. Nakata, and T. Ya- maguchi, who were most generous with their time, their collections, and their per- sonal solicitude. Figure | was drawn mostly by M. Tanaka. Literature Cited Fitzpatrick, J. F, Jr. 1987. Notes on the so-called “blue color phase” in North American Cambarid crawfishes (Decapoda, Astacoidea).—Crusta- ceana 52:316-319. . 1995. The Eurasian far-eastern crawfishes: a preliminary overview. Pp. 1-11 in R. P. Ro- maire, ed., Freshwater crayfish 8. Papers from the Eighth Symposium of the International As- sociation of Astacology, Baton Rouge, U.S.A. Gelder, S. R., & A. Ohtaka. 2000. Description of a new species and a redescription of Cirrodrilus aomorensis (Yamaguchi, 1934) with a detailed distribution of the branchiobdellidans (Anneli- da: Clitellata) in northern Honshu, Japan. — Proceedings of the Biological Society of Wash- ington 113:633-—643. Haan, W. de. 1841. Crustacea. Jn Ph. E von Siebold (1833-1850), Fauna Japonica sive descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava Imperium tenent, suscepto, annis 1823— 1830 collegit, notis, observationibus et adum- brationibus illustravit (Crustacea), i—xvii, 1— XXxi, ix—-xvi, 243 pp.+pls. A—-J, L—O, 1-S5, circ. tab. 2. Hart, C. W., Jr. 1953. Serial homologies among three pairs of abdominal appendages of certain male crayfishes (Decapoda, Astascidae).—Journal of Morphology 93:285—299. Kawai, T., & K. Saito. 1999. Taxonomic implication of the ‘Form’ and further morphological char- acters for the crayfish genus Cambaroides (Cambaridae). Pp. 82—89 in M. and M. M. Kel- ler, B. Oidtmann, R. Hoffmann, and G. Vogt, eds., Freshwater crayfish 12. Papers from the Twelfth Symposium of the International Asso- ciation of Astacology, Weltbild Verlag, Augs- burg, Germany. ,& . 2001. Observations on the mating behavior and season, with no form alternation, of the Japanese crayfish, Cambaroides japoni- cus (Decapoda, Cambaridae), in lake Koma- dome, Japan.—Journal of Crustacean Biology 21:885-—890. , K. Nakata, & T. Hamano. 2002. Temporal changes of the density in two crayfish species, the native Cambaroides japonicus (De Haan) and the alien Pacifastacus leniusculus (Dana), in natural habitats of Hokkaido, Japan. Pp. 198— 206 in G. Whisson and B. Knott, eds., Fresh- water crayfish 13. Papers from the Thirteenth Symposium of the International Association of Astacology, Curtin Print and Design, Perth, Australia. Kurimi, Z. 1811. Senchufu. Kouwa Shuppan, Tokyo, 534 pp. (rewritten in 1982) Ohtsuki, B. 1817. Ranwantekihou. Kouwa Shuppan, Tokyo, 524 pp. (rewritten in 1980) Okada, Y. 1933. Some observations of Japanese cray- fishes.—Science Reports of the Tokyo Bunrika Daigaku, Section B 1:155—-158 + pl. 14. Siebold, Ph. EF von. 1897. Nippon. Archiv zur Bes- chreibung von Japan und dessen Neben- und Schutzlandern Jezo mit den stidlichen Kurilen, Sachalin, Korea und den Liukiu-Inseln, 2nd edi- tion, vol. 1: 421 pp. + figs. 1-51, front 2 pls. and 1 map; vol. 2: 342 pp. + figs. 1—47. Unestam, T. 1969. Resistance to the crayfish plague in some American, Japanese and European cray- fishes.—Report, Institute of the Freshwater Re- search Drottningholm 49:202—209. Urita, T. 1942. Decapod crustaceans from Saghalien, Japan.—Bulletin of the Biogeographical Socie- ty of Japan 12:1—78. Yamaguchi, H. 1934. Studies on Japanese Brachiob- dellidae with some revisions on the classifica- tion.—Journal of the Faculty of Science, Hok- kaido University, Series VI, Zoology 3:177— 219. VOLUME 117, NUMBER 1 Appendix I.—Sampling data and geographic variation of morphology in Cambaroides japonicus. a OMANI ADUNAFSWNHFKFTOANIADAUMHKWNY AnmnrnnnbBPR HHH HHH HWWWWWWWWWWNNNNNNNN DN WV BWNF DOA AIANUAHEWNKTDOAWAIAIADAUNSWNHNHK CTOANIAUANKWNH OS Sampling site (No. ref. to Fig. 3) Rebun Rishiri Wakkanai Nakagawa Shibetsu Abashiri Utoro Tsubetsu Maruseppu Akan Shikaoi Memuro Ikeda Otofuke Obihiro Shintoku Erimo Monbetsu Kamikawa Biei Akabira Furano Iwamisawa Takikawa Ofuyu Teuri Yagishiri Hamamasu Sapporo Otaru Yoichi Kucchan Niseko Kyowa Rankoshi Iwanai Suttsu Kyogoku Oshamanbe Chitose Eniwa Soubetsu Shiraoi Sahara Assabu Kaminokuni Shikabe Kikonai Toi Todohokke Fukushima Matsumae Imabetsu Shiura Date 04 Sep 1968 17 Nov 1992 11 Aug 1998 16 Aug 1990 O01 Sep 1957 04 July 1982 18 Aug 1999 25 Apr 1987 10 Aug 1998 04 Aug 1933 14 May 1987 04 Aug 1986 25 Sep 2000 25 Sep 2000 08 May 1986 09 Aug 1998 19 Sep 2000 20 July 1990 20 Sep 2000 07 Aug 1998 08 Aug 1999 08 Aug 1999 27 May 1959 11 Aug 1992 23 Aug 1992 29 July 2002 29 July 2002 17 June 2000 03 Nov 1990 03 June 1999 28 Aug 2001 07 Oct 2000 08 Oct 2000 10 Oct 1998 21 July 2001 O01 July 2001 13 Sep 2001 28 Sep 2001 19 Sep 2001 02 Aug 1999 04 Noy 2001 02 Aug 1998 12 Aug 1990 O1 Aug 1998 23 June 1997 22 June 1997 23 Aug 1998 22 June 1997 20 Aug 1992 23 June 1997 22 June 1997 22 June 1997 05 Aug 1998 30 Aug 1997 Specimens 33 31 34 31 31 30 33 32 32 Jl 36 30 30 31 31 30 32 34 31 31 31 32 32 31 30 35 35 35 34 35 35 35 35 34 35 36 35 34 33 32 32 31 30 32 34 31 31 32 32 35 33 37 31 32 92 21 21 20 21 22 21 20 al 22 21 21 91 20 20 21 21 20 21 21 20 22 23 21 2 il 25) 25 23 2Q5 26 25 25 25 26 25 24 25) 26 25 Q5 Q? 20 23 91 23) 21 24 27 27 27 27 25 24 20 POCL + SD AIL3) 22 DS) 22.3 = 4.8 DQ 2 D§3 20.2 = 0.0 16.1 + 2.0 Dpdecd). 2 Pe) IQ/2 2= i) ASM 2 Of 21.0 + 4.1 26.8 + 10.8 26.9 + 84 24.5 + 0.0 13.2 = 0.0 1.97 = 0.0 19.2 + 0.0 15.4 + 0.0 18.8 + 0.5 IS}.5) 22 22,33 21.4 +49 D3) = 35 19.6 + 0.0 18.5 + 3.0 18.7 + 1.0 202 20.0 + 0.0 21.6 = 2.0 20.4 + 1.6 20.7 + 3.0 23a = 20 ADY) 2 27 USES =s1kS Dies) 28 2s QING 25 Doll PN 2 Dall 19.@ = 2A IO.2 2 If 20.5 + 4.2 p,3) 2 Il-9) I@.7/ = 2D 21.0 = 2.5 /DQ.3} 2 ZS 20.7 + 0.0 18.8 + 1.7 Alls) 22, 20) AVS 25 25) Doped 2 M5) Wp, = 2,1 20.4 = 1.7 19.9 + 1.0 20.7 = 2.5 Ailey 2= 2, DBA aD, 22.0 + 1.4 20:92 23 Rostrum 1.0 + IES = a= + OR S I+ 0.0 0.7 0.4 0.0 + 0.0 0.0 0.5 0.0 0.6 0.0 0.0 + 0.0 0.0 0.0 0.0 + 0.0 0.0 0.0 + 0.0 0.0 0.0 0.0 + 0.0 0.0 0.0 0.4 0.5 0.0 0.4 + 0.0 + 0.0 0.0 0.4 0.4 0.5 0.0 + 0.0 + 0.5 0.5 0.5 + 0.0 0.0 0.0 0.6 0.5 0.0 0.5 0.5 0.0 0.3 0.8 0.8 + 0.8 +14 Telson 33 Sternal plates 1.0 = 0.0 1.0 + 0.0 1.0 + 0.0 1.0 + 0.0 1.0 + 0.0 2.0 + 0.0 2.0 + 0.0 1.0 + 0.0 1.0 + 0.0 1.0 + 0.0 1.0 + 0.0 1.0 + 0.0 34 Appendix I.—Continued. Sampling site (No. ref. to Fig. 3) SS) Nakasato 56 Kanagi 57 Goshogawara 58 Kizukuri 59 Iwaki 60 Hirosaki 61 Ikarigaseki 62 Ajigasawa 63 Namioka 64 Aomori 65 Hiranai 66 Tenmabayashi 67 Shichinohe 68 Yokohama 69 Higashidouri 70 Mutsu 71 Ouhata 72 Kawauchi 73 Wakinosawa 74 Tashiro WD Ohdate 76 Kazuno Wi Ninohe —: not measured. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Date 21 Nov 1999 07 Oct 2000 19 Sep 1998 22 Aug 1999 26 July 1998 30 Sep 1997 04 Oct 1931 12 Aug 1998 22 June 2000 03 May 1998 10 Oct 2000 14 July 1994 22 June 1999 23 May 1999 27 Aug 1999 27 Aug 1999 22 June 1999 05 June 1998 17 May 1998 21 June 1991 24 Aug 2002 23 June 1990 22 June 1999 Specimens 31 30 37 38 36 32 32 32 34 37 32 30 32 31 32 31 31 30 33 30 33 37 31 al 92 24 22) 20 92 25 22 ont 23 23 92 91 20 QD 91 25 oD 20 22 27 22 QD POCL + SD DNS) = 2,5) 20.3 + 0.4 NZ se 4 DBI] 32 Af ANS) 2D W233 a= DD WYO 25 22 MO.i1 2.3) ANd} 2B NY) 20.7 + 3.4 IAL se 2a 18.6 + 1.7 IS) 22 ILO 19.4 + 0.0 19.0 + 1.4 23.0 = 0.7 18.3 + 0.8 AMA Se Mei 18.3 + 0.9 73 = 0.3 ANY = ZY 18.9 + 1.1 18.3) 2 12 Rostrum Ans) 25 (0),7/ 3.0 + 0.0 Des) = OLS) Ap 2 Os) 2.7 + 0.6 3.0 + 0.0 Mell 2 Dei 2.8 = 0.5 2.8 + 0.4 1.9 = 0.6 2.4 3.0 Zoll 3.0 = 0.0 2.5 + 0.6 It Ut 1+ 1 It dt Ut 1 Le IE It it Telson 1.5 2.0 2.0 De 1.5 2.0 2.6 eS) 2.2 2.8 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 3.0 1.5 3.0 3.0 2.7 0.7 1.4 + 0.9 xe O.7/ 0.8 1.0 0.5 1.0 + 0.8 0.4 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.6 I+ I+ Ine Ute We |e It It It It [+ It I+ It It it I+ I+ It I+ Sternal plates 3.0 = 0.0 Dis) 2 (0,7) 2.5 + 0.6 2.0 + 0.0 3.0 + 0.0 3.0 + 0.0 3.0 + 0.0 3.0 + 0.0 2.7 + 0.6 3.0 = 0.0 3.0 + 0.0 3.0 + 0.0 3.0 = 0.0 3.0 = 0.0 2.6 + 0.5 2) 22 (0,7) 3.0 + 0.0 3.0 = 0.0 1.0 + 0.0 3.0 = 0.0 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):35—41. 2004. Two new species of freshwater crabs of the genus Chaceus Pretzmann, 1965 from the Serrania de Perija of Colombia (Crustacea: Decapoda: Pseudothelphusidae) Martha R. Campos and Diego M. Valencia (MRC) Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 103698, Bogota, Colombia, S. A, e-mail: mhrocha@ciencias.unal.edu.co; (DMV) Universidad Nacional de Colombia, Departamento de Biologia, Apartado Aéreo 7495, Bogota, Colombia, S. A. Abstract.—Two new species of the genus Chaceus Pretzmann, 1965, C. cu- rumanensis and C. ibiricensis, are described and illustrated. The description of these two new species brings to nine the total number of species known in this genus, distributed in the Sierra de Santa Marta of Colombia, and Serrania de Perija of Colombia and Venezuela. A key for the identification of the species based on the morphology of the first male gonopod is presented. The genus Chaceus Pretzmann, 1965 comprises a group of freshwater crabs dis- tributed in the Sierra de Santa Marta in Co- lombia and the Serrania de Perija in Colom- bia and Venezuela. The systematics, cladis- tic and biogeography of the genus have been reviewed by Rodriguez (1982, 1992), Campos & Rodriguez (1984), Rodriguez & Campos (1989), Rodriguez & Bosque (1990), Rodriguez & Viloria (1992) and Rodriguez & Herrera (1994). With the dis- covery of two new species, described here- in, from the western slope of the Serrania de Perija of Colombia, the genus now con- tains nine species. Species of Chaceus are distinguished pri- marily by characteristics of the efferent branchial channel, the third maxilliped and the first male gonopod. The efferent bran- chial channel is partially closed by the spine of the jugal angle, and by the produced lat- eral lobe of the epistome. The exognath of the third maxilliped is 0.60 to 0.80 times as long as the ischium. The first male gonopod usually has the lateral process well devel- oped, its shape varying according to the species, and is either subtriangular, elon- gated, or rounded. The apex is formed by mesial and caudal processes. A key for the species of the genus is presented, based ex- clusively on the morphology of the first male gonopod. The terminology used for the different processes of the gonopod is that established by Smalley (1964), and Rodriguez (1982). The shape of the efferent branchial chan- nel, the length of the exognath of the third maxilliped, and the structure of the first male gonopod of the genus Chaceus sug- gest a close relationship with the genus Strengeriana Pretzmann, 1971. The first male gonopods in all species of Chaceus have the same basic elements as species of Strengeriana. Rodriguez (1982) has theo- rized on the possible derivation of the genus Hypolobocera Ortmann, 1897, from an an- cestral Chaceus based on the homology of the finger-like mesial process in the latter, and the triangular caudal process with the two papillae found near the spermatic chan- nel in the former. The morphology of the first gonopod in C. davidi Campos & Rod- riguez, 1984, for example, supports this the- ory since the mesial and caudal processes are surrounded by a ridge that somewhat resembles the shape of the apex in species of Hypolobocera. The material is deposited in Museo de 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Historia Natural, Instituto de Ciencias Na- turales, Universidad Nacional de Colombia, Bogota (ICN-MHN). The abbreviations cb and cl, reported as cl X cb, indicate cara- pace breadth and carapace length, respec- tively. Color nomenclature follows Smithe @sr)): Family Pseudothelphusidae Rathbun, 1893 Tribe Strengerianini Rodriguez, 1982 Genus Chaceus Pretzmann, 1965 Chaceus curumanensis, new species Fig. 1 Holotype.—Quebrada San Sebastian, Municipio Curumani, foothill of the Serran- ia de Perija, Cesar Department, Colombia, 100 m alt., 8 Dec 1978, leg. M. Tiirkay, male, 14.7 xX 24.5 mm, ICN-MHN-CR 1923). Paratype.—Same locality data as holo- type: | male, 13.2 x 23.4 mm, ICN-MHN- CR 1266. Type locality.—Quebrada San Sebastian, Municipio Curumani, foothill of the Serran- fa de Perijé, Cesar Department, Colombia, 100 m alt. Diagnosis.—Third maxilliped with ex- ognath 0.67 times length of ischium. First gonopod with lateral process elongated, with distal portion slightly rounded in cau- dal view, subtriangular in distal view; apex with needle-shaped mesial process, and tri- angular caudal process; disto-mesial margin curving below mesial and caudal processes. Description of holotype.—Carapace (Fig. 1F) with cervical groove straight, narrow and shallow distally, wide and deep proxi- mally, ending some distance from lateral margin. Anterolateral margin lacking de- pression behind external orbital angle, but with slight depression near middle followed by another near level of cervical groove. Lateral margin with series of tubercles. Postfrontal lobes small, oval, delimited an- teriorly by 2 depressions. Median groove lacking. Front without distinct upper border, frontal area regularly sloping downward, slightly bilobed in dorsal view, lower mar- gin sinuous in frontal view. Dorsal surface of carapace smooth, covered by small pa- pillae, regions not well demarcated. Third maxilliped with slight depression on distal half of external margin of merus, exognath 0.67 times length of ischium (Fig. 1H). Or- ifice of efferent branchial channel partially closed by spine of jugal angle, and by pro- duced lateral lobe of epistome (Fig. 1G). First pereiopods heterochelous; chelae with palms swollen, and fingers slightly gaping when closed (Fig. 11). Walking legs (pe- reiopods 2—5) slender, but not unusually elongated (total length 1.14 times breath of carapace). First male gonopod with lateral process elongated, with distal portion slightly rounded in caudal view (Fig. 1A), subtrian- gular in distal view (Fig. IE); apex with needle-shaped mesial process, directed ce- phalically, and triangular caudal process, directed transversely to mesial process in caudal and cephalic views, both processes surrounded by lateral process in distal view; disto-mesial margin curving below mesial and caudal processes (Fig. 1C—E); lateral side of gonopod expanded with irregular rows of short setae, caudal surface with long setae proximally (Fig. 1A, B, D). Color.—The holotype, preserved in al- cohol, is light brown (near 37, Antique Brown) with dark specks on the dorsal side of the carapace. The dorsal and ventral sur- faces of the chelae and walking legs are brown (near 139, True Cinnamon). The ventral surface of the carapace is brown (near 239, Ground Cinnamon). Etymology.—The specific name refers to the type locality, the Municipio Curumani. Remarks.—Comparison of this new spe- cies with descriptions and specimens of other species of the genus revealed that this new species is most similar to Chaceus pearsei (Rathbun, 1915). The two can be distinguished by differences in the gono- pods. The male first gonopod of C. pearsei has been described and illustrated by Rod- riguez (1982:37, fig. 12). The lateral pro- cess in this new species is elongated, with VOLUME 117, NUMBER 1 37 m UCFinwen Fig. 1. Chaceus curumanensis, new species, male holotype, ICN-MHN-CR 1993: A, left first gonopod, caudal view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view; E right side of carapace with eye, dorsal view; G, left orifice of efferent branchial channel; H, left third max- illiped, external view; I, left cheliped, external view. 1, lateral process; 2, mesial process; 3, caudal process. 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the distal portion slightly rounded in caudal view, whereas it is subtriangular in C. pear- sei. The mesial process in C. pearsei is fin- ger-like, blunt, whereas it is needle-shaped in C. curumanensis. The caudal process in this new species is slightly parallel to the mesial process in distal view, whereas it is recurved at its base in C. pearsei. Chaceus ibiricensis, new species Fig. 2 Holotype.—Los Laureles Farm, Vereda Alto del Tucuy, Corregimiento La Victoria de San Isidro, Municipio La Jagua de Ibir- ico, Serrania de Perija, Cesar Department, Colombia, 1100 m alt., 9°34'35.8’N, 73°6'26.0"W, 7 Mar 1996, leg. M. R. Cam- pos, male, 13.0 X 21.3 mm, ICN-MHN-CR IQQy. Paratypes.—Same locality data as holo- type: 19 males, size range 8.6 X 13.7 mm to 13.7 X 22.7 mm, 16 females, size range 8.2 X 12.8 mm to 12.4 X 20.1 mm, ICN- MHN-CR 1549. Additional non-paratypic material.—Be- tween Veredas Alto de las Flores, and Nue- vo Mundo, Corregimiento La Victoria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar Department, Co- lombia, 1350-1400 m alt., 7, 8 Mar 1996, leg. M. R. Campos, 23 males, size range 7.3 X 11.4 mm to 13.0 X 21.4 mm, 15 females, size range 8.4 X 13.1 mm, to 14.4 x 24.8 mm, ICN-MHN-CR 1550, 1552.— Tucuy River, Vereda Alto de las Flores, Corregimiento La Victoria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar Department, Colombia, 870 m alt., 11 Mar 1996, leg. M. R. Campos, 6 males, size range 9.9 X 15.9 mm to 10.9 x 17.6 mm, 3 females, size range 10.8 * 17.2 mm to 11.9 X 20.4 mm, 2 juveniles, ICN- MHN-CR 1559.—La Sorpresa Farm, Ver- eda Alto de las Flores, Corregimiento La Victoria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar De- partment, Colombia, 1280 m alt., 12 Mar 1996, leg. J. V. Rueda, 1 male, 12.4 X 21.1 mm, ICN-MHN-CR 1560. Type locality.—Los Laureles Farm, Ver- eda Alto del Tucuy, Corregimiento La Vic- toria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar Depart- ment, Colombia, 1100 m alt., 9°34’35.8’N, 73°6'26.0"W. Diagnosis.—Third maxilliped with ex- ognath 0.72 times length of ischium. First male gonopod with lateral process hood- like; mesial process prominent, subcylindri- cal, semicircular caudally with median con- striction and subdistal subtriangular papilla cephalically; caudal process subtriangular; disto-mesial margin forming semicircular projection in cephalo-lateral direction. Description of holotype.—Carapace (Fig. 2F) with cervical groove straight, narrow, shallow, ending some distance from lateral margin. Anterolateral margin with shallow depression behind external orbital angle fol- lowed by approximately 5 papillae. Lateral margin with series of approximately 10 tu- bercles. Postfrontal lobes small, oval, de- limited anteriorly by 2 depressions. Median groove shallow, and narrow. Front lacking distinct upper border, frontal area regularly sloping downward, bilobed in dorsal view, lower margin sinuous in frontal view. Dor- sal surface of carapace smooth, covered by small papillae, regions not well demarcated. Third maxilliped with external margin of merus straight, exognath 0.72 times length of ischium (Fig. 2H). Orifice of efferent branchial channel partially closed by spine of jugal angle, and by produced lateral lobe of epistome (Fig. 2G). First pereiopods het- erochelous; palm of larger chela strongly swollen, fingers gaping when closed (Fig. 21); palm of smaller chela moderately swol- len, fingers not gaping when closed. Walk- ing legs (pereiopods 2—5) slender and elon- gated (total length 1.25 times the breadth of carapace). First male gonopod with lateral process hood-like, lateral and cephalic outer surface covered with irregular papillae and spinules (Fig. 2A—C); apex with mesial and caudal VOLUME 117, NUMBER 1 39 OC Pinzen m mn A-D | G | Fig. 2. Chaceus ibiricensis, new species, male holotype, ICN-MHN-CR 1992: A, left first gonopod, caudal view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view; EF right side of carapace with eye, dorsal view; G, left orifice of efferent branchial channel; H, left third maxilliped, external view; I, right cheliped, external view. 1, lateral process; 2, mesial process; 3, caudal process. 3 3 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON processes; mesial process prominent, sub- cylindrical, semicircular caudally (Fig. 2A, B); with median constriction, and subdistal subtriangular papilla cephalically (Fig. 2C— E); caudal process subtriangular, both pro- cesses partially surrounded by lateral pro- cess in distal view; disto-mesial margin forming semicircular projection into ce- phalo-lateral direction (Fig. 2E); lateral ex- panded side of gonopod with rows of long, plumose setae, mesial side with row of spi- nules; caudal surface with conspicuous long setae proximally (Fig. 2A—D). Color.—The holotype, preserved in al- cohol, is brown (near 240, Kingfisher Ru- fous) on the dorsal side of the carapace. The dorsal and ventral surfaces of chelae and walking legs are brown (near 223B, Verona Brown). The ventral surface of the carapace is light brown (near 223C, Sayal Brown). Habitat.—The vegetation of the collec- tion areas is primary forest. The specimens were collected in shaded, moist banks of springs and streams, in soft mud under rocks. Etymology.—tThe specific name refers to the type locality, the Municipio La Jagua de Ibirico. Remarks.—Comparison of this new spe- cies with descriptions and specimens of other species of the genus revealed that it is most similar to Chaceus turikensis Rod- riguez & Herrera, 1994. The two can be distinguished by differences in the size of the eyes, and in the gonopod. The male first gonopod of C. turikensis has been described and illustrated by Rodriguez & Herrera (1994:123, fig. 2). In C. turikensis the eyes do not fill the orbital cavity, whereas in this new species they do fill the orbital cavity. In C. ibiricensis the lateral process of the gonopod is hood-like with the distal portion directed distally in caudal view (Fig. 2A— E), whereas the lateral lobe is foliose and the distal portion is directed transversely to the main axis of the appendage in C. turi- kensis. The mesial process is ellipsoidal in C. turikensis, whereas it is subcylindrical with a median constriction and subdistal subtriangular papilla cephalically in C. tbir- icensis. Key to Species of Chaceus 1. Lateral process of gonopod well devel- oped — Lateral process of gonopod reduced C. nasutus Rodriguez, 1980 2. Lateral process of gonopod subtriangular or elongated — Lateral process of gonopod rounded... 8 3. Lateral process of gonopod with semi- circular notch on lateral surface .. C. cesarensis Rodriguez & Viloria, 1992 — Lateral process of gonopod without semicircular notch on lateral surface .. 4 4. Mesial process of gonopod about same length as length of caudal process . C. dayidi Campos & Rodriguez, 1984 — Mesial process of gonopod longer than caudal process 5. Mesial process of gonopod with median constriction and subapical subtriangular papilla cephalically Peeters: Sos C. ibiricensis, new species — Mesial process of gonopod without me- dian constriction and subapical papilla cephalically 6. Mesial process of gonopod ellipsoidal ... .. C. turikensis Rodriguez & Herrera, 1994 — Mesial process of gonopod not ellipsoi- dal aco, To MR eee vi 7. Mesial process of gonopod finger-like, blunteaee ee ae C. pearsei (Rathbun, 1915) — Mesial process of gonopod needle- shaped C. curumanensis, new species 8. Mesial process of gonopod with round- ed, elongated papilla basally . C. caecus Rodriguez & Bosque, 1990 — Mesial process of gonopod lacking rounded, elongated papilla basally ia a eae C. motiloni Rodriguez, 1980 Acknowledgments I am indebted to R. Lemaitre, of the Na- tional Museum of Natural History, Smith- sonian Insitution, for his corrections and suggestions to improve the manuscript. I thank E G. Stiles, and the anonymous ref- erees for providing useful comments. The illustrations were prepared by Juan C. Pin- VOLUME 117, NUMBER 1 zon. The specimens of Chaceus curuma- nensis were donated by M. Tiirkay, of the Senckenberg Museum of Frankfurt to the ICN-MHN collection. Literature Cited Campos, M. R., & G. Rodriguez, 1984. New species of freshwater crabs (Crustacea: Decapoda: Pseudothelphusidae) from Colombia.—Pro- ceedings of the Biological Society of Washing- ton 97:538—543. Ortmann, A. 1897. Carcinologische Studien.—Zoolo- gische Jahrbiicher, Abtheilung fiir Systematik, Geographie und Biologie der Tiere 10:258-372. Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam- ilie Pseudothelphusidae.—Anzeiger der Oster- reichischen Akademie der Wissenschaften Mathematische Naturwissenschaftliche Klasse (1)1:1-10. . 1971. Fortschritte in der Klassifizierung der Pseudothelphusidae.—Anzeiger der Mathema- tisch Naturwissenschaftliche der Osterreichisch- en Akademie der Wissenschaften (1)179(1—4): 14-24. Rathbun, M. J. 1893. Descriptions of new species of American freshwater crabs.—Proceedings of the United States National Museum 16(959): 649-661. . 1915. New fresh-water crabs (Pseudothelphu- sa) from Colombia.—Proceedings of the Bio- logical Society of Washington 28:95—100. Rodriguez, G. 1980. Description préliminaire de quelques espéces et genres nouveaux de Crabes d'eau douce de 1’Amérique tropicale (Crusta- cea: Decapoda: Pseudothelphusidae).—Bulletin 41 du Muséum nationale d’ Histoire naturelle, Paris (4) 2 Section A (3):889—-894. . 1982. Les crabes d’eau douce d’ Amérique. Famille des Pseudothelphusidae.—Faune Trop- icale 22:1—223. . 1992. ‘The freshwater crabs of America. Fam- ily Trichodactylidae and supplement to the Family Pseudothelphusidae.—Faune Tropicale 31:1-189. , & C. Bosque. 1990. A stygobiont crab, Cha- ceus caecus n. sp. and its related stygophile spe- cies Chaceus motiloni Rodriguez, 1980, (Crus- tacea, Decapoda, Pseudothelphusidae) from a cave in the Cordillera de Perija, Venezuela.— Mémoires de Bioespéologie XVII:127—134. , & M. R. Campos. 1989. Cladistic relation- ships of freshwater crabs of the tribe Strenger- ianini (Decapoda: Pseudothelphusidae) from the northern Andes, with comments on their bio- geography and descriptions of new species.— Journal of Crustacean Biology 9:141—156. , & FE Herrera. 1994. A new troglophilic crab, Chaceus turikensis, from Venezuela, and addi- tional notes of the stygobiont crab Chaceus cae- cus Rodriguez & Bosque 1990, (Decapoda: Brachyura: Pseudothelphusidae)—Mémoires de Bioespéologie XXI:121—126. , & A. L. Viloria. 1992. Chaceus cesarensis, a new species of the fresh-water crab (Crustacea: Decapoda: Pseudothelphusidae) from Colombia with a key to the genus.—Proceedings of the Biological Society of Washington 105:77—80. Smalley, A. 1964. A terminology for the gonopods of the American river crabs.—Systematic Zoology 13:28-31. Smithe, FE B. 1975. Naturalist’s color guide. The Amer- ican Museum of Natural History, New York. Part 1: unnumbered pages. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):42—56. 2004. Reevaluation of the hermit crab genus Parapagurodes McLaughlin & Haig, 1973 (Decapoda: Anomura: Paguroidea: Paguridae) and a new genus for Parapagurodes doederleini (Doflein, 1902) Patsy A. McLaughlin and Akira Asakura* Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, WA 98221-9081B, U.S.A.; *Natural History Museum and Institute, Chiba 955-2, Aoba-cho, Chuo-ku, Chiba, 260-8682, Japan Abstract.—The question of polyphyly in the hermit crab genus, Parapagu- rodes McLaughlin & Haig, 1973, has been investigated by comparisons of a series of morphological characters among the eight species presently assigned to the genus. The results of the analysis have shown that the only mutually shared characters are an acutely developed rostrum and the presence, in males, of a short or very short right sexual tube. Consequently, the composition of Parapagurodes is herein restricted to the two species originally assigned, viz. P. makarovi McLaughlin & Haig, 1973, and P. laurentae McLaughlin & Haig, 1973. Parapagurodes hartae Mclaughlin & Jensen, 1996, is transferred to the genus Pagurus, and four species subsequently transferred from Pagurus to Parapagurodes, viz. P. gracilipes (Stimpson, 1858), P. nipponensis (Yokoya, 1933), P. constans (Stimpson, 1858), and P. imaii (Yokoya, 1939) are returned to Pagurus. A new genus, Dofleinia, is proposed for the species, Parapagu- rodes doederleini (Doflein). When first proposed, the genus Parapa- gurodes McLaughlin & Haig, 1973, was characterized, in part, as having 11 pairs of biserial gills; a moderately well developed, but not recurved external lobe of the max- illulary endopod; fifth pereopods with cox- ae symmetrical; males with a short right sexual tube, and biramous left pleopods ab- sent or weakly developed on pleomeres (cf. Schram & Koenmann 2003) 3—5; females lacking paired first pleopods, with biramous left pleopods 2—4 weakly to moderately well developed, left fifth pleopod weakly developed or absent; and a telson with ter- minal margins straight, slightly concave or slightly oblique. Additionally, the authors noted that while a right sexual tube was al- ways present in mature males, its length and orientation were variable, and in one specimen both right and left tubes were pre- sent. Variations also were observed in the number and development of male and fe- male pleopods in both P. makarovi Mc- Laughlin & Haig, 1973, the type species of the genus, and the second described spe- cies, P. laurentae McLaughlin & Haig, 1973. In recent years, one new species, P. hartae McLaughlin & Jensen, 1996, has been described in the genus, and five Jap- anese species have been transferred to it viz.: Pagurus gracilipes (Stimpson, 1858), P. nipponensis (Yokoya, 1933), P. constans (Stimpson, 1858), and P. imaii (Yokoya, 1939) by Komai (1998, 1999) and Cata- pagurus doederleini Doflein, 1902 by Asakura (2001). At the time of the establishment of Par- apagurodes McLaughlin & Haig, 1973, male sexual tube development had been re- ported in less than two dozen genera. McLaughlin & Haig (1973) could relate Parapagurodes to only two of those genera, VOLUME 117, NUMBER 1 Pagurodes Henderson, 1888 and Acantho- pagurus de Saint Laurent, 1969, but cited several characters by which the three genera could be separated. In the subsequent 30 years the number of genera with docu- mented sexual tube development has more than doubled (cf. McLaughlin 2003). None- theless, Parapagurodes still can be allied only to Pagurodes and Acanthopagurus and more remotely to Catapagurus A. Milne Edwards, 1880. However, recently the monophyly of Parapagurodes itself has come under question (Lemaitre & Mc- Laughlin 2003b). In their introductory remarks regarding Parapagurodes hartae, McLaughlin & Jen- sen (1996: 841) made the unfortunate state- ment that “‘... males have a small sexual tube on the coxa of the right fifth pereopod. This species, therefore, cannot be attributed to Pagurus, but must be assigned to Para- pagurodes ...” The comment was prompt- ed by the fact that prior to their description of P. hartae, this taxon had been reported from California and Washington, USA, and British Columbia, Canada, as Pagurus sp. (McLaughlin & Haig 1973, Hart 1982, Jen- sen 1995). Regrettably, McLaughlin & Jen- sen’s (1996) remark has been interpreted by some carcinologists to mean that species with papillae and/or very short sexual tubes are automatically excluded from the genus Pagurus (e.g., Komai 1998, 1999). Accord- ing to the views of McLaughlin & Lemaitre (2001) and Lemaitre & McLaughlin (2003a, 2003b), the presence or absence of very short male sexual tubes and/or papillae should not be seen as the single cause to transfer species from genera to which they are otherwise morphologically attributable, or assign species to genera that they are not otherwise morphologically allied. McLaughlin & Jensen (1996) justified their generic assignment on the basis of morphological and larval similarities among the three species then assigned to Parapagurodes. However, they also pointed out, as McLaughlin & Haig (1973) had for P. laurentae, that P. hartae had superficial 43 resemblances to a few northeastern Pacific species of Pagurus. Upon the observation of very short sex- ual tubes in Pagurus gracilipes and P. nip- ponensis, Komai (1998) provisionally transferred these two species to Parapagu- rodes, while noting their close similarities to species of McLaughlin’s (1974) bernhar- dus group of Pagurus. Komai (1998) also pointed out that while Pagurus gracilipes and P. nipponensis shared the presence of a small right male sexual tube, both species differed substantially from Parapagurodes makarovi, P. laurentae, and P. hartae. In the subsequent, continuing study of Japanese species of Pagurus, Komai (1999) transferred Pagurus constans and P. imaii to Parapagurodes because he found very short male sexual tubes on both fifth coxae in the former species, and a single right tube in the latter. He also provided a minor emendation to the generic diagnosis by call- ing attention to the slight median indenta- tion, cleft or concavity sometimes seen in the gill lamellae, and to the occurrence of the left sexual tube, although he acknowl- edged that McLaughlin & Haig (1973) sim- ilarly had reported the rare occurrence of a left tube in P. makarovi. Unfortunately, Ko- mai’s (1999) emendation, like McLaughlin & Jensen’s (1996) brief generic diagnosis, failed to acknowledge the absence or re- duction in number of male pleopods and the usual absence of the fifth left female pleo- pod in the type species. In his review of the genus Catapagurus A. Milne-Edwards, 1880, Asakura (2001) redescribed Catapagurus doederleini and found it to be markedly divergent from all other species that had been assigned to Ca- tapagurus. Asakura transferred Doflein’s (1902) taxon to Parapagurodes stating that it agreed with all the diagnostic characters proposed by McLaughlin & Haig (1973) for their genus; however, it was primarily the presence of a very short right sexual tube that prompted his action. He quite correctly acknowledged the dimorphic second pereo- pods of P. doederleini, as well as the lack 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of corneous spines on the ventral margins of the second right and both third pereo- pods. As previously indicated, Lemaitre & McLaughlin (2003b) expressed the opinion that Parapagurodes, as presently constitut- ed, represented a polyphyletic taxon. To evaluate the merits of their conclusion, we have critically reviewed the descriptions of each of the assigned taxa. We have supple- mented these reviews by reexamining spec- imens of Parapagurodes laurentae and P. hartae in the first author’s personal collec- tions (PMcL). Additionally we have ex- amined representatives of P. constans, P. doederleini, P. gracilipes, P. imaii, and P. nipponensis from the collections of the Nat- ural History Museum and Institute, Chiba (CBM-ZC), the Hilgendorf collection from the Museum fiir Naturkunde, Berlin, Ger- many (ZMB), and specimens donated to one of the authors by Dr. M. Imafuku, Kyo- to University. From our reviews and ex- aminations, we present the comparative di- agnoses of the eight species we have used to determine the validity of the current ge- neric assignments. Animal size is indicated by shield length (sl) as measured from the tip of the rostrum to the midpoint of the posterior margin of the shield. Reported sexual tube length cor- responds to the criterion of McLaughlin (2003): very short (<1 coxal length), short (1-2 coxal lengths), moderate (>2-—5 coxal lengths). The reference by McLaughlin & Haig (1973) to the fourth pereopod being subchelate or not subchelate is interpreted here according to McLaughlin (1997) who recognized three conditions in the propodal- dactyl articulation of this appendage. McLaughlin & Haig’s (1973) “‘subchelate”’ is viewed by McLaughlin (1997) as being semichelate, whereas McLaughlin & Haig’s (1973) “‘not subchelate” is now considered to actually be subchelate. The abbreviation Ovig. indicates ovigerous female. Previous- ly published illustrations used in this man- uscript are of specimens in the collections of the Los Angeles Country Natural His- tory Museum (LACM) [transferred to that Museum from the Allan Hancock Founda- tion (AHP)], Los Angeles, California; the Royal British Columbia Provincial Museum (RBCPM), Victoria, British Columbia; and Zoologische Staatssammlung Miinchen (ZSSM), Munich. Review and Reexamination Parapagurodes makarovi McLaughlin & Haig, 1973 Figs. 1A, 2A, 3A, 4A, B, 5A Description by McLaughlin & Haig (1973:119—120, figs. 4a, 5—8). No supple- mental material available. Diagnosis.—Gill lamellae essentially biserial but with or without very weak dis- tal indentation or concavity. Rostrum acute- ly triangular. Maxillule with somewhat pro- duced endopodal external lobe, not re- curved. Right cheliped elongate, more so in large individuals; dorsal surface of palm distinctly convex; dorsal surface of carpus with row of spines mesiad of midline. Left cheliped elongate dorsal surface of palm convex, elevated in midline proximally. Ambulatory legs similar, somewhat later- ally compressed; dactyls as long or longer than propodi, slender, in dorsal view straight, each with row of corneous spines on ventral margin; carpi each with dorso- distal spine. Fourth pereopods usually sub- chelate, occasionally weakly semichelate; preungual process very small; propodal rasp with 1—3 irregular rows of corneous scales. Sternite of third pereopods (sixth thoracomere) semi- or subsemicircular. Sternite of fifth pereopods (eighth thora- comere) separated into two broad lobes by weak median depression; coxae of fifth pe- reopods symmetrical. Males with short sex- ual tube developed from coxa of right fifth pereopod; left gonopore sometimes with pa- pilla, occasionally with short tube. No paired pleopods in either sex. Males usually without, occasionally with weakly biramous left pleopods on pleomeres 3 and 4. Fe- males with weakly developed, biramous left VOLUME 117, NUMBER 1 45 Fig. 1. Coxae and sternite of fifth pereopods. A, B, Parapagurodes makarovi McLaughlin & Haig, 1973, 3 (sl = 3.5 mm), 6 (sl = 3.6 mm), LACM; C, D, P. laurentae McLaughlin & Haig, 1973, ¢ (sl = 3.4 mm), 3 (sl = 3.4 mm), LACM; E, P. hartae McLaughlin & Jensen, 1996, ¢ (sl = 3.2 mm), RBCPM 974-00368- 22; E P. gracilipes (Stimpson, 1858), ¢ (sl = 9.0 mm), CBM-ZC 2977; G, P. nipponensis (Yokoya, 1933), 3 (sl = 7.3 mm), CBM-ZC 1162; H, P. constans (Stimpson, 1858), d (sl = 8.7 mm), CBM-ZC 59; I, P. imaii (Yokoya, 1939), 5 (sl = 2.5 mm), CBM-ZC 2699; J, P. doederleini (Doflein, 1902), 3 (sl = 8.6 mm), ZSSM 274/1. A—D redrawn from McLaughlin & Haig (1973); E redrawn from McLaughlin & Jensen (1996); J from Asakura (2001). 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Endopod of maxillule. A, Parapagurodes makarovi McLaughlin & Haig, 1973, 6 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, 3 (sl = 3.4 mm), LACM; C, P. hartae McLaughlin & Jensen, 1996, ¢ (sl = 3.2 mm), RBCPM 974-00368-22; D, P. gracilipes (Stimpson, 1858), ¢ (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), ¢ (sl = 7.3 mm), CBM-ZC 1162; E P. constans (Stimpson, 1858); 3d (sl = 8.7 mm), CBM-ZC 59; G, P. imaii (Yokoya, 1939), ¢ (sl = 2.5 mm), CBM-ZC 2699; H, P. doederleini (Doflein, 1902), d (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from McLaughlin & Haig (1973); C redrawn from McLaughlin & Jensen (1996); H from Asakura (2001). pleopods on pleomeres 2—4, pleopod 5 ab- sent or rudimentary. Telson with posterior lobes separated by very small, shallow me- dian cleft; terminal margins straight or somewhat concave, each with several to nu- merous spinules or small to very small spines. Parapagurodes laurentae McLaughlin & Haig, 1973 Figs. 1B, 2B, 3B, 4C, D, 5B Description by McLaughlin & Haig (1973:129—134, figs. 4b, 9-11). Supplemental material examined.— U.S.A.: 3 6 (sl = 1.4-2.9 mm), 3 @ (sl = 1.7—2.9 mm), 2.5 mi SE Seal Rocks, Santa Catalina I, CA, 159-174 m, 25 Oct 1941, PMcL. Diagnosis.—Gill lamellae essentially biserial but with or without very weak dis- tal indentation or concavity. Rostrum acute- ly triangular. Maxillule with moderately well developed endopodal external lobe, not recurved. Right cheliped usually elon- gate, more so in large individuals; dorsal surface of palm convex; dorsal surface of VOLUME 117, NUMBER 1 carpus with row of spines mesiad of mid- line. Left cheliped moderately long, dorsal surface of palm convex. Ambulatory legs similar, somewhat laterally compressed, dactyls as long or longer than propodi, slen- der, in dorsal view usually straight, with row of corneous spines on ventral margin; carpi each with dorsodistal spine. Fourth pereopods usually semichelate, occasional- ly subchelate; preungual process very small; propodal rasp with 1-3 irregular rows of corneous scales. Sternite of third pereopods (sixth thoracomere) subsemicir- cular. Sternite of fifth pereopods (eighth thoracomere) separated into two broad lobes by weak to moderate median depres- sion; coxae of fifth pereopods symmetrical. Males with short or very short sexual tube developed from coxa of right fifth pereo- pod; left gonopore sometimes with papilla. No paired pleopods in either sex. Males usually with weakly biramous left pleopods on pleomeres 3 and 4, occasionally without unpaired pleopods. Females usually with moderately well developed, biramous ple- opods on pleomere 2—4; pleopod 5 rudi- mentary, rarely absent. Telson with poste- rior lobes separated by very shallow me- dian cleft; terminal margins concave or slightly oblique, each with row of very small spinules and 1—4 small spines at pos- terolateral angles. Parapagurodes hartae McLaughlin & Jensen, 1996 PigsmlC2C3C4E 5C Description by McLaughlin & Jensen (1996:844—847, figs. 1-4). Supplemental material examined.—Can- ada: 1 ¢ (sl = 1.1 mm), 1 2 (sl = 1.5 mm), Taylor Inlet, Barkley Sound, British Colum- bia, 10 m, 10 Jun 1994, PMcL. Diagnosis.—Gill lamellae essentially biserial but with or without very weak dis- tal indentation or concavity. Rostrum acute- ly triangular. Maxillule with moderately well developed endopodal external lobe, not recurved. Right cheliped elongate in 47 large males; dorsal surface of palm convex; dorsal surface of carpus with row of spines mesiad of midline. Left cheliped with dac- tyl and fixed finger short and broad in small males and females, longer in large males; dorsal surface of palm convex. Ambulatory legs similar; dactyls slightly shorter to slightly longer than propodi, moderately slender, laterally compressed, in dorsal view straight, with row of corneous spines on ventral margin; carpi each with dorsodistal spine and row of low, sometimes spinulose protuberances on dorsal surface, rarely 1 dorsoproximal spines (second pereopods). Fourth pereopods usually semichelate, oc- casionally subchelate; preungual process very small; propodal rasp with 2—4 irregu- lar rows of corneous scales. Sternite of third pereopods (sixth thoracomere) subsemicir- cular to subrectangular. Sternite of fifth pe- reopods (eighth thoracomere) separated into two broad lobes by weak to moderate me- dian depression; coxae of fifth pereopods symmetrical. Males often with very short sexual tube developed from coxa of right fifth pereopod; left gonopore without papil- la. No paired pleopods in either sex. Males with unequally biramous left pleopods on pleomeres 3—5. Females with moderately well developed, left biramous pleopods on pleomeres 2—4; pleopod 5 as in male. Tel- son with posterior lobes separated by shal- low, U-shaped median cleft; terminal mar- gins rounded or slightly oblique, each with row of very small spinules and lor 2 small spines at posterolateral angles. Parapagurodes gracilipes (Stimpson, 1858) Figs. 1D, 2D, 3D, 4E 5D Redescription by Komai (1998:268—275, figs. LA, 2—5, 7). Supplemental material examined.—Ja- pan: 2 ¢ (sl = 5.4, 9.0 mm), 1 2 (sl = 6.7 mm), off Choshi, Chiba, 10—20 m, 3 Sep 1996, CBM-ZC 2977. Diagnosis.—Gill lamellae biserial. Ros- trum acutely triangular. Maxillule with 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 117, NUMBER 1 somewhat produced endopodal external lobe, slightly to distinctly recurved. Right cheliped moderately (small specimens) to considerably elongate in large individuals; dorsal surface of palm weakly convex but with dorsomesial portion somewhat elevat- ed; dorsal surface of carpus with row of spines mesiad of midline. Left cheliped with dorsal surface of palm somewhat flat- tened, dorsomesial and dorsolateral margins slightly elevated. Ambulatory legs similar; dactyls longer than propodi, strongly twist- ed; moderately broad, each with row of nu- Merous corneous spines on ventral margin; carpi each with single or double row of multifid spines. Fourth pereopods semiche- late; no preungual process; propodal rasp with several rows of corneous scales. Ster- nite of third pereopods (sixth thoracomere) subquadrate, weakly skewed, sulcate me- dially. Sternite of fifth pereopods (eighth thoracomere) separated into two subovate lobes by shallow median groove; coxae of fifth pereopods symmetrical. Males with very short sexual tube developed from coxa of right fifth pereopod; left gonopore with- out tube or papilla. No paired pleopods in either sex. Males with unequally biramous pleopods on pleomeres 3—5. Females with well developed, biramous pleopods on pleomeres 2—4; pleopod 5 with endopod noticeably reduced. Telson with posterior lobes separated by very small, or indistinct median cleft; terminal margins nearly hor- izontal, each with eight to ten small spines and two or three larger spines at postero- lateral angles, lateral margins occasionally with spinules. 49 Parapagurodes nipponensis (Yokoya, 1933) Figs. 1E, 2E, 3E, 4G, 5E Redescribed by Komai (1998:275—279, figs 1B, 6, 7) only as similar to P. gracili- pes with certain noted differences. Supplemental material examined.—Ja- pan: 4 6 (sl = 6.3-7.6 mm), 1 2 (sl = 5.3 mm), Kumano Nada, 50 m, Sep 1981, PMcL; 2 ¢ (sl = 8.0, 9.2 mm), off Kashi- ma, Irakaki, 65 m, 24 Apr 1991, CBM-ZC 50; 1 ¢ (sl = 7.3 mm), off Kii Minabe, Kii Peninsula, 80—100 M, 24 Mar 1995, CBM- ZC 1162. Diagnosis.—Gill lamellae biserial. Ros- trum acutely triangular. Maxillule with somewhat produced external lobe, slightly to distinctly recurved. Right cheliped mod- erately (small specimens) to considerably elongate in large individuals; dorsal surface of palm weakly convex but with dorsome- sial marginal area somewhat elevated; dor- sal surface of carpus with row of spines me- siad of midline. Left cheliped with dorsal surface of palm somewhat flattened, dor- somesial and dorsolateral margins slightly elevated. Ambulatory legs similar; dactyls longer than propodi, strongly twisted; mod- erately slender to moderately broad; each with prominent longitudinal sulcus on lat- eral face and row of numerous very tiny corneous spines on ventral margin; carpi each with single or double row of multifid spines. Fourth pereopods semichelate; no preungual process; propodal rasp with sev- eral rows of corneous scales. Sternite of third pereopods (sixth thoracomere) sub- < Fig. 3. Ambulatory dactyls. A—H, dactyl of left third pereopod (A-C lateral view, D-H, mesial view); I, dactyl of left second pereopod (mesial view); J, dactyl of right second pereopod (mesial view). A, Parapagurodes makarovi McLaughlin & Haig, 1973, 6 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, 3 (sl = 3.2 mm), LACM; C, P. hartae McLaughlin & Jensen, 1996, 3 (sl = 2.8 mm), RBCPM 974-00368-22; D, P. gracilipes (Stimpson, 1858), d (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), 3 (sl = 7.3 mm), CBM-ZC 1162; E P. constans (Stimpson, 1858), ¢ (sl = 8.7 mm), CBM-ZC 59; G, H, P. imaii (Yokoya, 1939), 5 (sl = 2.5 mm), CBM-ZC 2699, ovig. 2 (sl = 1.6 mm), CBM-ZC 1911; I, J, P. doederleini (Dofiein, 1902), 5 (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from McLaughlin & Haig (1973); C redrawn from McLaughlin & Jensen (1996); I, J from Asakura (2001). 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Dactyl and propodus of left fourth pereopod (lateral view). A, Parapagurodes makarovi McLaughlin & Haig, 1973, 3 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, ¢ (sl = 3.4 mm), LACM; C, P. hartae McLaughlin & Jensen, 1996, d (sl = 3.2 mm), RBCPM 974-00368-22; D, P. gracilipes (Stimpson, 1858), 5 (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), 3d (sl = 7.3 mm), CBM-ZC 1162; EP. constans (Stimpson, 1858), ¢ (sl = 8.7 mm), CBM-ZC 59; G, P. imaii (Yokoya, 1939), 3 (sl = 2.5 mm), CBM-ZC 2699; H, P. doederleini (Doflein, 1902), 3 (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from Mc- Laughlin & Haig (1973); C redrawn from McLaughlin & Jensen (1996); H redrawn from Asakura (2001). quadrate to subrectangular. Sternite of fifth pereopods (eighth thoracomere) separated into two subovate lobes by shallow median groove; coxae of fifth pereopods symmet- rical. Males with very short sexual tube de- veloped from coxa of right fifth pereopod; left gonopore without tube or papilla. No paired pleopods in either sex. Males with unequally biramous pleopods on pleomeres 3-5. Females with well developed, bira- mous pleopods on pleomeres 2—4; pleopod 5 with endopod noticeably reduced. Telson with posterior lobes separated by very small median cleft; terminal margins oblique, each with 8 or 9 small spines and 1 larger spine at posterolateral angles. Parapagurodes constans (Stimpson, 1858) Figs. 1E 2E 3K 4H, 5E G Redescription by Komai (1999:80-88, figs. 1-4). Supplemental material examined.—Ja- pan: Hilgendorf collection, 1 ovig. 2 (sl = 5.5 mm), ZMB 8650; 1 6 (sl = 11.1 mm), 1 ovig. 2 (sl = 10.2 mm), Sagami Bay, ZMB 17800; 1 3 (sl = 8.7 mm), off Tone River mouth, Choshi, Chiba, 60 m, 21 Oct 1991, CBM-ZC 59; 2 3 (sl = 6.3, 10.7 mm), Hakodate Bay, 10—20 m, 17 Mar 1995, CBM-ZC 2362. Diagnosis.—Gill lamellae biserial. Ros- trum triangular. Maxillule with moderately well developed external lobe, not recurved. Right cheliped somewhat suboval in dorsal view; dorsal surface of palm weakly con- vex; dorsal surface of carpus with scattered spines mesiad of midline. Left cheliped with dorsal surface of palm weakly convex. Ambulatory legs similar; dactyls slightly longer than propodi, moderately slender, laterally compressed, in dorsal view slightly to prominently twisted, with longitudinal VOLUME 117, NUMBER 1 sulcus on lateral face and row of corneous spines on ventral margin; carpi each with dorsodistal spine and row of low protuber- ances on dorsal surface. Fourth pereopods semichelate, preungual process apparently absent; propodal rasp with several rows of corneous scales. Sternite of third pereopods (sixth thoracomere) subrectangular. Sternite of fifth pereopods (eighth thoracomere) sep- arated into two somewhat flattened, round- ed lobes by shallow median depression; coxae of fifth pereopods symmetrical. Males with papilla or very short sexual tube developed from coxa of both right and left fifth pereopods. No paired pleopods in ei- ther sex. Males with unequally biramous left pleopods on pleomeres 3—5. Females with moderately well developed, biramous pleopods left on pleomeres 2—4; pleopod 5 reduced. Telson with posterior lobes sepa- rated by shallow median cleft; terminal margins broadly rounded, each unarmed or with few very small spinules adjacent to cleft. Parapagurodes imaii (Yokoya, 1939) Figs. 1G, 2G, H, 3G, 41, 5H Redescription by Komai (1994:33-38, figs. 1-3). Supplemental material examined.—Ja- pan: 1 ¢ (sl = 2.1 mm), 1 ovig. @ (sl = 1.6 mm), Funakoshi Bay, Iwate, Sanriku, 66 m, 25 May 1995, CM-ZC 1911; 1 ¢ (al = 2.5 mm), off Takeoka, Boso Peninsula, ca 80 m, 2 Mar 1995, CBM-ZC 2699. Diagnosis.—Gill lamellae biserial, but with slight terminal concavity, cleft or de- pression. Rostrum triangular. Maxillule with moderately well developed external lobe, not recurved. Right cheliped elongate in large males, dorsal surface of palm con- vex; dorsal surface of carpus with two lon- gitudinal rows of spines. Left cheliped with dorsal surface of palm elevated in midline. Ambulatory legs somewhat dissimilar, third sexually dimorphic; dactyls of second and third right slightly shorter to slightly longer than propodi, moderately slender, laterally 51 compressed, in dorsal view barely twisted, with row of corneous spines on ventral mar- gin, third left of females broadened, pro- podus with prominent ventral spine; carpi each with dorsodistal spine and row of low, sometimes spinulose protuberances on dor- sal surface. Fourth pereopods semichelate; preungual process absent; propodal rasp with 2 or 3 rows of corneous scales. Ster- nite of third pereopods (sixth thoracomere) subcircular to subovate, slightly skewed. Sternite of fifth pereopods (eighth thora- comere) separated into two somewhat flat- tened, rounded lobes by shallow median de- pression; coxae of fifth pereopods symmet- rical. Males with very short sexual tube de- veloped from coxa of both right and left fifth pereopods. No paired pleopods in ei- ther sex. Males with unequally biramous left pleopods on pleomeres 3—5. Females with moderately well developed, biramous left pleopods on pleomeres 2—4; pleopod 5 reduced. Telson with posterior lobes sepa- rated by shallow median cleft; terminal margins oblique, each with 3 or 4 moderate to strong spines. Parapagurodes doederleini (Doflein, 1902) Figs. 1H, 21, J, 3H, 4J, 5I Redescription by Asakura (2001:885— 888, figs. 45-47). Supplemental material examined.—Ja- pan: 2 6 (sl = 8.1, 9.2 mm), off Kochi, Tosa Bay, 190 m, 10 Aug 1991, CBM-ZC 184. Taiwan: 1 3 (sl = 9.3 mm), 1 @ (sl = 7.8 mm), Su-Aou, 100—200 m, 6 Aug 1996, CBM-ZC 2922. Diagnosis.—Gill lamellae biserial. Ros- trum triangular. Maxillule with moderately well developed endopodal external lobe, not recurved. Right cheliped stout, dorsal surface of palm slightly convex; dorsal sur- face of carpus with covering of spines and spinulose tubercles. Left cheliped with dor- sal surface of palm very slightly elevated in midline. Ambulatory legs dissimilar, dac- 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Telson (A-E H, I, dorsal view, G, ventral view of posterior portion). A, Parapagurodes makarovi McLaughlin & Haig, 1973 6 (sl = 3.5 mm), LACM; B, P. laurentae McLaughlin & Haig, 1973, d (sl = 3.4 mm), LACM; C, P. hartae McLaughlin & Jensen, 1996, 3 (sl = 3.2 mm), RBCPM 974-00368-22; D, P. gracilipes (Stimpson, 1858), d (sl = 9.0 mm), CBM-ZC 2977; E, P. nipponensis (Yokoya, 1933), 3 (sl = 7.3 mm), CBM-ZC 1162; E G, P. constans (Stimpson, 1858); ¢ (sl = 8.7 mm), CBM-ZC 59; H, P. imaii (Yokoya, 1939), 3 (sl = 2.5 mm), CBM-ZC 2699; I, P. doederleini (Doflein, 1902), 5 (sl = 8.6 mm), ZSSM 274/1. A, B redrawn from McLaughlin & Haig (1973); C redrawn from McLaughlin & Jensen (1996); I from Asakura (2001). tyls longer than propodi, strongly twisted, second left with row of 40—60 well devel- oped, comb-like corneous spines on ventral margin; second right and third each with longitudinal row of short transverse rows of setae; carpi each with row of spines on dor- sal surface. Fourth pereopods subchelate; preungual process absent; propodal rasp with 3 or 4 rows of corneous scales. Ster- nite of third pereopods (sixth thoracomere) rectangular. Sternite of fifth pereopods (eighth thoracomere) as narrow rod with pair of rounded lobes anteriorly; coxae of fifth pereopods asymmetrical. Males with very short sexual tube developed from coxa of right fifth pereopod, left sometimes with papilla. No paired pleopods in either sex. Males with unequally biramous left pleo- pods on pleomeres 3—5. Females with mod- erately well developed, biramous left pleo- pods on pleomeres 2—4; pleopod 5 reduced. Telson with posterior lobes separated by VOLUME 117, NUMBER 1 broad, deep median concavity; terminal margins oblique, each with 1—5 moderate to strong corneous spines. Results In her discussion of significant generic characters, de Saint Laurent-Dechancé (1966) listed three that are pertinent to our investigation: sexual tube development, de- velopment of the external lobe of the max- illulary endopod, and pleopod number and development. Perusal of the abbreviated di- agnoses of the eight species currently as- signed to Parapagurodes shows that attri- butes of these three characters are not uni- versally shared. While sexual tube length (Fig. 1) varies from very short to short in P. makarovi (Figs. 1A, B) and P. laurentae (Fig. 1C, D) only very short tubes develop in the other six species (Figs. 1E—J), and occasionally are not apparent at all. However, recent studies have shown that sexual tube devel- opment is known to vary within genera (e.g., McLaughlin 1997, 2003; McLaughlin & Lemaitre 2001; Lemaitre & McLaughlin 2003a, 2003b). Nevertheless, P. doederleini is more importantly distinguished from the other seven species because in addition to the very short right sexual tube, the coxae of the male fifth pereopods are asymmetri- cal (Fig. 1J). The external lobe of the maxillulary en- dopod (Fig. 2) is moderately well devel- oped in all eight species, but is slightly to distinctly recurved only in P. gracilipes and P. nipponensis (Figs. 2D, E). Parapagurodes was initially character- ized as having unpaired male pleopods varying from reduced on pleomeres 3—5 to completely absent, and female unpaired pleopods often being reduced on pleomeres 2—4 and absent on pleomere 5. All subse- quently assigned taxa are described as hay- ing at least moderately well developed un- paired, unequally biramous pleopods on male pleomeres 3—5 and on female pleo- meres 2—5. 53 Several other characters frequently in- cluded in generic diagnoses also have been examined. Rostral development, for exam- ple is generally similar among species with- in a single genus. All eight species have an acutely developed rostrum, but then so do many species assigned to other genera. With the exception of P. constans, all of the species under consideration herein are described as having an elongate right che- liped, at least in large males. In P. hartae and P. imaii this elongation is considered a sexually dimorphic character (McLaughlin & Jensen 1996, Komai 1999), whereas in P. gracilipes and P. nipponensis apparently the elongation is growth related (Komai 1998). Similar lengthening of the left che- liped is reported for these species. In con- trast, the chelipeds are typically elongate re- gardless of sex or size in P. makarovi, P. laurentae and P. doederleini. That cheliped elongation is comparable among the eight taxa is doubtful. Major differences among the eight spe- cies can be observed in the shape and ar- mature of the dactyls of the ambulatory legs (Fig. 3). In P. makarovi and P. laurentae the dactyls (Figs. 3A, B) are moderately long, slender, laterally compressed, and in dorsal view appear straight; the dorsal sur- faces of the carpi are armed only with a dorsodistal spine. The dactyls are similarly straight in P. hartae (Fig. 3C), but vary in length from shorter to only slightly longer than the propodi; the carpi each have a row of low protuberances on the dorsal surface in addition to the dorsodistal spine. The dactyls of P. gracilipes and P. nipponensis (Figs. 3D, E), although moderately long and laterally compressed, are moderate to broad and strongly twisted, the ventral mar- gins of each are provided with a row of numerous small corneous spines; the dorsal surfaces of the carpi are provided with one or more rows of small spines. In contrast, while the dactyls of P. constans (Fig. 3F) are longer than the propodi and slightly to noticeably twisted, the ventral margins each are armed with fewer and much larger cor- 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON neous spines; each carpus is armed only with a row of low protuberances in addition to the dorsodistal spine. The dactyls of P. imaii and P. doederleini are dimorphic, but do not represent comparable conditions. As reported by Komai (1999), the third left dactyl and propodus of females of P. imaii differ from those of males. In males the dactyl and propodus of the third left (Fig. 3G) are moderately long and slender as they are on the second and third right. The fe- male dactyl (Fig. 3H) is broad and promi- nently flattened; a well developed calcare- ous spine is present on the ventrodistal mar- gin of the propodus. The dimorphism in P. doederleini involves the dactyls of the sec- ond pereopods. The left is provided with a ventral row of closely-spaced, corneous spines that present a comb-like appearance (Fig. 31); the right, and the dactyls of the third pereopods completely lack spines, and instead are provided with short transverse rows of setae over the entire length of the mesial faces (Fig. 3J). The shape of the anterior lobe of the ster- nite of the third pereopods and the config- uration of the sternite of the fifth pereopods have been proposed as generic or at least group characters (e.g., McLaughlin 1981, 2003; Lemaitre et al. 1982). The anterior lobe of the sternite of the third pereopods is subsemicircular in P. makarovi, P. lau- rentae, and P. hartae, semicircular or su- bovate in P. imaii, but subquadrate to sub- rectangular in P. gracilipes and P. nippo- nensis and subrectangular in P. constans and P. doederleini. The sternites of the fifth pereopods are less clearly definable in these eight taxa. The fourth pereopods (Fig. 4) are sub- chelate or only very weakly semichelate in P. makarovi and P. laurentae and P. doe- derleini, but semichelate in the remaining species. The number of rows of corneous scales making up the propodal rasps of these appendages exhibit overlapping intra- specific variation in all eight species. The telsons of P. makarovi and P. lau- rentae (Figs. 5A, B) have straight to weakly concave or very slightly oblique terminal margins that are armed with small spines or spinules. Similar conformation and arma- ture are seen in P. gracilipes (Fig. 5D) and to a lesser extent in P. nipponensis (Fig. SE). In contrast, the terminal margins of the telsons of P. hartae (Fig. 5C) and P. con- stans (Fig. 5K G) are broadly rounded and unarmed or only weakly armed. The telson of P. imaii (Fig. 5H) differs in having dis- tinctly oblique terminal margins, each armed with prominent spines, and the telson of P. doederleini (Fig. 51) is plainly differ- ent from the other seven. Conclusions From the evidence presented, there can be little doubt that Parapagurodes, as pres- ently constituted, represents a heteroge- neous collection of taxa. Consequently, we restrict Parapagurodes to the two species initially assigned, P. makarovi and P. lau- rentae. Parapagurodes hartae is herein transferred to Pagurus and the four species formerly included in Pagurus are returned to it. We concur with Komai (1998) that P. gracilipes and P. nipponensis are closely al- lied to the bernhardus group of Pagurus, and undoubtedly should be included in that group. We do not advocate separating the bernhardus group from the admittedly polyphyletic Pagurus at this time, as Pa- gurus bernhardus (Linnaeus, 1758) is the type species of the genus. To remove P. bernhardus and its allied species would leave the remaining 80 or so species with- out generic union. Consequently, until such time as all species currently assigned to Pa- gurus have been thoroughly recognized and defined, this genus necessarily must remain a “‘catch-all’’. In contrast, there is ample justification to establish a new genus for the very distinctive P. doederleini as is done herein. Dofleinia gen. nov. Catapagurus: Doflein 1902:624 (in part).— Miyake 1978:78 (key, in part), 141 (Gn VOLUME 117, NUMBER 1 part); 1982:232 (key, in part); 1991:232 (key, in part); 1999:232 (key, in part). Parapagurodes: Asakura 2001:885 (in part). Diagnosis.—Gills biserial; 11 pairs. Ros- trum well developed, acute. Antennal pe- duncles with supernumerary segmentation. Maxillule with external lobe of endopod moderately well developed, not recurved. Third maxilliped with well developed crista dentata, 1 accessory tooth. Sternite of third maxillipeds unarmed. Chelipeds subequal, right stronger but not necessarily longer. Second pereopods dimorphic, left with row of closely-spaced comb-like corneous teeth on ventral margin, right with ventral margin unarmed. Third pereopods similar; sternite with subrectangular anterior lobe. Fourth pereopods subchelate; dactyl with well de- veloped preungual process; propodal rasp consisting of 3 or 4 rows of corneous scales. Fifth pereopods chelate; coxae of males asymmetrical. Males with very short sexual tube developed from right gonopore, papilla frequently produced from left. Abdomen well developed, twisted; colu- mellar muscle usually prominent. Males without paired first or second pleopods; with unequally biramous unpaired left ple- opods 3—5. Females without paired first ple- opods, with subequally biramous, unpaired, left pleopods 2—4, pleopod 5 as in male. Uropods asymmetrical. Telson with distinct lateral indentations; posterior lobes separat- ed by very broad median cleft. Type species.—Catapagurus doederleini Doflein, 1902. Etymology.—Named after E Doflein who first described the type species; gender fem- inine. Acknowledgements The authors acknowledge, with thanks, the gift of specimens to the first author by Dr. M. Imafuku, Kyoto University, and the loan of specimens by Dr. C. O. Coleman, Naturhistorisches Forschungsinstitut Muse- um ftir Naturkunde zu Berlin. This work 55 has been supported in part by a Grant-in- Aid for Scientific Research (C) from the Ministry of Education, Science, Culture and Sports of Japan to Akira Asakura (No. 14540654). This, in part, is also a scientific contribution from the Shannon Point Ma- rine Center, Western Washington Universi- ty. Literature Cited Asakura, A. 2001. A revision of the hermit crabs of the genera Catapagurus A. Milne-Edwards and Hemipagurus Smith from the Indo-West Pacific (Crustacea: Decapoda: Anomura: Paguridae).— Invertebrate Taxonomy 15:823-891. Doflein, FE 1902. Ostasiatische Dekapoden.—Abhan- dlungen der Kgl. Bayerischen Akademie der Wissenschaften Math.-Phys. Klassen. 21:613— 670. Hart, J. EF L. 1982. Crabs and their relatives of British Columbia.—British Columbia Provincial Mu- seum Handbook 40:1—266. Henderson, J. R. 1888. Report on the Anomura col- lected by H.M.S. Challenger during the years 1873-76. Scientific Results of the Exploratory Voyage of HMS Challenger, (Zoology) 27:1— 221. Her Majesty’s Stationary Office, Edin- burgh etc. Jensen, G. C. 1995. Pacific coast crabs and shrimps. viii + 87 pp. Sea Challengers, Monterey, CA. Komai, T. 1994. 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Addendum and taxonomic summary.— Proceedings of the Biological Society of Wash- ington 116:464—486. 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON » & . 2003b. New species of the genus Goreopagurus (Decapoda: Anomura: Paguri- dae) from Tasmania and reevaluation of sexual tubes in hermit crab systematics.—Memoirs of Museum Victoria 60:221—227. Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species Cum Characteribus, Differentiis, Syn- onymis, Locis, edition 10. 1, pp. 1-111, 1-824. Holmiae. McLaughlin, P. A. 1974. The hermit crabs (Crustacea Decapoda, Paguridea) of northwestern North America.—Zoologische Verhandelingen 130:1— 396. . 1981. Revision of Pylopagurus and Tomo- Pagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species: part I. Ten new genera of the Paguridae and a redescription of Tomopagurus A. Milne-Ed- wards and Bouvier.—Bulletin of Marine Sci- ence 31:1—30. . 1997. Crustacea Decapoda: hermit crabs of the family Paguridae from the KARUBAR cruise in Indonesia. Jn A. Crosnier & P. Bouch- et, eds., Résultats des Campagnes MUSOR- STOM, 16.—Mémoires du Muséum national d’ Histoire naturelle 172:433-572. . 2003. Illustrated keys to the families and gen- era of the superfamily Paguroidea (Crustacea: Decapoda; Anomura), with supplemental diag- noses of the genera of the Paguridae—Memoirs of Museum Victoria 60:111—144. , & J. Haig. 1973. On the status of Pagurus mertensii Brandt, with descriptions of a new ge- nus and two new species from California (Crus- tacea: Decapoda: Paguridae):—Bulletin of the Southern California Academy of Sciences 72: 113-136. , & G. S. Jensen. 1996. A new hermit crab species of the genus Parapagurodes from the eastern Pacific, with a description of its first zoeal stage.—Journal of Natural History 30: 841-854. , & R. Lemaitre. 2001. Revision of Pylopagu- rus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species, part VI. Pylopagurus Milne-Edwards and Bouvier, Haigia McLaughlin, and Pylopa- guridium new genus.—Proceedings of the Bio- logical Society of Washington 114:444—483. Milne-Edwards, A. 1880. Report on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico, and in the Ca- ribbean Sea, 1877, 78, 79, by the United States Coast Survey steamer “Blake”, Lieut.-Com- mander C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. VIII. Etudes préliminaires sur les Crustacés.—Bulletin of the Museum of Comparative Zoology, Harvard College, 8(1):1—68. Miyake, S. 1978. The crustacean Anomura of Sagami Bay: 1—200 (English), 1-161 (Japanese), Bio- logical Laboratory, Imperial Household, Tokyo. . 1982. Japanese crustacean decapods and sto- matopods in color, vol. 1. 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Koenemann. 2003. Developmental genetics and arthropod evolution: on body re- gions of crustaceans——Crustacean Issues 15: 75-92. Stimpson, W. 1858. Prodromus descriptionis animal- ium evertebratorum, quae in expeditione ad Oceanum Pacificum Septentrionalem, a Repub- lica Federate missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et des- cripsit. VII. [Preprint (December 1858) from] Proceedings of the Academy of Natural Scienc- es of Philadelphia 1858 [1859]:225—252. Yokoya, Y. 1933. On the distribution of decapod Crus- tacea inhabiting the continental shelf around Ja- pan, chiefly based upon the materials collected by S.S. “Soyo Maru” during the years 1923— 1930.—Journal of the College of Agriculture Tokyo Imperial University 12:1—236. 1939. Macrura and Anomura of decapod Crustacea found in the neighbourhood of Ona- gawa, Miyagi-ken.—Scientific Reports of To- hoku University 14:261—289. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):57—67. 2004. Pseudopaguristes bicolor, a new species of hermit crab (Crustacea: Decapoda: Diogenidae) from Japan, the third species of the genus Akira Asakura and Takeharu Kosuge (AA) Natural History Museum and Institute, Chiba. 955-2, Aoba-cho, Chuo-ku, Chiba 260-8682, Japan, asakura@chiba-muse.or.jp (TK) Ishigaki Tropical Station, Seikai National Fisheries research Institute, 148-446, Fukai Ota, Ishigaki, Okinawa, Japan Abstract.—Pseudopaguristes bicolor, a new species of the recently estab- lished diogenid genus Pseudopaguristes McLaughlin, is described and illus- trated from Okinawa, Japan. This is the third species assigned to this genus. The recently established diogenid genus Pseudopaguristes McLaughlin, 2002, is characterized by eight functional gills, male chelipeds with the right larger than the left and dissimilar in armature, female chelipeds similar from left to right, fourth pereopods with a clump of long capsulate setae on the carpi, and the paired first and second ple- opods modified as gonopods. The type spe- cies, P. janetkae McLaughlin, 2002, was re- corded from Guam, the Mariana Islands. A second species, P. bollandi Asakura & McLaughlin, 2003, was recorded from Oki- nawa, tropical Japan. The present authors recently found the third species of this ge- nus, again from Okinawa. The new species is very easily separated from both P. janet- kae and P. bollandi by its characteristic col- oration and morphology of the male cheli- peds. The holotype is deposited in the Natural History Museum and Institute, Chiba (CBM-ZC). The terminology used follows McLaughlin (1974, 2002) with the excep- tion of the fourth pereopods as defined by McLaughlin (1997), gill structure by McLaughlin & de Saint Laurent (1998), and the posterior carapace by McLaughlin (2000). Abbreviations used are; coll., col- lector; and SL, shield length as measured from the tip of the rostrum to the posterior margin of the shield. Pseudopaguristes bicolor, new species Figs. 1-8 Material.—Holotype: male, SL = 2.65 mm, 78 m, 24°25.5'’N, 124°03.3’E, off Yar- abu-zaki, Ishigaki-jima Island, Okinawa, 21 Nov. 2002, coll. T. Kosuge, CBM-ZC 6759. Description.—Eight functional pairs of quadriserial, phyllobranchiate gills (Fig. 1A). Shield (Fig. 1B) 1.30 times longer than broad; anterior margin between ros- trum and lateral projections concave; lateral projections triangular, with strong submar- ginal spine; anterolateral angles each with strong corneous spine; lateral margins con- vex; posterior margin truncate; dorsal sur- face slightly convex, with elevated area pre- sent on each anterolateral portion; scattered tufts of short setae. Rostrum (Fig. 1B) prominent, triangular, reaching nearly to apices of ocular acicles, with terminal spine. Posterior carapace lateral elements (Fig. 1B) small, well calcified, unarmed. Branchiostegites (Fig. 1C) each with row of spines on dorsal margin anteriorly. Ocular peduncles (Fig. 1B) moderately long, 0.75 length of shield. Corneas (Fig. 1B, C) very slightly dilated. Ocular acicles (Fig. 1B) each terminating in strong, bifid cormeous spines; separated basally by more than breadth of rostrum. Antennular peduncles (Fig. 1D) stout, with few setae on each segment; when fully 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SA SS YQ \ Fig. 1. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. A, arthrobranch gill lamella; B, shield and cephalic appendages, dorsal; C, distal half of cephalothorax and cephalic appendages, right, lateral; D, right antennule, lateral; E, right antennal peduncle, ventral; H, right antennal flagellum. Scales equal 0.5 mm (A) and 1 mm (B-F). extended, distal margins of ultimate seg- ments reaching distal margins of corneas; ultimate segments unarmed; penultimate segments with ventral margins each bearing acute spine; basal segments with ventrod- istal angles each bearing acute spine and dorsolateral margins each bearing acute subdistal spine. Antennal peduncles (Fig. 1B, C, E) mod- erately long, when fully extended, reaching VOLUME 117, NUMBER 1 Fig. 2. 59 Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. Right mouthparts: A, mandible, internal; B, maxillule, external; C, same, endopod; D, maxilla, external; E, first maxilliped, internal; E second maxilliped, external; G, third maxilliped, external; H, same, internal. distal 0.30 of ocular peduncles, scarcely se- tose; fifth segments with dorsal margins each bearing acute subproximal spine; fourth segments with dorsodistal margins each bearing acute spine and ventrodistal margins each bearing another acute spine; third segments with prominent spine at ven- trodistal margin; second segments with dor- solateral distal angles produced, terminating in prominent bifid spine, dorsomesial distal angles each with acute corneous spine; first segment unarmed. Antennal acicles mod- erately long, straight; dorsomesial margins each with 3 (right) or 4 (left) spines; dor- solateral margins each with 2 strong spines; distal margins each with 2 strong spines. Antennal flagella (Fig. 1F) consisting of about 18 articles, each article with several short setae. Mandible (Fig. 2A) without distinguish- 60 Fig. 3. i PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. Right cheliped: A, dorsal; B, mesial; C, lateral. ing characters. Maxillule (Fig. 2B, C) with external lobe of endopod well developed, articulated, and recurved; internal lobe with 2 bristles. Maxilla (Fig. 2D) with moder- ately narrow scaphognathite. First maxilli- ped (Fig. 2E) with well developed, setose epipod. Second maxilliped (Fig. 2F) with- out distinguishing characters. Third maxil- liped (Fig. 2G, H) with carpus bearing dor- sodistal spine; merus with dorsodistal spine, ventral margin bearing 4 spines; ischium with strong ventrodistal spine, crista dentata well-developed, no accessory tooth; basis with 2 sharp spines. Chelipeds subequal; right (Fig. 3) larger than left. Dactyl as long as palm; terminat- ing in broad corneous claw; dorsal face flat, with scattered large tubercles; cutting edge with several calcareous teeth. Fixed finger terminating in corneous claw; dorsal face VOLUME 117, NUMBER 1 61 Fig. 4. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- Zaki, Ishigaki-jima Is., Okinawa. Left cheliped: A, dorsal; B, mesial; C, lateral. flat, with scattered large tubercles; cutting edge with several calcareous teeth. Palm 1.07 length of carpus; dorsal surface flat, with scattered large tubercles; dorsomesial margin with row of very strong spines; dor- solateral margin of palm and fixed finger with row of strong spines. Carpus 0.50 length of merus; dorsal face with scattered large tubercles, dorsolateral margin with row of strong conical-shaped spines, dor- somesial margin with row of very strong spines. Merus with dorsal face bearing 2 distal spines, subdistal transverse row of several spines, and row of spines on re- mainder of dorsal margin, tips semitrans- parent; ventromesial margin with 3 widely- separated strong spines, tips semitranspar- ent, ventrolateral margin with row of spines or tubercles. Ischium unarmed. Coxa with acute spine ventromesially. Left cheliped (Fig. 4) slenderer than right. Dactyl with dorsal face without tu- 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. Second left pereopod: A, lateral; B, propodus, ventral; C, dactyl, propodus, and carpus, mesial; D, merus, mesial. bercles; number of tubercles or spines on dorsal faces of palm and carpus fewer; mer- us with ventromesial and ventrolateral mar- gins bearing 6 and 5 spines, respectively; other surfaces similar to right. Second pereopods (Fig. 5) with armature similar from left to right; right 1.10 length of left. Basically, spines on ambulatory pe- reopods with semitransparent tips. Dactyls 1.10 (eft) or 1.25 (ight) length of propodi, each terminating in strong corneous claw; dorsal margins each with row of strong spines; ventral margins each with row of 9 strong corneous spines and, on left, accom- panied with 2 tiny corneous spines mesial- ly. Propodi 1.60 (left) or 1.55 (right) length of carpi, each with row of 10 strong spines on dorsal margin; ventral faces each with 2 VOLUME 117, NUMBER 1 E 0.5 mm ee 63 |) Imm Fig. 6. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. Third left pereopod: A, lateral; B, dactyl and propodus, mesial (propodus slightly ventral view); C, carpus, merus and ischium, lateral. Fourth left pereopod: D, dactyl, propodus and carpus, lateral; E, ventral setae of carpus. irregular rows of widely-separated, tiny corneous spines, ventromesial distal mar- gins with | (right) or 2 (left) acute corneous spines. Carpi 0.55 (left) or 0.60 (right) length of meri, each with strong, corneous or corneous-tipped, slender spine at dorso- distal angle and row of 5 slender spines on dorsal face mesially. Meri with ventral mar- gins each with row of slender spines and, on left, accompanied with 2 small spines mesially; dorsal margins each with row of spines. Ischia each with few, slender cor- 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. A, right fifth pereopod. Left first pleopod: B, external; C, internal; D, distal portion, internal, enlarged. Left second pleopod: E, external; FE distal portion, enlarged, external; G, same, mesial. H, third pleopod. I, telson. Scales equal 1 mm (A, H, I) and 0.2 mm (B-E). VOLUME 117, NUMBER 1 A Fig. 8. 65 Pseudopaguristes bicolor, new species: holotype male (CBM-ZC 6759), SL = 2.65 mm, off Yarabu- zaki, Ishigaki-jima Is., Okinawa. A, dorsolateral view; B, right cheliped, mesial; C, left second pereopod, lateral; D, same, mesial; E, left third pereopod, lateral; K same, mesial. Photo by A. Asakura. neous-tipped spines dorsally and small spine at ventromesial distal angle. Coxae unarmed. Third pereopods (Figs. 6A—C) with ar- mature similar from left to mnght, nght 1.05 length of left. Dactyls 1.20 (left) or 1.25 (right) length of propodi, each terminating in strong corneous claw; mesial faces each with row of small corneous spines ventrally and 4 (left) or 2(right) small spines dorsal- ly; dorsal margins with few tiny spines on proximal 0.25; ventral margins each with row of 9 strong corneous spines. Propodi 1.60 length of carpi; dorsal faces unarmed (left) or row of small tubercles or spines (right); ventral faces each with row of 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON small, widely-separated corneous spines, ventromesial distal angles each with 1 (left) or 2 (right) acute corneous spines. Carpi 0.70 (eft) or 0.80 (right) length of meri, each with strong spine at dorsodistal angle; dorsal margin unarmed (left) or with minute subproximal spine (right). Meri with ventral margins each bearing 3 (left) or 2 (right) small spines; dorsal margins each with row of spines. Ischia each with small dorsodistal spine and another small ventrodistal spine. Coxae unarmed. Sternite of third pereopods with anterior lobe rectangular, unarmed. Fourth pereopod (Fig. 6D) subchelate. Dactyl terminating in strong corneous claw; prominent preungual process present at base of claw; ventral face with 1 corneous spine laterally. Propodal rasp with 2 rows of corneous scales. Carpus with large dor- sodistal spine; ventral face with clump of long capsulate setae (Fig. 6E). Fifth pereopod (Fig. 7A) chelate; dactyl and propodus with well-developed rasps. Male first pleopods (Fig. 7B—D) paired, modified as gonopods; basal lobe bearing few setae at superior mesial angle; inferior lamella with distal margin bearing row of short, hooked spines, and lateral margin with row of setae; internal lobe with row of setae on mesial margin; external lobe dis- tinctly exceeding inferior lamella in distal extension. Second pleopods (Fig. 7E—G) paired, modified as gonopods; basal seg- ment naked; endopod with several long se- tae; appendix masculina twisted; lateral and distal margins and inferior face with mod- erately long setae. Third (Fig. 7H) to fifth left pleopods each with exopod well devel- oped, endopod reduced. Uropods asymmetrical, left larger than right; rasps of exopods and endopods well developed; protopods each with row of spines posteriorly. Telson (Fig. 71) with lateral constrictions; anterior portion unarmed; posterior lobes separated by deep median cleft, left lobe larger than right, terminal margins fringed with spines. Female unknown. Color in life (Fig. 8).—Shield white; an- tennules with flagella and ultimate segment yellow, setae on flagella blue, penultimate and basal segments red; antennas with fla- gella bearing alternative red and white bands, fifth segment with middle red band, proximal half of third segment red, second segment red except for white distal spines, first segment red except for ventral face, an- tennal acicle with subdistal red band, other surfaces of antennas white; ocular pedun- cles yellow, each with red band on proximal 0.25; ocular acicles red except for white distal spines; third maxillipeds with propo- dus, carpus, and merus and penultimate segment of exopod each bearing middle red band, other surfaces white; second maxil- lipeds with middle red band on penultimate segment of exopod. Both chelipeds and sec- ond through fifth pereopods with irregular red area on each segment. Etymology.—From the Latin bicolor, two colors, in reference to the alternating red and white color bands on the pereopods characteristic to this species. Distribution.—Known only from the type locality. Remarks.—Despite their general similar- ities in morphology, the new species, P. bi- color, is readily distinguished from both P. jJanetkae and P. bollandi by differences in coloration in life. The chelipeds and the second and third pereopods in P. bicolor have alternating red and white bands. These appendages are uniformly red in P. bollan- di, and, in P. janetkae, the meri and carpi and proximal half of palm of the chelipeds are cranberry-red and the carpi, propodi and dactyls of the second and third pereopods are light cream, tinged with yellow. Morphologically, P. bicolor is similar to both P. janetkae and P. bollandi, but some differences are seen among them. The most striking difference that separates P. bicolor from both P. janetkae and P. bollandi is the degree of dissimilarity in the chelipeds in males. In male P. janetkae and P. bollandi, the chelipeds are very unequal, and arma- VOLUME 117, NUMBER 1 tures are much stronger in the right than in the left. However, in male P. bicolor, the chelipeds are subequal and the dissimilarity of the armature is not so large as in those in P. janetkae and P. bollandi. Furthermore, the dorsal surfaces of the chelae are provid- ed with tubercles in P. bicolor and P. bol- landi but with spines in P. janetkae. Other minor difference includes the fact that, although a preungual process is absent in P. bollandi, both P. bicolor and P. ja- netkae have a very prominent preungual process developed at the base of the claw, giving the dactyl a quasi-chelate appear- ance. However, so few specimens have been reported in any of these species (four specimens in P. janetkae, one in P. bollandi and one in P. bicolor), it is not possible to evaluate variability. Thus, we expect future collection efforts to provide more precise information on morphological descrimina- tion between the species. Acknowledgements The authors are most grateful to Dr. Patsy A. McLaughlin (Shannon Point Marine Center, Western Washington University) for her elaborate review of the manuscript and the captain Higa Koei (Okinawa) for the successful cruise to collect this important material. The comments by Jacques Forest (Muséum national d’ Histoire naturelle, Par- is), D. L. Rahayu (Research Center for Oceanography, Indonesia) and an anony- mous reviewer greatly improve the final 67 draft of the manuscript. This work was part- ly supported by a Grant-in-Aid for Scien- tific Research (C) from the Ministry of Ed- ucation, Science, Culture and Sports of Ja- pan awarded to Akira Asakura (No. 14540654). Literature Cited Asakura, A., & P. A. McLaughlin. 2003. Pseudopa- guristes bollandi, new species, a distinct hermit crab (Crustacea: Decapoda: Diogenidae) from Japan.—Proceedings of the Biological Society of Washington 116:453—463. McLaughlin, P. A. 1974. The hermit crabs (Crustacea, Decapoda, Paguridae) of northwestern North America.—Zoologische Verhandelingen 130:1— 396. . 1997. Crustacea Decapoda: hermit crabs of the family Paguridae from the KARUBAR cruise in Indonesia. In A. Crosnier & P. Bouch- et, eds., Résultats des Campagnes MUSOR- STOM, 16.—Mémoires du Muséum national d’Histoire naturelle 172:433-572. . 2000. Crustacea Decapoda: Porcellanopagu- rus Filhol and Solitariopagurus Viirkay (Pagur- idae), from the New Caledonian area, Vanuatu and the Marquesas: new records, new species. In A. Crosnier, ed., Résultats des Campagnes MUSORSTOM, volume 21.—Mémoires du Muséum national d’Histoire naturelle, Paris, volume 184:389—414. . 2002. Pseudopaguristes, a new and aberrant genus of hermit crabs (Anomura: Paguridea: Di- ogénidae).—Micronesica 34:185—-199. , & M. de Saint Laurent. 1998. A new genus for four species of hermit crabs formerly as- signed to the genus Pagurus Fabricius (Deca- poda: Anomura: Paguridae)—Proceedings of the Biological Society of Washington 111:158— 187. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):68-75. 2004. A new species of axiid shrimp from chemosynthetic communities of the Louisiana continental slope, Gulf of Mexico (Crustacea: Decapoda: Thalassinidea) Darryl L. Felder and Brian Kensley (DLF) Department of Biology, University of Louisiana at Lafayette, Louisiana 70504, U.S.A., e-mail: DLF4517 @louisiana.edu; (BK) Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A., e-mail: kensley.brian@nmnh.si.edu* Abstract.—Calaxius carneyi, new species (Axiidae), is described from two male specimens collected by manned submersibles working near hydrocarbon seeps in deep waters (544 m) on the continental slope off Louisiana, in the northern Gulf of Mexico. Both specimens were taken adjacent to communities of clams that comprise a major constituent of chemosynthetic assemblages at the collection site. The new species is characterized in part by ventrally truncate abdominal pleura, as opposed to the acutely triangular or broadly rounded pleura found in other known members of Calaxius, only one of which is known to occur in the Atlantic Ocean. The new species is readily distinguished from its congeners by unique dentition of its heavy triangular rostrum and postrostral carapace, its short eyestalks and antennal acicle, the absence of well-defined teeth on the massive chelipeds, and the narrow, subtriangular telson. Chelipeds, pleopods and uropods of the two known specimens herewith described are covered extensively by long setae, many of which are plumose and densely fouled by flocculent debris. Recent investigations of methane cold seeps in the Gulf of Mexico have discov- ered a number of previously undescribed taxa associated with chemosynthetic com- munities in deep waters of the continental slope (e.g., Gustafson et al. 1998). How- ever, some collections from these unique habitats consist of single specimens, and comparative studies have been deferred pending recovery of additional materials. One such case was presented by the collec- tion of a single, somewhat fragmented, molted integument from the male of an ap- parently undescribed axiid mud_ shrimp, collected in 1988 during a dive of the manned submersible Pisces IJ. A second, smaller, intact male specimen, was obtained * Deceased. from an adjacent site in 1992 with a shal- low core sampler deployed by the Johnson Sea-Link manned submersible. Collections on subsequent dives by submersibles and vessel-based box coring in this area have brought no additional materials to our at- tention. While the female of this species remains unknown, it is readily apparent that the spe- cies is undescribed. The marked size dif- ference between the intact specimen and the earlier recovered exuvia provides a glimpse of ontogenetic variation in characters, and allows us to select diagnostic characters that should apply to a wide size range. Also, from familiarity with typical sexual dimor- phism in congeneric species, we expect that the description here provided will serve ad- equately for identification of female speci- mens, if encountered. VOLUME 117, NUMBER 1 Upon arrival at the surface, specimens were fixed in 10% formalin (with Rose Bengal stain for only the 1992 collection), transferred to 80% ethyl] alcohol, and finally archived in 70% ethyl alcohol. Carapace length (CL) was measured from the poste- rior margin of the orbit to the posterior mar- gin of the carapace midline. Total length (TL) was measured from the tip of the ros- trum to the tip of the extended telson. Spec- imens are archived in the National Museum of Natural History (USNM), Smithsonian Institution, Washington, D.C. Family Axiidae Huxley, 1879 Calaxius Sakai & de Saint Laurent, 1989 Calaxius carneyi, new species Figs. 1 & 2 Material examined.—Holotype, USNM 1009165, male, CL 10.1 mm, TL 26.5 mm, Johnson Sea-Link submersible sta 3269, box core B, deployed from submersible about 2 m from chemosynthetic mussel community, Bush Hill site, 544 m, Louisi- ana continental slope, northern Gulf of Mexico, 27°46.904'N, 91°30.286'W, 11 Aug 1992. Paratype, USNM 1009166, exu- via of male, CL 18.3 mm, TL 50.5 mm, submersible Pisces I sta 880031 (8831), lo- cation and depth same as for holotype, Aug 1988. Diagnosis.—Rostrum heavy, triangular, extending slightly more than twice length of eyes, bearing flattened, upturned terminal spine and pair of similar upturned subter- minal spines. Antennal acicle short, over- reaching proximal third of penultimate (fourth) peduncular article. Chelipeds mas- sive, lacking well defined teeth. Carapace bearing dentate lateral and submedian ca- rina. Pereopodal epipods and pleurobranchs present. Abdomen with pleura 3—5 ventrally truncate, bearing small anterior and poste- rior marginal denticles; lacking male pleo- pod 1; appendices internae on pleopods 2— 5; uropodal exopod bearing tranverse su- ture; terminated by narrow, subtriangular telson. 69 Description of holotype.—Integument firm but pliable, with numerous clumps of elongate, plumose, fouled setae, often ob- scuring underlying structures on chelae, pleopods, uropods, and telson; calcification most heavy in carapace teeth and chelae. Carapace with posterior midline elevated, bracketed on either side by paired setose punctae, midline elevation becoming a rounded crest in cardiac region where sur- mounted by a slight but distinct prominence or tubercle, marked dorsally by translucent or worn area (Fig. 1a); rostrum heavily cal- cified, triangular, slightly more than twice length of eyes, terminal spine subacute, up- turned, dorsoventrally flattened, triangular in dorsal view; subterminal pair of spines similar to terminal, also upturned, imparting concave appearance to flattened dorsal sur- face of rostrum (Fig. 1b, c); supraocular spines (lacking in the paratype, a larger specimen) and supraorbital spines strong, similar in calcification, shape and orienta- tion to subterminal pair; lateral carina orig- inating from supraorbital spine, diminishing immediately anterior to second spine or tooth, continuing as a low ridge toward pos- terior; submedian carina originating from posteriormost of two slightly offset sub- median teeth, becoming ill-defined toward posterior; median carina a weak crest bear- ing a worn tubercle near its posterior end, and otherwise lacking ornamentation. Sternite of fourth pereopods (seventh thoracic somite) with deep median slit, tho- racic shield produced to form acute, mar- ginally sinuate, triangular flange to either side; 3-branched carina set between articu- lations of fourth pereopods (Fig. 2a). Ab- dominal pleuron | narrowed, acute ventral- ly (Fig. la); pleuron 2 ventrally broad, with an angular tooth or acute corner at the pos- teroventral end; pleura 3—5 ventrally trun- cate, with small acute tooth on anteroven- tral margin and another on posteroventral margin; pleuron 6 with small acute tooth on anteroventral margin and broad triangular flange at posteroventral margin. Eyestalks small, subcylindrical, reaching 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ a, f c, d e b Fig. 1. Calaxius carneyi, new species (where setation is shown, setules and flocculent coating of plumose appendages not fully illustrated). a—-e, holotype male, USNM 1009165: a, carapace, abdomen, left pereopods 1 and 3—5 in lateral view, with right pereopod 2 internal surface; b, anterior carapace, eyes and antennal peduncles, right side, lateral view, setation not shown; c, anterior of carapace in dorsal view; d, right pereopod 1 or major cheliped, internal surface; e, right pereopod 2, external surface. f, paratype male, USNM 1009166: right pereopod 1 or major cheliped, external surface. Scale bars indicate 2.0 mm. VOLUME 117, NUMBER 1 Fig. 2. 71 Calaxius carneyi, new species, holotype male, USNM 1009165 (where setation is shown, setules and flocculent coating of plumose appendages not fully illustrated). a, posterior thoracic sternites and coxae of pereopods 3—5, ventral view, setation not shown; b, left maxilliped 3, external surface; c, endopod of left maxilliped 3, internal surface; d, pereopod 3, external surface; e, pereopod 4, external surface; f, pereopod 5, external surface; g, abdominal somite 6, telson and uropods, dorsal surface. Scale bars indicate 2.0 mm. almost to midlength of rostrum (Fig. la, b, c). Cornea terminal, slightly globose, di- ameter equal to or slightly exceeding that of eyestalk. Antennular peduncle reaching well be- yond rostrum (Fig. 1b). Antennal peduncle bearing produced nephridiopore proximov- entrally, second article bearing dorsodistal spine overreaching much of acicle, acicle short, not bifid, overreaching proximal third of penultimate (fourth) peduncular article, third peduncular article distally bearing strong ventromesial spine. Maxilliped 3 ba- sis bearing short, acute mesial spine (Fig. 2b); ischium of endopod with strong, dis- tally elevated crista dentata on internal sur- face, bearing about 16 spines, distalmost of which are largest and most strongly direct- ed mesiad (Fig. 2c); merus with two mesial spines, one near or just short of midlength, the other larger and in distal third; carpus with short triangular, tooth at distal extreme of flexor margin; all articles of endopods bearing fields of long setae, many dense and heavily plumose, especially on mesiad and internal surfaces. Pereopods 1—4 bearing epipods. Pereo- pods 2—4 with pleurobranchs above coxae (on thoracic somites 5—7). Chelipeds (pereopod 1) similar in form and ornamentation on the 2 sides, right the heaviest (Fig. la, d); ischium with single well-defined spine on inferior margin; mer- us with single flattened spine near mid- 72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON length of inferior margin, which is weakly marked by adjacent sinuation or serration, distal corner of flexor margin on external side forming short, heavy raised spine (Fig. la); carpus very short, bearing numerous patches of long setae on external surface, dorsal margin weakly tuberculate, terminat- ing distally in blunt tooth, ventral carina of external surface forming flange distally which terminates in flattened, weakly hooked tooth; propodus very thick and heavily calcified, lacking well-defined teeth on weakly tuberculate dorsal margin, bear- ing numerous patches of long setae on ex- ternal surface, including among tubercles of dorsal surface, along well-marked carina of ventral margin, below cutting edge of fixed finger, and proximal to gape, external sur- face proximal to dactylus with scattered low tubercles and granules, fixed finger bearing two erect teeth on proximal half and single less erect tooth in distal half, ter- minus spiniform, internal surface with weak carina adjacent to and slightly below cut- ting edge; movable finger very thick and heavy, with dense patches of long setae ex- ternally and dorsally, cutting edge bearing rounded to lobiform tooth in proximal third, broad ill-defined tooth or sinuous lobe in distal two-thirds, terminus weakly hooked and subacute, a carina above cutting edge on internal surface. Pereopod 2 (Fig. le) merus lacking marginal spines, combined length of ischium and merus about equal to combined length of carpus and propodus, length of dactyl about half total length of propodus, opposable cutting edges of fin- gers corneous, finely pectinate, distinctly spooned distally. Pereopod 3 (Fig. 2d) mer- us lacking marginal spines, external surface of propodus bearing five sets of corneous spiniform setae, set individually or in short transverse rows of two or three near inferior margin, fewer sets and few such setae near superior margin, falcate dactylus with three distinct corneous spiniform setae on exter- nal surface, distally with two more very small ones set near flexor margin, and a sharp corneous spine forming terminus. Pe- reopod 4 (Fig. 2e) merus lacking marginal spines, external surface of propodus bearing six sets of corneous spiniform setae, set in- dividually or in transverse rows of two to four near inferior margin, four such sets of one to three setae near superior margin, fal- cate dactylus with five distinct corneous spiniform setae on external surface, distally with additional very small corneous seta, sharp corneous spine forming terminus. Pe- reopod 5 (Fig. 2f) merus and propodus lacking marginal spines, propodus bearing stiff bristles at distal inferior end of pro- podus, concealed by dense distal fields of setae; lanceolate dactylus twisted laterally, opposed to terminal bristles of propodus when flexed. Pleopod 1 absent, posterior pleopods all bearing dense cover of long, plumose, heavily fouled setae; appendix interna pre- sent on pleopods 2—5. Uropodal exopod (Fig. 2g) bearing four spines along external margin and an articulated spine where this margin meets the transverse suture, five ad- ditional spines along transverse suture, and no spines on dorsal surface, long setae forming dense fringe on margins, but on dorsal surface limited to few patches near external margin; endopod with single strong spine at distal end of external margin and another small spine overreaching distal margin at end of weak median ridge, long setae forming dense fringe on margins, and distributed in patches on dorsal surface near external margin and along medial ridge. Telson length distinctly greater than its bas- al width, tapering toward posterior, widest at lateral lobes in proximal one-quarter of length, single pair of strong fixed dorsal spines in anterior half, two to four fixed marginal serrations or spines posterior to proximal lobes, and two pairs of short, ar- ticulated marginal spines in distal third of lateral margins, distal margin evenly con- vex, densely setose. Variations.—Paratype: Spination and tu- berculation in the exuvia of this larger spec- imen differ in several minor ways from or- namentation in the holotype. There are few- VOLUME 117, NUMBER 1 er spines on the margin of the rostrum, as the supraoculars are not present. The mar- gin of the rostrum is somewhat broadly concave in this region on the paratype, al- though it retains an overall triangular shape. Dorsal tuberculation of the propodal palm is less evident than in the holotype. Spines on the cutting edge of the major chela differ slightly in shape from those on the holo- type, but the pattern and placement of this spination is conserved (Fig. If). Granula- tion on the internal surface of the propodus in the paratype is stronger than that in the holotype. The external margin of the uro- podal exopod in the paratype male bears five rather than four fixed spines, while the external margin of the endopod bears three spines rather than a single one. Lateral mar- gins the telson bear up to five serrations or fixed lateral spines in the paratype, and the pairs of articulated marginal spines are rel- atively smaller than in the holotype and very difficult to discern. Small angular, acute corners or teeth on the anteroventral margins of abdominal pleurae 3—5 are also more difficult to discern in the holotype. These appear to be somewhat worn or smoothed off in the paratype, although the acute posteroventral margins remain readily evident. Etymology.—The species is named for Robert S. Carney, Louisiana State Univer- sity, Baton Rouge, who oversaw collections of the specimens and made these materials available for our study. His own work on hydrocarbon vent communities of the Gulf of Mexico (see Carney 1994) has brought needed attention to these remarkable assem- blages of marine organisms. Remarks.—Yen species were listed in a recent review of Calaxius by Kensley & Hickman (2001). The present description accounts for the eleventh known member and only the second species to be found in the Atlantic Ocean. There seems little doubt as to the generic placement of this new spe- cies, given the dentate rostrum twice as long as the eyestalks, the dentate carapace carinae, the transverse suture on the uro- 73 podal exopod, presence of pereopodal epi- pods and pleurobranchs, the absence of ple- opod | in the male, and the presence of appendices internae on pleopods 2-5 (see Poore 1994:97). The specimens lack strong dentition on the dorsal surface of the first chelipeds, as seen in Acanthaxius Sakai & de Saint Laurent, 1989, which they some- what resemble. In contrast to the known congeners and the original generic definition (Sakai and de Saint Laurent 1989:84), the rostrum of C. carneyi is heavier and more broadly trian- gular, the eyestalks and antennal acicle are comparatively shorter, and the palm of the cheliped lacks well-separated and defined teeth dorsally, having at most a covering of low tubercles, and the telson is distinctly narrowed posteriorly or subtriangular. The abdominal pleura of C. carneyi resemble neither the acutely triangular plates seen in four other previously described species nor the broadly rounded plates seen in five oth- er species; rather, pleura 3—5 are ventrally truncate with small anterior and posterior marginal denticles. The only species of Calaxius previously reported from the broad geographic area of the Gulf of Mexico and contiguous regions is C. oxypleura (Williams, 1974), recorded from the Straits of Florida. This species has abdominal pleura 3—5 ventrally angular or acute, rather than truncate, and a narrow dentate rostrum unlike that of C. carneyi. Ecology.—A dense cover of flocculent materials on plumose setae of both the ho- lotype and paratype exuvia (much of which has disintegrated over time in alcohol, or which was brushed free in the course of morphological studies) may derive from the unique environments inhabited by these an- imals, but nothing is known of the burrow structure or feeding behavior. The floccu- lent coatings on the axiid setae could be a ‘passive result of these animals’ trophic ties to members of the chemosynthetic com- munity, at a primary or secondary consum- er level. Accumulations of mussels and worms near hydrocarbon seeps in the north- 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ern Gulf of Mexico do in varied ways de- pend upon methanotrophic or sulfur-oxidiz- ing bacteria for metabolic resources (see Van Dover 2000:363—366). These bacteria occur in animal tissues as endosymbiotic cells or as scattered mats immediately sur- rounding the seeps. They may directly ex- ploit methane as both a carbon and energy source or, much as in hydrothermal vent en- vironments, directly oxidize rich cold-seep sources of sulfides for metabolic energy. Precipitates and bacteria might simply ac- cumulate on or among the setules of highly plumose setae, perhaps as these axiids move about or ventilate burrows in these matted settings. While the flocculent mate- rials could simply mask movements among bacterial mats or mussel communities, we cannot rule out that the axiids themselves may directly consume accumulations of chemosynthetic bacteria, either by access- ing exposed mats or undertaking behaviors that favor the forming of such accumula- tions within and along walls of their bur- rows. It is suspected that other thalassini- deans engage in burrow-modulated feeding behaviors that are microbially based, albeit in reduced interstitial waters of shallow hypoxic environments (Felder 2001) where at least one species lives in apparent asso- ciation with lucinid bivalves harboring che- mosynthetic gill bacteria. Even if C. carneyi could be shown to de- pend upon the chemosynthetic community as a nutritional resource, perhaps by stable isotope measurements, this would not nec- essarily confirm its restriction to occurrence with chemosynthetic communities of hy- drocarbon seeps. As has recently been re- ported for infaunal worms associated with methane seeps off California (Levin et al. 2000), infaunal thalassinideans are also likely pre-adapted to organic-rich, reducing environments, and may in fact be widely distributed forms that do not strictly exhibit chemosynthesis-based trophic specializa- tions. Owing to very limited general sam- pling for infaunal macrocrustaceans from slope environments in the Gulf of Mexico to date, its occurrence in sediments other than those near cold seeps cannot be ruled out. Acknowledgements For providing the specimens, we thank R. Carney and his associates at Louisiana State University, Baton Rouge. Support for the present study was furnished to DLF un- der U.S. Department of Energy grant no. DE-FG02-97ER12220 for studies of ende- mism in the northern Gulf of Mexico. We are grateful to S. Brooke, C. Allen, and C. Young for field assistance and sharing ship time funded under support of NSF grant no 0243688-OCE (to C. Young), in the course of our continuing studies of hydrocarbon vent infaunal decapods. This is contribution no. 98 from the University of Louisiana Laboratory for Crustacean Research. Literature Cited Carney, R. S. 1994. Consideration of the oasis analogy for chemosynthetic communities at Gulf of Mexico hydrocarbon vents—Geo-Marine Let- ters 14:149-159. Felder, D. L. 2001. Diversity and ecological signifi- cance of deep-burrowing macrocrustaceans in coastal tropical waters of the Americas (Deca- poda: Thalassinidea).—Interciencias 26:2—12. Gustafson, R. G., R. D. Turner, R. A. Lutz, & R. C. Vrijenhoek. 1998. A new genus and five new species of mussels (Bivalvia, Mytilidae) from deep-sea sulfide/hydrocarbon seeps in the Gulf of Mexico.—Malacologia 40:63-113. Huxley, T. H. 1879. On the classification and distri- bution of the crayfishes.—Proceedings of the Zoological Society of London 1878:752-788. Kensley, B., & C. P. Hickman, Jr. 2001. A new species of Calaxius Sakai & Saint Laurent, 1989, from the Galapagos Islands (Crustacea: Decapoda: Axiidae).—Proceedings of the Biological Soci- ety of Washington 114:484—488. Levin, L. A., D. W. James, C. M. Martin, A. W. Rath- burn, L. H. Harris, & R. H. Michener. 2000. Do methane seeps support distinct macrofaunal as- semblages? Observations on community struc- ture and nutrition from the northern California slope and shelf.—Marine Ecology Progress Se- ries 208:21—39. Poore, G. C. B. 1984. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys VOLUME 117, NUMBER 1 75 to families and genera.—Memoirs of the Mu- Van Dover, C. L. 2000. The Ecology of Deep-Sea Hy- seum of Victoria 54:79—120. drothermal Vents. Princeton University Press, Sakai, K., & M. de Saint Laurent. 1989. A check list Princeton, New Jersey. 424p. of Axiidae (Decapoda, Crustacea, Thalassini- Williams, A. B. 1974. Two new axiids (Crustacea: De- dea, Anomura), with remarks and in addition capoda: Thalassinidea: Calocaris) from North descriptions of one new subfamily, eleven new Carolina and the Straits of Florida.—Proceed- genera and two new species.—Naturalists 3:1— ings of the Biological Society of Washington 104. 87(39):45 1-464. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):76-87. 2004. Description of a new Synidotea species (Crustacea: Isopoda: Valvifera: Idoteidae) from Hawaii Wendy Moore University of Arizona, Department of Entomology, Tucson, Arizona 85721-0036 U.S.A., email, wmoore @ag.arizona.edu Abstract.—This paper provides the first description of a Hawaiian isopod of the genus Synidotea, S. oahu n. sp. This species is most similar to S. laevi- dorsalis (Miers, 1881) and S. harfordi Benedict, 1897. A list of Synidotea species described to date with biogeographic information, and a list of all marine isopods described from the Hawaiian Islands, are provided. This paper provides the first description of a Synidotea species from the Hawaiian Islands. The isopod genus Synidotea Har- ger, 1878 currently contains 57 species, in- cluding the species herein described (see Table 1). The following characters define this genus: penes fused forming penial plate, fifth oostegites absent, and sexually dimorphic mouthparts (Poore 2001). In ad- dition, Synidotea species possess the fol- lowing combination of characters: antennae 2 flagellum multiarticulate, maxillipedal palp triarticulate, pleon with one partial su- ture, pereonites 2—4 coxal plates not visible in dorsal aspect and (unlike most other val- viferan genera) pereonites 5—7 tergite-coxal plate sutures can be either present or absent. The Californian species of Synidotea were reviewed by Menzies & Miller (1972), who also included a biogeographic account of the genus that, at the time, contained 36 species. The phylogeny and biogeography of the 22 idoteid genera, including Syni- dotea, were discussed by Brusca (1984). Poore (2001) redefined and inferred the phylogeny of the families within the Val- vifera. Most Synidotea species occur in the Arc- tic and in boreal waters (39 of the 57 de- scribed species); 13 species have been de- scribed from tropical/subtropical waters. To date, only one other Synidotea species has been described from the islands of the trop- ical Pacific, S. pacifica Nobili, 1906 from the Tuamotu Islands. Synidotea oahu n. sp. is one of only 29 marine isopods known from the Hawaiian islands (see Table 2). The only other known Hawaiian valviferan is Colidotea edmondsoni Miller, 1940. Nine species in this genus belong to the Synidotea hirtipes species-group (Monod 1931, Menzies & Miller 1972): S. hirtipes (H. Milne Edwards, 1840), S. laevidorsalis (Miers, 1881), S. laticauda Benedict, 1897, S. harfordi Benedict, 1897, S. marplatensis Giambiagi, 1922, S. brunnea Pires & Mor- eira, 1975, S. keablei Poore & Lew Ton, 1993, S. grisea Poore & Lew Ton, 1993, and §. oahu n. sp. Members of the S. hir- tipes species-group share the following dis- tinguishing characters: pereon smooth, frontal margin of head entire or slightly ex- cavate, and posterior border of pleotelson with median excavation. Because S. oahu n. sp. possesses these characters I herein consider it a member of this group. Species boundaries within the S. hirtipes group have been disputed in the literature. Chapman & Carlton (1991, 1994) argued that S. laevi- dorsalis is a widespread species, which has been widely introduced to many coastlines from Japan by the shipping industry. Chap- man & Carlton (1991, 1994) have thus sug- gested the synonymy of seven of the nine species within this group. However, their taxonomic justification for the synonymies VOLUME 117, NUMBER 1 Fig. 1. Holotype, dorsal view. was weak, based entirely on an analysis of length-width ratios of various body parts of the dorsal aspect of these species. Poore (1996) refuted the synonyms; through care- ful comparison of the pleotelson, penial plate and pereopod 1, he clearly demon- strated that the populations descibed from various Indo-Pacific coastlines represent valid and separate species. He also noted that the species boundaries are further sup- Ti ported by different ecological distributions of the species in this group. This case un- derscores the importance of detailed, accu- rate taxonomy in the pursuit of successfully identifying translocated species. Taxono- mists are accustomed to the challenging task of recognizing species boundaries within groups that contain many similar species; oftentimes differences between species, although solid and obvious once made explicit, are not apparent to the un- trained eye. Order Isopoda Latreille, 1817 Suborder Valvifera Sars, 1882 Family Idoteidae Samouelle, 1819 Genus Synidotea Harger, 1878 Synidotea oahu, new species Figs. 1-6 Type material examined.—Holotype, ovigerous female, USNM 1009176. Ha- wail: Oahu Is., 0.8 km from town of Kailua, collected from small batches of seaweed by Ray Greenfield, August 20, 1950. Paratype, female, USNM 99384. Hawaii: Oahu Is., Ewa Beach, 32 km from Honolulu, collect- ed from seaweed by Ray Greenfield, Au- gust 1, 1954. Etymology.—The specific epithet oahu derives from the poetic vowel-rich Hawai- ian language, providing this binomen, Syn- idotea oahu, with every vowel in the En- glish alphabet. In Hawaiian, oahu means “the gathering place.’ Oahu is also the name of the second largest island in the Ha- waiian archipelago and the type locality of this species. This word is used here as a noun in apposition. Diagnosis.—Cephalon dorsal surface with a weak, transverse depression in front of eyes. Pereonites 1—7 with mesial, broad- ly rounded grooves on dorsal surface. Max- illa 1 mesial lobe with two unique, stout, distally-serrate robust setae with mesial set- ules. Mandibles (both right and left) with four-toothed incisors and four-toothed laci- nia mobili with additional large serrate spine-like process. 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V9) nn, TC, Naa [epHiojur (Aequiog :eipuy) uerpuy [eordory, 6S61 Tea 2 lysor sisuariom “5 SE-L (BHOJIIA “eI[ePNsNY “AA\) UeIDQ UIEyINOS onorejueqns €66] “UOL MOT 2 100g apuosjpm ‘s (aeosesepeyy ‘onbiquiezopy Oc-I ‘BOLFY YINOG) ueIpuy MS “(exUeT Lg ‘eipuy) uerpuy Teordory, LI6L ‘S8ULIOD Vinsaliva “¢ Sel—-OCl QISIOYAO JO VIS) OYTO’, MAN SHS1TV 6061 “UOspreyory vipjno1eqni “Ss CZb-OS (SJ 9[uMy :eIssny) oy1OeVg MAN [es10g 6L6I “AOYUZII 27 ULYessny VIvsOWspUgns ‘*¢ O8-L (BoLIpYy “S) URIPUT AAS jeordory, VI6I ‘preureg safias ‘¢ V8C-09 CISIOUAO JO VIG) IYIOe MAN ONOTW SS6l ‘eAouelInD vidjnos -s [epiiojur (BIUIOJTeD) oytoed AN [Bolo g YO6I UOspreysTYy Mani “¢ LI6C-L887 oytoe’d MN [Boog €961 “UlolsIg V4yojnd “5 Or-Ol (Bysely) oytoed AN [Bolo g L681 “Ipsueg vjo1d “5 Cc-[episoqur (uoIsuTYyseM “eIquINTOD ysHIIg) oyloed AN [B00 Lr6l “YeH evau0giiad ‘5 Iv91-08e1 (eyseTy) oytoed AN [R010 L681 “OIpsusg vpyjod “s é, (s] Mouwlen]) sy1oeg [eotdory, jenued jeordory, 9061 “IIGON voifiond -¢ AOT[eYS (ueder) sy1oeg MAAN [eol0g $86] “vInUIOUNN sisualyonsjo «5 [epnsojur (SJ nyvO :HeMep) oyloeg [eordo1y, [enuad jeordonqns ‘ds ‘u nyvo -s (purfuseip) onuepYy N ‘oHory ‘(2[s]}OyxO Jo keg) EVES oytoed MN “(eIquinjoD ysniug “‘eysepy) oyloeg AN [esiog ‘ono1y (9p81 “1ohory) Vsojnpou ‘5 é (Aeg euro], :ueder) oytoeg MAN [eoi0g 686] ‘vInuUIOUNN sisuauoddiu -¢ C691 oytsed MN [eso g E961 “UlolsIIg BIoa]sau -§ O8e-[epHiejur (eysefVy) 9ytoed AN [Rolo g L681 WIpsueg vsojnqau ‘s OSI-St (eg SulIog) ytord N [eolog “ono1y (LL81 “prlojzeH]) vivojinu -¢ [epHsojur (BIquiInjoD Yysnig) oy1oed AN [B90 0661 “Ziqney] 2 yey vinuiwu ‘5 esl (etuIOIeD) oytoVd AN [e210 TL61 “UOSISAT DIPAU “¢ Oe-TepHiejur (eunuesrIy ‘[IzeIg §) onuElYy § qeordory, 7TH “WRIqQUIRID sisuajnjdinu “5 09¢E-VI (topeige]) onuepy N [eo10g (L981 “preyoed) Vin1oWIDUL “S 16-Ss¢ (BIUIOJTeD) oy1oed AN [eoIOg 6S6I ‘Pieuleg 2 solzucejy voyiuspu -¢ VOLUME 117, NUMBER 1 (sioj}owl) yidoq (UOTNGINSIP pep10se1) uLZ9O uoIse1 o1ydeisoos01g ee ee 2 eR Re ee OE eee a se ake Se ey ene ee tes Seo ee OM ‘ponunuoy— | 219qeL Joyine pue saroadg 80 mS B Cn i ry ay Fig. 2. Paratype. A, lateral view; B, dorsal view. pereonite 4 width is 0.69. Pleotelson (fused pleonites and telson) 1.26 times longer than wide (measured along lateral margin, from posterior margin of coxa of pereonite 7 to distal-most tip of telson). Description.—Body length: ovigerous female holotype, 8 mm, non-ovigerous fe- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON male paratype, 7.5 mm. Body yellowish tan in alcohol. Cephalon dorsal surface with a weak, transverse depression in front of eyes. Fron- tal margin straight. Eyes bulge outward, forming part of contour of lateral margin of head. Ratio of head width to pereonite 4 VOLUME 117, NUMBER 1 81 Fig. 3. Holotype. A, right antennna 1; B, left maxilla 2 close-up of inner lobe; C, left maxilla 2: D, head, ventral view; E, head, lateral view; E left mandible, dorsal view: G, left mandible, mesial view: H, left mandible, ventral view, I, right antenna 2. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 82 Bag pod POLY S$ ‘onbiquiezoy] ‘ueder ‘eijensny § ‘spuvisy uenemepy Spurs] ULIIeEMeyy uvderp ‘puvjsusangd ‘eljensny AA ‘Spuvysy uenemey Spurs] UviIeMey Spurs] uelieMeyy Spue]s] uelieMey Spur]s] uviemeyy UvOURLIOIpsy] “ST soyTuuy ‘Sol0ZY ‘SoT[oyoAes “eoLyy AWN ‘euynD ep uejsity ‘apia, odes ‘uvder ‘eory eisop oyloeg ‘spuvys] uenemepy spuvys] uelieMey spuv]s] uelieMey spur]s] ueliemey Spurs] uelieMeyy spuvys] uelieMey Ieosesepey] ‘Spurys] uelemey spuv]s] uelieMey spurs] ueneMey spuejs] uelieMey spuv]s] uelieMey snnuneyy “vag poy “s] O1OUWIOD [OV eIqG -BPlV ‘eIpuy ‘Oorxey] JO FIND ‘uvoqqrie ‘spuvysy uenemey purypreyy ‘eqnD ‘emios1eD ‘spuvysy uenemeyy spuv]s] uvieMeyy spuvjs] uvieMey spuvjs] uvieMeyy (P88I “MSUlsy 2 Wps01yIS) YIIYsoUuDjawU YDKIOYIOP| (Z96| ‘URWIMOG) 14nd snuaxokyjYyoT 686] ‘URWIMOg 2 soNIg sMjoUD snigossoj]H €SQI ‘eueq Vj9a4 vOYIOWKD [881 ‘Moule 2 Apso1yss) sdadiaasg vjoluasD PO61 ‘UOspreyory sisuanpupy Djaulz0y €06] “UOsiVYorYy snuispjpippnb psay (OP8I ‘SprlemMpy UTA “H) Vuviskoysap vsay €06] ‘UOSpIeyoTY suaposas snxkKsydouoz O€6I ‘Sipuvrig ev IOUSIg ZSeNSISIN DLQDIS DqDA O16] “UOspreyory sisuanpMpy uodaons4K95 OL6I “UMoyued mMosiyoinw pj]auol LO6I ‘YNoJued sisuaipmoy auolupsiyH E061 ‘UOSpreyoTY snjoapumo snjiydoyuq (LP6L “ISMN wnofiovdipaw wniuasunzy (LP61 “ISLA BUlivop puuniuo01Q IP6L “IONA Sisuapmpy sisdosaor L961 “IOAN 202Jad vuupipMDy (Lr61 “STA 2/0218]0 spidind IS6] ‘Salzusy 1yyDALO0Y DAéadlizavD TS6I ‘SOIZUDI 2 IO[IAl IP4VVSsajso DinyUDID TS6[ ‘SOIZUDPY JOYA VPNYIYJaq DANYJUDIDG TSG ‘SOIZUDI 2 IO[UYA VI1YdK]sosa1y DiAnyJuvsSapy oeploujoulAZD depisoy Vaal Td dv 14 ovpileq oeprlosiuoj\dAia aepuAdog VadIeVlda depiLyaua}S oepruunyyy oepisdois0f oppure VLOTIOSV oepLnyueieg Spurs] uviiIeMey (ZS6I ‘SOIZUDII 2 IJO[[NA) VIVUsOUI DANYJUDSNYDULY oepunyiuy VAECIeNHINV UONNIISIP pop1osoay Joyjne pue saisadg Ayruuey Japiogng “SpUPISs] ULIIEMLH{ OY) SUIPUNOLINS s1o}eM dy} UI 1N990 0} poytoder spodosr sue [fe JO ISI] Y—Z 2IGe.L VOLUME 117, NUMBER 1 Table 2.—Continued. Recorded distribution Species and author Suborder family Hawaiian Islands, Samoa, Japan, Christmas Is., Andaman Paralimnoria andrewsi (Calman, 1910) Limnoriidae Is., Aldabra Atoll, Caribbean Hawaiian Islands Cymodocella hawaiiensis Bruce, 1994 Neonaesa rugosa Harrison & Holdich, 1982 Sphaeromatidae Hawaiian Islands, Society Islands, New Guinea, Queensland VALVIFERA Hawaiian Islands Idoteidae Colidotea edmondsoni Miller, 1940 Synidotea oahu n. sp. Hawaiian Islands 83 width is 0.69. Antenna 1 with triarticulate peduncle and uniarticulate flagellum with six pairs of jointed aesthetascs. Antennae 2 extended to third pereonite; with five-artic- ulate peduncle, article 5 at least twice as long as any other peduncular article; fla- gellum with 15—17 articles, terminal two ar- ticles very small. Maxilliped with a triarticulate maxilli- pedal palp, single coupling hook on the left maxilliped only (holotype). The paratype has one coupling hook on both left and right maxilliped. Maxilla | mesial lobe with two stout distally-serrate robust setae with mesial setules; lateral lobe with ten serrate robust setae and many simple setae along lateral and mesial margins. Maxilla 2 with plumose, simple and comb setae as figured. Mandibles with four-toothed incisors and large molar processes with short spines sur- rounding margins. Lacinia mobilis of left and right mandible four-toothed with an ad- ditional large serrate spine-like process. Pereonites 1-7 with mesial, broadly rounded grooves on dorsal surface, other- wise dorsal surface and lateral margins smooth, without rugae, tubercules, or scales. Pereonite 3 widest. Lateral margins of pereonite 1-3 evenly convex, 4-7 straighter but not sharply angulate. Pereo- pods setose. Pereopod 1 with dactyl as long as propodus; two stout setae arise from base of unguis; distal lateral surface of propodus covered with serrate setae. Pereopods 2—7 with setation patterns as figured. Pleotelson 1.26 times longer than wide; dorsal surface evenly convex; posterior bor- der with median excavation. Pleopods 1 and 2 with plumose marginal setae on endopods and exopods, both rami without sutures. Pleopods 1—3 with coupling setae on mesial margin of peduncle. Pleopods 3—5 with plu- mose marginal setae on exopods only; the number of setae decrease from pleopods 3 to pleopods 5; exopods with partial sutures on lateral margins, Uropod with an oblique ridge and 3 plumose setae at mesial junc- tion of protopod and exopod. Uropod exo- pod length to width ratio is 0.96. 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Discussion.—Synidotea oahu males are unknown. This species superficially resem- bles other members of the S. hirtipes spe- cies-group, particularly S. laevidorsalis and the widely distributed species S. harfordi. These three species are possibly closely re- lated, however, a phylogenetic analysis of this large genus is needed to test this hy- pothesis. Synidotea oahu differs from S. harfordi and S. laevidorsalis most strikingly in its smaller body size (S. oahu, 7.5—8.0 mm; S. laevidorsalis, 12.3-—35 mm, S. harfordi, 18 mm). Menzies & Miller (1972) noted that Synidotea species follow a general trend of increasing body size with increasing lati- tude. Wallerstein & Brusca (1982) showed the same trend for all intertidal idoteids oc- curring in the northeast Pacific. Species within Synidotea range in length from the 3 mm tropical Pacific S. pacifica, to the 32 mm S. bicuspida and 35 mm S. laevidor- salis from Arctic and boreal waters. Syni- dotea oahu fits this pattern, with a body size Holotype. A, left maxilliped; B, left maxilla 1. of 7.5-8 mm, the average body size for tropical Synidotea (Menzies & Miller IQ7/2)). S. oahu also differs from other members of the S. hirtipes species-group in the fol- lowing characters: S. oahu has unique stout distally-serrate robust setae with mesial set- ules on the mesial lobe of maxilla 1 and a four-toothed mandibular incisor, whereas S. harfordi and S. laticauda both have a two- toothed mandibular incisor. Synidotea oahu also differs from S$. harfordi in its broadly rounded median dorsal impressed lines on pereonites 2—4, whereas in S. harfordi these lines are distinctly triangulate. Also, the dactyl of pereopod 1 in S. oahu is nearly as long as the propodus, whereas in S. harfordi it is much longer than the propodus. Acknowledgments I am grateful to Marilyn Schotte for loan- ing specimens from the USNM collections. I also thank R. C. Brusca, G. C. B. Poore, VOLUME 117, NUMBER 1 85 Fig. 5. Holotype. A, right pereopod 5; B, right pereopod 7; C, right pereopod 6; D, right pereopod 1; E, right pereopod 1, close-up of propodus and dactyl; EK right pereopod 2; G, right pereopod 3; H, right pereo- pod 4. 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Holotype, A, right pleopod 5; B, right pleopod 4; C, right pleopod 3; D, right uropod; E, right pleopod 1; E right pleopod 2. VOLUME 117, NUMBER 1 B. Kensley, M. Schotte and 2 anonymous reviewers for commenting on the manu- script. Chip Griffin did the illustrations. Bri- an Kensley, Marilyn Schotte and Steve Schilling’s World List of Marine, Freshwater and Terrestrial Crustacea Isopoda website fa- cilitated the compilation of both tables. Literature Cited Benedict, J. E. 1897. A revision of the genus Synido- tea.—Proceedings of the Academy of Sciences of Philadelphia 1897:387-404. Brusca, R. C. 1984. Phylogeny, evolution and bioge- ography of the marine isopod Subfamily Ido- teinae (Crustacea: Isopoda: Idoteniae).—Trans- actions of the San Diego Society of Natural His- tory 20(7):99-134. Chapman, J. W., & J. T. Carlton. 1991. A test of cri- teria for introduced species; the global invasion by the isopod Synidotea laevidorsalis (Miers, 1881).— Journal of Crustacean Biology 11:386- 400. & . 1994. Predicted discoveries of the introduced isopod Synidotea laevidorsalis (Miers, 1881).—Journal of Crustacean Biology 14:700-7 14. Giambiagi, D. 1922. Cuatro nuevos isopodos de la Ar- gentina.—Physis Buenos Aires 5(20):230-244. Harger, O. 1878. Descriptions of new genera and spe- cies of Isopoda, from New England and adja- cent regions.—American Journal of Science 15(3):373-379. Latreille, P. A. 1817. Les Crustaces, les Arachnides, et les Insectes. In G.L.C.ED. Cuvier Le Regne Animal, distribue d’ apres son organisation, pour servrir de base a l’histoire naturelle des ani- maux et d’introduction a l’anatomie comparee. Volume 3 Paris. Menzies, R. J., & M. A. Miller. 1972. Systematics and zoogeography of the genus Synidotea (Crusta- cea: Isopoda) with an account of Californian 87 species—Smithsonian Contributions to Zoology 102:1-33. Miers, E. J. 1881. Revision of the Idoteidae, a family of sessile-eyed Crustacea.—Journal of the Lin- nean Society of London 16:1-88. Miller, M. A. 1940. The isopod Crustacea of the Ha- waiian Islands (Chelifera and Valvifera)—Oc- casional Papers of the Bernice P. Bishop Mu- seum, Honolulu 15(26):295-361. Milne Edwards, H. 1840. Histoire Naturelle des Crus- taces, comprenant Il’anatomie, la physiologie et la classification de ces animaux. Paris: Roret. Monod, T. 1931. Tanaidaces et Isopodes subantarctique de la collection Kohl-Larsen du Senckenberg Museum.—Senckenbergiana 13(1):10-30. Nobili, G. 1906. Diagnoses preliminaires de Crustaces, Decapodes et Isopodes nouveaux recueillis par M. le Dr. G. Seurat aux iles Tuamotou.—Bul- letin du Museum National d’ Histoire Naturelle, Paris 12:256-270. Pires, A. M. S., & P. S. Moreira. 1975. Two new spe- cies of Synidotea (Crustacea, Isopoda, Valvi- fera) from Brazil.—Boletim do Instituto Ocean- ografico 24:45-67. Poore, G. C. B. 1996. Species differentiation in Syni- dotea (Isopoda: Idoteidae) and recognition of introduced marine species: a reply to Chapman and Carlton.—Journal of Crustacean Biology 16:384-394. . 2001. Isopoda Valvifera: diagnoses and rela- tionships of the families—Journal of Crusta- cean Biology 21:213-238. , & H. M. Lew Ton. 1993. Idoteidae of Aus- tralia and New Zealand (Crustacea: Isopoda: Valvifera).—Invertebrate Taxonomy 7:197-278. Samouelle, G. 1819. The entomologists’ useful com- pendium, or an introduction to the knowledge of British Insects. London: Boys. 496 pp. Sars, G. O. 1882. Oversigt af Norges Crustacea.—For- handlinger 1 Videnskaps-selskabet 1 Christiania 18:1-124. Wallerstein, B. R., & R. C. Brusca. 1982. Fish preda- tion: a preliminary study of its role in the zoo- geography and evolution of shallow water ido- teid isopods (Crustacea: Isopoda: Idoteidae).— Journal of Biogeography 9:135-150. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):88—94. 2004. A new species of Synidotea (Crustacea: Isopoda: Valvifera) from the northern Gulf of Mexico Marilyn Schotte and Richard Heard (MS) Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A.; (RH) Department of Coastal Sciences, Gulf Coast Laboratory, PO. Box 7000, Ocean Springs, Mississippi 39566, U.S.A. Abstract.—Synidotea fosteri, n. sp., the sixth known member of the genus Synidotea from the western Atlantic Ocean, is described from shallow waters (1—2 m) adjacent to open beaches in the northern Gulf of Mexico. Its current range extends from western Florida westward to Texas. The new species is distinguished from other related species by small size, fairly straight lateral margins of first pereonite, having the posterior margin of pleotelson straight to very slightly emarginate and by details of the appendix masculina. A key to the known western Atlantic species of the genus Synidotea is also given. Introduction The presence of an undescribed species belonging to the valviferan genus Synidotea has been known from the Gulf of Mexico for over 20 years. Although there are only two published records listed as “Synidotea sp.’ and “Synidotea sp. A’ from Texas and Florida, respectively (Clark & Robertson 1982, Rakocinski et al. 1996), it has also been observed in beach habitats at Grand Island, Louisiana and Gulf Shores, Ala- bama (R. Heard, pers. obs.). More recent collections of Synidotea made near Panama City, Florida, have made possible the de- termination of a new species, which is the subject of this report. In the most recent dis- cussion of the 56 nominal world species of Synidotea, Poore (1996) lists the relevant characters used to differentiate several spe- cies in this genus which closely resemble S. laevidorsalis (Miers, 1881), as does the present new species. Family Idoteidae Samouelle, 1819 Genus Synidotea Harger, 1878 Synidotea Harger, 1878:374; Richardson, 1905:376; Rafi and Laubitz, 1990:2672; Poore and Lew Ton, 1993:261—262. Diagnosis.—Body about twice as long as wide, integument sometimes setose or with sculpturing; cephalon narrower than per- eonite 1; body width greatest at pereonite 4. Pleon lacking articulating pleonites, pleonite 1 indicated by single, small ventro- lateral suture; apex acute, rounded or ex- cavate. Antenna 2 multiarticulate. Mandible with secondary tooth on lacinia mobilis. Maxillipedal palp, with articles 2 and 3 fused, 4 and 5 also fused. Coxae 2—4 with- out dorsal coxal plates; coxae 5—7 with ex- panded dorsal plates. Penes fused complete- ly and swollen distally, attached to posterior margin of pleonite 1. Oostegites forming brood-pouch on pereonites 1—4. Key to the Species of Synidotea from the Western Atlantic Region la. Pleotelson tapers to narrowly rounded, produced apex S. nodulosa lb. Pleotelon faintly to deeply emarginate at apex, not produced 2a. Cephalon bearing two convexities sep- arated by narrow groove and 2 small, medial tubercles anteriorly; lateral mar- gins of pereonites 1—4 angular .... S. littoralis VOLUME 117, NUMBER 1 2b. Cephalon smooth, lacking sculpturing; lateral margins of pereonites not angu- EVR a6 Tate eset ce SN ere ee ree ere one eee 3 3a. Lateral margins of pleotelson almost parallel for first 4 of length; angled me- dially in distal one-third with broad, shallow emargination on distal margin 3 Sea Betirera peeetin eee sey cee eee ears S. brunnea 3b. Lateral and distal margins of pleotelson NOLASTADONE Se en hake ee POR op eee 4 4a. Cephalon with deep medial notch; pleo- telson tapering to very narrow, emar- GMAW BOOK ocooacocccses S. marmorata 4b. Cephalon without deep medial notch, faintly emarginate at most; pleotelson not tapering to narrow apex 5a. Antennal flagellum with 13—20 articles; body length of mature male ca. 12.5 mm; appendix masculina of male not extending beyond apex of endopod of IODOG! D> socssaaacacac S. marplatensis 5b. Antennal flagellum with 7—8 articles; body length of mature male ca. 4.2 mm; appendix masculina extending beyond apex of endopod of pleopod 2 S. fosteri n.sp. Synidotea fosteri, n. sp. Figs. 1, 2 Synidotea sp. Clark & Robertson, 1982:46 (key), 49-50 (Table & Text), 57 (Fig. 5); Rakocinski, et al. 1996:351 (Table). Material examined.—Holotype male USNM 1022910, TL 6.5 mm, from sea grass clumps (origin unknown) in surf/ swash zone, “Bid-a-wee’’ Beach, Panama City) Beachy sElonday 30712727 N- 085°52.5’'W, sal. 34 ppt., coll. R. Heard and J. Foster, 23 Nov 1996; Allotype female USNM 1022911, TL 6.0 mm, same data. Paratypes: 7 males, 6 ovig. females, 53 fe- males, 1 juv., USNM 1022912, same local- ity data. Other material: | male, 7 ovig. females, 2 females, 2 juvs., open gulf off Santa Rosa Beach, northwest Florida, 1—1.5 m, coarse sand with detritus, Sargassum and algae, coll. R. Heard, 15 July 1991. Description.—Male: body length 2.7 89 times greatest width (at pereonite 3) with minutely spinulose integument (Figs. 3C, D) and faint dorsolateral sculpturing on all tergites. Cephalon with faint curved ante- rior groove above slight dome and faint lat- eral grooves, transverse posterior groove deeper; anterior margin of cephalon straight, sometimes with minute medial emargination. Width of head to width of pereonite 4 ratio 0.79. Eyes prominent. Lat- eral angles of first pereonite nearly straight; lateral angles of pereonites 2—4 convex and 5-7 nearly straight, making continuous margin. Lunette on pereonites 2—4 with broadly rounded posterior margin. Coxal plates not discernible dorsally. Sutures sep- arating tergites from coxae faintly visible on tergites 2—4. Pleotelson length to width ratio 1.29; length of pleotelson 0.32 times body length, lateral margins tapering slight- ly to broad apex with straight or very slightly emarginate posterior margin. Uro- podal peduncle with single, oblique ridge; length to width ratio of exopod 0.94, with curve between lateral margin and truncate apex. Antenna 1 with 4 articles, terminal article bearing aesthetascs; antenna 2, single distal plumose seta on 5th peduncle article; fla- gellum with 8 articles. Mouthparts typical of genus, with secondary tooth noted on la- cinia mobilis of mandible. Pereopod 1, palmate propodus bearing many pectinate spine-like setae; end of dac- tyl reaching carpus-merus suture. Pereo- pods 2—4 similar with posterior margins of propodi, carpi, and meri bearing several long and short simple setae. Pereopod 7 with spine-like setae, some pectinate on an- terodistal margins of propodus, carpus and merus. Pereopods lacking dense pads of se- tae. Pleopod 1, peduncle with 5 coupling hooks, 23 and 25 plumose marginal setae on endopod and exopod, respectively. Ple- opod 2, appendix masculina parallel-sided through 90% of its length, tapering to near- ly acute apex, curving laterad, extending slightly beyond apex of endopod; endopod 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. A, male habitus; B, female habitus; C, antenna; D, antennule; E, lateral aspect of male; EK mght mandible; G, left mandible; H, maxilliped; I, second maxilla; J, first maxilla. Scale = 1 mm. bearing 9 plumose marginal setae, exopod bearing 22. Pleopods 3—5 with partial suture on exterior margin of exopod; exopods bearing few setae, endopods none. Fused penial plate weakly waisted, widening somewhat distally with apex evenly, broad- ly rounded. Ovigerous female.—As in male except for sexual characters and length/width pro- portions. Length of body 2.3 times width. VOLUME 117, NUMBER 1 91 Fig. 2. A, ventral pleon; B, pereopod 1; C, pereopod 2; D, pereopod 4; E, pereopod 7; E uropod; G, penial papilla; H, pleopod 1; I, pleopod 2 of male; J, apex of appendix masculina. Length of pleotelson 0.29 times body Etymology—The species is named for length. Length to width ratio of pleotelson Mr. John M. Foster of Gulf Coast Labora- de 32. tory, who collected the new species in the Color.—Specimens in preservation a company of the second author. light red-brown color, pigmentation subtly Remarks.—The Synidotea species S. hir- reticulated overall. tipes H. Milne Edwards, 1940, S. brunnea 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Pires & Moreira, 1975, S. marplatensis Giambiagi, 1922, S. laticauda Benedict, 1897, S. harfordi Benedict, 1897, S. laevi- dorsalis Miers, 1881, S. keablei Poore and Lew Ton 1993, and S. fosteri n. sp. resem- ble each other closely. Based on morpho- logical differences, Poore, 1996 concluded that S. hirtipes, S. laticauda and S. laevi- dorsalis, all from Indo-Pacific coasts, are valid and separate species, not synonyms of the earliest described member of the group (S. laevidorsalis), and do not represent a global invasion thereof, as suggested by Chapman and Carlton, 1991. Poore and Lew Ton, 1993 described S. keablei from Australia, which also superficially resem- bles S. laevidorsalis. But consistently dif- ferent character states again allowed these authors to call into question the conclusion of Chapman and Carlton and their resulting synonymies. Of the western Atlantic species, S. fosteri Scanning Electron Micrographs: A, dorsal view of cephalon, pereonites | and 2; B, lateral margins of pereonites 1—3; C, integument of dorsal pereon; D, close-up of integument. most resembles S. marplatensis and S. brunnea, neither of which were available for direct observation. S$. marplatensis and S. fosteri can be separated by the number of articles in the antennal flagellum (7—8 in S. fosteri, 13-20 in S. marplatensis); rela- tive length of the appendix masculina (ex- tending beyond apex of pleopodal endopod in S. fosteri, shorter than the apex in S. mar- platensis); and the larger size of mature male specimens, e.g., 12.5 mm in the latter vs. 6.5 mm in the new species. Chief dif- ferences separating S. brunnea from S. fos- teri include 13 articles in the antennal fla- gellum (7-8 in S. fosteri), convex margin of pereonite | lateral margin (nearly straight in the new species) and distinct difference in shape of the pleotelson. In S. brunnea these lateral margins are nearly straight then angled medially in the distal third, joined by a broad but shallowly emarginate apex on the distal margin. In S. fosteri the pleo- VOLUME 117, NUMBER 1 93 Table 1.—Comparison of two additional Synidotea species from North America, as addendum to Poore, 1996. Data from our own observations. Maximum length of ovigerous fe- male Maximum length of adult male Color in alcohol Pleotelson length : width in males (number of specimens) Pereon margin Frontal margin of cephalon; dorsal sculpture Head width: pereonite 4 width Pereopod 1 of male Setation of ischium-propodus of pereopods of female Setation of ischium-propodus of pereopods of male Fused penial plate Uropodal peduncle Uropodal exopod: length/width S. fosteri 6.0 mm 6.5 mm red-brown, reticulated 1.29 (6) pereonite 1 nearly straight; 2 and 3 convex, 4—7 straight straight; weak depression in front of eyes 0.79 palm of propodus concave; dactyl reaching carpus-merus suture long and short setae along lower margins long and short setae along lower margins weakly waisted; length: width 1.85; broadly rounded apically 1 oblique ridge curve between lateral margin lat- eral and truncate apex; 0.94 S. harfordi 17.0 mm blotchy yellow-brown; darker me- dial stripe on pereon 1.21 (1) pereonite 1 with subtle curved an- gle; 2—7 making continuous line straight; obvious depression in front of eyes 0.62 palm of propodus concave; dactyl reaching carpus-merus suture dense pads of short setae not waisted; length: width 2.14; rounded apically no oblique ridge curve between lateral margin and truncate apex; 0.88 telson is broadly rounded apically with little or no emargination. The male of S$. brunnea is as yet unknown. S. fosteri can be distinguished from all others in this group by the combination of the nearly rectilinear lateral margins of the first pereonite, which are rounded or convex in most of the others. It is readily separated from S. laevidorsalis by the shape of the fused penial plate, and the longer, narrower pleon in the latter. Mature males of the new species measure 4.2 to 6.5 mm in length, whereas Miers’ type specimens of S. lae- vidorsalis (also male) are longer than 25 mm. Table 1, patterned after Poore’s 1996 comparison of five Synidotea species Indo- Pacific coasts, lists the same morphological data for S. fosteri and S. harfordi to help distinguish this group of similar animals. Ecological notes.—Synidotea fosteri was collected on sand substrata at depths of 1— 2m. All of our records came from sites adjacent to high energy beaches facing the open Gulf of Mexico. Specimens collected and observed during our study occurred be- tween the beach and first or second seaward sand bar. The specimens were always found associated with unattached macro-plant de- tritus or algae. Other peracarids commonly found associated with S. fosteri included the amphipods Micropotopus raneyi Wigely and Atylus urocarinatus Mc Kinney. Acknowledgments We wish to thank John Foster, Sara LeCroy, and Jerry McClelland for making material available for study. Our sincere ap- preciation goes to Scott D. Whittaker, SEM Lab Manager in the Laboratories of Ana- lytical Biology, National Museum of Nat- ural History for technical assistance with the Scanning Electron Micrographs. We also thank Dr. Brian Kensley of the Nation- al Museum of Natural History and two anonymous reviewers for helpful comments on the manuscript. 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Benedict, J. E. 1897. A revision of the genus Synido- tea.—Proceedings of the Academy of Sciences of Philadelphia 1897:387—404. Chapman, J. W., & J. T. Carlton. 1991. A test of cri- teria for introduced species: the global invasion by the isopod Synidotea laevidorsalis.—Journal of Crustacean Biology 11(3):386—400. Clark, S. T., & R. B. Robertson. 1982. Shallow water marine isopods of Texas.—Contributions in Marine Science 25:45—59. Giambiagi, D. 1922. Cuatro nuevos isopodos de la Ar- gentina.—Physis 5:230—244. Harger, O. 1878. Descriptions of new genera and spe- cies of Isopoda, from New England and adja- cent regions.—American Journal of Sciences and Arts (series 3) 15:373—-379. Miers, E. J. 1881. Revision of the Idoteidae, a family of sessile-eyed Crustacea.—Journal of the Lin- nean Society 16(89):1—87. Milne Edwards, H. 1840. Histoire Naturelle des Crus- tacés, comprenant |’anatomie, la physiologie et la classification de ces animaux, vol. 3. Paris. Pires, A. M. S., & P. S. Moreira. 1975. Two new spe- cies of Synidotea (Crustacea, Isopoda, Valvi- fera) from Brazil.—Boletim Instituto Oceano- grafico da Universidade de Sao Paulo 24:45— 67. Poore, G. C. B. 1996. Species differentiation in Syni- dotea ({sopoda: Idoteidae) and recognition of introduced marine species: a reply to Chapman and Carlton.—Journal of Crustacean Biology 16:384—394. Poore, G. C. B., & H. Lew Ton. 1993. Idoteidae of Australia and New Zealand (Crustacea: Isopo- da: Valvifera).—Invertebrate Taxonomy 7:197— 278. Rafi, EF, & D. R. Laubitz. 1990. The Idoteidae (Crus- tacea, Isopoda, Valvifera) of the shallow waters of the northeastern Pacific Ocean.—Canadian Journal of Zoology 68:2649—2689. Rakocinski, C., R. W. Heard, S. E. LeCroy, J. A. McClelland, & T. Simmons. 1996. Responses by macrobenthic assemblages to extensive beach restoration at Perdidi Key, Florida, U.S.A.—Journal of Coastal Research 12:326— 35/2). Richardson, H. 1905. A monograph on the isopods of North America.—Bulletin of the United States Museum 54:1—727. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):95—-105. 2004. A new genus of the Clausidiidae (Copepoda: Poecilostomatoida) associated with a polychaete from Korea, with discussion of the taxonomic status of Hersiliodes Canu, 1888 Ju-shey Ho and Il-Hoi Kim (JSH) Department of Biological Sciences, California State University, Long Beach, California 90840-3702, U.S.A., e-mail: jsho@csulb.edu; (IHK) Department of Biology, Kangreung National University, Kangreung, Kangwondo, 210-702, Korea, e-mail: ihkim@kangnung.ac.kr Abstract.—A new genus and new species of the Clausidiidae (Copepoda: Poecilostomatoida), Hemadona clavicrura, is described based on the specimens obtained from the washings of the polychaete, Dasybranchus caudatus Grube, collected from Namhae-do Island in Korea. The new genus is characteristic in having (1) the 3rd segment of the antenna drawn out to form a sharp claw, (2) a 3-segmented maxilliped in female, and (3) an armature formula of IJ-4 on the third endopodal segment of leg 1. Phylogenetic analysis on the genera of the Clausidiidae shows that Hersiliodes can not be relegated to a synonym of Hemicyclops as proposed in the recent past. It is a sister-taxon with the new genus, Hemadoma, and separated from Hemicyclops in having a 6-segmented antennule, an armature formula of IJ,4 on the distal segment of the endopod of leg 1, and a medial protrusion on the proximal segment of the male max- illiped. Interestingly, the phylogenetic analysis shows also that the three genera (Conchyliurus, Leptinogaster, and Pholadicola) living in bivalve mollusks are monophyletic. Poecilostome copepods of the family Clausidiidae are known to live largely in symbiosis with various marine inverte- brates, such as alcyonarians, polychaetes, mollusks, and callianassid crustaceans. Cur- rently, the family comprises nearly 80 spe- cies in 9 genera. One of its genera, Hersi- liodes Canu, 1888, living in association with polychaetes and bivalves, has been considered almost impossible to separate from Hemicyclops Boeck, 1872 by Bocquet et al. (1963) and Vervoort & Ramirez (1966). Furthermore, Gooding (1963) as well as Humes & Huys (1992) had even advocated the doubtfulness of keeping the genus Hersiliodes as a valid taxon in the Clausidiidae. Nevertheless, in his book on the copepods associated with the marine in- vertebrates of the British Isles, Gotto (1993) treated Hersiliodes as a valid genus of the Clausidiidae and, furthermore, in their re- port on a new species of Hersiliodes from Korea, Kim & Stock (1996) alleged that the genus differs from Hemicyclops in bearing a 6-segmented (instead of 7-segmented) an- tennule and an armature formula of II,4 Gn- stead of I,5) on the third endopodal segment of leg 1. However, it should be pointed out that the former character state is also found in one (out of 38) species of Hemicyclops and the latter, in all nine species of Con- chyliurus. Recently, one of us (IHK) discovered, during his general survey of the symbiotic copepods on Namhae-do Island in Korea Strait, a new genus and species of clausidiid associated with a polychaete. The new form carries, interestingly, some characteristic features of both Conchyliurus and Hersili- odes. Thus, in this paper, in addition to de- 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON scribing this new clausidiid, a phylogenetic analysis of the ten genera of the Clausidi- idae will be conducted to investigate the taxonomic status of the genus Hersiliodes. Materials and Methods The polychaetes, Dasybranchus caudatus Grube (Capitellidae), were dug out from the mud flat and were placed in a plastic bag and fixed with 70% ethanol. Back at the laboratory, water was added into the bag containing the worm fixed in alcohol and then shaken hard to dislodge the copepods. The water together with the sediment and debris were examined under a dissection microscope for associated copepods. The copepods were removed and preserved in 70% ethanol. In studying the preserved co- pepods, the specimens were cleared in lac- tic acid, dissected on a wooden slide (Hu- mes & Gooding 1964), and examined under a compound microscope. All drawings were made with the aid of a camera lucida. For formula of armature, ““A’’ represents aes- thete; Roman numeral, spine; and Arabic numeral, seta. Seven genera were recognized by Humes & Huys (1996) in the family Clausidiidae. They are Hemicyclops Boeck, 1873; Clau- sidium Kossmann, 1874; Hippomolgus G. O. Sars, 1917; Leptinogaster Pelseneer, 1929; Conchyliurus Bocquet & Stock, 1957; Doviella Rocha, 1986; and Hyphal- ion Humes, 1987. However, several chang- es have been made since then; two new genera (Foliomolgus Kim and Pholadicola Ho & Wardle) were added, respectively, by Kim (2001) and Ho & Wardle (1992), Her- siliodes Canu, 1888 was suggested to be resurrected by Kim & Stock (1996), and Doviella was relegated to a synonym of a clausiid genus by Ho & Kim (in press). Thus, including a new genus to be de- scribed below, there are now 10 genera in the Clausidiidae to be considered. The data used in the character analysis to prepare for construction of a matrix were taken from the type species of each of the 10 clausidiid genera. Since Ho’s (1992) phylogenetic analysis of the Poecilostoma- toida shows that Erebonasteridae Humes, 1987 occurs in sister-taxa relationship with a monophyletic clade comprising Clausidi- idae + (Oncaeidae + Paralubbockiidae), Erebonasteridae was accordingly employed as an outgroup to polarize the 14 characters selected and also to root the cladogram(s) in reconstruction of the phylogeny. Al- though Centobnaster Huys & Boxshall, 1990 is generally considered the most prim- itive erebonasterid copepod (Huys & Boxshall 1990), some features in Tychidion Humes, 1973 were found to be even more primitive. Therefore, both Centobnaster and Tychidion were used as outgroup in the polarization of the selected characters shown in Appendix A. Also, in coding mul- tistate characters, when a transformation se- ries containing a single basal bifurcation (dichotomous transformation) was encoun- tered, the method of “‘internal rooting” pro- posed by O’Grady & Deets (1987) was em- ployed. In this case, as shown in Characters 3, 4 and 8 in Appendix B, the coding of “0” indicates apomorphy, not plesiomor- phy. The computer program HENNIGS86 Ver- sion 1.5 (Farris 1988) was employed to an- alyze the phylogenetic relationships among the genera of the Clausidiidae. The com- mand “‘ie*’’ (implicit enumeration) was used to produce multiple, shortest trees through performance of exhaustive search and use all available tree space to find all shortest trees. In order to avoid predeter- mination of the topology of the resultant cladogram(s), all multistate characters were changed to nonadditive (unordered) before employing the command to reconstruct the phylogeny. Description Order Poecilostomatoida Thorell, 1859 Family Clausidiidae Embleton, 1901 Hemadona, new genus Diagnosis.—Body elongate, 9-segment- ed in female and 10-segmented in male. VOLUME 117, NUMBER 1 First pediger fused to cephalosome. Anten- nule short, 6-segmented, with 2nd and 3rd segments incompletely separated. Antenna 4-segmented, with 3rd segment (middle segment of endopod) drawn out into a large claw, distal segment tipped with 7 elements. Labrum well-developed. Mandible tipped with 2 large spiniform elements and 2 setae. Paragnath a lobe with spinules. Maxillule bilobate distally, both lobes tipped with se- tae. Maxilla 2-segmented, with armature formula of 2, 4. Maxilliped 3-segmented in female and 4-segmented in male; proximal segment in male with medial outgrowth. Legs 1—4 biramous with 3-segmented rami; armature formulae generally as in Hemicy- clops, except 3rd segment of leg 1 endopod with II,4 and 3rd segment of leg 4 exopod with III,I,5. Leg 5 2-segmented, armature formula as in Hemicyclops. Basal segment of leg 5 in male fused to pediger. Leg 6 in male a single seta on genital operculum. Caudal ramus with usual 6 elements. Egg sac elongate, multiseriate. Etymology.—TYhe generic name Hema- dona is an anagram of the island Namhae- do located in the Korean Strait from where the new genus was discovered. Gender fem- inine. Type species.—Hemadona clavicrura new species. Hemadona clavicrura, new species Figs. 1-3 Material examined.—3 22 and 7 dd collected from washings of Dasybranchus caudatus Grube collected from intertidal mud flat on Nambhae-do Island (34°49'N 128°03’E) in Korea Strait on 22 July 2001. Holotype 2 (USNM 1013731), allotype ¢ (USNM 1013732), and 6 paratypes (USNM 1013733, including 1 2 and 5 6 <) are de- posited in the U.S. National Museum of Natural History in Washington, D.C. Dis- sected paratypes (1 2 and | ¢G) are kept in the author’s ([HK) collection. Female.—Body (Fig. 1A) elongate, 6.34 mm long (excluding setae on caudal rami). 97 Cephalothorax semicircular and containing Ist pediger. Second pediger widest of body, 1.04 mm; width of 3rd and 4th pedigers de- creasing only slightly from that of 2nd pe- diger. Urosome 5-segmented, 2.37 times longer than prosome. Genital double somite longer than wide, 877 X 693 wm, with ali- form dorsolateral protrusion in anterior half of somite covering area of egg sac attach- ment (Fig. 1A). Abdomen 3-segmented, with all segments longer than wide, 833 X 553 pm, 798 X 508 wm, and 880 X 430 yum. Caudal ramus (Fig. 1C) 4.44 times lon- ger than wide (720 X 162 wm), armed with 1 short, outer seta at about midlength of lateral margin, | short, medial, subterminal seta, and 2 short and 2 long terminal setae; longest terminal seta (830 wm) 1.15 times as long as ramus. Egg sac greatly elongated (7.05 mm), longer than body and cylindri- cal. Rostrum subquadrate in dorsal view, pro- duced forward, and well demarcated from cephalothorax (Fig. 1A). Antennule (Fig. 1B) short and robust, 6-segmented; formula of armature: 5, 16, 10, 4, 2 + A, and 7 + A. Antenna (Fig. 1C) 4-segmented; first segment (coxobasis) longer than wide, with long outer-distal seta; second segment (Ist endopodal segment) shorter than proximal segment, with small subterminal seta; third segment (2nd endopodal segments) drawn out into a large uncinate claw, with basal patch of spinules on outer surface and 2 un- equal setae plus | blunt tip, bent, spiniform seta bearing terminal row of spinules on medial margin; terminal segment 2.75 times longer than wide, tipped with 3 unequal se- tae and 4 spiniform setae structured as that one on 3rd segment. Labrum (Fig. 1D) well-developed, with submarginal, inner, central process, and 2 disjunct, marginal rows of spinules on either side of this pro- cess. Gnathobase of mandible (Fig. 1E) armed terminally with 1 stout, pinnate ele- ment, | stout, spinulose element, and | pin- nate and | naked setae. Paragnath (Fig. 1F) an obtuse lobe fringed with spinules on dis- tal margin. Maxillule (Fig. 1G) bilobate, 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Hemadona clavicrura, new genus, new species, female. A, habitus, dorsal; B, antennule; C, antenna; D, labrum; E, mandible; E paragnath; G, maxillule; H, maxilla. Scale bars: A, 1 mm; B, C, 0.02 mm; D, E 0.02 mm; E, G, H, 0.05 mm. VOLUME 117, NUMBER 1 small outer lobe tipped with | long and 2 short setae and larger inner lobe with 5 un- equal setae. Maxilla (Fig. 1H) 2-segmented; robust proximal segment (syncoxa) armed with | large spiniform and 1 small pinnate setae; distal segment (allobasis) tipped with 2 spiniform elements bearing spinules on one side and 2 pinnate setae. Maxilliped (Fig. 2A) 3-segmented; proximal segment (syncoxa) with 2 unequal medial setae; middle segment (basis) greatly expanded laterally and carrying 2 unequal medial se- tae; terminal segment (endopod) tiny, bear- ing | spiniform and 2 setiform elements. Legs 1—4 (Figs. 2B—D, 3A) biramous, with 3-segmented rami. Formula of spines and setae as follows: Coxa Basis Exopod Endopod Leg 1 O-1 1-1 I-0; 1-1; O-1; 0-1; I,1,4 11,4 Leg 2 O-I 1-0 [-0; I-1; O—1; 0-2; II,15 I1,1,3 Leg 3 O-I 1-0 I-0; I-1; O-1; 0-2; III,15 IL,0,2 Leg 4 O-1 1-0 1J1-0; I-1; O-1; O-2; I,1,5 ILU,1 Outer surface of all segments on rami fringed with spinules. Outer spines on all legs club-shaped, with swollen tip covered with fine denticles. Leg 5 (Fig. 3B) 2-seg- mented; proximal segment small, carrying simple, outer seta; distal segment elongate, about 4 times longer than wide (750 X 187 zm), armed with 3 club-like spines and 1 thin, simple seta. Male.—Body (Fig. 3C) elongate as in fe- male, 3.40 mm long (excluding setae on caudal rami). Cephalothorax semi-ellipsoid shaped and containing Ist pediger. Second pediger widest of body, 532 wm wide; width of 3rd and 4th pedigers decreasing only slightly from that of 2nd pediger. Uro- some 6-segmented, 1.51 times longer than prosome. Ventrally, proximal segment of leg 5 indistinctly separated from its pediger (Fig. 3D). Genital somite slightly longer than wide, 310 X 300 um; genital opercu- lum (Fig. 3D) small. Abdomen 4-segment- 99 ed, with following measurements (proceed- ing from anterior to posterior): 295 x 282 pm, 366 X 275 pm, 317 X 254 pm, and 423 X 246m. Caudal ramus 4.08 times longer than wide (408 Xx 100 wm) and armed as in female. Maxilliped (Fig. 3E) 4- segmented; proximal segment (syncoxa) with large, medial protrusion tipped with 3 sharp tines; second segment (basis) largest, armed with small patch of subterminal den- ticles on lateral surface, a seta in distome- dial corner followed by a row of spinules on medial margin; third segment (lst en- dopodal segment) smallest and naked; distal segment (2nd endopodal segment) drawn out into a long claw with accessory tine and 2 simple setae on medial surface of basal region. Etymology.—The species name is a com- bination of Latin, clava (= a club) and crus or cruris (= leg), alluding to the club- shaped outer and terminal spines on all five pairs of legs. Remarks.—The general appearance of Hemadona clavicrura resembles the species of Conchyliurus in having an elongated (non-cyclopiform) body. They are further alike in having a 6-segmented antennule, an armature formula of II,4 on the terminal segment of the endopod of leg 1, and a prominent medial, basal protuberance on the proximal segment (syncoxa) of the male maxilliped. These four features are also shared with Hersiliodes. However, H. clay- icrura cannot be placed in Conchyliurus due to the presence of the following char- acter states: (1) the hook on the 3rd seg- ment of the antenna is completely fused to its segment proper, (2) the gnathobase of the mandible carries four (instead of three) terminal elements, (3) the proximal seg- ment (syncoxa) of the maxilla bearing two (instead of none) elements at outer-distal corner, and (4) the maxilliped in female is 3-segmented (instead of 2-segmented). Moreover, 10 species of Conchyliurus are known and, unlike H. clavicrura living in association with polychaetes, they were all PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 100 KS LORD ere > VIE £ SS IWS SRS QS Ja = SN if » IANS ZEKXYWNN x Uy Za WN SS KX Ye NOs = \ YY \ = MOO IOS MON a SSS SY QS S SH. ; O 4 — 55S <2 ZEISS SOS SEB SSS NNN OES OSS \ g 2; D, leg 3. Hemadona clavicrura, new genus, new species, female. A, maxilliped; B, leg 1; C, le Fig. 2. Scale bars: A, 0.05 mm; B—D, 0.2 mm. 101 VOLUME 117, NUMBER 1 Weta ae. Zz SS. OO SKE SS ss ers SS KS GE: SSS SS & << SRR KK SSS Z SRE 2 SSS SSX < SoS oss LE x Ss NY, Hemadona clavicrura, new genus, new species. Female: A, leg 4; B, leg 5. Male: C, habitus, dorsal: D, first three somites of urosome, ventral; E, maxilliped. Scale bars: A, B, D, 0.2 mm; C, 0.5 mm: E, 0.05 mm. Fig. 3. 102 Tree 1 f— Foliomolgus j=1 64 f= Clausidium L144 Hemicyclops f— Hemadona | lk-124— Hersiliodes | f= Hippomolgus Tease f= Hyphalion f— Conchyliurus L134 j= Leptinogaster 114— Pholadicola Tree 9 f— Hyphalion PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tree 7 {(— Hippomolgus f=184| (= Conchyliurus | 16 [= Leptinogaster 194| L141 Pholadicola | f= Foliomolgus L174] [= Hemicyclops 1 54| f= Clausidium 134 f= Hyphalion 124) f= Hemadona 414— Hersiliodes (= Pholadicola pel2i Leptinogaster | f=144_ Conchyliurus | j=-16 Hippomolgus L154 7 f= Foliomolgus f— Clausidium L134 Hemicyclops Fig. 4. [= Hemadona L314— Hersiliodes Clausidiid phylogeny produced through analysis of nonadditive (unordered) coding. Showing three representatives from three patterns of phylograms. (Other 15 phylograms are available from JSH upon request.) reported from the mantle cavities of the bi- valve mollusks. Of the four differences mentioned above between H. clavicrura and Conchyliurus, only items (1) and (4) also apply to the dis- tinction between it and Hersiliodes. So far two species of Hersiliodes are known from either a polychaete (Bocquet et al. 1963) or a bivalve (Kim & Stock 1996). Thus, it seems Hemadona is closer to Hersiliodes than to Conchyliurus. In general, H. clavicrura is most char- acteristic in having an unusually long uro- some (2.37 times longer than its prosome) and club-shaped outer and/or terminal spines on all five pairs of legs. Phylogenetic Analysis A total of 18 equally parsimonious trees (cladograms, phylograms) were obtained with a length of 37 steps, a consistency in- dex (CI) of 64 and a retention index of 62. A close comparison of these 18 trees shows that there are three patterns of tree accord- ing to the grouping of the 10 genera. In Pattern I, as Tree 1 in Fig. 4, the 10 genera are separated into two clades, with one clade (Clade 16) containing Clausidium, Foliomolgus, Hemadona, Hemicyclops, and Hersiliodes and the other clade (Clade 17), Conchyliurus, Hippomolgus, Hyphalion, Leptinogaster, and Pholadicola. There are 10 phylograms belonging to this catego- ry—Tree 1, 2, 3, 4, 5, 6, 11, 12, 13 and 14 (authors’ enumeration; unpublished data). Phylograms in Pattern I, as Tree 9 in Fig. 4, have Hyphalion set aside on a clade of its own and the remaining nine genera di- vided into two groups, with Clausidium, Foliomolgus, Hemadona, Hemicyclops, and Hersiliodes in one clade (Clade 15) and Conchyliurus, Hippomolgus, Leptinogaster, and Pholadicola in the other clade (Clade 16). There are six phylograms belonging to VOLUME 117, NUMBER 1 this category—Tree 8, 9, 10, 16, 17 and 18 (authors’ enumeration; unpublished data). One of the two remaining phylograms, Tree 7, belonging to Pattern III, is shown in Fig. 4. It has the 10 clausidiid genera divided into two groups, with one comprising Con- chyliurus, Hippomolgus, Leptinogaster, and Pholadicola and another one, Clausidium, Foliomolgus, Hemadona, Hemicyclops, Hy- phalion and Hersiliodes. The difference among the three patterns mentioned above is chiefly due to the in- consistent positions of Hyphalion. In Pat- tern I (see Tree 1 in Fig. 4), it is a member of the group comprising Conchyliurus, Hip- pomolgus, Leptinogaster, and Pholadicola; in Pattern II (see Tree 9 in Fig. 4) it is by itself; and in Pattern III (see Tree 7 in Fig. 4) it is a member of the group comprising Clausidium, Foliomolgus, Hemadona, Hemicyclops, Hyphalion and Hersiliodes, which is entirely different from the one that it is affiliated with in Pattern I. There are two monophyletic taxa that maintain identical relationships in all 18 phylograms. They are Hemadona + Her- siliodes and Conchyliurus + (Pholadicola + Leptinogaster). The former two genera are held together by sharing characters 1 (with 6-segmented antennule), 12 (proximal segment of male maxilliped with medial protrusion) and 13 (with an armature for- mula of II,4 on distal segment of leg 1 en- dopod), and the latter three genera, by shar- ing characters 3 (with 2 elements on 3rd segment of antenna), 5 (mandible tipped with 3 elements) and 10 (with 3-segmented maxilliped in female). It is noteworthy that both Hemadona and Hersiliodes are char- acteristic in having a 6-segmented anten- nule (Character 1). They were not placed in the same group (clade) with Conchyliurus + Leptinogaster + Pholadicola on any of the 18 phylograms. In other words, the two constant monophyletic taxa are remotely re- lated. It is interesting to point out that the latter three genera comprise parasites of bi- valve mollusks, while species of Pholadi- cola inhabit in the host’s intestine, and 103 those of Conchyliurus and Leptinogaster are found in the host’s mantle cavities. Five of the 18 obtained phylograms con- tain a clade with trichotomy, three of these phylograms (Trees 1, 5 and 12) are in Cat- egory I and the other two (Trees 9 and 17), in Category II. Hemicyclops appears as one of the three terminal clades in all five phy- lograms showing trichotomy. In four of these five phylograms, 1.e., Trees 1, 5, 12 and 9, the branch embracing Hemadona + Hersiliodes appears as another terminal clade, with either Clausidium or Foliomol- gus as the third terminal clade. In addition, Trees 2 and 13 in Category I have a topol- ogy showing Hemicyclops in a sister-taxon relationship with Hemadona + Hersiliodes. These six phylograms indicate that both Hemadona and Hersiliodes are closely af- filiated with Hemicyclops. However, since none of the 18 phylogram shows Hersili- odes in a sister-taxa relationship with Hem- icyclops, the former, accordingly, cannot be relegated to a synonym of the latter. Thus, the present phylogenetic analysis supports Kim and Stock’s (1996) notion that Hersi- liodes is a valid genus in the Clausidiidae and cannot be synonymized with Hemicy- clops. Key to the Genera of the Clausidiidae A key to the genera of the Clausidiidae was provided by Humes and Huys (1992). Since only six of the ten genera currently recognized were dealt with in that key, a new key is provided below. 1. Formula of armature on terminal seg- ment of leg 1 endopod I,5........... 2 — Formula of armature on terminal seg- ment of leg 1 endopod otherwise ..... 5 2. Antenna 3-segmented ........ Ayphalion — Antenna 4-segmented .............. 3 3. Maxilliped in female 4-segmented .. airieaeacdescn: «nea eo aan «ces a Hemicyclops — Maxilliped in female reduced or absent 4 4. Antennule 7-segmented; 3rd segment of antenna with 4 elements ..... Foliomolgus — Antennule 6-segmented; 3rd segment of antenna with 2 elements .... Leptinogaster 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5. Endopods of legs 1—4 with sucking discs; middle exopodal segment of leg | without inner seta Clausidium — No sucking discs on legs; middle exo- podal segment of leg 1 with inner seta ATMO SO Oe oon Oe oo hoe OS 6 6. Proximal segment of maxilla armed with SCtAG! fn, ary 3 oe eR Mi hocle seen entrees 7 — Proximal segment of maxilla unarmed.. 8 7. Maxilliped in female 4-segmented and well-developed; armature formula for terminal segment of leg 1 exopod I,I,5 EES ROR: TRAINS oe Hersiliodes — Maxilliped in female 3-segmented and reduced; armature formula for terminal segment of leg] exopod III,I,5 .. Hemadona 8. Armature formula for terminal segment of female leg 1 endopod II,4 ...... Se Oe LenS otto ate Conchyliurus — Armature formula for terminal segment of female leg 1 endopod otherwise.... 9 9. Maxilliped in female rudimentary ... RNA. ee ENE REAP EN NE ORE 2, Pholadicola — Maxilliped in female well developed, at least 3-segmented Hippomolgus Acknowledgment Studies on this project were aided by a grant from the Paramitas Foundation to the senior author (JSH) and from the Korea Science and Engineering Foundation (2000- 1-20200-003-3) to the junior author (HK). Literature Cited Bocquet, C., J. H. Stock, & G. Kleeton. 1963. Copé- podes parasites d’invertébrés des cotes de la Manche. X. Cyclopoides poecilostomes asso- ciés aux Annélides polychetes dans la région de Roscoff.—Archives de Zoologie expérimentale et générale 102 (Notes et Revue 1):20—40. Farris, J. S. 1988. Hennig86 Reference, Version 1.5.— Published by the author, Port Jefferson, New York, 18 pp. Gooding, R. U. 1963. External morphology and clas- sification of marine poecilostome copepods be- longing to the families Clausidiidae, Clausiidae, Nereicolidae, Eunicicolidae, Synaptiphilidae, Catiniidae, Anomopsyllidae, and Echiurophili- dae.—Unpublished Ph.D. thesis, University of Washington, Seattle, 276 pp. Gotto, V. 1993. Commensal and parasitic copepods as- sociated with marine invertebrates (and whales).—The Linnean Society (London), 264 pp. Ho, J.-s. 1992. Phylogeny of Poecilostomatoida: a ma- jor order of symbiotic copepods.—Bulletin of Plankton Society of Japan, Special Volume Pp. 25-48. , & I.-H. Kim. (2003). New clausiid copepods (Poecilostomatoida) associated with poly- chaetes of Korea, with cladistic analysis of the family Clausiidae.—Journal of Crustacean Bi- ology 22:568-581. , & W. J. Wardle. 1992. Pholadicola intestin- alis, new genus and species, a clausidiid cope- pod parasitic in a deep-burrowing clam from Texas.—Bulletin of Marine Science 51:37—44. Humes, A. G., & R. U. Gooding. 1964. A method for studying the external anatomy of copepods.— Crustaceana 6:238—240. , & R. Huys. 1992. Copepoda (Poecilostoma- toida and Siphonostomatoida) from deep-sea hydrothermal vent areas off British Columbia, including Amphicrossus altalis, a new species of Erebonasteridae, with notes on the taxonomic position of the genus Tychidion Humes.—Ca- nadian Journal of Zoology 70:1369—1380. Huys, R., & G. A. Boxshall. 1990. Discovery of Cen- tobnaster humesi, new genus, new species (Er- ebonasteridae), the most primitive poecilosto- matoid copepod known, in New Caledonian deep waters.—Journal of Crustacean Biology 10:504—519. Kim, I.-H. 2001. Foliomolgus cucullus, a new genus and species of Clausidiidae (Crustacea: Copep- oda: Poecilostomatoida) associated with a poly- chaete in Korea.—Proceedings of the Biological Society of Washington 114:660—666. , & J. H. Stock. 1996. A new species of Clau- sidiidae (Copepoda, Poecilostomatoida) associ- ated with the bivalve Ruditapes philippinarum in Korea. Cahiers de Biologie marine 14:1—6. O’Grady, R. T., & G. B. Deets. 1987. Coding multi- state characters, with special reference to the use of parasites as characters of their hosts.— Systematic Zoology 36:268—279. Vervoort, W., & E Ramirez. 1966. Hemicyclops thal- assius nov. spec. (Copepoda, Cyclopoida) from Mar del Plata, with revisionary notes on the family Clausidiidae.—Zodlogische Mededelin- gen (Leiden) 41:195—220. VOLUME 117, NUMBER 1 105 Appendix 1.—Characters and character states used in the phylogenetic analysis of the Clausidiidae. Numbers in parentheses denote the numerical coding of the character states. Coding in Characters 3, 4, and 8 employed “‘internal rooting’? proposed by O’Grady & Deets (1987). Centobnaster and Tychidion were utilized as the outgroup in polarization of the character state transformations. Ol 3rd and 4th segments of antennule separated (0) or fused (1) 02 Aesthetasc on antepenultimate segment of antennule absent (0) or present (1) 03 3rd segment of antenna with 3 elements (1), 4 elements (0) or 2 elements (2) 04 Terminal segment of antenna with 6 elements (1), 7 elements (0), 5 elements (2) or 4 elements (3) 05 Mandible tipped with 4 elements (0) or 3 elements (1) 06 Inner lobe of maxillule carrying 2 elements (0) or 3 elements (1) 07 Outer lobe of maxillule carrying 3 elements (0) or 4 elements (1) 08 Proximal segment of maxilla without seta (0), with 1 seta (1), 2 setae (2) or 3 setae (3) 09 Distal segment of maxilla with 4 elements (0), 3 elements (1), 2 elements (2) or 1 element (3) 10 Maxilliped in female 4-segmented (0), 3-segmented (1), 2-segmented (2) or absent (3) 11 Proximal segment of female maxilliped with 2 setae (0), 1 seta (1) or none (2) 12 Proximal segment of male maxilliped without protrusion (0) or with medial protrusion (1) 13 Distal segment of endopod on leg 1 with an armature formula of I, 5 (O) or Il, 4 (1) 14 Distal segment of exopod on leg 4 with an armature formula of II, I, 5 (O) or II, I, 5 (1) Appendix 2.—Data matrix of 14 characters and their states in ten genera of Clausidiidae as used in the cladistic (phylogenetic) analysis. The question mark “*?’’ indicates an unknown state. Due to the application of “internal rooting’’ (O'Grady & Deets, 1987) those characters coded with “‘1°’ in the outgroup are treated as plesiomorphic and *‘O”’ in the ingroup, apomorphic. Characters Genus (OTU) 1 5 10 14 Outgroup 0) 0) 1 1 0 0) 0) 1 0 0) 0 0) 0) 0) Clausidium 0 0 0 0 1 1 1 3 0 0) 0) 0 ? 0 Conchyliurus 1 1 2 0) 1 1 1 0) 0 2 1 ] 1 1 Foliomolgus 0) 1 0) (0) 0) 1 1 1 1 3 ey 0) 0) 0) Hemadona ! 0 0) 0) 0) 1 1 2 0 I 0) 1 1 1 Hemicyclops 0 1 0) 0 0) 1 1 2 0) 0) 0) 0) 0) 0) Hersiliodes 1 1 0) 0) 0) 1 1 D 0 0) 0) 1 1 0) Hippomolgus 1 1 i 0 0) i 1 0) 0) 0) 1 ? 0) 0) Hyphalion 1 0 2 v 0) ? ? 1 0 2 2 0) O 0) Leptinogaster 1 1 2 2 1 0 0) 0 D 3 ? 0) 0 0) Pholadicola 1 0) D 3 1 0 0 0 3 3 ? 0 ? ? PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(1):106—113. 2004. Vesicomyicola trifurcatus, a new genus and species of commensal polychaete (Annelida: Polychaeta: Nautiliniellidae) found in deep-sea clams from the Blake Ridge cold seep Jennifer Dreyer, Tomoyuki Miura, and Cindy Lee Van Dover (JD, CLVD) The College of William and Mary, Department of Biology, Millington Hall, Williamsburg, Virginia 23187, U.S.A., email: jcdrey@wm.edu; clvand@wm.edu (ITM) Department of Biological Production and Environmental Science, Faculty of Agriculture, Miyazaki University, 1-1 Gakuen-Kibanadai-Nishi, Miyazaki 889-2192, Japan, email: miura@cc.miyazaki-u.ac.jp Abstract.—A new genus and species of deep-sea polychaete belonging to the family Nautiliniellidae is described from the Blake Ridge cold seep off the coast of South Carolina at a depth of 2155 m. This species is commensal within the mantle cavity of ~60% of the vesicomyid clams collected at the seep site. Vesicomyicola trifurcatus is distinguished from previously described nautili- niellid genera and species by the presence of two pairs of tentacular cirri and up to seven trifurcate hooked chaetae on the posterior parapodia. The new species resembles [heyomytilidicola tridentatus in having trifurcate hooks, but the arrangement and number of chaetae differs. Only two types of chaetae are present in V. trifurcatus: four to seven stout, simple hooks anteriorly to mid- body, and up to seven trifurcate hooks posteriorly. In contrast, there are three types of chaetae in /. tridentatus: up to five stout hooks per parapodium, each with a minute projection on cutting edge of the main fang, 10—20 simple, slender tridenate chaetae, and numerous minute mucronate chaetae. A key to species of Nautiliniellidae is included. The Nautiliniellidae is a small group of deep-sea polychaetes that live in the mantle cavity of a clam or mussel host. Nautili- niellids have been collected from chemo- synthetically based deep-sea habitats, in- cluding cold seeps and hydrothermal vents. Since nautiliniellids were first reported by Miura & Laubier (1989), 10 genera and 14 species have been described (Table 1). Two undescribed species have also been report- ed, one from a cold seep at Barbados Trench (4960 m; Olu et al. 1996) and one off the Pacific coast of Mexico (3221 m; Olu, pers. comm.). An additional genus, Santelma, has been assigned to the family Nautiliniellidae (Blake 1993, Glasby 1993), but its affilia- tion with the Nautiliniellidae remains ques- tionable. The only known species, Santelma miraseta (Fauchald, 1972), was first placed in the family Pilargidae and the genus Pi- largis. Blake (1993) redescribed the species and assigned it to Santelma, a new nautili- niellid genus, based on chaetal similarities. Unlike nautiliniellids, S. miraseta has ex- truded neuroaciculae, a median antenna (or its trace), and it lacks neuropodial hooks and parapodial cirri. Based on these fea- tures, S. miraseta fits better within the orig- inal family Pilargidae (Salazar- Vallejo, pers. comm.). We follow the precedent of Miura & Hashimoto (1996) and exclude S. miraseta from the Nautiliniellidae. Nautiliniellids have reduced and simpli- fied body structures that are associated with a commensal or parasitic life. These modi- fications include a less developed anterior region, the presence of only simple hooked VOLUME 117, NUMBER 1 Table 1—Family Nautiliniellidae: list of genera and species, host bivalve genus and family, collection depth of type specimen, location where type specimen was collected and author reference. Reference Location Depth (m) Host bivalve Genus (Family) Genus and species Blake (1993) Florida Escarpment Okinawa Trough 3303 Unknown 1 Flascarpia alvinae Miura & Hashimoto (1996) Blake (1993) Blake (1993) 1395 3243 Bathymodiolus (Mytilidae) (Mytilidae) Unknown 2 Iheyomytilidicola tridentatus 3 Laubierus mucronatus 4 Miura spinosa Florida Escarpment Santa Maria Basin _ Okinawa Trough 565 625 701 1114 Miura & Hashimoto (1993) Miura & Hashimoto (1993) Miura & Laubier (1990) near Adula (Mytilidae) 5 Mytilidiphila enseiensis Okinawa Trough Sagami Bay Bathymodiolus (Mytilidae) Solemya (Solemyidae) 6 Mytilidiphila okinawaensis 7 Natsushima bifurcata Miura & Hashimoto (1996) Miura & Laubier (1989) Blake (1990) Kagoshima Bay Japan Trench 98 5650: 3700 Solemya (Solemyidae) 8 Natsushima graciliceps Calyptogena (Vesicomyidae) Thyasira (Thyasiridae) 9 Nautiliniella calyptogenicola 10 Petrecca thyasira Laurentian Fan Miura & Ohta (1991) Okinawa Trough Sagami Bay 1400 1170 625 1160 2155 Calyptogena (Vesicomyidae) Calyptogena (Vesicomyidae) Calyptogena (Vesicomyidae) Conchocele (Thyasiridae) 11 Shinkai longipedata 12 Shinkai sagamiensis 13 Shinkai semilonga Miura & Laubier (1990) Miura & Hashimoto (1996) Miura & Hashimoto (1996) Present study Okinawa Trough Sagami Bay 14 Thyasiridicola branchiatus Blake Ridge Diapir Vesicomya (Vesicomyidae) 15 Vesicomyicola trifurcatus 107 chaetae modified to grasp host tissue, and the absence of anal cirri on the pygidium. Diagnostic characters of the family include the number of prostomial appendages, num- ber of tentacular cirri, and chaetal mor- phology and number. These characters are specific to each genus but are useful for species identifications since seven of the ten nautiliniellid genera are monospecific. Based on these morphological characters, we determined that the specimens collected from the Blake Ridge cold seep belong to a new genus and species described herein. Material and Methods Biological samples were collected at the Blake Ridge Diapir site (ODP Site 996; 32°30'N, 76°11'W; 2155 m) on 25 to 28 Sep 2001, using the DSV Alvin. A descrip- tion of the study site can be found in Van Dover et al. (2003). Although geological and chemical properties of this site have been explored during the past decade, the Alvin 2001 samples represent the first col- lections of megafauna and macrofauna from this area. Host clams were collected using a suc- tion sampler. The clams were identified as a new genus and species in the Family Ves- icomyidae, based on morphological char- acters, molecular differences in comparison to described species, and geographic and bathymetric location (E. Kryolora, pers. comm.). Clams were dissected and nautili- niellids were removed and placed into ei- ther 10% buffered formalin or 3% glutar- aldehyde and 0.1 M phosphate buffer with 0.25 M sucrose (pH 7.4). After 24 hours, formalin-fixed nautiliniellids were rinsed and stored in 70% ethanol. Photographs of the external morphology were taken with a compound light micro- scope (LM) and a scanning electron micro- scope (SEM). Specimens for LM were mounted in glycerol and ethanol and ob- served with a Zeiss Axioskop 2 binocular compound microscope. Specimens for SEM were dehydrated through a graded series of 108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ethanol, terminating with 100% ethanol. Samples were then critical-point dried, gold sputter coated (20 nm thick), and observed with an Amray SEM 1810. Images were captured using a Spot camera (Diagnostic Instruments) or a DPI11 digital camera (Olympus). Line illustrations were prepared using a camera lucida attached to a Wild Heerbrugg compound microscope. Systematics Family Nautiliniellidae Miura & Laubier, 1989 Vesicomyicola, new genus Type species.—Vesicomyicola trifurca- tus, new species, by present designation. Diagnosis.—Body with strong dorsal arch, ventrally flattened. Prostomium with one pair of palps, without eyes. Tentacular segment fused with prostomium, with dor- sal and ventral cirri, neuroacicula, and neu- ropodial hooked chaetae. Parapodia sub-bi- ramous, with dorsal and ventral cirri. Noto- and neuropodia each with one embedded acicula. Chaetae absent on notopodia. Two types of chaetae present on neuropodia: simple hooked chaetae on anterior segments (some with single subapical tooth present on anterior to mid-body segments), and tri- curcate hooked chaetae on posterior seg- ments. Pygidium cylindrical, without anal cirri. Gender.—Masculine. Etymology.—TYhe generic name is de- rived from the name of the host vesicomyid clams these polychaetes inhabit. Vesicomyicola trifurcatus, new species (Figs. 1—4) Type material.—Holotype (ODP Site Woe S2SOMIN, WO We QiSS ma, Bs} Seo 2001, Alvin Dive 3712; USNM 1016220) and five paratypes (USNM 1016221) from same dive and date were deposited in the collections of the National Museum of Nat- ural History, Smithsonian Institution, Washington, District of Columbia. An ad- ditional five paratypes, each from the same dive and date, were deposited in the Mu- seum National d’Histoire Naturelle, Paris (MNHN POLY TYPE 1405) and the Na- tional Science Museum, Tokyo (NSMT— Pol P 458). Additional material.—Voucher speci- mens were retained in the collection of CLVD in the Department of Biology at The College of William and Mary. Description.—Holotype female, oviger- ous, measuring 8.4 mm long, 1.3 mm wide, including parapodia, with 37 segments. Paratypes ranging from 4.4—12.7 mm long, 0.8—1.6 mm wide, including parapodia, and with 28—41 segments. Body flattened ven- trally, arched dorsally. Some live specimens with green pigment in parapodia, others with pale pink color; preserved specimens in alcohol pink to white in color. Some pre- served females pale green; internal oocytes evident through transparent parapodial epi- dermis. Preserved holotye and paratypes curled (Fig. 1A). Prostomium rounded, with palps (Fig. 2A-C). Eyes absent. Tentacular segment fused with prostomium, with one pair of dorsal and ventral cirri, neuroacicula, and neuropodial hooked chaetae (Fig. 2C). Foregut with well-developed muscular re- gion (Fig. 2A—C). Pygidium rounded, with- out anal cirm (Fig. 2D). Parapodia subbiramous, with dorsal and ventral cirri. Dorsal cirri with inflated base and tapering tip, twice as long as ventral cirri. Notopodia with single embedded acic- ula, lacking chaetae (Fig. 3A). Neuropodia with a single bent acicula and hooked chae- tae (Fig. 3B). Neuropodial hooks of two types. Ante- rior neuropodia with simple stout hooks with recurved tips, four to seven on each parapodium (Fig. 4A, B), some anterior to mid-body chaetae with single small apical tooth near tip, appearing slightly bifid (Fig. 4C). Posterior neuropodia with thinner, sim- ple hooks with trifurcate tips, up to seven per neuropodium (Fig. 4D, E). Etymology.—The specific name comes VOLUME 117, NUMBER 1 Fig. 1. body. from fri- = three times, + furcatus = forked, in reference to the trifurcate chaetae present on the posterior segments. Biology.—The mantle cavities of ~60% of the Blake Ridge clams sampled contained one to five nautiliniellid polychaetes. Carbon and nitrogen stable isotope compositions of worm and clam tissues were consistent with a parasitic life-style for the worm, but the sulfur isotope composition of the worms was so distinct from that of the clams that an alternative diet must be inferred (Van Dover et al. 2003). Van Dover et al. (2003) pro- posed a feeding strategy whereby ciliary ac- tivity of the clam gills moves sufficient vol- umes of seawater to allow the polychaetes to collect and consume suspended organic particles either from gill mucus or from a worm-generated mucus net. Discussion Vesicomyicola trifurcatus resembles spe- cies in the genera Nautiliniella, Natsushi- 109 Vesicomyicola trifurcatus new genus, new species. Scanning electron micrograph (SEM) of whole ma, Shinkai, and Thyasiridicola, based on shared characters of the tentacular segment, which in these four genera includes dorsal and ventral cirri and neurochaetae (with the exception of the genus Thyasiridicola, which lacks neurochaetae). The genus Ves- icomyicola differs from these four genera in the number and morphology of the neuro- podial chaetae. Vesicomyicola trifurcatus resembles Iheyomytilidicola tridentatus Miura & Hashimoto, 1996 based on the trifurcate chaetal morphology, but the arrangement and number of chaetae on the parapodia dif- fers. There are only two types of chaetae present in V. trifurcatus: stout, simple hooks (four to seven; sometimes bifid) on the anterior to mid-body parapodia, and tri- furcate hooks (up to seven) on the posterior parapodia. In contrast, there are three types of chaetae in /. tridentatus: stout hooks (up to five), each with a minute projection on the cutting edge of the main fang; simple, 110 Fig. 2. view. B. Drawing of anterior end, dorsal view. C. Drawing of anterior end, ventral view. D. LM of pygidium, dorsal view. slender tridentate chaetae (10—20); and nu- merous minute chaetae with mucronate tips (Miura & Hashimoto 1996). Based on its unique set of morphological characters, we consider V. trifurcatus to be a new genus and species. A key to nautili- niellid species is provided to aid in identi- fication; most species are location and host specific. The terminology and interpretation of prostomial appendages in this family is the subject of some debate (Blake 1993, Miura & Hashimoto 1996), suggesting the need for a re-evaluation and revision of this fam- ily and its genera once a consistent diag- nosis of prostomial appendages can be ap- plied. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 100pm B SF Vi O cae e Fy me SSS \\ SS 100pm G Vesicomyicola trifurcatus new genus, new species. A. Light micrograph (LM) of anterior end, dorsal Color dimorphism was a distinctive char- acter of live V. trifurcatus, but on preser- vation the color variation was lost. Poly- chaetes with green parapodia in new col- lections (2003) were all gravid females. In other nautiliniellid species, color dimor- phism corresponds to sexual dimorphism (Miura & Hashimoto 1996, Miura 1998). We have yet to confirm that the pale colored specimens are males. With the discovery of each new species in the Nautiliniellidae, we learn more about the ecology of these worms and their relationship with their host bivalves; we still know little about the in- ternal anatomy, reproductive biology and larval characteristics, or the trophic ecology of this polychaete family. VOLUME 117, NUMBER 1 Fig. 3. 100pm A Vesicomyicola trifurcatus new genus, new species. A. Drawing of mid-body parapodium with em- bedded aciculum and dorsal cirrus; lateral view. B. Drawing of mid-body neuropodium and ventral cirrus; ventral view. Key to the species of Nautiliniellidae la. 1b. 2a. 2b. 3a. 3b. Aa. Ab. Sa. Sb. Prostomial appendages (palps or an- tennae) absent BECUASE cid dtten ee sei Miura spinosa Blake, 1993 One or two pairs of prostomial ap- pendages present ................ 2 Tentacular segment with only one pair GUGTIUO A Se Bian GG Eee Orono olan rets 3 Tentacular segment with one pair of dorsal and ventral cirri ........... 7 Tentacular segment with or without neurochaetae; all neuropodial hooks SIEM SLE a a en eet ae alee tis le eis 4 Tentacular segment without neuro- chaetae; some neuropodial hooks stout ote Graken ero ot orem acne ee 5 Neurochaetae 220 (up to 35) per parapodium; neurochaetae with inflat- ed, subdistal stems and _ slightly curved, pointed distal ends ...... Mytilidiphila enseiensis Miura & Hashimoto, 1993 Neurochaetae =20 per parapodium; neurochaetae with rounded tips and slightly curved, distal ends Mytilidiphila okinawaensis Miura & Hashimoto, 1993 Only one type of neurochaeta present: large, stout hooks Two types of neurochaetae present: Yon 6a. 6b. 7a. 7b. 8a. 8b. 8c. 111 100pm B One to two large, stout hooks and 15— 20 small, mucronate tipped chaetae Gin crows of 2) ..... Laubierus mucronatus Blake, 1993 Three types of neurochaetae present: 8 mm long, not oblanceolate 2. Cymule bracts ovate, elliptic or obovate, scarcely broader than long, not leaf-like; corollaideepy pina eee eee 3 2. Cymule bracts linear to linear-(ob-)lan- ceolate, sometimes with a leaf-like apex, always much broader than long; corolla red, orange or yellow 3. Cymule bracts ovate, 6-15 mm wide; leaves pubescent below D. scutellata 3. Cymule bracts elliptic to obovate, 3-6 mm wide; leaves almost glabrous below D. cochabambensis 4. Cymule bracts usually leafy at the apex; flowers usually in dense, sessile clusters or heads in the leaf axils .... D. squarrosa 4. Cymule bracts not leafy at the apex; flowers in pedunculate, axillary cymes, often forming a leafy panicle 5. Cymule bracts narrowly linear-elliptic, broadest in the middle; leaves softly pu- bescent above D. palmariensis 5. Cymule bracts linear-lanceolate, broad- VOLUME 117, NUMBER 1 est at the base; leaves soon glabrescent above 6. Inflorescence branches leafless, usually short; bracts linear-oblong, acute; corolla lobes almost half as long as the tube .... Schatten Amare de eike a) ea ein D. jujuyensis 6. Inflorescence branches often with sub- tending leaves, often well-developed; bracts lanceolate, finely acuminate; co- rolla lobes less than one fourth as long as the tube D. purpurascens Dicliptera palmariensis Wassh. & J. R. I. Wood, sp. nov. Fig. 1 A-H Quoad formam bractearum cymulorum Diclipteram garciae Leonard tangit, ob fo- lia pilis lanatis induta, bracteas acutas, non apiculatas ab ea removendum. Ascending or weakly erect, much- branched, probably perennial herb to 0.75 m; stems dark purplish-green, obscurely ridged with paler vertical lines along the de- pressions, densely pilose with long, patent, straggly, multicellular trichomes; leaves petiolate, the petioles 0.3—1.6 cm long, pi- lose, the blades ovate or elliptic, acute at apex, narrowed to the base and = attenuate on the petiole, 3—9 cm long, 1—4 cm wide, both surfaces pilose with large-celled tri- chomes, especially on the veins, cystoliths abundant above, the margin entire or ob- scurely repand, ciliate; inflorescence of pe- dunculate cymes in the axils of the upper leaves, the cymes typically few-flowered and the flowers often aborting, the inflores- cence thus rather lax and open; peduncles 1-10 cm long, subtending bracts leaf-like, shortly petiolate, the petioles 3—5 mm long, the blades lanceolate or lanceolate-elliptic, acute, 0.7—2 cm long; cymules pedicellate, the pedicels ca. 0.5 mm long; cymule bracts slightly unequal, 8-15 mm long, 3 mm wide, narrowly oblong-elliptic, acute at both ends, pilose, the base often pale; inner bracts 6—10 mm long, lanceolate, ciliate on the upper margins; bracteoles lanceolate, ca. 4 mm long; calyx 2.5—3 mm long, 5- lobed to just above the base, the lobes 141 equal, ca. 2 mm long, lanceolate, acute, cil- iolate; corolla red, 25—28 mm long, cylin- drical from a slightly bulbous base, gradu- ally widened to ca. 3 mm, pilose without, 2-lipped, the lips ca. 3 mm long; anthers equaling the corolla; filaments 14 mm long, sparsely pilose, inserted ca. 14 mm above the base of the corolla, anther thecae at dif- ferent heights, glabrous, ca. 1.25 mm long; ovary pubescent; style ca. 25 mm long, with a few scattered trichomes; capsule 6 mm long, obovoid, pubescent, 2-seeded; seeds with a few, short trichomes, lenticu- lar, ca. 2.25 mm wide. Type.—BOLIVIA: Tiraque, 1—2 km above El Palmar along the old road from the Chaparé to Cochambamba, 900 m, 6 Jul 1997, J. R. I. Wood 12403 (holotype K!; isotypes LPB, US!). Additional specimens.—BOLIVIA: Ti- raque, 1—2 km above El Palmar along the old road from the Chaparé to Cochabamba, 1200 m, 6 Jun 1998, J. R. I. Wood 13674 (K, LPB, US); El Palmar, 155 km along old road from Cochabamba to Villa Tunari [LSS GAB WO tn, 2 Seo ISIS Kessler et al. 8115 (GOET, LPB, US). The only possible Bolivian species D. palmariensis could be confused with is D. purpurascens but it can readily be distin- guished by its diffuse ascending habit, very pubescent indumentum, pedunculate cymes, smaller corollas and above all by the short- er, narrowly oblong-elliptic bracts, broadest in the middle and narrowed to both an acute apex and base. However, there are two col- lections from San Martin Department in Central Peru (Schunke 3349 and 4370, both at K and F), which are in many ways in- termediate between D. palmariensis and D. purpurascens. The bracts are similar in shape to those of D. palmariensis but the apex is acuminate and apiculate and the specimens lack the distinct pilose indumen- tum of D. palmariensis. Given the wide variation found in many species of Diclip- tera these specimens might suggest D. pal- mariensis should be included in a very var- lable D. purpurascens but they are far re- 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a \\ y NY y q WY j ves PA WN IPNYIVE \ i a AWA NWA We ~ I: NV LEZ Fig. 1. A-H, Dicliptera palmariensis (J. R. I. Wood 12403). A, Habit; B, Pedunculate cymes; C, Calyx and corolla; D, Inner bract, bracteoles, calyx and aborted flower; E, Calyx and pistil; K Calyx lobes and nectar disk; G, capsule; H, Capsule dehisced. VOLUME 117, NUMBER 1 moved geographically from D. palmarien- sis and are not exactly intermediate between the two recognized species. It seems best, therefore, to recognize the two species particularly as there are no inter- mediates in Bolivia. Dicliptera purpurascens Wassh. & J. R. I. Wood, sp. nov. Fig. 2 A-F Species nova plerumque purpurascens bracteis longis (usque 2.5 mm) lanceolatis, long-acuminatis bene distincta. Annual or short-lived perennial herb, 0.5—2.5 m high, usually erect in open situ- ations but commonly ascending or even de- cumbent in moister, shady conditions; stems stout, somewhat woody below, strongly an- gled, usually purplish, scurfy-pubescent, much branched; leaves petiolate, the peti- oles 0.5—4 cm long, scurfy-pubescent, the blades equal or nearly so, ovate or ovate- elliptic, 4-15 cm long, 2—7 cm wide, acute or shortly acuminate at apex, tapering at the base, entire, often purplish, darker green above than below, glabrous except for the usually ciliolate margins and a few scat- tered, usually multicellular trichomes, es- pecially on the veins, cystoliths scattered on both surfaces; inflorescence of shortly pe- dunculate or subsessile, axillary and ter- minal cymes, these becoming very dense on older plants with 1—3 cymes arising from each axil, commonly purplish and glandu- lar-pilose but sometimes greenish and very thinly pilose; peduncles 0-3 cm long, scurfy-pubescent; subtending bracts leaf- like, petiolate, the petioles 0-3 mm long, the blades typically narrowly oblong-ellip- tic, to 3 cm long; cymules pedicellate, the pedicels 0-4 mm long; cymule bracts slightly unequal, 20—25 mm long, 2-5 mm wide, lanceolate, long-acuminate; inner bracts linear-acuminate, 12-15 mm long; bracteoles similar but only to 10 mm long; calyx 3—4 mm long, 5-lobed to just above the base, the lobes subulate, minutely cili- olate; corolla orange-red, 34-40 mm long, 143 cylindrical from a slightly bulbous base, gradually widened to 3—4 mm, sparsely pi- lose and minutely gland-dotted without, 2- lipped, the lips ca. 4 mm long; anthers equaling the corolla; filaments 17 mm long, sparsely pilose, inserted ca. 13 mm above base of corolla, anther thecae at different heights, glabrous, ca. 1.5 mm long; ovary pubescent; style ca. 29 mm long, glabrous; stigma globose; capsule 7 mm long, 4 mm wide, obovoid, pubescent, 2-seeded; seeds papillose, lenticular, ca. 1.25 mm wide. Type.—BOLIVIA: Carrasco, ca. 5 km E of Valle de Sajta on main road from Chi- moré to Santa Cruz, 240 m, 29 May 1996, Wasshausen, Brummitt, Wood & Ritter 2067 (holotype US!; isotypes K!, LPB). Habitat and distribution.—Dicliptera purpurascens is locally frequent in moist lowland rain forest between 200 and 600 m in Bolivia and Peru. It is essentially a plant of the SW basin of the Amazon River with an outlying population in a very moist area of the Andean foothills in Bolivia. It has not yet been found in Brazil but is likely to occur in Acre as well. This disjunct distri- bution is shared with a number of other Acanthaceae species, notably Pachystachys spicata (Ruiz & Pavon) Wassh., Ruellia in- flata Rich., R. yurimaguensis Lindau, Jus- ticia megalantha Wassh. & J. R. I. Wood (in press), J. pilosa (Ruiz & Pavon) Lindau and J. riedeliana (Nees) V. A. W. Graham and appears to be a common pattern. Additional specimens.—BOLIVIA: San- ta Cruz, Ichilo, by track from Escuela Ichilo to Campamento Ichilo on E side of Rio Ich- ilo, Ambor6 Park, 400 m, 27 Jul 1999, J. R. I. Wood 14943 (K, LPB); Cochabamba, Carrasco, Valle de Sajta, 1 Jul 1988, Hen- sen 6 (BOL, US); km 228, Santa Cruz road, Rio Murillo, Valle de Sajta, 212 m, 18 Jul 1990, Sigle 510 (US); Experimental Sta- tion, Valle de Sajta, 280 m, 11 Aug 1990, I. Vargas 673 (LPB, USZ); Valle de Sajta, ca. 235 km NW of Santa Cruz, 400 m, J. R. I. Wood 10072 (K, LPB, US); 0.5 km E of Valle de Sajta, 250 m, 29 May 1996, J. R. I. Wood 11178 (K, LPB); 12 de Julio, 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ——— EE 2 = << J Fig. 2. A-K Dicliptera purpurpascens (Wasshausen 2067). A, Habit; B, Pedunculate cymes; C, Inner bracts, bracteoles, calyx and pistil; D, Calyx and pistil; E, Corolla; R Nectar disk, pistil and calyx lobes. VOLUME 117, NUMBER 1 ca. 9 km S of Israel, on E side of Rio Sajta, 400 m, 24 Jul 1999, J. R. I. Wood 14893 (K, LPB); Zona del Arroyo de 6 de Agosto, Cerro de la Concordia, E bank of Rio Ichoa, 600 m, 27 Jul 1999, J. R. I. Wood 14935 (K, LPB); Pando, Abuna, Nacebe, Rio Or- ton, 11 Oct 1989, Beck et al. 19283 (LPB, US); Gentry et al. 77583 (MO, US). PERU; Cuzco, Convencion, along Rio Pichari, 2 km E of Colonizacion Pichari, 620 m, 13 Jun 1975, Wasshausen & Encarnacion 544 (K, US); Paucartambo, Kosfipata District, along trail behind and W of Pilcopata, 580 m, 26 Jun 1975, Wasshausen & Encarna- ci6n 583 (K, US); Quispicanthis, 3 km E of Quincimil, 960 m, 7 Oct 1976, Wasshausen & Encarnacion 735 (US); Madre de Dios, Manu, Adan Rajo, km 225, Shintuya-Pil- copata, 520 m, 26 Jun 1975, Wasshausen & Encarnacion 575 (K, US); Talmamanu, Chilifas, km 18 on Iberia-IMapari road, 1 Jun 1978, Encarnacion 1169 (K, US); near Shintuya, along Alto Rio Madre de Dios, 450 m, 13 Oct 1979, Gentry et al. 26736 (MO, US); Manu, Parque Nacional de Manu, Est. Cocha Cashu [11°50’S,71°25’ W], 350 m, 4 Aug 1984, Foster 9746 (MO, US); Explorer’s Inn, near confluence of Rio Tambopata and Rio La Torre, 29 km SW of Puerto Maldonado [12°50’S, 69°20’W], 9 Jul 1987, Smith, Smith & Condon 938 (K, US); Rio Tambopata [12°48’S, 69°17'W], 200 m, 9 Jul, 1998, Michelangeli 477 (US); Ayacucho, La Mar, on trail between Santa Rosa and Sanabamba along Rio Santa Rosa, 700 m, 9 Jun 1975, Wasshausen & Encar- nacion 531 (K, US). There is considerable variation in the in- dumentum and color of Dicliptera purpur- ascens. The purple colored form is the only form found in central Bolivia in the De- partments of Cochabamba and Santa Cruz while only green forms are known from Pando. Both forms occur in Peru but the purple one is a good deal more common. All plants from Bolivia are densely glan- dular-pilose. In Peru plants are more com- monly glandular-pilose but thinly pilose forms also occur. 145 Dicliptera purpurascens is obviously re- lated to D. palmariensis but the two species are immediately distinguished by the dif- ferent bracts. Dicliptera squarrosa Nees Dicliptera squarrosa Nees, in Mart., Fl. Bras. 9:161. 1847. Type: Brazil, Minas Ger- ais, Reidel 34 (lectotype, here chosen, GZU!; isolectotype NY!); sin loc., Schiich S.n. (Syntype W, not seen). Dicliptera sericea Nees, in Mart., FI. Bras. 9:162. 1847. Type: Brazil, Sao Paulo, Sorocoba, Riedel & Lund 1984 (lectotype, here chosen, LE!; isolectotype NY!). Dicliptera pohliana Nees, in Martt., FI. Bras. 9:162. 1847. Type: Brazil, Minas Ger- ais, Tazenda de Roma, Pohl 2973 (lecto- type, here chosen, W!). Dicliptera tweediana Nees, in DC., Prodr. 11:482. 1847. Type: Uruguay, Porto Alegre, Sellow 13 (d585) (syntype B, de- stroyed); ibid, Sellow 16 (d531) (syntype B, destroyed); Argentina, Buenos Aires, 7Twee- die s.n. (syntype K!). Dicliptera niederleiniana Lindau, Bot. Jahrb. 19, Beibl. 48:18. 1894. Type: Argen- tina, Entre Rios, Primer Misionero de Her- nandez, Puck & Fernandez 42 (holotype B, destroyed ?). Dicliptera imminuta Rizzini, Arquiv. Jard. Bot. Rio de Janeiro, 8:348. 1948. Type: Brazil, Santa Catarina, Reitz, C861 (holotype RB). Dicliptera rauhii Wassh., Beitr. Biol. Pflanzen 63:425. 1988. Type: Peru, Cuzco, prov. Urubamba, Machu Picchu, Rauh & Hirsch P804 (holotype HEID!). Dicliptera squarrosa is an exceptionally widespread species extending from Brazil south of the Amazon region westward to the eastern slopes of the Andes in Bolivia and then southward to Uruguay and central Argentina. Its occurrence further north is uncertain although we feel that Dicliptera rauhii Wassh. from Peru belongs to this species and probably also several species described by Leonard from Colombia. D. 146 squarrosa is very variable with a welter of different forms throughout its range all in- tergrading with each other and forming no discrete units except perhaps at a very local level. We can make out the following rather imprecise geographical forms: Form 1J.—Plants from Argentina, Uru- guay and Paraguay corresponding to the types of D. tweediana and D. pohliana have glabrous, narrowly lanceolate, obtuse leaves and relatively few-flowered axillary cymes, which become congested above into a terminal thyrse. This form does not occur in Bolivia but some Argentinian plants, es- pecially from the Tucuman region have broader leaves which approach form 4 (be- low) found in Bolivia although the leaves always appear to be glabrous. Form 2.—Fig. 3 A—G. Some populations in the Rio Unduavi Valley along the road from La Paz to Sud Yungas appear very distinct. These plants have subglabrous leaves and a relatively long inflorescence of axillary cymes forming many distinct pseu- doverticels, which are not confluent above. The corollas are yellow and the cymule bracts are oblong, gray-pubescent and cili- ate-margined with distinct squarrose tips. Collections corresponding to this form in- clude: BOLIVIA: La Paz, Nor Yungas, on N side of Rio Unduavi valley, on road to Sud Yungas, 2200-2400 m, 9 Jul 1974, Wood 8596 (K, LPB, US); ibid, 2400 m, 1 Jul 1995, Wood 9952 (K, LPB); ca. 2 km above El Velo de la Novia on the Sud Yun- gas road, 2400 m, 14 Jun 1998, Wood 13716 (K, LPB); Sud Yungas, km 66 on Sud Yungas road to Puente Villa ca. 50 m from El Castillo, 1830 m, 12 Jun 1996, Wasshausen & Brummitt 2123 (CAS, GOET, K, LPB, US). However, forms similar to form 2 but with reddish-orange corollas and bracts with few or no cilia occur elsewhere in the La Paz region and also in Peru. All these forms are difficult to distinguish from Di- cliptera scandens Leonard from Colombia except that they bear no field notes to sug- gest they are scandent. Even D. scandens PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON itself is not always scandent. Collections which conform to this more broadly-de- fined form 2 include: BOLIVIA: La Paz, Tamayo, on descent into Rio Yuyo, ca. 60 km S of Apolo on road to Charazani, 1150 m, 12 Jun 2000, Wood & Wendleberger 16438 (K, LPB); Murillo, 29.3 km NE of the summit along Zongo Valley, 2200—2300 m, Solomon, Luteyn & Dorr 19068 (LPB, MO, US); Zongo Valley, 1900 m, 28 Jun 1997, Wood 12349 (K, LPB); Sud Yungas, ca. 15 km from Huancané on road to San Isidro, 2300 m, | Jul 1995, Wood 9964 (K, LPB, US); 2 km E of Puente Villa, 1200 m, 12 Jun 1996, Wasshausen & Brummitt 2124 (K, LPB, US); Inquisivi, Lewis 39127 (LPB, MO, US); Cochabamba, Ayopaya, 1 km above Independencia, 2500 m, 13 May 2000, Wood & Zaraté 16339 (K, LPB); Co- chabamba, Ayopaya, 4 km S of Saila Pata, Kessler 12364 (LPB, US). PERU: La Mer- ced-Oxapampa, 2300 m, 17 Aug 1976, Palmer 44 (K); San Martin, Zepalacio near Moyobamba, 1200-1600 m, Mar 1934, Klug 3601 (& K). Form 3.—In the northern Bolivian An- des, mostly at lower altitudes and particu- larly in areas of high rainfall, there is an- other form. This also has glabrous leaves but the bracts are relatively broad, leaf-like and mucronate, usually elliptic or obovate, never ciliate or squarrose but commonly pubescent to subglabrous. The axillary cymes are relatively few-flowered. Speci- mens that conform to this form include: BOLIVIA: Pando, W bank of Rio Madeira, 3 km above Riberao, 27 Jul 1968, Prance et al. 6539 (K, NY, US); Beni, Ballivian, 10 km S of Rurrenabaque, 250 m, 29 Jul 1998, Wasshausen & Wood 2162 (US, LPB); La Paz, Caranavi, 2 km up road be- hind Caranavi, 640 m, 10 Jun 1996, Was- shausen et al. 2118 (K, LPB, US); Sud Yungas, Santa Ana de Alto Beni, 580 m, 20 Aug 1963, Holliday 26 (K); 7.5 km N of end of Road to San José, 26.5 km along road to La Asunta, 1040 m, 5 Aug 1991, Acevado et al. 4451 (K, US); stream at bot- tom of ascent to Huancané, ca. 5 km from 147 VOLUME 117, NUMBER 1 ———- AVY 4 Annual loss in value of Endowment. ik me ee AE ops ne) eT Ve To 4 hoe tao x Nes 1 ' ) oe y 4 a is a _) el, ‘ie eng : ‘si st Nz Mi " - i i LL Pu! é a) ce # Pai Hct Ay Met i aa eaigetom ee ' 9 shames ae iii mig el Whe gitinny hon’ oy iN Tigcagiie Penna ngedarcetl) gb ae in wht oilirenen ‘wan ieee i) N a ° \ a er es a tits 4 mm; antenna 1 distal article with Isr GESINEAHSES 5 cccccnnn B. cavernarum (caves, southcentral Texas, unconfirmed in southeastern New Mexico) 4a. Male pleopod 1 exopod tapering to a single point (figs. 5a, 6c) 4b. Male pleopod | exopod tip with two processes (fig. 6d) 5a. Male pleopod 1 exopod tapering to a digitiform process, antenna | aesthe- tascs 8 B. villalobosi (caves, Veracruz) 5b. Male pleopod 1 exopod tapering to a point (figs. 5a, 6c), antenna | aesthe- tascs 5—6 (fig. 2c) 6a. Maxilliped palp basal segment broad, obscuring endite (fig. 3d); male pleo- pod 1 exopod without subapical knobs (fig. 6c) B. heroldi (humus and caves, southern California) 184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Brackenridgia acostai, Cueva del Ticho, Chiapas: (a) head and pereonites 1-3; B. cavernarum, Kappelman Salamander Cave, Comal Co., Texas: (b) pleopod 1 exopod tip; B. heroldi, Crystal Cave, Sequoia National Park, California: (c) pleopod 1 exopod tip; B. reddelli, Valdina Farms Sinkhole, Medina Co., Texas: (d) pleopod | exopod tip. 6b. Maxilliped palp basal segment narrow, not obscuring endite (fig. 3c); male ple- opod 1 exopod with low, subapical MOOS (UG, SA) scoososccvcvcs B. ashleyi (Tumbling Creek Cave, Missouri) 7a. Male pleopod 1 exopod tapering to two acute spines, antenna | aesthetascs 14 .. B. bridgesi (caves, northeastern Mexico) 7b. Male pleopod 1 exopod tapering to an acute spine and a second brush-like process (6d); antenna | aesthetascs 9— 10 (fig. 2f) B. reddelli (caves, southcentral Texas) Acknowledgments I thank Mr. Thomas Aley, Dr. David Ashley, Dr. William Elliott, Dr. John R. Holsinger, Dr. Joan Jass, Dr. Brian Kensley, Dr. William Muchmore, Dr. Christian Schmidt, Ms. Marilyn Schotte, Dr. George Schultz and Dr. Stefano Taiti for reading the manuscript and making suggestions for its improvement. The loan of specimens from the collections of the Smithsonian Institu- tion was kindly provided by Ms. Marilyn Schotte and Dr. Brian Kensley. The descrip- tion of B. ashleyi was funded by the U.S. Fish & Wildlife Service, The Nature Con- servancy and the Ozark Underground Lab- oratory. Literature Cited Aley, T. 1975. Biology.—Ozark Underground Labo- ratory Newsletter, 7(4)/8(1):1. Arcangeli, A. 1929. Isopodi terrestri raccolti in Cuba dal Prof. E Silvestri—Bollettino del Laborato- rio di Zoologia Generale e Agraria della R. Scu- ola Superiore di Agricoltura di Portici 23:129— 148. Black, J. H. 1971. Cave life of Oklahoma.—Oklahoma VOLUME 117, NUMBER 2 Underground (Central Oklahoma Grotto, Na- tional Speleological Society) 4(1 & 2):1—5S6. Craig, J. L. 1975. A checklist of the invertebrate spe- cies recorded from Missouri subterranean hab- itats —Missouri Speleology 15(2):1—10. Dearolf, K. 1953. The invertebrates of 75 caves in the United States Pennsylvania Academy of Sci- ences 27:225—241. Fletcher, M. W. 1982. Microbial ecology of a bat gua- no community. Unpublished M.S. thesis, South- west Missouri State University, Springfield, 425 Pp- Gardner, J. E. 1986. Invertebrate fauna from Missouri caves and springs.—Missouri Department of Conservation Natural History Series number 3: 1-72. Jass, J., & B. Klausmeier. 2001. Terrestrial isopod (Crustacea: Isopoda) atlas for Canada, Alaska and the contiguous United States —Mulwaukee Public Museum Contributions in Biology and Geology 95:1—105. Leistikow, A., & J. W. Wagele. 1999. Checklist of ter- restrial isopods of the new world (Crustacea, Isopoda, Oniscoidea).—Revista Brasileira de Zoologia 16(1):1—72. Martin, B. J. 1980. The community structure of ar- thropods on bat guano and bat carcasses in Tumbling Creek Cave. Unpublished M.S. the- sis, University of Illinois at Chicago Circle, 178 pp. Mitchell, R. W., & J. R. Reddell. 1971. The inverte- brate fauna of Texas caves. Pp. 35—90 in E. L. Lundelius, & B. H. Slaughter, Natural History of Texas Caves. Gulf Natural History, Dallas. Mulaik, S. B. 1960. Contribucion el conocimiento de lost isopodos terrestres de Mexico (lospoda, Oniscoidea).—Revista de la Sociedad Mexicana de Historia Natural 21(1):79—292. Reddell, J. R. 1981. A review of the cavernicole fauna of Mexico, Guatamala, and Belize. Bulletin 27, Texas Memorial Museum, 327 pp. Rioja, E. 1950. Estudios Carcinologicos. XXII. Los tri- coniscidos cavernicolas de México del género Protrichoniscus y descripcion de una nueva es- pecie del Mismo.—Anales del Instituto de Biol- ogia 21(1):127—146. . 1951. Estudios Carcinol6gicos. XXVI. Des- cripcion de Protrichoniscus acostai n. sp. (Crust. Is6podo) de Comitan, Chiapas.—Anales del Instituto de Biologia 22(1):181—189. . 1953. Estudios Carcinologicos. XXIX. Un nuevo género de isOpodo triconiscido de la Cue- va de Ojo de Aguan Grande, Parje Nuevo, Cor- 185 doba, Ver.—Anales del Instituto de Biologia 23(1—2):227-241. 1955. Triconiscidae cavernicolas de Méxi- co.—Revista de la Sociedad Mexicana de En- tomolgia 1(1—2):39-62. . 1957. Estudios Carcinolo6gicos. XXXVI. Des- cripcion y estudio de una nueva especie del gé- nero Cylindroniscus (Is6poda Triconiscido) de Yucatdn.—Anales del Instituto de Biologia 28(1—2):267-278. Schultz, G. A. 1970. Cylindroniscus vallesensis sp. nov.: description with review of genus (Isopoda, Trichoniscidae).—Transactions of the American Microscopical Society 89(3):407—412. . 1982. Amerigoniscus malheurensis, new spe- cies, from a cave in western Oregon (Crustacea: Isopoda: Trichoniscidae).—Proceedings of the Biological Society of Washington 95(1):89—92. . 1984. Brackenridgia sphinxensis n. sp. from a cave with notes on other species from Arizona and California (Isopoda, Oniscoidea).—South- western Naturalist 29(3):309-319. . 1994. Typhlotricholigioides and Mexiconiscus from Mexico and Cylindroniscus from North America (Isopoda: Oniscidea: Trichonisci- dae).—Journal of Crustacean Biology 14(4): 763-770. Tabacaru, I. 1993. Sur la classification des Trichonis- cidae et la position systématique de Thauma- toniscellus orghidani Tabacaru, 1973 (Crusta- cea, Isopoda, Oniscidea).—Travaux Institute Spéleologique Emile Racovitza 32:43—85. Ullrich, C. J. 1902. A contribution to the subterranean fauna of Texas.—Transactions of the American Microscopical Society 23:83—100. U.S. Fish & Wildlife Service. 2001. Listing the Tum- bling Creek Cavesnail as endangered.—Federal Register 66(248):66803—66811. Vandel, A. 1950. Campagne spéleologique de C. Bo- livar et R. Jeannel dans 1’Amérique du Nord (1928). 14. Isopodes terrestres recueillis par C. Bolivar et R. Jeannel (1928) et le Dr. Henrot (1946).—Archives de Zoologie Expérimentale et Génerale 87:183—210. . 1953. A new terrestrial isopod from Oregon, Caucasonethes rothi n. sp.—Pacific Science 7(2):175-178. . 1965. Les Trichoniscidae cavernicoles (Iso- poda Terrestria: Crustacea) de L- Amérique du Nord.—Annales de Spéléologie 20(3):347-389. . 1978. Les espéces appartenant au genre Amer- igoniscus Vandel, 1950 (Crustacés, Isopodes, Oniscoides).—Bulletin de la Société d’ Histoire Naturelle de Toulouse 113:303-—310. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(2):186-198. 2004. New species and records of Bopyridae (Crustacea: Isopoda) infesting species of the genus Upogebia (Crustacea: Decapoda: Upogebiidae): the genera Orthione Markham, 1988, and Gyge Cornalia & Panceri, 1861 John C. Markham Arch Cape Marine Laboratory, Arch Cape, Oregon 97102-0105 U.S.A. Abstract.—Two bopyrid genera whose species parasitize only Upogebia spp. are reviewed and revised. Orthione Markham, 1988, heretofore known only from its type-species, O. furcata (Richardson, 1904), is rediagnosed and en- larged. Orthione griffenis, new species, infests Upogebia pugettensis (Dana, 1852) in Oregon, U.S.A. Orthione mesoamericana, new species, infests U. spinigera (Smith, 1871) on the Pacific coasts of Costa Rica and Colombia. The genus Metabopyrus Shiino, 1939, is incorporated into Gyge Cornalia and Pan- ceri, 1861, which is rediagnosed, and a key is given to the four known species. Gyge ovalis (Shiino, 1939), formerly Metabopyrus ovalis Shiino, 1939, is re- described on the basis of new material found infesting U. edulis Ngoc-Ho & Chan, 1992, in Taiwan, a new host and geographical record. In a recent compilation of the known bo- pyrid parasites of thalassinidean decapod crustaceans throughout the world (Mark- ham 2001), I listed 26 species of the genus Upogebia known to harbor a total of 27 species of bopyrid isopods. Since then, ad- ditional material of parasites of species of Upogebia has become available for exami- nation or has been reliably reported to me. It includes two new species of Orthione Markham, 1988, described herein, as well as new host and geographic records for Me- tabopyrus ovalis Shiino, 1939, which is re- described and reassigned to the genus Gyge Cornalia & Panceri, 1861. Materials and Methods Host specimens bearing parasites, or par- asites that had been removed from their hosts, have become available for study from various sources over a period of several years. Some were already in scientific col- lections, while others are being newly do- nated to institutions housing such collec- tions. Those institutions are indicated thus: Museo de Zoologia, Universidad de Costa Rica, MZUCR; Naturhistorisches Museum, Wien, Austria, NHMW; Naturhistoriska Rijksmuseet, Sweden, SMNH; and Natural History Museum, Smithsonian Institution, USNM. Results Family Bopyridae Rafinesque-Schmaltz, 1815 Subfamily Pseudioninae Codreanu, 1967 Genus Orthione Markham, 1988 Type-species, by original designation, Pseudione furcata Richardson, 1904. Num- ber of previously known species: 1, O. fur- cata (Richardson, 1904), infesting Upoge- bia affinis (Say, 1818), Massachusetts to North Carolina, U.S.A. Revised generic diagnosis, based on three known species.-Female. Body outline oblong, about twice as long as wide, sides nearly parallel, axis only slightly distorted, all body regions and segments distinct dor- sally. Head deeply set into pereon, its an- VOLUME 117, NUMBER 2 terior margin completely covered by frontal lamina and forming continuous curve with pereon; maxilliped lacking palp; barbula with single prominent lanceolate process on each side, rarely minute process lateral to it. Pereopods slightly to much larger pos- teriorly; oostegites generously enclosing brood pouch, first one with prominent but unadorned internal ridge, no posterolateral point. Pleon of six pleomeres, much broad- er than long, final pleomere deeply enclosed by fifth; five pairs of biramous pleopods and similar uniramous uropods completely covering lateral margins and all but center of dorsal surface of pleon, endopodites of first pair larger and medially extended. Male. Body oblong, at least three times as long as wide; all body regions and seg- ments distinct. Head nearly semicircular; second antennae prominently extended. Pe- reopods relatively small, though overall larger and with smaller dactyli posteriorly, all clustered medially. Pleon about % of to- tal body length, of six pleomeres; pleopods absent or as low incomplete oval uniramous flaps; final pleomere largely surrounded by fifth, ending in pair of uniramous flaplike uropods. Hosts. All in genus Upogebia. Key to Three Species of Orthione, Based on Mature Females 1. Pereopods with propodal cups receiving tips of dactyli, bases produced into large carinae; pleopodal rami somewhat ovate O. griffenis new species [Oregon, U.S.A.]. —. Pereopods lacking propodal cups to re- ceive tips of dactyli, bases lacking cari- nae; pleopodal rami lanceolate 2. Head much broader than long, with mi- nute barbular projection lateral to main process; final pleomere visible dorsally ... O. mesoamericana new species [Pacific coast from Costa Rica to Colombia]. —. Head about as broad as long, only single process on each side of barbula; final pleomere more or less hidden dorsally O. furcata (Richardson, 1904) [Atlantic coast of U.S.A.]. 187 Orthione griffenis, new species Figs. 1-3 ““New species .. . [of] . . . Orthione.” —Da- vid, 2001:6. Material examined.—Infesting Upogebia pugettensis (Dana, 1852). Collected and hosts det. by B. D. Griffen. Mudflats, Idaho Inlet, Yaquina Bay, Oregon, USA, 44° 35.4'N, 124°01.5'W, unspecified date, 2000: 1 2, holotype, USNM 1008784, 1 6, allotype, USNM 1008785. Same locality, 23 June 2001, 9 22, 7 Sd, paratypes, USNM 1008786. Collected and hosts det. by T. H. DeWitt: Riverbend, Yaquina Bay, Oregon, 12 November 1999, Sample S59MDF-U31: 1 @, dextral, 1 35, paratypes, USNM 1008787. Idaho Flat, Yaquina Bay, Oregon, 24 September 1999, Sample 59UU104M: 1 2°, sinistral, immature, para- type, USNM 1008788. Description.—Holotype female (Fig. 1). Length 11.0 mm, maximal width 9.2 mm, head length 2.2 mm, head width 2.1 mm, pleon length 2.9 mm; distortion 15°, dex- trally. Outline oval, nowhere abruptly broader or narrower; all body regions and segments distinct. No pigmentation (Fig. 1A, B). Head almost square, deeply set into pe- reon, its anterior edge continuous with per- eonal margin. Distinct rather long frontal lamina extending completely across front of head but not beyond its sides. First anten- nae long and extended beyond margin of head, of 5 articles, distal two terminally se- tose; second antennae greatly reduced, of 1 to 3 articles (Fig. 1C). Barbula (Fig. 1D) with single long falcate process on each side, central region entire. Maxilliped (Fig. 1E) subtriangular, lacking palp, with plec- tron short and blunt; anterior article nearly rectangular, much longer than triangular posterior article. Pereon widest across pereomeres 4—5. Pereomere | curved strongly around head, it and pereomere 2 markedly concave an- teriorly; pereomeres 3—4 nearly straight across; pereomeres 5—7 concave posterior- 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. side of barbula. E. Right maxilliped. EF Right oostegite 1, external. G. Same, internal. H. Right pereopod 1. I. End carpus and dactylus of same. J. Right pereopod 7. K. End carpus and dactylus of same. Scale: 3.60 mm for A, B, E—G; 1.20 mm for C; 2.40 mm for D; 1.00 mm for H, J; 0.4 mm for I, K. ly. Pereomere | shortest, all others about same length. Shallow elongate depression near each side of dorsal surface of pereo- meres 2—5. Pereomeres 1—6 bordered by coxal plates on long side of body, those on pereomeres 5—6 with crenulate margins; smaller coxal plates on short sides of per- eomeres | and 5. First oostegite (Fig. 1E G) subcircular, its articles of about same size, separated by deep but narrow groove externally; internal ridge smoothly curved and lacking ornamentation. Oostegites 2—5 all long and relatively slender, each reach- ing about % of distance across brood pouch and together completely enclosing it. Fifth oostegite with fringe of long setae along posterior margin. Pereopods (Fig. 1H, J) with all articles distinct, more than doubling in size posteriorly; all bases produced into broadly rounded carinae; short comma- Orthione griffenis, new species. Holotype female. A. Dorsal. B. Ventral. C. Right antennae. D. Right shaped dactyli (Fig. 11, K) with sharp tips fitting into lip-like receptacles on distal cor- ners of propodi; all carpi densely setose dis- tally. Pleon of 6 pleomeres, all sharply concave posteriorly, its posterior margin almost straight across sides of all pleomeres. Ven- trally, sides of pleon completely covered by 5 pairs of overlapping lanceolate uniramous lateral plates and uropods, and its middle region equally covered by 5 pairs of bira- mous pleopods, all of their rami of size and structure similar to lateral plates, except that endopodites of first pleopods much larger than others and crossing each other in middle of pleon. Allotype male (Fig. 2). Length 8.0 mm, maximal width 3.0 mm, head length 1.1 mm, head width 1.9 mm, pleon length 2.7 mm. Body straight on both sides, rounded VOLUME 117, NUMBER 2 189 Fig. 2. antenna 2. E. Right pereopod 1. EK End carpus and dactylus of same. G. Right pereopod 7. H. End carpus and dactylus of same. I. End of pleon, ventral. Scale: 2.00 mm for A, B; 0.84 mm for E, G; 0.24 mm for C, D, EH, I. at both ends. All body regions and seg- ments distinctly separated (Fig. 2A, B). No pigmentation. Head almost semicircular, markedly nar- rower than any pereomeres. Antennae (Fig. 2C, D) of 3 and 5 articles, respectively, both pairs directed laterally; first antennae distally setose; second antennae extending beyond margins of head. Pereon broadest across pereomere 6, but only slightly so. Most pereomeres deeply separated by anterolateral notches. Low broad middorsal ridge along full length of pereon. Pereopods (Fig. 2E, G) relatively small, clustered medially under body; all of about same size, but their dactyli smaller posteriorly. All propodi bearing corneous ridges on surfaces met by folded dactyli (Fig. 2F H); distal corner of propodus of Orthione griffenis, new species. Allotype male. A. Dorsal. B. Ventral. C. Right antenna 1. D. Left pereopod 7 extended into receptacle for end of dactylus (Fig. 2G, H); all carpi distally setose. Pleon of 6 distinct pleomeres, each nar- rower than that before it. Five pairs of dis- tinct but sessile oval pleopods, progressive- ly smaller posteriorly. Final pleomere (Fig. 21) deeply set into fifth, produced into pair of stubby pointed uniramous uropods, their margins bearing many short setae. Remarks on paratypes (Fig. 3).—Of the nine paratype females, five are dextrally distorted, as is the holotype, three are sinis- tral, and one is too immature for assess- ment. They range in length from 6.2 mm to 18.8 mm and in width from 2.7 to 13.3 mm (Fig. 3A—G). Most of the mature females have the endopodites of the first pair of ple- opods prominently visible (Fig. 3A). One 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. pleon, ventral. C. End of pleon, ventral. D. Immature, dorsal. E. Same, ventral. E Late larva, dorsal. G. Same, with male attached, ventral. H. Pleon, ventral. I. Pleon, ventral. J. Immature, end of pleon, ventral. Scale: 4.20 mm for C; 2.10 mm for A, B, D, E; 1.00 mm for F—J. has long slender extended uropods (Fig. 3B); one mature female has the fifth pleo- mere deeply separated from the preceding one (Fig. 3C), as does one immature female (Fig. 3D) In a very immature female (Fig. 3E G), all oostegites are absent, and pleo- pods are only uniramous flaps. At a slightly later stage, an immature female (Fig. 3D, E) has rudimentary oostegites and small but distinctly pleopods, the endopodites of the first pair already pointing medially. The seven paratype males (Fig. 3G—J) are 6.1 to 10.7 mm in length, and 2.0 to 3.2 mm in width. All are very similar to the allotype, but one has a more elongate pleon, its pleomeres deeply separated (Fig. 3H); one has a deformed fifth pleomere (Fig. 31); and one has only very tiny traces of pleo- pods (Fig. 3J). Etymology.—The name griffenis, geni- tive singular of a name regarded as a Latin Orthione griffenis, new species. Paratypes. A-G, females. H—J, males. A. Pleon, ventral. B. End of third-declension noun, is selected to honor Blaine D. Griffen, who, in the course of an ecological study of the host, Upogebia pug- ettensis, collected most of the material, called it to my attention and furnished it and collection data for this description. Remarks.—Upogebia pugettensis has been reported many times as the host of Phyllodurus abdominalis Stimpson, 1857, which attaches to its abdomen throughout the host’s geographic range from British Columbia to central California (Williams 1986, Markham 1992), though the coast of Oregon remains a gap in the known distri- bution of P. abdominalis. This is the first record of infestation of U. pugettensis by a branchial bopyrid. So far this new species, Orthione griffenis, is known only from Ya- quina Bay, Oregon, in the middle of the range of U. pugettensis, but there it appears to be fairly common. Upogebia pugettensis VOLUME 117, NUMBER 2 191 Fig. 4. D. Right maxilliped. E. Plectron of same. FE Right side of barbula. G. Right oostegite 1, external. H. Same, internal. J. Right pereopod 1. I. J. Right pereopod 7. Scale: 2.00 mm for A, B, D, F—H; 1.00 mm for E; 0.36 mm for C, I, J. attains densities of up to 300 burrows per square meter in Yaquina Bay and occurs throughout the lower region of that estuary (Griffen 2002). For comparison of Orthione griffenis with other species in the genus, see the remarks on the following species. Orthione mesoamericana, new species Figs. 4, 5 ““Bopyrid Isopod.’’—Holthuis, 1952:9 [Buenaventura, Colombia; infesting Upo- gebia spinigera (Smith)]. Material examined.—Infesting Upogebia spinigera (Smith, 1871). Puerto Jiménez, Golfo Dulce, Puntarenas, Costa Rica, 08°32'30"N, 83°18'20"W, 13 January 1977. 1 2, holotype, 1 3d, allotype, MZUCR 2194-04. Lund University Chile Expedi- tion. Buenaventura, Colombia, 03°77'N, 77°02'W, on beach, under lump of clay, 30 Orthione mesoamericana, new species. Holotype female. A. Dorsal. B. Ventral. C. Right antennae. August 1948. H. Brattstrom and E. Dahl, colls., L. B. Holthuis det. of host: 1 ¢, para- type, SMNH 5325. Description.—Holotype female (Fig. 4). Length 6.5 mm, maximal width 5.4 mm, head length 1.3 mm, head width 1.8 mm, pleon length 2.2 mm. Body axis distortion 4°, dextrally. Body nearly oval, all regions and segments distinct. No pigmentation ex- cept for dark eyespots (Fig. 4A, B). Head slightly convex anteriorly, nearly semicircular posteriorly, deeply embedded into pereon. Frontal lamina very short but extending completely across anterior of head but not beyond. Eyes as prominent slender slashes near anterolateral corners of head. Antennae (Fig. 4C) well-developed, of 3 and 6 articles, respectively, first ones extending plainly beyond front margin of head. Barbula (Fig. 4F) with two projec- tions on each side, outer one minute, inner 192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON one extended and curved, both with entire margins; no decoration in middle of bar- bula. Maxilliped (Fig. 4D) suboval, lacking palp but with notch in anterior margin; plectron (Fig. 4E) small and slender, point- ing anteriorly. Pereon broadest across pereomere 4, all pereomeres distinctly separated laterally. Tergal plates on both sides of pereomeres 1—4, though only faint on first one. Ooste- gite 1 (Fig. 4G, H) with nearly parallel sides, slightly convex ends, both segments about equally wide and long, separated by deep external groove, no posterolateral pro- jection; internal ridge unornamented, pro- duced into long right-angled flap. Oostegi- tes 2—5 overlapping and reaching nearly across and completely enclosing brood pouch. Pereopods (Fig. 41) more than dou- bling in size posteriorly, but none extending beyond body margin; all dactyli short and fairly blunt and retracting into anterior notches of distally extended propodi; carpi all sparsely setose distally, meri and carpi of anterior pereopods fused. Pleon of 6 distinct pleomeres, final one deeply embedded in preceding one. Five pairs of biramous pleopods, uniramous lat- eral plates and uniramous uropods. Endop- odites of first pair of pleopods large, inflat- ed and extending medially, touching each other and overlapping fifth oostegites an- teriorly. Other pleopodal rami, lateral plates and uropods all similar to each other, all as lanceolate flaps with entire edges, com- pletely covering margins of pleomeres. Allotype male (Fig. 5).—Length 3.3 mm, maximal width 1.2 mm, head length 0.3 mm, head width 0.9 mm, pleon length 1.0 mm. Sides of body nearly parallel, rounded at each end. All body regions and segments distinctly separated. No pigmentation (Fig. 5A, B). Head semicircular, lacking eyes. Anten- nae (Fig. 5C) prominent, first ones of 3 ar- ticles, second ones of 6 or 7 articles; second antennae extending far beyond margins of head. Pereomeres separated by anterior notches reaching inward nearly % of body width. Pereopods (Fig. 5D, E) all of nearly same size, all their articles distinct; dactyli of pe- reopods | and 2 long and sharply pointed, others short and blunt; carpi of pereopods 5—7 much longer than others. Pleon of 6 pleomeres, first one as wide as pereon, others tapering rapidly posteri- orly. Pleopods (Fig. 5F) as sessile plates fairly conspicuous on pleomere 1, much fainter on pleomeres 2—5, absent behind. Sixth pleomere embedded in fifth, produced posteriorly into blunt clublike uropods ex- tending rearward unequal distances, both sparsely fringed by minute setae. Comparison of paratype male.—The oth- er male is considerably larger, with these dimensions: length 4.2 mm, maximal width 1.4 mm, head length 0.6 mm, head width 0.9 mm, pleon length 1.7 mm. It is the same as the allotype in all respects except that both second antennae are 7-articled, and its uropods are equal in length. Etymology.—Adjective mesoamericana selected to indicate the known range of the new species, along the Pacific coast of Cen- tral America. Remarks.—The paratype male from Co- lombia is the unidentified bopyrid reported by Holthuis (1952). Because there was no female accompanying it, it could not be identified until the allotype was described here. The hosts are the same species in both collections. Comparison of three known species of Orthione.—Females. One important addi- tion to the original generic diagnosis (Markham 1988) is that the endopodites of the first pair of pleopods are markedly en- larged and medially directed in a manner highly distinctive for Orthione. This is the case in the type-species, O. furcata, as well as in the two new species, but I did not recognize its importance as a diagnostic character earlier. Of the two new species, O. mesoamericana is much more similar to O. furcata than is O. griffenis. Females of the first two species have heads wider than long, bearing slit-shaped eyes, and the in- VOLUME 117, NUMBER 2 193 Fig. 5. Right pereopod 1. E. Right pereopod 7. E Right pleopods 1. G. Pleomeres 5, 6, ventral. Scale: 1.4 mm for A, B; 0.5 mm for C-G. ternal ridge of the first oostegite is produced into a broad angled flap reaching far pos- teriorly, though that of O. mesoamericana is less extended. Also, females of both of those species have very slender pleopodal appendages, those of O. mesoamericana be- ing relatively somewhat broader. All pereo- pods of O. furcata are about the same size, while those in the two new species more than double in size posteriorly. The minute flap lateral to the large projection on the barbula is unique to O. mesoamericana. Fe- males of O. griffenis are distinctive in hav- ing heads longer than broad, margins of coxal plates crenulate rather than entire, pereopodal bases carinate, and pleopodal appendages ovate rather than lanceolate. Males. All three species are very similar. Pereopods of O. furcata are much longer posteriorly, while those of the two new spe- cies are of nearly the same size throughout. Orthione mesoamericana, new species. Allotype male. A. Dorsal. B. Ventral. C. Right antennae. D. Similarly, uropods of O. furcata are much smaller than in either of the other two spe- cies. In O. mesoamericana, second anten- nae are greatly extended, and pleopods are mostly absent, in contrast with the other two species. The head of O. furcata has a medial region extending posteriorly, that of O. griffenis is smoothly rounded posterior- ly, and that of O. mesoamericana is straight posteriorly. Genus Gyge Cornalia & Panceri, 1861 Type-species, by monotypy, Gyge bran- chialis Cornalia & Panceri, 1861. Revised diagnosis.—Female. Body oval to squarish, at least %4 as broad as long, body axis only slightly distorted either dex- trally or sinistrally, angle of distortion far forward. Head deeply set into pereon, its sides diverging slightly to greatly anterior- 194 ly, frontal lamina completely covering an- terior, its posterior end rounded to pointed. Eyes usually absent. Antennae reduced. Barbula with two lateral processes on each side, they and middle region with deeply digitate margins. Maxilliped usually nearly straight across anterior margin, lacking palp, with slender forward-pointing plec- tron and at most only small posterior point. Pereon broadest across pereomere 4, smoothly rounded both ways, sides of first 4 or 5 pereomeres covered by conspicuous coxal plates. Pereopods all equally small, anterior ones with meri and carpi fused, ba- ses large and often carinate. First oostegite produced onto long slender and usually curved posterior point. Other oostegites narrowly pointed and incompletely enclos- ing brood pouch. Pleon of 5 pleomeres, fi- nal one usually notched posteriorly. Three or four pairs of reduced biramous (or, intra- specifically, uniramous) pleopods not ex- tending beyond pleonal edges, their leaflike rami all of same size and generally with digitately divided margins. Uniramous uro- pods tiny to quite large, flaplike, extended posteriorly and with entire margins. Male. Body long and slender, its head well-extended and separated from pereon, with or without eyes. Antennae well-devel- oped. Pereopods uniformly small, first two with proportionately longer dactyli, all with fused meri and carpi. No midventral tuber- cles. Pleon of 6 pleomeres, final one em- bedded in fifth. Pleopods absent or as ses- sile oval scars. No uropods. Hosts: In genus Upogebia. Four species known, from Britain through Mediterranean to Black Sea; New Zealand; Japan and Tai- wan; and Thailand. Discussion.—I am hereby incorporating Metabopyrus Shiino, 1939, and its two spe- cies, the type-species M. ovalis Shiino, 1939, and M. irregularis Markham, 1985, into Gyge Cornalia & Panceri, 1861, which contained two species, G. branchialis Cor- nalia & Panceri, 1861, and G. angularis Page, 1985. The new diagnosis above is based on all four species. Bourdon (1968: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 151) observed that “Ce genre [Gyge] res- semble beaucoup, en vue dorsale, a Meta- bopyrus Shiino (1939), également parasite d’Upogebia ...” Similarly, Page (1985: 196) asserted that “... Gyge and Metabo- pyrus should be united,” but he did not for- mally make such a combination. In addition to the four species herein included in Gyge, two originally in Gyge and two in Meta- bopyrus, four other species have been cited as members of the genus, but all of them are either synonyms of G. branchialis or now considered to belong to other genera. The date of publication of the paper by Cornalia & Panceri (1861), in which they established the genus Gyge and described its type-species G. branchialis, has been subject to some confusion. Bate & West- wood (1868) and Richardson (1905) listed the date as 1861, while Bonnier (1900) and Bourdon (1968) cited it as 1858. Reference to the original publication indicates that, while the volume in which the report ap- peared was for the year 1858, it actually appeared in 1861. Thus I am citing the date of publication for both the genus Gyge and its type-species Gyge branchialis as 1861. Key to Four Species of Gyge Cornalia & Panceri, 1861, Based on Mature Females 1. Body smoothly rounded, oval; body axis distorted more than 30°; final pleomere extending farther rearward than any oth- er pleomeres —. Body with indistinct corners, subrectan- gular; body axis distorted less than 15°; final pleomere at least partly embedded in fifth pleomere and exceeded by one or more of other pleomeres 2. Long sides of pereomeres distinctly set apart by extended posterolateral angles; posterior margin of pleon entire, large uropods not visible in dorsal view .... PRT ce ee ere whee oak G. irregularis (Markham, 1985), n. comb. [Thailand] —. Long sides of pereomeres continuously curved; posterior margin of pleon deeply cleft, revealing minute uropods in dorsal VIEW") 3 Payee tee eens Sie G. branchialis Cornalia & Panceri, 1861 [Europe] VOLUME 117, NUMBER 2 3. Body segments all distinctly separated; margins of barbula projections digitately subdivided; internal ridge of oostegite | cisitatem ere G. ovalis (Shiino, 1939), n. comb. [Japan, Korea, Taiwan] —. Body segments only obscurely separat- ed; margins of barbula projections smooth; internal ridge of oostegite | smooth G. angularis Page, 1985 [New Zealand] Gyge branchialis Cornalia & Panceri, 1861 Abbreviated synonymy. (See Bourdon, 1968, for complete synonymy to 1968.) Gyge branchialis Cornalia & Panceri, 1861: 87-111; plates I, II [Estuary of Venice, Italy; infesting Upogebia pusilla (Petag- na, 1792)].—Bourdon, 1968:147, 151— 159, 169, 322, 410; figs. 28-32; tables 23, 24 [synonymy, summaries of previ- ous accounts, including records from Britain and Channel Islands through France to Adriatic and Black Seas, in- festing U. deltaura Leach, 1815, U. pus- illa and U. stellata (Montagu, 1808); re- description. Arcachon, France, and Na- poli, Italy; infesting U. pusilla. Roscoff, France, and Plymouth, England; infesting U. deltaura. Roscoff, France; infesting U. stellata. Jersey, Channel Islands; no host].—Restivo, 1968:506 [Napoli,; in- festing U. pusilla|.—Restivo, 1975:152, 153, 161-163; table | [Golfo di Napoli; infesting U. pusilla; study of hyperpara- sitism by Paracabirops marsupialis (Car- oli, 1953)].—Dworschak, 1988:68 [Gra- do, north Adriatic Sea, Italy; near Trieste, Italy; Rovinj, Slovenia; infesting U. pus- illa|.—Astall et al., 1996:821—823: table 1 [Clyde Sea, Scotland, and Irish Sea; in- festing U. deltaura and U. stellata. Ar- cachon Basin, France; infesting U. pus- illa). Gyges [sic] branchialis.—Grube, 1864:77 [Lussin Island, Croatia, Adriatic Sea; in- festing U. pusilla). Gyge galatheae Bate & Westwood, 1868: 225-229 [Guernsey, Channel Islands, in- 195 festing Galathea squamifera Leach, 1814 {subsequently reidentified as Upogebia stellata by Norman, 1905:86}]. Not Gyge branchialis var. arcassonensis Carayon, 1943:46—47 [=Progebiophilus euxinicus (Popov, 1929)]. Material.—All identified and reported by Peter Dworschak.—Infesting Upogebia pusilla. Punta Spin, Grado, Adriatic Sea, It- aly, 45°40'’N, 13°23’E, D. Abed-Navandi coll., August 2000. 1 2 (ovigerous), 1 d, NHMW 19521. Infesting U. tipica (Nardo, 1868), off Isola Rossa, Rovinj, Croatia, Adriatic Sea, 45°05'N, 13°40’E, 18 m, D. Abed-Navandi coll. 4 July 2000, P. Dwor- schak det., | 2, NHMW 19523. Remarks.—This is the first record of bo- pyrid infestation of Upogebia tipica, and thus a new record for Gyge branchialis. Gyge branchialis is already known from the Croatian coast of the Adriatic Sea, and it does not need further redescription beyond the detailed accounts presented by Bonnier (1900) and Bourdon (1968). As indicated in the synonymy above, G. branchialis has been reported many times from Britain through the Mediterranean to the Black Sea as a parasite of three other species of Upo- gebia. Gyge ovalis (Shiino, 1939), new combination Fig. 6 Metabopyrus ovalis Shiino, 1939:88—91; figs. 7, 8 [Hakata Bay, Kytsyt, Japan; infesting Upogebia major (de Haan, 1839) {subsequently corrected to U. is- saeffi Balss, 1913}].—Shiino, 1958:48— 49, fig. 10 [unknown specific locality, Ja- pan; infesting unknown host; further de- scriptive notes].—Codreanu, 1941: 140.—Codreanu, 1961:140; fig. 1.— Codreanu & Codreanu, 1963:283.— Shino, 1972:7.—Markham, 1982:340.— Markham, 1985:14.—Page, 1985:196.— Kim & Kwon, 1988:199, 201—203, 220; fig. 2 [Komso, southwest Korea; infesting U. major|.—Markham, 2001:198, 201; 196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON at aS) G ‘a i. {2 eo cae 3 N Fig. 6. Gyge ovalis (Shiino, 1939), new combination. A—L, female. M—S, male. A. Dorsal. B. Ventral. C. Left antennae. D. Left side of barbula. E. Left maxilliped, external. EK Plectron of same. G. Right pereopod 1. H. Distal end of same. I. Left pereopod 7. J. Distal end of same. K. Left oostegite 1, external. L. Same, internal. M. Dorsal. N. Ventral. O. Left antenna 1. P. Left antenna 2. Q. Left pereopod 1. R. Right pereopod 7. S. End of pleon, ventral. Scale: 4.00 mm for A, B, K, L; 1.93 mm for D, E, M, N; 1.00 mm for C, F; 0.88 mm for G, I; 0.35 mm for H, J, Q—S; 0.18 mm for O, P. tables 1, 2.—Itani et al., 2002:72; fig. la9 [Yamaguchi Bay, Seto, Inland Sea, Japan; infesting U. major, study of response to host’s molting]. Material examined.—Infesting Upogebia edulis Ngoc-Ho & Chan, 1992. Shan-kong mudflat, Chang-Hua County, southwest Tai- wan, 24°06'25"N, 120°25'30"E, Tin-Yam Chan collector and det. of host: 1 2, 1 d, USNM 1008790. Descriptive notes.—Gyge ovalis has now been collected five times, but no lot has been large. The original description (Shiino 1939) consisted of two pairs, the next two collections (Shino 1958, Kim & Kwon 1988) were single females, and the present material is one pair. The size of the most recent Japanese collection (Itani et al. 2002) was not indicated; the photograph in that report, derived from a frame of a videotape which was published only to show the fe- male’s orientation on its host, lacks recog- nizable details. Variations among the spec- imens are slight, but all are noticeably dif- ferent. The present female (Fig. 6A—L) most resembles the figured syntype in pro- portions and shapes of body parts, though it lacks the prominent tergal plates seen on the long side of pereomeres 1-3 of all pre- viously recorded females (Fig. 6A, B). The body of one female (Shiino 1958) is pro- portionately shorter, and another (Kim & Kwon 1988) lacks the posterior notch on the final pleomere. The barbula (Fig. 6D) is the same as in other females. The maxilli- ped (Fig. 6E) is less distinctly segmented than previously seen. The propodus of the first pereopod (Fig. 6G, H) is produced into VOLUME 117, NUMBER 2 a helmet-like shape not previously seen. The first oostegite (Fig. 6K, L) is very much like that reported by Shiino (1958), while the one from Korea (Kim & Kwon 1988) was straight posteriorly. The male (Fig. 6M-—S) has a much more extended head and an embedded final pleomere (Fig. 6S), in contrast to the figured syntype (Shi- ino 1939), whose head was little longer than any pereomere, and whose final pleo- mere was extended behind the preceding one. Remarks.—The present material repre- sents both a new host, Upogebia edulis, and new locality, Taiwan, for Gyge ovalis, al- though Tin-Yam Chan (pers. comm.) re- ports that it is commonly collected there. Gyo Itani (pers. comm.) informs me that he has found Gyge ovalis infesting five differ- ent species of Upogebia in Japan, although so far there are published records of only two host species there. Acknowledgments Tin-Yam Chan, National Taiwan Ocean University, provided material of Gyge oval- is that he had collected and information about it. Peter C. Dworschak, NHMW, pro- vided information about his collections of G. branchialis and granted me permission to report on them. Christer Erséus and Kar- in Sindemark, SMNH, lent the paratype male of Orthione mesoamericana, to which Lipke B. Holthuis, Nationaal Natuurhisto- risch Museum, The Netherlands, had re- ferred me. Blaine D. Griffen and Theodore H. DeWitt, Oregon State University, col- lected and furnished the type material of O. griffenis and provided information on its collection. Gyo Itani, Seto Marine Labora- tory, Japan, sent me reprints of his papers and provided information about collections he had made. Becky S. Jordan, Iowa State University Library, confirmed the date of publication of the paper by Cornalia & Pan- ceri (1861). Nguyen Ngoc-Ho, Muséum National d’Histoire Naturelle, Paris, con- firmed the current usage of names of Upo- 197 gebia spp. Marilyn Schotte, USNM, provid- ed essential curatorial services and infor- mation on collections, lent much material for examination and description, furnished elusive references and information about them. Rita Vargas, MZUCR, lent type-spec- imens of O. mesoamericana and furnished details of their collection. Three anonymous reviewers provided helpful remarks. This report is a scientific contribution of the Arch Cape Marine Laboratory (number 27) and of the College of Oceanic and At- mospheric Sciences, Oregon State Univer- sity. Literature Cited Astall, C. M., A. C. Taylor, & R. J. A. Atkinson. 1996. Notes on some branchial isopods parasitic on upogebiid mudshrimps.—Journal of the Marine Biological Association of the United Kingdom 76:821—824. Bate, C. S., & J. O. Westwood. 1868. A history of the British sessile-eyed Crustacea. Volume II. John van Voort, London. lvi + 536 pp. Bonnier, J. 1900. Contribution a |’ étude des épicarides. Les Bopyridae.—Travaux de 1’Institut Zoolo- gique de Lille et du Laboratoire de Zoologie Maritime de Wimereux 8:1—478. Bourdon, R. 1968. Les Bopyridae des mers Européen- nes.—Meémoires du Muséum National d’Histoire Naturelle de Paris, Nouvelle Série (A) 50(2):77-424. Carayon, J. 1943. Sur les épicarides du Bassin d’ Arcachon (2e note).—Bulletin de la Société Zoologique de France 68:43—48. Codreanu, R. 1941. Sur les pagures du littoral Rou- main de la Mer Noire et leurs crustacés para- sites —Academia Romane, Analele, Bucaresti (3)16:1095—1133. . 1961. Crustacei paraziti cu afinitati indo-pa- cifice in Marea Neagra.—Hidrobiologia, Aca- demia Republicii Socialiste Romine 3:133—-146. , & M. Codreanu. 1963. Sur plusieurs bopy- riens parasites branchiaux des anomoures de la Mer Noire, de la Méditerranée et du Viet- Nam.—Commission Internationale pour l’Exploration de la Mer Méditerranée: Rapports et Procés Verbaux, Réunion 17:283—-285. Cornalia, E., and P. Panceri. 1861. Osservazioni zool- ogische ed anatomische sopra un nuovo genere di isopodi sedentarii (Gyge branchialis).—Me- morie della Reale Accademia di Scienze di To- rino (2)19(1858):85—-118. David, A. 2001. Research Highlights.—Streamlines, the newsletter of Oregon State University’s Col- 198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lege of Oceanic and Atmospheric Sciences 7(20):6. Dworschak, P. C. 1988. The biology of Upogebia pus- illa (Petagna) (Decapoda, Thalassinidea). III. Growth and production Marine Ecology 9: 51-77. Griffen, B. D. 2002. Feeding rates of the mud shrimp Upogebia pugettensis and implications for es- tuarine phytoplankton abundance. Master of Science thesis, Oregon State University, 102 pp. Grube, E. A. 1864. Die Insel Lussin und ihre Meer- esfauna. Ferdinand Hirt, Breslau, 116 pp. Holthuis, L. B. 1952. On two species of Crustacea De- capoda Macrura from the N. W. coast of South America. Reports of the Lund University Chile Expedition, 1948—49.—Lunds Universitets Arsskrift. N. E (Avdeling 2), Bind 47 (Number 9):1-11. Itani, G., M. Kato, & Y. Shirayama. 2002. Behaviour of the shrimp ectosymbionts, Peregrinamor ohshimai (Mollusca: Bivalvia) and Phyllodurus sp. (Crustacea: Isopoda) through host ecdy- ses.—Journal of the Marine Biological Associ- ation of the United Kingdom 82:69-78. Kim, H. S., & D. H. Kwon. 1988. Bopyrid isopods parasitic on decapod crustaceans in Korea.— The Korean Journal of Systematic Zoology. Special Issue No. 2:199—221. Markham, J. C. 1982. Bopyrid isopods parasitic on decapod crustaceans in Hong Kong and south- ern China. 1. Pp. 375-391 in B. S. Morton & C. K. Tseng, eds., Proceedings of the First In- ternational Marine Biological Workshop. The Marine Fauna and Flora of Hong Kong and southern China, Hong Kong, 1980. Hong Kong University Press, Hong Kong, 554 pp. . 1985. Additions to the bopyrid fauna of Thai- land.—Zoologische Verhandelingen 224:1—63. . 1988. Descriptions and revisions of some spe- cies of Isopoda Bopyridae of the north western Atlantic Ocean.—Zoologische Verhandelingen 246: 1-63. . 1992. The Isopoda Bopyridae of the eastern Pacific—missing or just hiding?—Proceedings of the San Diego Society of Natural History 17: 1-4. . 2001. A review of the bopyrid isopods para- sitic on thalassinidean decapods. Pp. 195-204 in B. Kensley & R. C. Brusca, eds., Isopod sys- tematics and evolution. Crustacean Issues 13. A. Balkema, Rotterdam. Norman, A. M. 1905. Museum Normanianum, or a catalogue of the Invertebrata of the Arctic and North Atlantic, Temperate Ocean and Palearctic Region, which are contained in the collection of the Rev. Canon A. M. Norman, M. AS DNC. Ly Lak Dik Re St. EF asset uille Crustacea (2nd edit.). Thos. Caldcleugh & Son, Durham, vi + 47 pp. Page, R. D. M. 1985. Review of the New Zealand Bopyridae (Crustacea: Isopoda: Epicaridea).— New Zealand Journal of Zoology 12:185—212. Restivo, F 1968. Alcuni nuovi dati sul parassitismo da bopiridi in alcuni decapodi del Golfo di Napo- li.—Pubblicazioni della Stazione Zoologica di Napoli 36:505—506. . 1975. Nuovi dati su Paracabirops (n. d. Ca- birops) marsupialis Caroli, parassita di Gyge branchialis.—Pubblicazioni della Stazione Zoologica di Napoli 39:150—168. Richardson, H. 1905. A monograph on the isopods of North America.—Bulletin of the United States National Museum 54:liii + 727 pp. Shiino, S. M. 1939. Bopyrids from Kyisyt and Ryt- kya.—Records of Oceanographic Works in Ja- pan 10:79—99. . 1958. Note on the bopyrid fauna of Japan.— Report, Faculty of Fisheries. Prefectural Uni- versity of Mie 3:29—73. . 1972. [The Epicaridea (list of species) from Japan].—Kansai Shizenkagaku 24:7-10. [In Japanese] Williams, A. B. 1986. Mud shrimps, Upogebia, from the eastern Pacific (Thalassinoidea: Upogebi- idae).—San Diego Society of Natural History, Memoir 14:1—60. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(2):199-212. 2004. Three new species and a new genus of Farreidae (Porifera: Hexactinellida: Hexactinosida) Kirk Duplessis and Henry M. Reiswig* (KD) Redpath Museum, McGill University, Montreal, Quebec, H3A 2K6 Canada, e-mail: olodrin@hotmail.com; (HMR) Department of Biology, University of Victoria, and Natural History Section, Royal British Columbia Museum, Victoria, British Columbia, V8W 3N5 Canada, e-mail: hmreiswig@shaw.ca Abstract.—Two new species of Farrea and a single new species of a closely related new genus, Asceptrulum, all members of the hexactinosan family Far- reidae, are described from three widely distant locations. Farrea herdendorfi, new species, with a type series of eight specimens obtained from 2200 m off Charleston, South Carolina, U.S.A., NW Atlantic Ocean, is distinguished by two anchorate clavule forms (umbellate and thimblate or thimble-shaped), both without spiral arrangement of claws. Farrea seiri, new species, represented by 3 fragments of a single specimen from 1450 m near the South East Indian Ridge, mid Indian Ocean, is characterized by only anchorate clavules of thim- blate form, a moderate proportion of which have claws spiralled. Asceptrulum axialis, new genus, new species, is represented by several fragments from a single specimen collected from 2387 m on the Juan de Fuca Ridge, northern Oregon, U.S.A., NE Pacific Ocean. It is distinguished by the combination of complete absence of sceptrules and a one-layered farreoid framework. The diagnosis of Farreidae is emended to encompass the new genus. Although recognized 132 years ago, hex- actinellid sponges, more familiarly referred to as “glass sponges’’, are still obscure members of the deep sea invertebrate fauna. Members of the dictyonine family Farreidae Gray are among the most commonly en- countered hexactinellids, distributed mainly on continental shelves and slopes, but ex- tending into deep water to over 5200 m depth. This hexactinosan family presently includes 21 species distributed unevenly in 5 genera, 17 of those in the genus Farrea. The most recent family diagnosis (Reiswig 2002) focused on one class of free spicules, sceptrules, which here consist of at least one form of clavule or sarule, with or with- out lonchiole or aspidoscopule. Forms with narrow-head scopule were excluded from the family. A second common feature of most, but not all, members of the family, not included in that recent diagnosis, is the minimal, one-layered dictyonal framework at the growing margin. Specimens of three forms, obtained from moderately deep water and submitted to our laboratory for identification, have proven to be undescribed species of this family. Two of them are easily incorporated in the spe- ciose genus Farrea Bowerbank, 1862, but the third entirely lacks sceptrules and thus cannot be assigned to a family on the basis of present diagnoses. Its assignment to a new genus erected within Farreidae requires emendation of that family diagnosis, pro- posed here. Materials and Methods Most submitted specimens (all F. her- dendorfi, new species, and Asceptrulum ax- ialis, new genus, new species) were col- lected by robot submersible and were ac- 200 companied by videotape of the collection process. Only F. seiri new species, was col- lected by dredge. Small fragments of the sponge body wall were either whole-mounted in Canada bal- sam for light microscopy (LM), or were dissolved in hot nitric acid. The acid- cleaned skeletal frameworks were removed, rinsed and dried; the remaining spicule sus- pensions were filtered through 25 mm di- ameter, 0.2 mm-pore-size, nitrocellulose fil- ters; filters with spicules were thoroughly rinsed with distilled water, dried, cleared with xylene, and mounted in Canada bal- sam on microscope slides. Characters of frameworks and spicules were measured by computer using a microscope-coupled dig- itizer. Data are reported as mean + standard deviation (range, number of measure- ments). Spicules for scanning electron mi- croscopy (SEM) were similarly nitric acid cleaned, rinsed in distilled water and then directly deposited onto cover glasses mounted on SEM stubs. Acid-cleaned and rinsed fragments of body wall skeletal frameworks were mounted directly on stubs with epoxy. Following gold-palladium coat- ing, specimens were viewed and photo- graphed with a JEOL JSM-840 SEM. Spic- ule drawings were the made by importing LM or SEM images into a computer image- processing program, and then tracing on screen. Type specimens of the new species have been deposited in the National Mu- seum of Natural History, Smithsonian In- stitution, Washington D.C. (USNM). Family Farreidae Gray, 1872 Diagnosis (emended).—Hexactinosida typically with sceptrules in the form of cla- vules, or their derivatives, sarules, lonchi- oles or aspidoscopules, and typically with a farreoid dictyonal framework. Where scep- trules are lacking the framework is farreoid. Where the framework is euretoid, sarules are present. Remarks.—Emendation of the diagnosis PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Farreidae is required for inclusion of the new genus Asceptrulum as proposed below. Genus Farrea Bowerbank, 1862 Type species.—Farrea occa Bowerbank, 1862, by monotypy. Farrea herdendorfi, new species (Figs. 1-3; Table 1) Holotype.—USNM 1001596; S.S. “Cen- tral America’ wreck, 300 km S. of Charles- ton, S.C., 31.5°N, 77°W, 12 Sep 1989, 2200 m depth, coll. C.E. Herdendorf, R/S ‘Nemo’ from R/V ‘Arctic Explorer’, dive UA. Paratypes.—All from above sampling location and vessels; USNM 1001597, USNM 1001600, USNM 1001601, USNM 1001602, USNM 1001603, all 12 Sep 1989, col. C.E. Herdendorf, Dive UA; USNM 1001598, USNM 1001599, 21 Sep 1990, col. B. Evans, dive AC. Diagnosis.—Farrea with only anchorate clavules, the heads of which vary between two extreme forms—an umbellate (hemi- spherical) form and a thimblate (nearly cy- lindric) form with straight claws nearly par- allel to shaft. Shafts of all clavules are rough but lack conspicuous spines. Description.—Size and shape: The ho- lotype is 30 cm in height, with a branching element arising 16 cm above the lower end (base is missing), extending 11 cm at 60° from the primary axis (Fig. 1E). At widest points, the main body and lateral branch are, respectively, 5.0 and 4.2 cm thick. The central body of the sponge is cryptically bi- laterally symmetical (see below), composed of an original flat blade or stipe incorporat- ed as one side of an axial tube by medial fusion of lateral undulations or ruffles. Through further lateral extension and tight curvature, the lateral ruffles fuse to form tubes appended onto the axial tube. The di- ameter of outer exposed tubular apertures of the holotype are 6.7 + 2.0 mm (range 5-10 mm, n = 12). A sequence of age/maturation stages is VOLUME 117, NUMBER 2 201 A oat. nat - Suber Fig. 1. Farrea herdendorfi, new species; body form. A. Paratype USNM 1001601 in lateral view. B. Same in frontal view. C. Paratype USNM 1001602 in frontal view, asterisks indicate fusion points of lateral pleats to form axial tube. D. Same viewed from distal end with axial tube closed by fusion series at asterisk. E. Holotype USNM 1001596, lateral view. EK Paratype USNM 1001597, with thickened walls, in frontal view. Scale bars = 2 cm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. apparent in the type series as inferred from wall thickness and degree of expansion and fusion of lateral edges—from a simple blade to the complex fused structure of the lateral tubes. The simplest paratype avail- able (Figs. 1A, B) is an axial blade, 18.6 cm long, with undulatory expansion of lat- eral margins as ruffles but without fusion between them. A more complex and older but shorter specimen, 8.2 cm long (Fig. IC), exhibits logitudinal fusion between ruffles along one side to form an axial tube on the axial blade (Fig. 1D). The holotype (Fig. 1E) exhibits the next stage with fusion and branching of ruffles to form a complex bed of lateral tubes supported upon the ax- ial tube. Beyond the branching point, the axial blade and tube of both extremities arise de-novo from the lateral ruffles—con- tinuity does not exist between axial com- ponents of the basal axis and the two distal shoots. By wall thickness, paratype CA6, 14.7 cm long, is next oldest of the series (Fig. 1F) but its gross morphology has ap- parently been simplified by abrasion. It con- sists of the axial tube bearing only the thick-walled bases of lateral tubes in alter- nating offset pairs. The marginal ruffles and tubes have apparently been torn off during Farrea herdendorfi, new species, framework (SEM). A. Single-layer primary framework in marginal area of paratype USNM 1001601, bearing long, usually straight spurs. B. Thickened main body area of holotype USNM 1001596 with irregular dictyonal layers added on dermal side. Scale bars = 1 mm. collection. The oldest specimen, not fig- ured, is an extremely thick-walled and dense basal part, 6.6 X 4.9 cm, of a much larger specimen. That this constructed se- ries of body shapes represents a sequence in growth of a single species is confirmed by identical loose spicules in all specimens. Framework (Fig. 2, Table 1): The frame- work is an unchannelized dictyonal lattice, consisting of, in the thinnest-walled (young) specimens and at growing edges of thick-walled (older) specimens, a one-layer, two-dimensional lattice of somewhat irreg- ular rectangular meshwork with easily rec- ognizable longitudinal dictyonal strands (Fig. 2A). Most of the framework of all but the thinnest specimens is augmented by ad- dition of secondary dictyonalia in irregular- mesh network mainly on dermal, but in some places on the atrial side to form a three-dimensional framework (Fig. 2B). Small hexactins attached to beams of pri- mary and secondary dictyonalia are abun- dant. Wall thickening and increasing rigid- ity of the framework with aging is due to addition of both more secondary dictyona- lia and small hexactins, but only slightly accompanied by thickening of primary dic- tyonal strands. Spurs are long, thin, rough VOLUME 117, NUMBER 2 203 Fig. 3. B. Thimblate anchorate clavule, whole and head. C. Umbellate anchorate clavule head. D. Uncinate, whole and magnified segment. E. Oxyhexaster. EF Onychexaster with magnified ray tip. and spine-like, usually straight but occa- sionally slightly curved. Spicules (Fig. 3, Table 1): Pentactins (Fig. 3A), lining both dermal and atrial sur- faces, have tangential rays heavily spined on outer surfaces, and long proximal ray with heavy spination only on upper third. Uncinates are very long, exceptionally thin and moderately barbed. Two forms of an- chorate clavules occur intermixed on both surfaces, both forms having a thin, smooth shaft ending in a slightly rough, bluntly pointed tip. The thimblate (thimble-shaped) Farrea herdendorfi, new species, spicules of holotype USNM 1001596 (SEM). A. Surface pentactin. form (Fig. 3B) has approximately 15 spines projecting down from a discoid cap, flaring slightly outward at the lower edge. The an- chorate form (Fig. 3C) has approximately 10 spines projecting out and down from a smoothly rounded cap, continuing on the angle of curvature without reflexion. Mi- croscleres consist of two types of smooth hexasters distributed throughout the speci- men. Oxyhexasters (Fig. 3E) have six long primary rays, each bearing 2—3 secondary rays ending in sharp tips. Onychexasters (Fig. 3F), alternately interpretable as dis- 204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Spicule and framework dimensions (in 1m) of Farrea herdendorfi, new species, holotype USNM 1001596, from off South Carolina, USA. Item Mean A. Surface pentactin: tangential ray length 231 tangential ray width 13.1 proximal ray length 298 proximal ray width 2 B. Thimblate clavule length 353 head length SOP head diameter 34.9 C. Umbellate clavule length 454 head length 36.9 head diameter 48.2 D. Uncinate length 1368 width 6.1 E. Oxyhexaster diameter 120 primary ray length 30.7 secondary ray length 33.1 F. Onychexaster diameter 114 primary ray length 27.9 secondary ray length XO} G. Framework beam length 352 beam width 28.2 H. Framework spur length 287.0 cohexasters with reduced discs, have 3—4 irregularly lumpy secondary rays (without sharp spines) each ending in a flat disc with 2—4 short blunt claws. Etymology.—The species is named after the collector of the holotype, Prof. Charles E. Herdendorf, who also served as coordi- nator of the Adjunct Science and Education Program, S.S. ‘Central America’ Project, Columbus-America Discovery Group. Gen- der of the species name is female. Remarks.—Herdendorf et al. (1995) re- ported this form as ““Farrea new species” (p. 86) and figured the specimen designated here as holotype being collected (their Fig. 45). The species differs from all known Farrea in the form of its clavules. The dis- tinctive thimblate clavule head is most sim- ilar to that of: form B of F. kurilensis (Oka- da, 1932), but those clavules have coarsely thorned shafts and are. accompanied by a St. dey. Range N 29 179-299 50 2.9 5.2—20.0 50 81 141-464 50 DED, 6.6—17.2 50 WD 130—490 50 35) 20-45 50 eS 19.7-56.2 50 55 313-581 50 4.4 29-49 50 5.3 32-66 50 308 830-2086 50 1.4 3.5-9.9 50 13 86-146 50 4.6 21.6-43.1 50 47 20.7-44.3 50 12 87-141 50 3.6 22.2-39.7 50 4.0 19.3-37.8 50 129 125-748 50 9.2 14.5-57.7 50 110 124-727 50 pileate clavule not present in F. herden- dorfi. Farrea seiri, new species (Figs. 4, 5; Table 2) Holotype.—USNM_ 1001594, Southeast Indian Ridge, Indian Ocean, 39°12.83’S, 77°52.88'W, 22 Mar 1996, 1450 m depth, coll. D.S. Scheirer and K. Johnson, Boo- merang Expedition, Leg 6, R/V ‘Melville’, biosample #7, Site 48, Dredge 58, dive# BMRG 06 MV. Diagnosis.—Farrea with only anchorate clavules, all thimblate in form without shaft spines. The most abundant clavule type has straight claws while the less common form has claws spiralled either dextrally or si- nestrally. Description.—Size and shape: The ho- lotype and only sample consists of three VOLUME 117, NUMBER 2 205 Fig. 4. Farrea seiri, new species, body form and framework of holotype USNM 1001594. A. Body form with epirhyses evident in magnified inset. B. Outer surface with shallow epirhyses indicated by arrowheads (SEM). C. Probable primary layer in middle frontal layer of framework exposed by dissection (SEM). D. Two transverse sections of body wall showing thickened main longitudinal strands deep within framework (SEM). Scale bars = 0.5 mm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Straight thimblate anchorate clavule, whole and head. C. Spiral umbellate anchorate clavule head. D. Uncinate, whole and magnified segment. E. Onychexaster with magnified ray tips. very stout fragments from the basal part of a single specimen (Fig. 4A). The specimen was severely damaged during dredge col- lection, all distal parts with thin body wall having been lost. The largest fragment mea- sures 9.6 X 4.3 cm, the second largest 3.9 x 1.5 cm, and the smallest 1.8 X 1.7 cm. All fragments are white in colour, with fair- ly thick walls (for Farreidae), 2.08 + 1.04 mm (range 0.95—3.80 mm, n = 10), though all are quite fragile, easily crushed and crumbled. The main fragment is composed of two fused tubes, the younger attached obliquely along the side of the older. The older tu- bular element provided basal attachment for the specimen and was dead at the time of collection. The younger tube has a series of lateral openings arranged in sets of two in alternating offset pairs, aperture length 9.2 Farrea seiri, new species, spicules of holotype USNM 1001594 (SEM). A. Surface pentactin. B. + 0.9 mm (range 8.3—-10.4 mm, n = 5) width 5.1 + 0.9 mm (range 4.1—7.0 mm, n = 5). Attempts to map dermal and atrial surface topology showed that there is no consistent distinction between surfaces rel- ative to tubular walls. Framework (Fig. 4, Table 2): The outer layers of the rigid dictyonal framework (Fig. 4B), are composed of a highly irreg- ular mesh of hexactins, many with polyra- dial nodes, outlining shallow extradictyonal epirhyses and aporhyses as surface pits (Figs. 4A insert, 4B). Pits have ovoid ap- ertures, length 0.33 + 0.038 mm (range 0.26—0.38 mm, n = 8), width 0.24 + 0.045 mm (range = 0.18—0.32 mm, n = 8). Dis- tances between pits 0.66 + 0.18 mm (range 0.33-1.20 mm, n = 32). Beam thickening has occurred through- out the entire specimen, and there is no sin- VOLUME 117, NUMBER 2 207 Table 2.—Spicule and framework dimensions (in 1m) of Farrea seiri, new species, holotype USNM 1001594, from mid Indian Ocean. Item Mean A. Surface pentactin: tangential ray length 155 tangential ray width 8.1 proximal ray length 193 proximal ray width eo) B. Straight clavule length 243 head length D3) head diameter 17.0 C. Spiral clavule length 265 head length D3).7) head diameter 21.4 D. Uncinate length 802 width 6.5 E. Onychexaster diameter 91 primary ray length . 26.2 secondary ray length 21.9 F. Framework beam length 287 beam width 51 gle layer which can be identified as a buried farreoid two-dimensional grid (Fig. 4C). Long stretches of smooth dictyonal strands located deep within the wall (Fig. 4D) are hypersilicified, obscuring original hexac- tins. Both outer and inner meshes are fur- ther obscured by large numbers of small in- tercalated hexactins. Spurs are moderately common on both surfaces, and within the internal meshwork, but these are not di- rectly comparable to spurs on the primary dictyonalia of other farreids. Spicules (Fig. 5, Table 2): Pentactins have strong spination on outer surface of tangential rays, extending almost to the tips (Fig. 5A). The proximal ray is heavily spined near the centrum, and entirely rough throughout its length. Clavules are all an- chorate and thimblate in form and occur in two types, a straight thimblate type (Fig. 5B) and spiro-thimblate type (Fig. 5D). Both have a thin, smooth shaft, ending in a bluntly pointed tip. The head of the straight thimblate type has approx. 25 claws pro- jecting down from a discoid cap, either St. dev. Range N 37 67-253 50 2.6 3.9-12.5 50 68 99-379 50 2.3 3.1-13.3 50 36 184-347 50 4.9 16.3-39.3 50 3.8 12.8-29.5 50 36 182-345 50 43 21.0-39.3 50 3.4 13.4-28.1 50 376 520-1610 8 DS) 3.3-10.1 8 11 Wag 50 4.0 IQIBSS 50 3.7 13.2—29.0 50 107 89-503 50 16 22-96 83 straight and parallel or flaring slightly out- ward. The spiro-thimblate type has similar cap and claws, but claws curve distally ei- ther to the left (sinistral) or right (dextral). Pentactins and both clavule types occur on all surfaces without distinction. Uncinates are typically long and thin with moderately developed barbs but without a distinguish- able centrum (Fig. 5D). The only micros- clere type is an onychexaster (Fig. 5E) dis- tributed evenly throughout the specimen. The finely rough primary rays each bear four similarly rough secondary rays, each of which ends in a button margined by 3— 6 short, slightly reclined claws. Etymology.—The species name, seiri, is formed from the acronym of its collection locale, the Southeast Indian Ridge. Gender of the species name is female. Remarks.—This species is most similar and closely related to F. herdendorfi de- scribed above, but differs in having only thimblate anchorate clavules and lacking oxyhexaster microscleres. The unavailabil- ity of distal portions of the specimen, and 208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Spicule and framework dimensions (in wm) of Asceptrulum axialis, new genus, new species, ho- lotype USNM 1001604, from NE Pacific. Item Mean A. Surface pentactin: tangential ray length 217 tangential ray width 16.8 proximal ray length 446 proximal ray width 15.1 B. Uncinate length 1641 width 10.4 C. Discohexaster diameter 66 primary ray length 10.0 » secondary ray length 25.4 D. Framework beam length* 346 beam width* 43.8 E. Spur length? 335 * In marginal areas of framework. its presumable two-dimensional farreoid framework do not seriously compromise its placement in Farrea since its spiculation is compelling evidence of its relationship to other Farrea species. Thickness of the basal skeleton and presence of shallow epirhyses and aporhyses are rarely encountered in the genus but are not unique to this species. Genus Asceptrulum, new genus Type species.—Asceptrulum axialis, here designated. Diagnosis.—Farreidae lacking sceptru- les. Etymology.—The genus name, Asceptru- lum, is formed from Greek a = without, plus sceptrula, from Greek skeptron and Latin sceptrum (neuter) = royal wand or staff, in allusion to the absence of sceptrule spicules. Gender of the genus name is neu- ter. Remarks.—See under species below. St. dev. Range N 29 141-279 50 4.4 7.9-25.1 50 108 261-752 50 4.5 8.6—27.1 50 250 1173-1967 12 Zell 5§.5-15.2 23 7 53-81 50 1.7 6.2—13.5 50 3.0 19.8-31.2 50 75 204-584 50 9.1 28.4—77.7 50 719 147-528 50 Asceptrulum axialis, new species (Figs. 6, 7; Table 3) Holotype.-—USNM_ 1001604: North CoAxial segment, Juan de Fuca ridge, northern Oregon, 46°29.83'N, 129°35.79'W, 19 Jul 1993, 2387 m depth, coll. V. Tun- nicliffe, R/S ‘“ROPOS’ dive HYS 221. Diagnosis.—Asceptrulum with axial con- densation of its farreoid framework. Description.—Size and shape: The single specimen encountered and recorded in situ by video, was broken during collection; about one-half was recovered as four frag- ments (Fig. 6A). The intact specimen was attached to hard substrate in a region of re- cently formed basalt blocks sparsely clothed in bacterial mats and strands. Be- fore collection, the organism was frond-like or Y-shaped, 14 cm tall, with a branch point 9 cm from the basal attachment. The four recovered pieces are all thin ribbons or blades with clear axial thickening, thin mar- Fig. 6. => Asceptrulum axialis, new genus, new species, body form and framework of holotype USNM 1001604. A. Body form of recovered fragments with cross-section of one fragment. B. Frontal view of single-layer primary framework in marginal area (LM). C. Same in slightly oblique transverse view showing long, straight spurs VOLUME 117, NUMBER 2 209 (SEM). D. Cross-section of axial region of blade, atrial surface down, with longitudinal primary strands seen at extreme left (SEM). E. Atrial surface of axial region showing thickened primary longitudinal strands (SEM). FE Dermal surface of same (SEM). Scale bars of B—F = 0.5 mm. 210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. ginal fringes with low-amplitude undulation or ruffles. At one point the curved lateral margins are extended and undergo self-fu- sion resulting in a short lateral tube 1.4 cm in diameter. Intact areas of blades are 9.2— 26.2 mm wide and 1.63 + 0.32 mm (range 0.86-1.88 mm, n = 10) thick at axis cen- ters. Framework (Fig. 6, Table 3): The pri- mary framework is a typical farreoid one- layer, two-dimensional mesh of smooth beams (Figs. 6B, C) most obvious in the marginal areas. Blade thickness increases gradually toward the axis by addition of up to nine layers of secondary dictyonalia in irregular arrangement on one side, assumed dermal, of the primary framework (Fig. 6D). On the two surfaces of the blade axes, beams are twice as thick on the atrial side with exposed old primary frame (Fig. 6E), 81.3 + 31.2 pm (range 54-134 wm, n = 5), than on the dermal side (Fig. 6F), 41.1 Asceptrulum axialis, new genus, new species, spicules of holotype USNM 1001604 (SEM). A. Surface pentactins. B. Uncinate, whole and magnified segment. C. Discohexaster with magnified ray tips. + 14.8 wm (range 28.0—64.5 wm, n = 5). Small hexactins fused to framework beams are present but sparse. Spurs of the primary frame are long and straight on both surfaces (Fig. 6C), but while those on the atrial side are rough, those of the dermal side are smooth and often extended and variable in texture. Many of the dermal spurs are fused to centres of secondary dictyonalia or tips of their rays. The secondary structures are a mixture of true and false nodes, with con- nections occurring between grid levels by synapticula. Channelization is absent. Spicules (Fig. 7, Table 3): The species has both low diversity and density of loose spicules. Megascleres consist of large, ro- bust pentactins (Fig. 7A) and long, thin un- cinates (Fig. 7B). Pentactins, present on dermal and gastral surfaces, have tangential rays with heavy spination on outer and lat- eral surfaces and proximal rays with coarse tubercles on the upper third of the ray and VOLUME 117, NUMBER 2 very sparse low spines over the remainder. Uncinates are typical with well-developed barbs, brackets, and no detectable central tyle. The only microsclere type is a rela- tively scarce, robust discohexaster (Fig. 7C), distributed evenly throughout the wall. The six primary rays are short, thick and smooth, each supporting three heavily spined secondary rays which end in hemi- spherically arched discs bearing 5—6 re- curved marginal spines. Etymology.—The species name, axialis, is originally derived from Latin axis = rod or pole. It is here formed from the English adjective axial to preserve its euphonious spelling and reflect the easily visible, dense, axial skeletal framework. Gender of the species name is neuter. Remarks.—Absence of sceptrules in this specimen cannot be attributed to patholog- ical condition, damage during collection or inadequate sampling of spicules. The spec- imen was almost certainly alive at collec- tion, with surface pentactins arrayed in the normal rectangular lattice in places. It is ex- tremely unlikely that disease or the collec- tion process would result in loss of only the one spicule type had sceptrules been pre- sent. Occasionally sceptrules may be diffi- cult to obtain in very small samples of far- reids, but the use of filtration for spicule collection from cm-size fragments has nev- er failed to find scopules. When the first searches for sceptrules in this specimen proved negative, the entire set of fragments was eventually extracted for spicules and examined; not one part of a sceptrule was found. We are very confident that sceptrules were neither lost nor overlooked. They must have been intrinsically absent. Since sceptrules are lacking in Asceptru- lum, its assignment to Farreidae rather than Euretidae is based its one-layered farreoid framework as its primary dictyonal skele- ton. A farreoid framework (Reid 1964) con- sists of a two-dimensional primary grid-like scaffold with dictyonalia, fused in parallel longitudinal strands, cross-linked to adja- cent strands by tangential rays fused side- 211 to-side, resulting in a grid-like layer of fused framework. It is the single layer, or two-dimensional, character of this structure that is considered by some authors to be distinctive for the family Farreidae. This al- ternate definition of Farreidae is extremely important for paleontologists, since loose spicules are unavailable in fossil material. Reiswig (2002) did not include the farreoid framework as a diagnostic feature of Far- reidae since it is absent in one of its five extant genera, Sarostegia Topsent, 1904 (with euretoid framework). He did, how- ever, note that it has historically been an important diagnostic feature, and thus it is included here in the emended diagnosis. The alternative assignment of Asceptru- lum to Euretidae is poorly supported by similar overall spiculation (excepting scep- trule) and presence of a farreoid framework in one of its genera, the monospecific ge- nus, Bathyxiphus Schulze, 1899. Position of Bathyxiphus cannot be used to support as- signment of Asceptrulum to Euretidae since its (Bathxiphus) sceptrule type is not known with complete certainty and its own assign- ment is both provisional and precarious. Based upon the firm relationship of the far- reoid framework with Farreidae, Asceptru- lum is best assigned to that family. Within Farreidae, A. axialis has no ob- vious close relatives. Axial thickening of a blade-form body is unknown in the family and absence of oxy-tip microscleres (pres- ence of only disc- or onych-tip forms) is known only in four Farrea, all of which have a body form of branching and usually anastomosing tubes: F. woodwardi Kent, 1870; F. sollasi Schulze, 1886; F. weltneri Topsent, 1901; F. occa polyclavula Ta- bachnick, 1988. None of these are likely an- cestral forms which could have given rise to A. axialis through the one-step loss of sceptrules. Acknowledgments We thank the following for providing ac- cess to the specimens, permission for their AND processing, and/or collection data: Drs. C. E. Herdendorf, R. D. Evans, V. Tunnicliffe, M. Tsurumi, R. Toll, R. W. Embley, S. K. Juniper, D. Scheirer, M. K. Harper, and the Columbus-America Discovery Group, Inc. This work was supported by a Research Grant from the Natural Sciences and En- gineering Research Council of Canada to HMR. Literature Cited Bowerbank, J. S. 1862. On the anatomy and physiol- ogy of the Spongiadae, part III. On the generic characters, the specific characters, and on the method of examination.—Philosophical Trans- actions of the Royal Society of London 152(2): 1087-1135, pls 72-74. Gray, J. E. 1872. Notes on the classification of the sponges.—Annals and Magazine of Natural History (4) 9(54):442—461. Herdendorf, C. E., T. G. Thompson, & R. D. Evans. 1995. Science on a deep-ocean shipwreck.— Ohio Journal of Science 95(1):4—224. Kent, W. S. 1870. On the Hexactinellidae or hexradiate spiculed siliceous sponges taken in the “Norna’ Expedition off the coast of Spain and Portugal with description of new species and revision of the order—Monthly Microscopical Journal 4: 241-252, pls 63-65. Okada, Y. 1932. Report on the hexactinellid sponges PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON collected by the United States Fisheries steamer ‘Albatross’ in the northwestern Pacific during the summer of 1906.—Proceedings of the Unit- ed States National Museum 81(12):1—118, pls 1-6. Reid, R. E. H. 1964. A monograph of the Upper Cre- taceous Hexactinellida of Great Britain and Northern Ireland, part I1V.—Paleontographical Society Monographs 117(3):cxlix—cliv. Reiswig, H. M. 2002. Family Farreidae Gray. Pp. 1332-1340 in J. N. A. Hooper & R. M. W. van Soest, eds., Systema Porifera: A Guide to the Supraspecific Classification of the Phylum Por- ifera, vol. 2. Klewer Academic/Plenum Publish- ers, New York, 1708 pp. Schulze, F E. 1886. Uber den Bau und das System der Hexactinelliden.—Abhandlungen der Ko6nig- lichen Akademie der wissenschaften zu Berlin (Physikalisch-Mathematisch Classe) 1886:1—97. . 1899. Amerikanische Hexactinelliden nach dem materiale der Albatross-Expedition bear- beitet. Gustav Fischer, Jena, 126 pp. Tabachnick, K. R. 1988. Hexactinellid sponges from the mountains of West Pacific. Pp. 49—64 in P. P. Shirshov, ed., Structural and Functional Re- searches of the Marine Benthos. Academy of Sciences of the USSR, Moscow (in Russian). Topsent, E. 1901. Eponges nouvelles des Agores.— Mémoires de la Société zoologique de France 14:448—466. . 1904. Sarostegia oculata, Hexactinellide nou- velle des iles du Cap-Vert.—Bulletin du Musée Océanographique de Monaco 1904(10):1—8. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(2):213—239. 2004. The origin of biological information and the higher taxonomic categories Stephen C. Meyer Palm Beach Atlantic University, 901 S. Flagler Dr., West Palm Beach, Florida 33401 e-mail: stevemeyer@discovery.org Introduction In a recent volume of the Vienna Series in Theoretical Biology (2003), Gerd B. Miiller and Stuart Newman argue that what they call the “origination of organismal form” remains an unsolved problem. In making this claim, Miiller and Newman (2003:3—10) distinguish two distinct issues, namely, (1) the causes of form generation in the individual organism during embryo- logical development and (2) the causes re- sponsible for the production of novel or- ganismal forms in the first place during the history of life. To distinguish the latter case (phylogeny) from the former (ontogeny), Miiller and Newman use the term “origi- nation”’ to designate the causal processes by which biological form first arose during the evolution of life. They insist that “the mo- lecular mechanisms that bring about biolog- ical form in modern day embryos should not be confused”? with the causes respon- sible for the origin (or “‘origination”’) of novel biological forms during the history of life (p. 3). They further argue that we know more about the causes of ontogenesis, due to advances in molecular biology, molecu- lar genetics and developmental biology, than we do about the causes of phylogen- esis—the ultimate origination of new bio- logical forms during the remote past. In making this claim, Miiller and New- man are careful to affirm that evolutionary biology has succeeded in explaining how pre-existing forms diversify under the twin influences of natural selection and variation of genetic traits. Sophisticated mathemati- cally-based models of population genetics have proven adequate for mapping and un- derstanding quantitative variability and populational changes in organisms. Yet Miiller and Newman insist that population genetics, and thus evolutionary biology, has not identified a specifically causal expla- nation for the origin of true morphological novelty during the history of life. Central to their concern is what they see as the in- adequacy of the variation of genetic traits as a source of new form and structure. They note, following Darwin himself, that the sources of new form and structure must pre- cede the action of natural selection (2003: 3)—that selection must act on what already exists. Yet, in their view, the ““genocentric- ity’’ and “‘incrementalism”’ of the neo-Dar- winian mechanism has meant that an ade- quate source of new form and structure has yet to be identified by theoretical biologists. Instead, Miiller and Newman see the need to identify epigenetic sources of morpho- logical innovation during the evolution of life. In the meantime, however, they insist neo-Darwinism lacks any “theory of the generative” (p. 7). As it happens, Miiller and Newman are not alone in this judgment. In the last de- cade or so a host of scientific essays and books have questioned the efficacy of se- lection and mutation as a mechanism for generating morphological novelty, as even a brief literature survey will establish. Thomson (1992:107) expressed doubt that large-scale morphological changes could accumulate via minor phenotypic changes at the population genetic level. Miklos (1993:29) argued that neo-Darwinism fails to provide a mechanism that can produce 214 large-scale innovations in form and com- plexity. Gilbert et al. (1996) attempted to develop a new theory of evolutionary mechanisms to supplement classical neo- Darwinism, which, they argued, could not adequately explain macroevolution. As they put it in a memorable summary of the sit- uation: “starting in the 1970s, many biol- ogists began questioning its [neo-Darwin- ism’s] adequacy in explaining evolution. Genetics might be adequate for explaining microevolution, but microevolutionary changes in gene frequency were not seen as able to turn a reptile into a mammal or to convert a fish into an amphibian. Microevo- lution looks at adaptations that concern the survival of the fittest, not the arrival of the fittest. As Goodwin (1995) points out, “the origin of species—Darwin’s problem—re- mains unsolved’ ”’ (p. 361). Though Gilbert et al. (1996) attempted to solve the problem of the origin of form by proposing a greater role for developmental genetics within an otherwise neo-Darwinian framework,' nu- merous recent authors have continued to raise questions about the adequacy of that framework itself or about the problem of the origination of form generally (Webster & Goodwin 1996; Shubin & Marshall 2000; Erwin 2000; Conway Morris 2000, 2003b; Carrol 2000; Wagner 2001; Becker & Loénnig 2001; Stadler et al. 2001; Lonnig & Saedler 2002; Wagner & Stadler 2003; Valentine 2004:189—194). What lies behind this skepticism? Is it warranted? Is a new and specifically causal theory needed to explain the origination of biological form? This review will address these questions. It will do so by analyzing the problem of the origination of organismal form (and the ' Specifically, Gilbert et al. (1996) argued that changes in morphogenetic fields might produce large- scale changes in the developmental programs and, ul- timately, body plans of organisms. Yet they offered no evidence that such fields—if indeed they exist—can be altered to produce advantageous variations in body plan, though this is a necessary condition of any suc- cessful causal theory of macroevolution. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON corresponding emergence of higher taxa) from a particular theoretical standpoint. Specifically, it will treat the problem of the origination of the higher taxonomic groups as a manifestation of a deeper problem, namely, the problem of the origin of the information (whether genetic or epigenetic) that, as it will be argued, is necessary to generate morphological novelty. In order to perform this analysis, and to make it relevant and tractable to systema- tists and paleontologists, this paper will ex- amine a paradigmatic example of the origin of biological form and information during the history of life: the Cambrian explosion. During the Cambrian, many novel animal forms and body plans (representing new phyla, sub-phyla and classes) arose in a geologically brief period of time. The fol- lowing information-based analysis of the Cambrian explosion will support the claim of recent authors such as Miiller and New- man that the mechanism of selection and genetic mutation does not constitute an ad- equate causal explanation of the origination of biological form in the higher taxonomic groups. It will also suggest the need to ex- plore other possible causal factors for the origin of form and information during the evolution of life and will examine some other possibilities that have been proposed. The Cambrian Explosion The “‘Cambrian explosion”’ refers to the geologically sudden appearance of many new animal body plans about 530 million years ago. At this time, at least nineteen, and perhaps as many as thirty-five phyla of forty total (Meyer et al. 2003), made their first appearance on Earth within a narrow five- to ten-million-year window of geolog- ic time (Bowring et al. 1993, 1998a:1, 1998b:40; Kerr 1993; Monastersky 1993; Aris-Brosou & Yang 2003). Many new sub- phyla, between 32 and 48 of 56 total (Mey- er et al. 2003), and classes of animals also arose at this time with representatives of these new higher taxa manifesting signifi- VOLUME 117, NUMBER 2 cant morphological innovations. The Cam- brian explosion thus marked a major epi- sode of morphogenesis in which many new and disparate organismal forms arose in a geologically brief period of time. To say that the fauna of the Cambrian period appeared in a geologically sudden manner also implies the absence of clear transitional intermediate forms connecting Cambrian animals with simpler pre-Cam- brian forms. And, indeed, in almost all cas- es, the Cambrian animals have no clear morphological antecedents in earlier Ven- dian or Precambrian fauna (Miklos 1993, Erwin et al. 1997:132, Steiner & Reitner 2001, Conway Morris 2003b:510, Valentine et al. 2003:519—520). Further, several re- cent discoveries and analyses suggest that these morphological gaps may not be mere- ly an artifact of incomplete sampling of the fossil record (Foote 1997, Foote et al. 1999, Benton & Ayala 2003, Meyer et al. 2003), suggesting that the fossil record is at least approximately reliable (Conway Morris 2003b:505). As a result, debate now exists about the extent to which this pattern of evidence comports with a strictly monophyletic view of evolution (Conway Morris 1998a, 2003a, 2003b:510; Willmer 1990, 2003). Further, among those who accept a monophyletic view of the history of life, debate exists about whether to privilege fossil or molec- ular data and analyses. Those who think the fossil data provide a more reliable picture of the origin of the Metazoan tend to think these animals arose relatively quickly—that the Cambrian explosion had a “short fuse.” (Conway Morris 2003b:505—506, Valentine & Jablonski 2003). Some (Wray et al. 1996), but not all (Ayala et al. 1998), who think that molecular phylogenies establish reliable divergence times from pre-Cambrian ances- tors think that the Cambrian animals evolved over a very long period of time—that the Cambrian explosion had a “long fuse.”’ This review will not address these questions of historical pattern. Instead, it will analyze whether the neo-Darwinian process of mu- 215 tation and selection, or other processes of evolutionary change, can generate the form and information necessary to produce the animals that arise in the Cambrian. This analysis will, for the most part,? therefore, not depend upon assumptions of either a long or short fuse for the Cambrian explo- sion, or upon a monophyletic or polyphyletic view of the early history of life. Defining Biological Form and Information Form, like life itself, is easy to recognize but often hard to define precisely. Yet, a rea- sonable working definition of form will suf- fice for our present purposes. Form can be defined as the four-dimensional topological relations of anatomical parts. This means that one can understand form as a unified arrange- ment of body parts or material components in a distinct shape or pattern (topology )—one that exists in three spatial dimensions and which arises in time during ontogeny. Insofar as any particular biological form constitutes something like a distinct ar- rangement of constituent body parts, form can be seen as arising from constraints that limit the possible arrangements of matter. Specifically, organismal form arises (both in phylogeny and ontogeny) as possible ar- rangements of material parts are con- strained to establish a specific or particular arrangement with an identifiable three di- mensional topography—one that we would recognize as a particular protein, cell type, organ, body plan or organism. A particular ?If one takes the fossil record at face value and assumes that the Cambrian explosion took place within a relatively narrow 5—10 million year window, explain- ing the origin of the information necessary to produce new proteins, for example, becomes more acute in part because mutation rates would not have been sufficient to generate the number of changes in the genome nec- essary to build the new proteins for more complex Cambrian animals (Ohno 1996:8475—8478). This re- view will argue that, even if one allows several hun- dred million years for the origin of the metazoan, sig- nificant probabilistic and other difficulties remain with the neo-Darwinian explanation of the origin of form and information. 216 “form,” therefore, represents a highly spe- cific and constrained arrangement of mate- rial components (among a much larger set of possible arrangements). Understanding form in this way suggests a connection to the notion of information in its most theoretically general sense. When Shannon (1948) first developed a mathe- matical theory of information he equated the amount of information transmitted with the amount of uncertainty reduced or elim- inated in a series of symbols or characters. Information, in Shannon’s theory, is thus imparted as some options are excluded and others are actualized. The greater the num- ber of options excluded, the greater the amount of information conveyed. Further, constraining a set of possible material ar- rangements by whatever process or means involves excluding some options and actu- alizing others. Thus, to constrain a set of possible material states is to generate infor- mation in Shannon’s sense. It follows that the constraints that produce biological form also impart information. Or conversely, one might say that producing organismal form by definition requires the generation of in- formation. In classical Shannon information theory, the amount of information in a system is also inversely related to the probability of the arrangement of constituents in a system or the characters along a communication channel (Shannon 1948). The more improb- able (or complex) the arrangement, the more Shannon information, or information- carrying capacity, a string or system pos- sesses. Since the 1960s, mathematical biologists have realized that Shannon’s theory could be applied to the analysis of DNA and pro- teins to measure the information-carrying capacity of these macromolecules. Since DNA contains the assembly instructions for building proteins, the information-process- ing system in the cell represents a kind of communication channel (Yockey 1992: 110). Further, DNA conveys information via specifically arranged sequences of nu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cleotide bases. Since each of the four bases has a roughly equal chance of occurring at each site along the spine of the DNA mol- ecule, biologists can calculate the probabil- ity, and thus the information-carrying ca- pacity, of any particular sequence n bases long. The ease with which information theory applies to molecular biology has created confusion about the type of information that DNA and proteins possess. Sequences of nucleotide bases in DNA, or amino acids in a protein, are highly improbable and thus have large information-carrying capacities. But, like meaningful sentences or lines of computer code, genes and proteins are also specified with respect to function. Just as the meaning of a sentence depends upon the specific arrangement of the letters in a sen- tence, so too does the function of a gene sequence depend upon the specific arrange- ment of the nucleotide bases in a gene. Thus, molecular biologists beginning with Crick equated information not only with complexity but also with “‘specificity,” where ‘‘specificity’’ or “‘specified’’ has meant “necessary to function” (Crick 1958:144, 153; Sarkar, 1996:191).> Molec- ular biologists such as Monod and Crick understood biological information—the in- formation stored in DNA and proteins—as something more than mere complexity (or improbability). Their notion of information associated both biochemical contingency and combinatorial complexity with DNA sequences (allowing DNA’s carrying capac- ity to be calculated), but it also affirmed that sequences of nucleotides and amino ac- ids in functioning macromolecules pos- sessed a high degree of specificity relative to the maintenance of cellular function. The ease with which information theory applies to molecular biology has also cre- ated confusion about the location of infor- 3 As Crick put it, “‘information means here the pre- cise determination of sequence, either of bases in the nucleic acid or on amino acid residues in the protein” (Crick 1958:144, 153). VOLUME 117, NUMBER 2 mation in organisms. Perhaps because the information carrying capacity of the gene could be so easily measured, it has been easy to treat DNA, RNA and proteins as the sole repositories of biological information. Neo-Darwinists in particular have assumed that the origination of biological form could be explained by recourse to processes of ge- netic variation and mutation alone (Levin- ton 1988:485). Yet if one understands or- ganismal form as resulting from constraints on the possible arrangements of matter at many levels in the biological hierarchy— from genes and proteins to cell types and tissues to organs and body plans—then clearly biological organisms exhibit many levels of information-rich structure. Thus, we can pose a question, not only about the origin of genetic information, but also about the origin of the information nec- essary to generate form and structure at lev- els higher than that present in individual proteins. We must also ask about the origin of the “specified complexity,” as opposed to mere complexity, that characterizes the new genes, proteins, cell types and body plans that arose in the Cambrian explosion. Dembski (2002) has used the term “‘com- plex specified information”’ (CSI) as a syn- onym for “specified complexity” to help distinguish functional biological informa- tion from mere Shannon information—that is, specified complexity from mere com- plexity. This review will use this term as well. The Cambrian Information Explosion The Cambrian explosion represents a re- markable jump in the specified complexity or “complex specified information” (CSI) of the biological world. For over three bil- lion years, the biological realm included lit- tle more than bacteria and algae (Brocks et al. 1999). Then, beginning about 570—565 million years ago (mya), the first complex multicellular organisms appeared in the rock strata, including sponges, cnidarians, and the peculiar Ediacaran biota (Grotzin- 217 ger et al. 1995). Forty million years later, the Cambrian explosion occurred (Bowring et al. 1993). The emergence of the Edi- acaran biota (570 mya), and then to a much greater extent the Cambrian explosion (530 mya), represented steep climbs up the bio- logical complexity gradient. One way to estimate the amount of new CSI that appeared with the Cambrian ani- mals is to count the number of new cell types that emerged with them (Valentine 1995:91—93). Studies of modern animals suggest that the sponges that appeared in the late Precambrian, for example, would have required five cell types, whereas the more complex animals that appeared in the Cambrian (e.g., arthropods) would have re- quired fifty or more cell types. Functionally more complex animals require more cell types to perform their more diverse func- tions. New cell types require many new and specialized proteins. New proteins, in turn, require new genetic information. Thus an increase in the number of cell types implies (at a minimum) a considerable increase in the amount of specified genetic informa- tion. Molecular biologists have recently es- timated that a minimally complex single- celled organism would require between 318 and 562 kilobase pairs of DNA to produce the proteins necessary to maintain life (Koonin 2000). More complex single cells might require upward of a million base pairs. Yet to build the proteins necessary to sustain a complex arthropod such as a tri- lobite would require orders of magnitude more coding instructions. The genome size of a modern arthropod, the fruitfly Dro- sophila melanogaster, 1s approximately 180 million base pairs (Gerhart & Kirschner 1997:121, Adams et al. 2000). Transitions from a single cell to colonies of cells to complex animals represent significant (and, in principle, measurable) increases in CSI. Building a new animal from a single- celled organism requires a vast amount of new genetic information. It also requires a way of arranging gene products—pro- teins—into higher levels of organization. 218 New proteins are required to service new cell types. But new proteins must be orga- nized into new systems within the cell; new cell types must be organized into new tis- sues, organs, and body parts. These, in turn, must be organized to form body plans. New animals, therefore, embody hierarchically organized systems of lower-level parts within a functional whole. Such hierarchi- cal organization itself represents a type of information, since body plans comprise both highly improbable and functionally specified arrangements of lower-level parts. The specified complexity of new body plans requires explanation in any account of the Cambrian explosion. Can neo-Darwinism explain the discon- tinuous increase in CSI that appears in the Cambrian explosion—either in the form of new genetic information or in the form of hierarchically organized systems of parts? We will now examine the two parts of this question. Novel Genes and Proteins Many scientists and mathematicians have questioned the ability of mutation and se- lection to generate information in the form of novel genes and proteins. Such skepti- cism often derives from consideration of the extreme improbability (and specificity) of functional genes and proteins. A typical gene contains over one thousand precisely arranged bases. For any specific ar- rangement of four nucleotide bases of length n, there is a corresponding number of pos- sible arrangements of bases, 4”. For any pro- tein, there are 20" possible arrangements of protein-forming amino acids. A gene 999 bases in length represents one of 4°”? possi- ble nucleotide sequences; a protein of 333 amino acids is one of 20%*? possibilities. Since the 1960s, some biologists have thought functional proteins to be rare among the set of possible amino acid sequences. Some have used an analogy with human lan- guage to illustrate why this should be the case. Denton (1986, 309-311), for example, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON has shown that meaningful words and sen- tences are extremely rare among the set of possible combinations of English letters, es- pecially as sequence length grows. (The ra- tio of meaningful 12-letter words to 12-letter sequences is 1/10'4; the ratio of 100-letter sentences to possible 100-letter strings is 1/10!°°.) Further, Denton shows that most meaningful sentences are highly isolated from one another in the space of possible combinations, so that random substitutions of letters will, after a very few changes, in- evitably degrade meaning. Apart from a few closely clustered sentences accessible by random substitution, the overwhelming ma- jority of meaningful sentences lie, probabi- listically speaking, beyond the reach of ran- dom search. Denton (1986:301—324) and others have argued that similar constraints apply to genes and proteins. They have questioned whether an undirected search via mutation and selection would have a reasonable chance of locating new islands of func- tion—representing fundamentally new genes or proteins—within the time avail- able (Eden 1967, Shtitzenberger 1967, Lgvtrup 1979). Some have also argued that alterations in sequencing would likely result in loss of protein function before funda- mentally new function could arise (Eden 1967, Denton 1986). Nevertheless, neither the extent to which genes and proteins are sensitive to functional loss as a result of sequence change, nor the extent to which functional proteins are isolated within se- quence space, has been fully known. Recently, experiments in molecular biol- ogy have shed light on these questions. A variety of mutagenesis techniques have shown that proteins (and thus the genes that produce them) are indeed highly specified relative to biological function (Bowie & Sauer 1989, Reidhaar-Olson & Sauer 1990, Taylor et al. 2001). Mutagenesis research tests the sensitivity of proteins (and, by im- plication, DNA) to functional loss as a result of alterations in sequencing. Studies of pro- teins have long shown that amino acid res- VOLUME 117, NUMBER 2 idues at many active positions cannot vary without functional loss (Perutz & Lehmann 1968). More recent protein studies (often us- ing mutagenesis experiments) have shown that functional requirements place significant constraints on sequencing even at non-active site positions (Bowie & Sauer 1989, Reid- haar-Olson & Sauer 1990, Chothia et al. 1998, Axe 2000, Taylor et al. 2001). In par- ticular, Axe (2000) has shown that multiple as opposed to single position amino acid substitutions inevitably result in loss of pro- tein function, even when these changes oc- cur at sites that allow variation when altered in isolation. Cumulatively, these constraints imply that proteins are highly sensitive to functional loss as a result of alterations in sequencing, and that functional proteins rep- resent highly isolated and improbable ar- rangements of amino acids—arrangements that are far more improbable, in fact, than would be likely to arise by chance alone in the time available (Reidhaar-Olson & Sauer 1990; Behe 1992; Kauffman 1995:44; Dembski 1998:175—223; Axe 2000, 2004). (See below the discussion of the neutral the- ory of evolution for a precise quantitative assessment.) Of course, neo-Darwinists do not envi- sion a completely random search through the set of all possible nucleotide sequenc- es—so-called ““sequence space.’’ They en- vision natural selection acting to preserve small advantageous variations in genetic se- quences and their corresponding protein products. Dawkins (1996), for example, lik- ens an organism to a high mountain peak. He compares climbing the sheer precipice up the front side of the mountain to build- ing a new organism by chance. He ac- knowledges that this approach up ““Mount Improbable” will not succeed. Neverthe- less, he suggests that there is a gradual slope up the backside of the mountain that could be climbed in small incremental steps. In his analogy, the backside climb up “Mount Improbable” corresponds to the process of natural selection acting on ran- dom changes in the genetic text. What 219 chance alone cannot accomplish blindly or in one leap, selection (acting on mutations) can accomplish through the cumulative ef- fect of many slight successive steps. Yet the extreme specificity and complex- ity of proteins presents a difficulty, not only for the chance origin of specified biological information (i.e., for random mutations act- ing alone), but also for selection and muta- tion acting in concert. Indeed, mutagenesis experiments cast doubt on each of the two scenarios by which neo-Darwinists envision new information arising from the mutation/ selection mechanism (for review, see LOnnig 2001). For neo-Darwinism, new functional genes either arise from non-coding sections in the genome or from preexisting genes. Both scenarios are problematic. In the first scenario, neo-Darwinists en- vision new genetic information arising from those sections of the genetic text that can presumably vary freely without conse- quence to the organism. According to this scenario, non-coding sections of the ge- nome, or duplicated sections of coding re- gions, can experience a protracted period of “neutral evolution”’ (Kimura 1983) during which alterations in nucleotide sequences have no discernible effect on the function of the organism. Eventually, however, a new gene sequence will arise that can code for a novel protein. At that point, natural selection can favor the new gene and its functional protein product, thus securing the preservation and heritability of both. This scenario has the advantage of allow- ing the genome to vary through many gen- erations, aS mutations “‘search”’ the space of possible base sequences. The scenario has an overriding problem, however: the size of the combinatorial space (i.e., the number of possible amino acid sequences) and the extreme rarity and isolation of the functional sequences within that space of possibilities. Since natural selection can do nothing to help generate new functional se- quences, but rather can only preserve such sequences once they have arisen, chance alone—random variation—must do_ the 220 work of information generation—that is, of finding the exceedingly rare functional se- quences within the set of combinatorial possibilities. Yet the probability of random- ly assembling (or “‘finding,”’ in the previous sense) a functional sequence is extremely small. Cassette mutagenesis experiments per- formed during the early 1990s suggest that the probability of attaining (at random) the correct sequencing for a short protein 100 amino acids long is about 1 in 10° (Reid- haar-Olson & Sauer 1990, Behe 1992:65— 69). This result agreed closely with earlier calculations that Yockey (1978) had per- formed based upon the known sequence variability of cytochrome c in different spe- cies and other theoretical considerations. More recent mutagenesis research has pro- vided additional support for the conclusion that functional proteins are exceedingly rare among possible amino acid sequences (Axe 2000, 2004). Axe (2004) has performed site directed mutagenesis experiments on a 150- residue protein-folding domain within a B- lactamase enzyme. His experimental meth- od improves upon earlier mutagenesis tech- niques and corrects for several sources of possible estimation error inherent in them. On the basis of these experiments, Axe has estimated the ratio of (a) proteins of typical size (150 residues) that perform a specified function via any folded structure to (b) the whole set of possible amino acids sequenc- es of that size. Based on his experiments, Axe has estimated this ratio to be | to 1077. Thus, the probability of finding a functional protein among the possible amino acid se- quences corresponding to a 150-residue protein is similarly 1 in 1077. Other considerations imply additional improbabilities. First, new Cambrian ani- mals would require proteins much longer than 100 residues to perform many neces- sary specialized functions. Ohno (1996) has noted that Cambrian animals would have required complex proteins such as lysyl ox- idase in order to support their stout body structures. Lysyl oxidase molecules in ex- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tant organisms comprise over 400 amino acids. These molecules are both highly complex (non-repetitive) and functionally specified. Reasonable extrapolation from mutagenesis experiments done on shorter protein molecules suggests that the proba- bility of producing functionally sequenced proteins of this length at random is so small as to make appeals to chance absurd, even granting the duration of the entire universe. (See Dembski 1998:175—223 for a rigorous calculation of this “Universal Probability Bound’’; See also Axe 2004.) Yet, second, fossil data (Bowring et al. 1993, 1998a:1, 1998b:40; Kerr 1993; Monastersky 1993), and even molecular analyses supporting deep divergence (Wray et al. 1996), suggest that the duration of the Cambrian explosion (between 5—10 X 10° and, at most, 7 X 107 years) is far smaller than that of the entire universe (1.3—2 X 10!° years). Third, DNA mutation rates are far too low to generate the novel genes and proteins necessary to building the Cambrian animals, given the most probable duration of the explosion as determined by fossil studies (Conway Mor- ris 1998b). As Ohno (1996:8475) notes, even a mutation rate of 10~° per base pair per year results in only a 1% change in the sequence of a given section of DNA in 10 million years. Thus, he argues that muta- tional divergence of pre-existing genes can- not explain the origin of the Cambrian forms in that time.* *To solve this problem Ohno himself proposes the existence of a hypothetical ancestral form that pos- sessed virtually all the genetic information necessary to produce the new body plans of the Cambrian ani- mals. He asserts that this ancestor and its “‘panani- malian genome” might have arisen several hundred million years before the Cambrian explosion. On this view, each of the different Cambrian animals would have possessed virtually identical genomes, albeit with considerable latent and unexpressed capacity in the case of each individual form (Ohno 1996:8475—8478). While this proposal might help explain the origin of the Cambrian animal forms by reference to pre-exist- ing genetic information, it does not solve, but instead merely displaces, the problem of the origin of the ge- netic information necessary to produce these new forms. VOLUME 117, NUMBER 2 The selection/mutation mechanism faces another probabilistic obstacle. The animals that arise in the Cambrian exhibit struc- tures that would have required many new types of cells, each of which would have required many novel proteins to perform their specialized functions. Further, new cell types require systems of proteins that must, as a condition of functioning, act in close coordination with one another. The unit of selection in such systems ascends to the system as a whole. Natural selection selects for functional advantage. But new cell types require whole systems of pro- teins to perform their distinctive functions. In such cases, natural selection cannot con- tribute to the process of information gen- eration until after the information neces- sary to build the requisite system of pro- teins has arisen. Thus random variations must, again, do the work of information generation—and now not simply for one protein, but for many proteins arising at nearly the same time. Yet the odds of this occurring by chance alone are, of course, far smaller than the odds of the chance or- igin of a single gene or protein—so small in fact as to render the chance origin of the genetic information necessary to build a new cell type (a necessary but not suffi- cient condition of building a new body plan) problematic given even the most op- timistic estimates for the duration of the Cambrian explosion. Dawkins (1986:139) has noted that sci- entific theories can rely on only so much “‘luck”’ before they cease to be credible. The neutral theory of evolution, which, by its own logic, prevents natural selection from playing a role in generating genetic information until after the fact, relies on entirely too much luck. The sensitivity of proteins to functional loss, the need for long proteins to build new cell types and animals, the need for whole new systems of proteins to service new cell types, the probable brevity of the Cambrian explo- sion relative to mutation rates—all suggest the immense improbability (and implausi- 221 bility) of any scenario for the origination of Cambrian genetic information that relies upon random variation alone unassisted by natural selection. Yet the neutral theory requires novel genes and proteins to arise—essentially— by random mutation alone. Adaptive advan- tage accrues after the generation of new functional genes and proteins. Thus, natural selection cannot play a role until new in- formation-bearing molecules have indepen- dently arisen. Thus neutral theorists envi- sion the need to scale the steep face of a Dawkins-style precipice of which there is no gradually sloping backside—a situation that, by Dawkins’ own logic, is probabilis- tically untenable. In the second scenario, neo-Darwinists envision novel genes and proteins arising by numerous successive mutations in the preexisting genetic text that codes for pro- teins. To adapt Dawkins’s metaphor, this scenario envisions gradually climbing down one functional peak and then as- cending another. Yet mutagenesis experi- ments again suggest a difficulty. Recent experiments show that, even when explor- ing a region of sequence space populated by proteins of a single fold and function, most multiple-position changes quickly lead to loss of function (Axe 2000). Yet to turn one protein into another with a com- pletely novel structure and function re- quires specified changes at many sites. In- deed, the number of changes necessary to produce a new protein greatly exceeds the number of changes that will typically pro- duce functional losses. Given this, the probability of escaping total functional loss during a random search for the chang- es needed to produce a new function is ex- tremely small—and this probability dimin- ishes exponentially with each additional requisite change (Axe 2000). Thus, Axe’s results imply that, in all probability, ran- dom searches for novel proteins (through sequence space) will result in functional loss long before any novel functional pro- tein will emerge. 222 Blanco et al. have come to a similar con- clusion. Using directed mutagenesis, they have determined that residues in both the hydrophobic core and on the surface of the protein play essential roles in determining protein structure. By sampling intermediate sequences between two naturally occurring sequences that adopt different folds, they found that the intermediate sequences “‘lack a well defined three-dimensional structure.” Thus, they conclude that it is unlikely that a new protein fold would evolve from a pre-existing fold via a series of folded in- termediates sequences (Blanco et al. 1999: 741). Thus, although this second neo-Darwin- ian scenario has the advantage of starting with functional genes and proteins, it also has a lethal disadvantage: any process of random mutation or rearrangement in the genome would in all probability generate nonfunctional intermediate sequences be- fore fundamentally new functional genes or proteins would arise. Clearly, nonfunctional intermediate sequences confer no survival advantage on their host organisms. Natural selection favors only functional advantage. It cannot select or favor nucleotide se- quences or polypeptide chains that do not yet perform biological functions, and still less will it favor sequences that efface or destroy preexisting function. Evolving genes and proteins will range through a series of nonfunctional interme- diate sequences that natural selection will not favor or preserve but will, in all prob- ability, eliminate (Blanco et al. 1999, Axe 2000). When this happens, selection-driven evolution will cease. At this point, neutral evolution of the genome (unhinged from se- lective pressure) may ensue, but, as we have seen, such a process must overcome immense probabilistic hurdles, even grant- ing cosmic time. Thus, whether one envisions the evolu- tionary process beginning with a noncoding region of the genome or a preexisting func- tional gene, the functional specificity and complexity of proteins impose very strin- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gent limitations on the efficacy of mutation and selection. In the first case, function must arise first, before natural selection can act to favor a novel variation. In the second case, function must be continuously main- tained in order to prevent deleterious (or le- thal) consequences to the organism and to allow further evolution. Yet the complexity and functional specificity of proteins im- plies that both these conditions will be ex- tremely difficult to meet. Therefore, the neo-Darwinian mechanism appears to be inadequate to generate the new information present in the novel genes and proteins that arise with the Cambrian animals. Novel Body Plans The problems with the neo-Darwinian mechanism run deeper still. In order to ex- plain the origin of the Cambrian animals, one must account not only for new proteins and cell types, but also for the origin of new body plans. Within the past decade, devel- opmental biology has dramatically ad- vanced our understanding of how body plans are built during ontogeny. In the pro- cess, it has also uncovered a profound dif- ficulty for neo-Darwinism. Significant morphological change in or- ganisms requires attention to timing. Mu- tations in genes that are expressed late in the development of an organism will not affect the body plan. Mutations expressed early in development, however, could con- ceivably produce significant morphological change (Arthur 1997:21). Thus, events ex- pressed early in the development of organ- isms have the only realistic chance of pro- ducing large-scale macroevolutionary change (Thomson 1992). As John and Mik- los (1988:309) explain, macroevolutionary change requires alterations in the very early stages of ontogenesis. Yet recent studies in developmental bi- ology make clear that mutations expressed early in development typically have dele- terious effects (Arthur 1997:21). For ex- ample, when early-acting body plan mol- VOLUME 117, NUMBER 2 ecules, or morphogens such as_ bicoid (which helps to set up the anterior-poste- rior head-to-tail axis in Drosophila), are perturbed, development shuts down (Niis- slein-Volhard & Wieschaus 1980, Lawr- ence & Struhl 1996, Miiller & Newman 2003).° The resulting embryos die. More- over, there is a good reason for this. If an engineer modifies the length of the piston rods in an internal combustion engine without modifying the crankshaft accord- ingly, the engine won’t start. Similarly, processes of development are tightly inte- grated spatially and temporally such that changes early in development will require a host of other coordinated changes in sep- arate but functionally interrelated devel- opmental processes downstream. For this reason, mutations will be much more likely to be deadly if they disrupt a functionally deeply-embedded structure such as a spinal column than if they affect more isolated anatomical features such as fingers (Kauff- man 1995:200). This problem has led to what McDonald (1983) has called ‘“‘a great Darwinian par- adox”’ (p. 93). McDonald notes that genes that are observed to vary within natural populations do not lead to major adaptive changes, while genes that could cause ma- jor changes—the very stuff of macroevo- lution—apparently do not vary. In other words, mutations of the kind that macro- evolution doesn’t need (namely, viable ge- netic mutations in DNA expressed late in development) do occur, but those that it does need (namely, beneficial body plan mutations expressed early in development) >Some have suggested that mutations in “master regulator’ Hox genes might provide the raw material for body plan morphogenesis. Yet there are two prob- lems with this proposal. First, Hox gene expression begins only after the foundation of the body plan has been established in early embryogenesis (Davidson 2001:66). Second, Hox genes are highly conserved across many disparate phyla and so cannot account for the morphological differences that exist between the phyla (Valentine 2004:88). 223 apparently don’t occur.° According to Dar- win (1859:108) natural selection cannot act until favorable variations arise in a popu- lation. Yet there is no evidence from de- velopmental genetics that the kind of vari- ations required by neo-Darwinism—name- ly, favorable body plan mutations—ever occur. Developmental biology has raised anoth- er formidable problem for the mutation/se- lection mechanism. Embryological evi- dence has long shown that DNA does not wholly determine morphological form (Goodwin 1985, Niyhout 1990, Sapp 1987, Miller & Newman 2003), suggesting that mutations in DNA alone cannot account for the morphological changes required to build a new body plan. DNA helps directs protein synthesis.’ It also helps to regulate the timing and ex- pression of the synthesis of various proteins within cells. Yet, DNA alone does not de- termine how individual proteins assemble themselves into larger systems of proteins; still less does it solely determine how cell types, tissue types, and organs arrange themselves into body plans (Harold 1995: © Notable differences in the developmental pathways of similar organisms have been observed. For exam- ple, congeneric species of sea urchins (from genus He- liocidaris) exhibit striking differences in their devel- opmental pathways (Raff 1999:110—121). Thus, it might be argued that such differences show that early developmental programs can in fact be mutated to pro- duce new forms. Nevertheless, there are two problems with this claim. First, there is no direct evidence that existing differences in sea urchin development arose by mutation. Second, the observed differences in the developmental programs of different species of sea ur- chins do not result in new body plans, but instead in highly conserved structures. Despite differences in de- velopmental patterns, the endpoints are the same. Thus, even if it can be assumed that mutations pro- duced the differences in developmental pathways, it must be acknowledged that such changes did not result in novel form. 7Of course, many post-translation processes of modification also play a role in producing a functional protein. Such processes make it impossible to predict a protein’s final sequencing from its corresponding gene sequence alone (Sarkar 1996:199—202). 224 2774, Moss 2004). Instead, other factors— such as the three-dimensional structure and organization of the cell membrane and cy- toskeleton and the spatial architecture of the fertilized egg—play important roles in de- termining body plan formation during em- bryogenesis. For example, the structure and location of the cytoskeleton influence the patterning of embryos. Arrays of microtubules help to distribute the essential proteins used during development to their correct locations in the cell. Of course, microtubules themselves are made of many protein subunits. Nev- ertheless, like bricks that can be used to as- semble many different structures, the tu- bulin subunits in the cell’s microtubules are identical to one another. Thus, neither the tubulin subunits nor the genes that produce them account for the different shape of mi- crotubule arrays that distinguish different kinds of embryos and developmental path- ways. Instead, the structure of the micro- tubule array itself is determined by the lo- cation and arrangement of its subunits, not the properties of the subunits themselves. For this reason, it is not possible to predict the structure of the cytoskeleton of the cell from the characteristics of the protein con- stituents that form that structure (Harold 2001:125). Two analogies may help further clarify the point. At a building site, builders will make use of many materials: lumber, wires, nails, drywall, piping, and windows. Yet building materials do not determine the floor plan of the house, or the arrangement of houses in a neighborhood. Similarly, electronic circuits are composed of many components, such as resistors, capacitors, and transistors. But such lower-level com- ponents do not determine their own ar- rangement in an integrated circuit. Biolog- ical systems also depend on hierarchical ar- rangements of parts. Genes and proteins are made from simple building blocks—nucle- otide bases and amino acids—arranged in specific ways. Cell types are made of, among other things, systems of specialized PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON proteins. Organs are made of specialized ar- rangements of cell types and tissues. And body plans comprise specific arrangements of specialized organs. Yet, clearly, the prop- erties of individual proteins (or, indeed, the lower-level parts in the hierarchy generally) do not fully determine the organization of the higher-level structures and organization- al patterns (Harold 2001:125). It follows that the genetic information that codes for proteins does not determine these higher- level structures either. These considerations pose another chal- lenge to the sufficiency of the neo-Darwin- ian mechanism. Neo-Darwinism seeks to explain the origin of new information, form, and structure as a result of selection acting on randomly arising variation at a very low level within the biological hier- archy, namely, within the genetic text. Yet major morphological innovations depend on a specificity of arrangement at a much higher level of the organizational hierarchy, a level that DNA alone does not determine. Yet if DNA is not wholly responsible for body plan morphogenesis, then DNA se- quences can mutate indefinitely, without re- gard to realistic probabilistic limits, and still not produce a new body plan. Thus, the mechanism of natural selection acting on random mutations in DNA cannot in prin- ciple generate novel body plans, including those that first arose in the Cambrian ex- plosion. Of course, it could be argued that, while many single proteins do not by themselves determine cellular structures and/or body plans, proteins acting in concert with other proteins or suites of proteins could deter- mine such higher-level form. For example, it might be pointed out that the tubulin sub- units (cited above) are assembled by other helper proteins—gene products—called Mi- crotubule Associated Proteins (MAPS). This might seem to suggest that genes and gene products alone do suffice to determine the development of the three-dimensional structure of the cytoskeleton. Yet, MAPS, and indeed many other nec- VOLUME 117, NUMBER 2 essary proteins, are only part of the story. The location of specified target sites on the interior of the cell membrane also helps to determine the shape of the cytoskeleton. Similarly, so does the position and structure of the centrosome which nucleates the mi- crotubules that form the cytoskeleton. While both the membrane targets and the centrosomes are made of proteins, the lo- cation and form of these structures is not wholly determined by the proteins that form them. Indeed, centrosome structure and membrane patterns as a whole convey three-dimensional structural information that helps determine the structure of the cy- toskeleton and the location of its subunits (McNiven & Porter 1992:313—329). More- over, the centrioles that compose the cen- trosomes replicate independently of DNA replication (Lange et al. 2000:235—249, Marshall & Rosenbaum 2000:187—205). The daughter centriole receives its form from the overall structure of the mother centriole, not from the individual gene products that constitute it (Lange et al. 2000). In ciliates, microsurgery on cell membranes can produce heritable changes in membrane patterns, even though the DNA of the ciliates has not been altered (Sonneborn 1970:1—13, Frankel 1980:607— 623; Nanney 1983:163—170). This suggests that membrane patterns (as opposed to membrane constituents) are impressed di- rectly on daughter cells. In both cases, form is transmitted from parent three-dimension- al structures to daughter three-dimensional structures directly and is not wholly con- tained in constituent proteins or genetic in- formation (Moss 2004). Thus, in each new generation, the form and structure of the cell arises as the result of both gene products and pre-existing three-dimensional structure and organiza- tion. Cellular structures are built from pro- teins, but proteins find their way to correct locations in part because of pre-existing three-dimensional patterns and organization inherent in cellular structures. Pre-existing three-dimensional form present in the pre- 225 ceding generation (whether inherent in the cell membrane, the centrosomes, the cyto- skeleton or other features of the fertilized egg) contributes to the production of form in the next generation. Neither structural proteins alone, nor the genes that code for them, are sufficient to determine the three- dimensional shape and structure of the en- tities they form. Gene products provide nec- essary, but not sufficient conditions, for the development of three-dimensional structure within cells, organs and body plans (Harold 1995:2767). But if this is so, then natural selection acting on genetic variation alone cannot produce the new forms that arise in history of life. Self-Organizational Models Of course, neo-Darwinism is not the only evolutionary theory for explaining the ori- gin of novel biological form. Kauffman (1995) doubts the efficacy of the mutation/ selection mechanism. Nevertheless, he has advanced a self-organizational theory to ac- count for the emergence of new form, and presumably the information necessary to generate it. Whereas neo-Darwinism at- tempts to explain new form as the conse- quence of selection acting on random mu- tation, Kauffman suggests that selection acts, not mainly on random variations, but on emergent patterns of order that self- organize via the laws of nature. Kauffman (1995:47—92) illustrates how this might work with various model sys- tems in a computer environment. In one, he conceives a system of buttons connected by strings. Buttons represent novel genes or gene products; strings represent the law-like forces of interaction that obtain between gene products—i.e., proteins. Kauffman suggests that when the complexity of the system (as represented by the number of buttons and strings) reaches a critical threshold, new modes of organization can arise in the system “‘for free’? —that is, nat- urally and spontaneously—after the manner of a phase transition in chemistry. 226 Another model that Kauffman develops is a system of interconnected lights. Each light can flash in a variety of states—on, off, twinkling, etc. Since there is more than one possible state for each light, and many lights, there are a vast number of possible states that the system can adopt. Further, in his system, rules determine how past states will influence future states. Kauffman as- serts that, as a result of these rules, the sys- tem will, if properly tuned, eventually pro- duce a kind of order in which a few basic patterns of light activity recur with greater- than-random frequency. Since these actual patterns of light activity represent a small portion of the total number of possible states in which the system can reside, Kauf- man seems to imply that self-organizational laws might similarly result in highly im- probable biological outcomes—perhaps even sequences (of bases or amino acids) within a much larger sequence space of possibilities. Do these simulations of self-organiza- tional processes accurately model the origin of novel genetic information? It is hard to think so. First, in both examples, Kaufmann pre- supposes but does not explain significant sources of preexisting information. In his buttons-and-strings system, the buttons rep- resent proteins, themselves packets of CSI, and the result of pre-existing genetic infor- mation. Where does this information come from? Kauffman (1995) doesn’t say, but the origin of such information is an essential part of what needs to be explained in the history of life. Similarly, in his light sys- tem, the order that allegedly arises for “‘for free’’ actually arises only if the programmer of the model system “tunes” it in such a way as to keep it from either (a) generating an excessively rigid order or (b) devolving into chaos (pp. 86-88). Yet this necessary tuning involves an intelligent programmer selecting certain parameters and excluding others—that is, inputting information. Second, Kauffman’s model systems are not constrained by functional consider- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ations and thus are not analogous to biolog- ical systems. A system of interconnected lights governed by pre-programmed rules may well settle into a small number of pat- terns within a much larger space of possi- bilities. But because these patterns have no function, and need not meet any functional requirements, they have no specificity anal- ogous to that present in actual organisms. Instead, examination of Kauffman’s (1995) model systems shows that they do not pro- duce sequences or systems characterized by specified complexity, but instead by large amounts of symmetrical order or internal redundancy interspersed with aperiodicity or (mere) complexity (pp. 53, 89, 102). Get- ting a law-governed system to generate re- petitive patterns of flashing lights, even with a certain amount of variation, is clearly interesting, but not biologically relevant. On the other hand, a system of lights flash- ing the tithe of a Broadway play would model a biologically relevant self-organi- zational process, at least if such a meaning- ful or functionally specified sequence arose without intelligent agents previously pro- gramming the system with equivalent amounts of CSI. In any case, Kauffman’s systems do not produce specified complex- ity, and thus do not offer promising models for explaining the new genes and proteins that arose in the Cambrian. Even so, Kauffman suggests that his self- organizational models can specifically elu- cidate aspects of the Cambrian explosion. According to Kauffman (1995:199—201), new Cambrian animals emerged as the re- sult of “long jump”? mutations that estab- lished new body plans in a discrete rather than gradual fashion. He also recognizes that mutations affecting early development are almost inevitably harmful. Thus, he concludes that body plans, once established, will not change, and that any subsequent evolution must occur within an established body plan (Kauffman 1995:201). And in- deed, the fossil record does show a curious (from a neo-Darwinian point of view) top- down pattern of appearance, in which high- VOLUME 117, NUMBER 2 er taxa (and the body plans they represent) appear first, only later to be followed by the multiplication of lower taxa representing variations within those original body de- signs (Erwin et al. 1987, Lewin 1988, Val- entine & Jablonski 2003:518). Further, as Kauffman expects, body plans appear sud- denly and persist without significant modi- fication over time. But here, again, Kauffman begs the most important question, which is: what produc- es the new Cambrian body plans in the first place? Granted, he invokes “long jump mu- tations” to explain this, but he identifies no specific self-organizational process that can produce such mutations. Moreover, he con- cedes a principle that undermines the plau- sibility of his own proposal. Kauffman ac- knowledges that mutations that occur early in development are almost inevitably dele- terious. Yet developmental biologists know that these are the only kind of mutations that have a realistic chance of producing large-scale evolutionary change—1i.e., the big jumps that Kauffman invokes. Though Kauffman repudiates the neo-Darwinian re- liance upon random mutations in favor of self-organizing order, in the end, he must invoke the most implausible kind of ran- dom mutation in order to provide a self- organizational account of the new Cambri- an body plans. Clearly, his model is not suf- ficient. Punctuated Equilibrium Of course, still other causal explanations have been proposed. During the 1970s, the paleontologists Eldredge and Gould (1972) proposed the theory of evolution by punc- tuated equilibrium in order to account for a pervasive pattern of ““sudden appearance”’ and “‘stasis’”’ in the fossil record. Though advocates of punctuated equilibrium were mainly seeking to describe the fossil record more accurately than earlier gradualist neo- Darwinian models had done, they did also propose a mechanism—known as species selection—by which the large morphologi- 227 cal jumps evident in fossil record might have been produced. According to punctua- tionalists, natural selection functions more as a mechanism for selecting the fittest spe- cies rather than the most-fit individual among a species. Accordingly, on this mod- el, morphological change should occur in larger, more discrete intervals than it would given a traditional neo-Darwinian under- standing. Despite its virtues as a descriptive model of the history of life, punctuated equilibri- um has been widely criticized for failing to provide a mechanism sufficient to produce the novel form characteristic of higher tax- onomic groups. For one thing, critics have noted that the proposed mechanism of punctuated evolutionary change simply lacked the raw material upon which to work. As Valentine and Erwin (1987) note, the fossil record fails to document a large pool of species prior to the Cambrian. Yet the proposed mechanism of species selec- tion requires just such a pool of species upon which to act. Thus, they conclude that the mechanism of species selection proba- bly does not resolve the problem of the or- igin of the higher taxonomic groups (p. 96).° Further, punctuated equilibrium has not addressed the more specific and fun- damental problem of explaining the origin of the new biological information (whether genetic or epigenetic) necessary to produce novel biological form. Advocates of punc- tuated equilibrium might assume that the new species (upon which natural selection acts) arise by known micro-evolutionary processes of speciation (such as founder ef- 8 Erwin (2004:21), although friendly to the possibil- ity of species selection, argues that Gould provides lit- tle evidence for its existence. ““The difficulty” writes Erwin of species selection, “... is that we must rely on Gould’s arguments for theoretical plausibility and sufficient relative frequency. Rarely is a mass of data presented to justify and support Gould’s conclusion.” Indeed, Gould (2002) himself admitted that species se- lection remains largely a hypothetical construct: “I freely admit that well-documented cases of species se- lection do not permeate the literature” (p. 710). 228 fect, genetic drift or bottleneck effect) that do not necessarily depend upon mutations to produce adaptive changes. But, in that case, the theory lacks an account of how the specifically higher taxa arise. Species selection will only produce more fit species. On the other hand, if punctuationalists as- sume that processes of genetic mutation can produce more fundamental morphological changes and variations, then their model be- comes subject to the same problems as neo- Darwinism (see above). This dilemma is evident in Gould (2002:710) insofar as his attempts to explain adaptive complexity in- evitably employ classical neo-Darwinian modes of explanation.’ Structuralism Another attempt to explain the origin of form has been proposed by the structuralists such as Gerry Webster and Brian Goodwin (1984, 1996). These biologists, drawing on the earlier work of D’Arcy Thompson (1942), view biological form as the result of structural constraints imposed upon mat- ter by morphogenetic rules or laws. For rea- sons similar to those discussed above, the structuralists have insisted that these gen- erative or morphogenetic rules do not reside in the lower level building materials of or- ° “T do not deny either the wonder, or the powerful importance, of organized adaptive complexity. I rec- ognize that we know no mechanism for the origin of such organismal features other than conventional nat- ural selection at the organismic level—for the sheer intricacy and elaboration of good biomechanical de- sign surely precludes either random production, or in- cidental origin as a side consequence of active pro- cesses at other levels’’ (Gould 2002:710). “Thus, we do not challenge the efficacy or the cardinal impor- tance of organismal selection. As previously discussed, I fully agree with Dawkins (1986) and others that one cannot invoke a higher-level force like species selec- tion to explain ‘things that organisms do’—in partic- ular, the stunning panoply of organismic adaptations that has always motivated our sense of wonder about the natural world, and that Darwin (1859) described, in one of his most famous lines (3), as ‘that perfection of structure and coadaptation which most justly excites our admiration’ (Gould 2002:886). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ganisms, whether in genes or proteins. Webster and Goodwin (1984:510—511) fur- ther envision morphogenetic rules or laws operating ahistorically, similar to the way in which gravitational or electro-magnetic laws operate. For this reason, structuralists see phylogeny as of secondary importance in understanding the origin of the higher taxa, though they think that transformations of form can occur. For structuralists, con- straints on the arrangement of matter arise not mainly as the result of historical contin- gencies—such as environmental changes or genetic mutations—but instead because of the continuous ahistorical operation of fun- damental laws of form—laws that organize or inform matter. While this approach avoids many of the difficulties currently afflicting neo-Darwin- ism (in particular those associated with its ““genocentricity’’), critics (such as Maynard Smith 1986) of structuralism have argued that the structuralist explanation of form lacks specificity. They note that structural- ists have been unable to say just where laws of form reside—whether in the universe, or in every possible world, or in organisms as a whole, or in just some part of organisms. Further, according to structuralists, morpho- genetic laws are mathematical in character. Yet, structuralists have yet to specify the mathematical formulae that determine bio- logical forms. Others (Yockey 1992; Polanyi 1967, 1968; Meyer 2003) have questioned wheth- er physical laws could in principle generate the kind of complexity that characterizes bi- ological systems. Structuralists envision the existence of biological laws that produce form in much the same way that physical laws produce form. Yet the forms that phys- icists regard as manifestations of underlying laws are characterized by large amounts of symmetric or redundant order, by relatively simple patterns such as vortices or gravi- tational fields or magnetic lines of force. In- deed, physical laws are typically expressed as differential equations (or algorithms) that almost by definition describe recurring phe- VOLUME 117, NUMBER 2 nomena—patterns of compressible “order” not “complexity” as defined by algorithmic information theory (Yockey 1992:77-—83). Biological forms, by contrast, manifest greater complexity and derive in ontogeny from highly complex initial conditions— i.e., non-redundant sequences of nucleotide bases in the genome and other forms of in- formation expressed in the complex and ir- regular three-dimensional topography of the organism or the fertilized egg. Thus, the kind of form that physical laws produce is not analogous to biological form—at least not when compared from the standpoint of (algorithmic) complexity. Further, physical laws lack the information content to specify biology systems. As Polanyi (1967, 1968) and Yockey (1992:290) have shown, the laws of physics and chemistry allow, but do not determine, distinctively biological modes of organization. In other words, liv- ing systems are consistent with, but not de- ducible, from physical-chemical laws (1992:290). Of course, biological systems do manifest some reoccurring patterns, processes and be- haviors. The same type of organism devel- ops repeatedly from similar ontogenetic pro- cesses in the same species. Similar processes of cell division re-occur in many organisms. Thus, one might describe certain biological processes as law-governed. Even so, the ex- istence of such biological regularities does not solve the problem of the origin of form and information, since the recurring process- es described by such biological laws (if there be such laws) only occur as the result of pre- existing stores of (genetic and/or epigenetic) information and these information-rich ini- tial conditions impose the constraints that produce the recurring behavior in biological systems. (For example, processes of cell di- vision recur with great frequency in organ- isms, but depend upon information-rich DNA and proteins molecules.) In other words, distinctively biological regularities depend upon pre-existing biological infor- mation. Thus, appeals to higher-level biolog- ical laws presuppose, but do not explain, the 229 origination of the information necessary to morphogenesis. Thus, structuralism faces a difficult in principle dilemma. On the one hand, phys- ical laws produce very simple redundant patterns that lack the complexity character- istic of biological systems. On the other hand, distinctively biological laws—if there are such laws—depend upon pre-existing information-rich structures. In either case, laws are not good candidates for explaining the origination of biological form or the in- formation necessary to produce it. Cladism: An Artifact of Classification? Some cladists have advanced another ap- proach to the problem of the origin of form, specifically as it arises in the Cambrian. They have argued that the problem of the origin of the phyla is an artifact of the clas- sification system, and therefore, does not require explanation. Budd and Jensen (2000), for example, argue that the problem of the Cambrian explosion resolves itself if one keeps in mind the cladistic distinction between “‘stem’’ and “‘crown’”’ groups. Since crown groups arise whenever new characters are added to simpler more an- cestral stem groups during the evolutionary process, new phyla will inevitably arise once a new stem group has arisen. Thus, for Budd and Jensen what requires expla- nation is not the crown groups correspond- ing to the new Cambrian phyla, but the ear- lier more primitive stem groups that pre- sumably arose deep in the Proterozoic. Yet since these earlier stem groups are by def- inition less derived, explaining them will be considerably easier than explaining the or- igin of the Cambrian animals de novo. In any case, for Budd and Jensen the explo- sion of new phyla in the Cambrian does not require explanation. As they put it, “given that the early branching points of major clades is an inevitable result of clade di- versification, the alleged phenomenon of the phyla appearing early and remaining morphologically static is not seen to require 230 particular explanation” (Budd & Jensen 2000:253). While superficially plausible, perhaps, Budd and Jensen’s attempt to explain away the Cambrian explosion begs crucial ques- tions. Granted, as new characters are added to existing forms, novel morphology and greater morphological disparity will likely result. But what causes new characters to arise? And how does the information nec- essary to produce new characters originate? Budd and Jensen do not specify. Nor can they say how derived the ancestral forms are likely to have been, and what processes, might have been sufficient to produce them. Instead, they simply assume the sufficiency of known neo-Darwinian mechanisms (Budd & Jensen 2000:288). Yet, as shown above, this assumption is now problematic. In any case, Budd and Jensen do not ex- plain what causes the origination of biolog- ical form and information. Convergence and Teleological Evolution More recently, Conway Morris (2000, 2003c) has suggested another possible ex- planation based on the tendency for evolu- tion to converge on the same structural forms during the history of life. Conway Morris cites numerous examples of organ- isms that possess very similar forms and structures, even though such structures are often built from different material sub- strates and arise (in ontogeny) by the ex- pression of very different genes. Given the extreme improbability of the same struc- tures arising by random mutation and se- lection in disparate phylogenies, Conway Morris argues that the pervasiveness of convergent structures suggests that evolu- tion may be in some way “channeled”’ to- ward similar functional and/or structural endpoints. Such an end-directed under- standing of evolution, he admits, raises the controversial prospect of a teleological or purposive element in the history of life. For this reason, he argues that the phenomenon of convergence has received less attention PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON than it might have otherwise. Nevertheless, he argues that just as physicists have re- opened the question of design in their dis- cussions of anthropic fine-tuning, the ubiq- uity of convergent structures in the history of life has led some biologists (Denton 1998) to consider extending teleological thinking to biology. And, indeed, Conway Morris himself intimates that the evolution- ary process might be “underpinned by a purpose”’ (2000:8, 2003b:511). Conway Morris, of course, considers this possibility in relation to a very specific as- pect of the problem of organismal form, namely, the problem of explaining why the same forms arise repeatedly in so many dis- parate lines of decent. But this raises a question. Could a similar approach shed ex- planatory light on the more general causal question that has been addressed in this re- view? Could the notion of purposive design help provide a more adequate explanation for the origin of organismal form generally? Are there reasons to consider design as an explanation for the origin of the biological information necessary to produce the higher taxa and their corresponding morphological novelty? The remainder of this review will suggest that there are such reasons. In so doing, it may also help explain why the issue of tel- eology or design has re-emerged within the scientific discussion of biological origins (Denton 1986, 1998: Thaxton et al. 1992; Kenyon & Mills 1996; Behe 1996, 2004; Dembski 1998, 2002, 2004; Conway Mor- ris 2000, 2003a, 2003b; Lonnig 2001; Lon- nig & Saedler 2002; Nelson & Wells 2003; Meyer 2003, 2004; Bradley 2004) and why some scientists and philosophers of science have considered teleological explanations for the origin of form and information de- spite strong methodological prohibitions against design as a scientific hypothesis (Gillespie 1979, Lenior 1982:4). First, the possibility of design as an ex- planation follows logically from a consid- eration of the deficiencies of neo-Darwin- ism and other current theories as explana- VOLUME 117, NUMBER 2 tions for some of the more striking “‘ap- pearances of design” in biological systems. Neo-Darwinists such as Ayala (1994:5), Dawkins (1986:1), Mayr (1982:xi—x11) and Lewontin (1978) have long acknowledged that organisms appear to have been de- signed. Of course, neo-Darwinists assert that what Ayala (1994:5) calls the “‘obvious design” of living things is only apparent since the selection/mutation mechanism can explain the origin of complex form and or- ganization in living systems without an ap- peal to a designing agent. Indeed, neo-Dar- winists affirm that mutation and selection— and perhaps other similarly undirected mechanisms—are fully sufficient to explain the appearance of design in biology. Self- organizational theorists and punctuational- ists modify this claim, but affirm its essen- tial tenet. Self-organization theorists argue that natural selection acting on self-organiz- ing order can explain the complexity of liv- ing things—again, without any appeal to design. Punctuationalists similarly envision natural selection acting on newly arising species with no actual design involved. And clearly, the neo-Darwinian mecha- nism does explain many appearances of de- sign, such as the adaptation of organisms to specialized environments that attracted the interest of 19th century biologists. More specifically, known micro-evolutionary pro- cesses appear quite sufficient to account for changes in the size of Galapagos finch beaks that have occurred in response to var- iations in annual rainfall and available food supplies (Weiner 1994, Grant 1999). But does neo-Darwinism, or any other fully materialistic model, explain all ap- pearances of design in biology, including the body plans and information that char- acterize living systems? Arguably, biologi- cal forms—such as the structure of a cham- bered nautilus, the organization of a trilo- bite, the functional integration of parts in an eye or molecular machine—attract our attention in part because the organized complexity of such systems seems reminis- cent of our own designs. Yet, this review 231 has argued that neo-Darwinism does not adequately account for the origin of all ap- pearances of design, especially if one con- siders animal body plans, and the informa- tion necessary to construct them, as espe- cially striking examples of the appearance of design in living systems. Indeed, Dawk- ins (1995:11) and Gates (1996:228) have noted that genetic information bears an un- canny resemblance to computer software or machine code. For this reason, the presence of CSI in living organisms, and the discon- tinuous increases of CSI that occurred dur- ing events such as the Cambrian explosion, appears at least suggestive of design. Does neo-Darwinism or any other purely materialistic model of morphogenesis ac- count for the origin of the genetic and other forms of CSI necessary to produce novel organismal form? If not, as this review has argued, could the emergence of novel in- formation-rich genes, proteins, cell types and body plans have resulted from actual design, rather than a purposeless process that merely mimics the powers of a design- ing intelligence? The logic of neo-Darwin- ism, with its specific claim to have account- ed for the appearance of design, would it- self seem to open the door to this possibil- ity. Indeed, the historical formulation of Darwinism in dialectical opposition to the design hypothesis (Gillespie 1979), coupled with neo-Darwinism’s inability to account for many salient appearances of design in- cluding the emergence of form and infor- mation, would seem logically to re-open the possibility of actual (as opposed to appar- ent) design in the history of life. A second reason for considering design as an explanation for these phenomena fol- lows from the importance of explanatory power to scientific theory evaluation and from a consideration of the potential ex- planatory power of the design hypothesis. Studies in the methodology and philosophy of science have shown that many scientific theories, particularly in the historical sci- ences, are formulated and justified as infer- ences to the best explanation (Lipton 1991: 232 32-88, Brush 1989:1124-1129, Sober 2000:44). Historical scientists, in particular, assess or test competing hypotheses by evaluating which hypothesis would, if true, provide the best explanation for some set of relevant data (Meyer 1991, 2002; Cleland 2001:987-989, 2002:474—496).'° Those '0 Theories in the historical sciences typically make claims about what happened in the past, or what hap- pened in the past to cause particular events to occur (Meyer 1991:57—72). For this reason, historical sci- entific theories are rarely tested by making predictions about what will occur under controlled laboratory con- ditions (Cleland 2001:987, 2002:474—496). Instead, such theories are usually tested by comparing their ex- planatory power against that of their competitors with respect to already known facts. Even in the case in which historical theories make claims about past caus- es they usually do so on the basis of pre-existing knowledge of cause and effect relationships. Neverthe- less, prediction may play a limited role in testing his- torical scientific theories since such theories may have implications as to what kind of evidence is likely to emerge in the future. For example, neo-Darwinism af- firms that new functional sections of the genome arise by trial and error process of mutation and subsequent selection. For this reason, historically many neo-Dar- winists expected or predicted that the large non-coding regions of the genome—so-called “junk DNA—would lack function altogether (Orgel & Crick 1980). On this line of thinking, the non-functional sections of the ge- nome represent nature’s failed experiments that remain in the genome as a kind of artifact of the past activity of the mutation and selection process. Advocates of the design hypotheses on the other hand, would have predicted that non-coding regions of the genome might well reveal hidden functions, not only because design theorists do not think that new genetic information arises by a trial and error process of mutation and se- lection, but also because designed systems are often functionally polyvalent. Even so, as new studies reveal more about the functions performed by the non-coding regions of the genome (Gibbs 2003), the design hy- pothesis can no longer be said to make this claim in the form of a specifically future-oriented prediction. Instead, the design hypothesis might be said to gain confirmation or support from its ability to explain this now known evidence, albeit after the fact. Of course, neo-Darwinists might also amend their original pre- diction using various auxiliary hypotheses to explain away the presence of newly discovered functions in the non-coding regions of DNA. In both cases, consid- erations of ex post facto explanatory power re-emerge as central to assessing and testing competing historical theories. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with greater explanatory power are typical- ly judged to be better, more probably true, theories. Darwin (1896:437) used this method of reasoning in defending his the- ory of universal common descent. More- over, contemporary studies on the method of “inference to the best explanation” have shown that determining which among a set of competing possible explanations consti- tutes the best depends upon judgments about the causal adequacy, or “causal pow- ers,” of competing explanatory entities (Lipton 1991:32—88). In the historical sci- ences, uniformitarian and/or actualistic (Gould 1965, Simpson 1970, Rutten 1971, Hooykaas 1975) canons of method suggest that judgments about causal adequacy should derive from our present knowledge of cause and effect relationships. For his- torical scientists, “‘the present is the key to the past’ means that present experience- based knowledge of cause and effect rela- tionships typically guides the assessment of the plausibility of proposed causes of past events. Yet it is precisely for this reason that cur- rent advocates of the design hypothesis want to reconsider design as an explanation for the origin of biological form and infor- mation. This review, and much of the lit- erature it has surveyed, suggests that four of the most prominent models for explain- ing the origin of biological form fail to pro- vide adequate causal explanations for the discontinuous increases of CSI that are re- quired to produce novel morphologies. Yet, we have repeated experience of rational and conscious agents—in particular ourselves— generating or causing increases in complex specified information, both in the form of sequence-specific lines of code and in the form of hierarchically arranged systems of parts. In the first place, intelligent human agents—in virtue of their rationality and consciousness—have demonstrated the power to produce information in the form of linear sequence-specific arrangements of characters. Indeed, experience affirms that VOLUME 117, NUMBER 2 information of this type routinely arises from the activity of intelligent agents. A computer user who traces the information on a screen back to its source invariably comes to a mind—that of a software engi- neer or programmer. The information in a book or inscription ultimately derives from a writer or scribe—from a mental, rather than a strictly material, cause. Our experi- ence-based knowledge of information-flow confirms that systems with large amounts of specified complexity (especially codes and languages) invariably originate from an in- telligent source—from a mind or personal agent. As Quastler (1964) put it, the “‘cre- ation of new information is habitually as- sociated with conscious activity” (p. 16). Experience teaches this obvious truth. Further, the highly specified hierarchical arrangements of parts in animal body plans also suggest design, again because of our experience of the kinds of features and sys- tems that designers can and do produce. At every level of the biological hierarchy, or- ganisms require specified and highly im- probable arrangements of lower-level con- stituents in order to maintain their form and function. Genes require specified arrange- ments of nucleotide bases; proteins require specified arrangements of amino acids; new cell types require specified arrangements of systems of proteins; body plans require spe- cialized arrangements of cell types and or- gans. Organisms not only contain informa- tion-rich components (such as proteins and genes), but they comprise information-rich arrangements of those components and the systems that comprise them. Yet we know, based on our present experience of cause and effect relationships, that design engi- neers—possessing purposive intelligence and rationality—have the ability to produce information-rich hierarchies in which both individual modules and the arrangements of those modules exhibit complexity and spec- ificity—information so defined. Individual transistors, resistors, and capacitors exhibit considerable complexity and specificity of design; at a higher level of organization, 233 their specific arrangement within an inte- grated circuit represents additional infor- mation and reflects further design. Con- scious and rational agents have, as part of their powers of purposive intelligence, the capacity to design information-rich parts and to organize those parts into functional information-rich systems and _ hierarchies. Further, we know of no other causal entity or process that has this capacity. Clearly, we have good reason to doubt that mutation and selection, self-organizational processes or laws of nature, can produce the infor- mation-rich components, systems, and body plans necessary to explain the origination of morphological novelty such as that which arises in the Cambrian period. There is a third reason to consider pur- pose or design as an explanation for the or- igin of biological form and information: purposive agents have just those necessary powers that natural selection lacks as a con- dition of its causal adequacy. At several points in the previous analysis, we saw that natural selection lacked the ability to gen- erate novel information precisely because it can only act after new functional CSI has arisen. Natural selection can favor new pro- teins, and genes, but only after they per- form some function. The job of generating new functional genes, proteins and systems of proteins therefore falls entirely to ran- dom mutations. Yet without functional cri- teria to guide a search through the space of possible sequences, random variation is probabilistically doomed. What is needed is not just a source of variation (i.e., the free- dom to search a space of possibilities) or a mode of selection that can operate after the fact of a successful search, but instead a means of selection that (a) operates during a search—before success—and that (b) is guided by information about, or knowledge of, a functional target. Demonstration of this requirement has come from an unlikely quarter: genetic al- gorithms. Genetic algorithms are programs that allegedly simulate the creative power of mutation and selection. Dawkins and 234 Ktippers, for example, have developed computer programs that putatively simulate the production of genetic information by mutation and natural selection (Dawkins 1986:47—49, Ktippers 1987:355—369). Nev- ertheless, as shown elsewhere (Meyer 1998: 127-128, 2003:247—248), these programs only succeed by the illicit expedient of pro- viding the computer with a “target se- quence”’ and then treating relatively greater proximity to future function (i.e., the target sequence), not actual present function, as a selection criterion. As Berlinski (2000) has argued, genetic algorithms need something akin to a “forward looking memory” in or- der to succeed. Yet such foresighted selec- tion has no analogue in nature. In biology, where differential survival depends upon maintaining function, selection cannot oc- cur before new functional sequences arise. Natural selection lacks foresight. What natural selection lacks, intelligent selection—purposive or goal-directed de- sign—provides. Rational agents can arrange both matter and symbols with distant goals in mind. In using language, the human mind routinely “‘finds’” or generates highly improbable linguistic sequences to convey an intended or preconceived idea. In the process of thought, functional objectives precede and constrain the selection of words, sounds and symbols to generate functional (and indeed meaningful) se- quences from among a vast ensemble of meaningless alternative combinations of sound or symbol (Denton 1986:309-311). Similarly, the construction of complex tech- nological objects and products, such as bridges, circuit boards, engines and soft- ware, result from the application of goal- directed constraints (Polanyi 1967, 1968). Indeed, in all functionally integrated com- plex systems where the cause is known by experience or observation, design engineers or other intelligent agents applied boundary constraints to limit possibilities in order to produce improbable forms, sequences or structures. Rational agents have repeatedly demonstrated the capacity to constrain the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON possible to actualize improbable but initial- ly unrealized future functions. Repeated ex- perience affirms that intelligent agents (minds) uniquely possess such causal pow- ers. Analysis of the problem of the origin of biological information, therefore, exposes a deficiency in the causal powers of natural selection that corresponds precisely to pow- ers that agents are uniquely known to pos- sess. Intelligent agents have foresight. Such agents can select functional goals before they exist. They can devise or select mate- rial means to accomplish those ends from among an array of possibilities and then ac- tualize those goals in accord with a precon- ceived design plan or set of functional re- quirements. Rational agents can constrain combinatorial space with distant outcomes in mind. The causal powers that natural se- lection lacks—almost by definition—are as- sociated with the attributes of conscious- ness and rationality—with purposive intel- ligence. Thus, by invoking design to ex- plain the origin of new _ biological information, contemporary design theorists are not positing an arbitrary explanatory el- ement unmotivated by a consideration of the evidence. Instead, they are positing an entity possessing precisely the attributes and causal powers that the phenomenon in question requires as a condition of its pro- duction and explanation. Conclusion An experience-based analysis of the causal powers of various explanatory hy- potheses suggests purposive or intelligent design as a causally adequate—and perhaps the most causally adequate—explanation for the origin of the complex specified in- formation required to build the Cambrian animals and the novel forms they represent. 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Cambridge University Press, Cambridge, United Kingdom. i ere Nh ae .. ere » arpa eT bOom-we's aus, et wath fs A —_p Me # Po a. om pO le, nee te Waele af rors = mare rete | { sik hydra y J Nad 4 ; - (hoes al, a were mys jimmie ie? eal Verda Pee <— ngs . ye rybiery * — . % i oat Cee Nee + | eels ited Law et: a lids f J ‘Ae age pha he, ~ ae i 9 i +i. ae we t ‘ a a 1 ay jer MLN it is pay rouse/Sue ¥ ‘1 2 mad ‘ = ‘ mabe é oh : te 7 @ a | ie . i ao { ’ hk vi id i . aus Lane 6 el So peaneial alae 7 7@ saerTt heaton teh i @¢ ieee ih 4 prey ee »@ a ie ry ? ie 4 : -” as ¥ a 5 | ' j D r q fi L . ¢ af 4 f i i 4 t ‘a ‘ a a ° on. bern ing end, AS i iP Oui 1 INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. 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Please include email address on all correspondence. Costs.—Printed pages @ $65.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. Front cover—from this issue, p. 206. CONTENTS Pseudopaguristes shidarai, a new species of hermit crab (Crustacea: Decapoda: Diogenidae) from Japan, the fourth species of the genus Akira Asakura A new species of Procambarus (Crustacea: Decapoda: Cambaridae) from Veracruz, Mexico Marilu Lopez-Mejia, Fernando Alvarez, and Luis M. Mejia-Ortiz Brackenridgia ashleyi, a new species of terrestrial isopod from Tumbling Creek Cave, Missouri (Isopoda: Oniscidea: Trichoniscidae) Julian J. Lewis New species and records of Bopyridae (Crustacea: Isopoda) infesting species of the genus Upogebia (Crustacea: Decapoda: Upogebiidae): the genera Orthione Markham, 1988, and Gyge Cornalia & Panceri, 1861 John C. Markham Three new species and a new genus of Farreidae (Porifera: Hexatinellida: Hexactinosida) Kirk Duplessis and Henry M. Reiswig The origin of biological information and the higher taxonomic categories Stephen C. Meyer TITUTION LIBRARIES iN WN | | 153 169 176 186 199 213 a ~ a t | yw ear 324X a BUX \) 4 PROCEEDINGS of THE BIOLOGICAL SOCIETY or WASHINGTON 7 DECEMBER 2004 VOLUME 117 NUMBER 3 THE BIOLOGICAL SOCIETY OF WASHINGTON 2003-2004 Officers President: Roy W. McDiarmid Secretary: Carole C. Baldwin President-elect: W. Ronald Heyer Treasurer: T. Chad Walter Elected Council Michael D. Carleton G. David Johnson Clyde Roper Michael Vecchione Marilyn Schotte Don Wilson Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: Richard C. Banks Associate Editors Plants: Carol Hotton Invertebrates: Stephen L. Gardiner Insects: Wayne N. Mathis Christopher B. Boyko Vertebrates: Gary R. Graves Janet W. Reid Ed Murdy Invertebrate Paleontology: Gale A. 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Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. Manuscripts, corrected proofs, and editorial questions should be sent to: EDITOR, RICHARD C. BANKS DEPT. OF ZOOLOGY MRC-116 NATIONAL MUSEUM OF NATURAL HISTORY WASHINGTON, D.C. 20013-7012 Known office of publication: Biological Society of Washington, National Museum of Natural History, Washington, D.C. 20013-7012. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Periodicals postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):241. 2004. STATEMENT FROM THE COUNCIL OF THE BIOLOGICAL SOCIETY OF WASHINGTON The paper by Stephen C. Meyer, “The origin of biological information and the higher taxonomic categories,” in vol. 117, no. 2, pp. 213-239 of the Proceedings of the Biological Society of Washington, was published at the discretion of the former ed- itor, Richard v. Sternberg. Contrary to typ- ical editorial practices, the paper was pub- lished without review by any associate ed- itor; Sternberg handled the entire review process. The Council, which includes offi- cers, elected councilors, and past presidents, and the associate editors would have deemed the paper inappropriate for the pag- es of the Proceedings because the subject matter represents such a significant depar- ture from the nearly purely systematic con- tent for which this journal has been known throughout its 122-year history. For the same reason, the journal will not publish a rebuttal to the thesis of the paper, the su- periority of intelligent design (ID) over evolution as an explanation of the emer- gence of Cambrian body-plan diversity. The Council endorses a resolution on ID pub- lished by the American Association for the Advancement of Science (www.aaas.org/ news/releases/2002/1 106id2.shtml), which observes that there is no credible scientific evidence supporting ID as a testable hy- pothesis to explain the origin of organic di- versity. Accordingly, the Meyer paper does not meet the scientific standards of the Pro- ceedings. We have reviewed and revised editorial policies to ensure that the goals of the So- ciety, as reflected in its journal, are clearly understood by all. Through a web presence (www.biolsocwash.org) and improvements in the journal, the Society hopes not only to continue but to increase its service to the world community of systematic biologists. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):242—250. 2004. A review of the North American subspecies of the Great Blue Heron (Ardea herodias) Robert W. Dickerman Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131, U.S.A., e-mail: bobdickm @unm.edu Abstract.—Geographic variation in the great Blue Heron (Ardea herodias) was comprehensively reviewed by H. C. Oberholser (1912), who recognized nine North American subspecies—excluding the so-called Great White Heron (A. occidentalis = A. h. occidentalis). Oberholser’s revision provided the frame- work generally followed in subsequent subspecific treatments of this species. However, Payne’s (1979) brief general summary of this species’ geographic variation rejects most of these North American taxa, recognizing as valid only the nominate subspecies and those of the Pacific northwest [A. h. fannini] and Florida [A. h. occidentalis]. My studies verify that A. h. herodias and A. h. fannini are taxonomically distinct, along with A. h. wardi in which I include A. h. treganzai, A. h. hyperonca, and A. h. sanctilucae. In addition, I regard A. h. lessoni, A. h. adoxa, and A. h. olgista as synonymous with A. h. herodias, as all are based on migrant specimens of this form. In addition, I suspect Payne is justified as recognizing the Caribbean A. h. occidentalis as valid, based on its white plumage and shorter head plumes. The Great Blue Heron (Ardea herodias) nests in North America from southeastern Alaska, southern British Columbia, north- ern Alberta, central Saskatchewan, northern Manitoba, northern Ontario, southern Que- bec, New Brunswick, and Nova Scotia southward to the Gulf states, southern Flor- ida; on the coastal lowlands of Mexico south to Tabasco, Nayarit, and Baja Cali- fornia; and locally in the Caribbean Basin (A.O.U. 1998). There are no nesting season specimens of Great Blue Herons taken be- tween the Yucatan Peninsula of Mexico and Venezuela. Oberholser (1912) recognized nine subspecies over this extensive area, these being A. h. herodias L., 1758 (type locality: America [= Hudson Bay, Cana- da]); A. h. wardi Ridgway, 1882 (Oyster [= Estero] Bay, Florida; A. h. treganzai Court, 1908 (Egg Island, Great Salt Lake, Utah); A. h. fannini Chapman, 1901 (Skidegate [Graham Island], Queen Charlotte Islands, British Columbia); A. h. hyperonca Ober- holser, 1912 (Baird [Shasta Co.], Califor- nia); A. h. sanctilucae Thayer and Bangs, 1912 (Espiritu Santo Island, Baja Califor- nia); A. h. lessoni Wagler, 1831 (Mexico); A. h. adoxa Oberholser, 1912 (Curacao); and A. h. olgista Oberholser, 1912 (San Clemente Island, California). Not included in Oberholser’s revision was the so-called Great White Heron (A. occidentalis), which is now widely regarded as a white morph of A. herodias (e.g., A.O.U. 1998). In ad- dition, he did include the endemic A. h. cognatus of the Galapago Islands, which is the only nesting population of this species outside North America. Oberholser’s (1912) subspecies were based on differences in plumage coloration and measurements among populations, which in some cases included migrants from other areas. In fact, although Ober- holser was aware of both migration and oth- er forms of dispersal in this species, he ap- pears to have underestimated the extent of VOLUME 117, NUMBER 3 this phenomenon. For example, Bond (1935) found that Oberholser’s A. h. adoxa from Curacao is based on a series of eight specimens, all of which are southward mi- grants of A. h. herodias. In addition, the adult female holotype (examined) for Ob- erholser’s (abid.) A. h. olgista from San Cle- mente Island is also an example of the nom- inate form, based on its dark coloration and a wing chord of 433 mm, even though pre- viously synonomized with the locally nest- ing A. h. “‘hyperonca” (= A. h. wardi) by Grinnell and Miller (1944) and Hellmayr and Conover (1948). Hellmayr and Con- over (1948) also listed A. lessoni Wagler as a synomym of A. h. herodias, simply noting “type in Munich Museum examined.” Oberholser’s revision provided the framework generally followed in subse- quent subspecific treatments such as A. O. U. (1931, 1957), Peters (1931), Friedmann et al. (1950), Palmer (1962) and Hancock and Elliot (1978). More recently, Payne (1979) has treated overall geographic variation in the A. her- odias complex (including A. occidentalis), although he did so only briefly, generally, and without measurements or references to subspecific names. He recognized only three taxa in North America: the wide- spread A. h. herodias, A. h. fannini of the north Pacific Coast, and the white-plum- aged A. h. occidentalis of the Caribbean Ba- sin. My revisionary work on the Great Blue Heron began in an attempt to identify then recently collected Mexican specimens in the 1960’s. Since then I have examined most of the available adult specimens in North American collections. My findings generally agree with those of Payne, except that I also recognize the populations of pal- er and larger birds of southern and western North America as A. h. wardi. I did not examine plumage variation in nesting pop- ulations of the Caribbean Basin, but these may constitute a valid subspecies (A. h. oc- cidentalis) based on the dominance of the white morph (rare elsewhere). If not rec- ognizable, then these populations and those 243 of A. h. wardi should be merged under the older name of A. h. occidentalis. Methods Several caveats apply to the museum specimens used in this study, the first being the dearth of properly labeled and prepared adult nesting season skins for studies of geographic variation among Great Blue Herons in North America. For example, I found no nesting season adult males from Delaware, Virginia, West Virginia, and Kentucky; only single males from Mary- land, Tennessee, South Carolina, and Ala- bama; and only two from North Carolina! Secondly, many specimens lack informa- tion on gonad size, weight, and fat condi- tion, making it difficult to ascertain whether such birds are likely nesting or are mi- grants. As a result, one must often assume that birds are nesting on the basis of col- lection localities and dates, which can be complicated by (a) regional differences in the timing of breeding activities and (b) the migration and other forms of dispersal in this species. For example, we know post- nesting southern populations (A. h. wardi) can be dispersing northward in the north- eastern U.S. while northern birds (A. h. her- odias) are in the process of nesting (Dick- erman 2002). Whereas coloration and mea- surements do distinguish these subspecies, some specimens overlap or intergrade be- tween the two. As a result, these may be either included in or excluded from nesting samples, thus introducing some degree of bias into the data. In any case, I have ar- bitrarily set the nesting season for most North American populations of this species as April to July, subject to modification based on specimens’ gonadal condition, weight, fat levels, coloration, and measure- ments. A second caveat with Great Blue Heron specimens is that the plumage coloration can be altered by a variety of factors, in- cluding wear, bleaching, molt stage, chem- icals used to preserve or protect skins, mu- 244 seum age, and especially staining due to the leakage and oxidation of body fat. In ad- dition, winter-taken specimens in the north may be under greater nutritional stress, so that they may produce less powder down to coat the feathers. This in turn would greatly affect feather color, as the powder-down coating produces a pale bloom that makes the plumage appear lighter. In fact, this same effect can be extreme when the plum- age is washed and the powder-down is re- moved (Dickerman 2004). For my final comparisons of plumage coloration in Great Blue Heron populations, I borrowed 26 adult skins taken throughout North America, representing all of the mainland forms. All were clean but un- washed specimens, taken as early in the nesting season and chronologically recently as possible. As I found no differences in plumage color between males and females, I combined the sexes for these comparisons. In addition, I measured 214 males and 189 females for the following characters: wing chord, tail length, exposed culmen, and tar- sus length (all in mm). After a preliminary analysis, I have variously grouped these measurements by subspecies, area, and sometimes type specimens (Table 1). I then calculated the sample sizes, ranges, means, and standard deviations for the four men- sural characters, as well as performing two- sample t-tests to determine the significances (P = 0.05) of differences. I did not analyze either plumage or mensural variation in oth- er age classes, because sample sizes were too small for juveniles and nestlings. Im- matures were not analyzed. Results As did Oberholser (1912), I find Great Blue Herons can be aggregated into three distinct North American nesting popula- tions on the basis of plumage coloration, exclusive of the white-phased birds of the Caribbean Basin (A. h. occidentalis). More specifically, this variation involves the col- oration of the upper-parts, neck, and wing PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON feathers in adult birds, which ranges from pale to darker gray. The first of these ag- gregates consists of the moderately gray populations to which the name A. h. hero- dias can be applied. These nest in southern Canada west to interior southern British Columbia, then southward in the United States to eastern Washington, North Dako- ta, Wisconsin, Indiana, Maryland, and South Carolina. The second aggregate of paler populations in the southeastern, cen- tral, and western U.S. and Mexico that Ob- erholser (ibid.) assigned to four subspecies, of which the oldest name is A. h. wardi with A. h. treganzai, A. h. hyperonca and A. h. sanctilucae here considered synonyms. And the third is the darker gray A. h. fannini, whose range I have recommended be re- stricted to the coastal region of northwest- ern British Columbia and adjacent Alaska, specifically the Queen Charlotte Islands north to Prince William Sound (Dickerman 2004). However, as noted earlier, the slaty- black coloration of the holotype (Chapman 1901) is abnormally dark, apparently due to washing that removed the powder down coating and thus the paler bloom of the plumage (Dickerman 2004). Oberholser (1912) further characterized nesting Great Blue Herons on the basis of measurements, which he particularly em- phasized in allotting pale populations to four subspecies. Given this, I also assessed measurements in this species, in which males generally average larger than females in nesting populations (Table 1). For ex- ample, my overall samples reveal that males are 4.1% larger in wind chord, 3.5% in tail length, 6.5% in exposed culmen, and 6.5% in tarsus length (including only “‘typ- ical populations of named forms, excluding fannini because of small sample size). However, the sexes overlap in all of these mensural characters, and t-tests often show the differences are not significant at the P = 0.05 level. Nonetheless, it is important to segregate the sexes when using measure- ments to allocate specimens to subspecies VOLUME 117, NUMBER 3 and populations. As for the mensural char- acters themselves, I found the following: Wing chord.—Nesting populations with the longest wings are 4.8% and 5.2% great- er than those with the shortest in males and females, respectively. Means are smallest in A. h. herodias, generally becoming pro- gressively larger through the populations of the interior western U.S., the Pacific Coast region, and Mexico to the southeastern U.S. (Table 1). However, a notable departure from this is that A. h. occidentalis has the wing chord intermediate, as opposed to be- ing among the largest in the species. T-tests reveal that A. h. herodias averages signifi- cantly shorter in wing length than all but two other North American populations, the exceptions being males of A. h. fannini and A. h. “‘treganzai.”” By contrast, the latter is significantly shorter-winged than all but one of the A. h. wardi populations, that being the small Texas sample. All other popula- tional differences in this character are insig- nificant, with clinal intergradation being smoother among females than males. Tail length.—Nesting populations with the longest tails are 7.7% and 8.8% greater than those with the shortest in males and females, respectively. Means are smallest in A. h. herodias and become progressively and significantly larger in Texas/Florida populations of A. h. wardi, A. h. “‘hyperon- ca” X A. h. fannini, and A. h. fannini (Table 1). All other populational differences in this character are insignificant, with rather mo- saic intergradation occurring among both males and females. Exposed culmen.—Nesting populations with the longest culmens are 29.4% and 31.9% greater than those with the shortest in males and females, respectively. Means are smallest in A. h. fannini and then A. h. “hyperonca”’ X A. h. fannini, each of which has a significantly shorter culmen than all other populations of the species (Table 1). Elsewhere, males average small- est in A. h. herodias, which differ signifi- cantly from those with the longest culmens in Florida, Texas, and eastern Mexican A. 245 h. wardi and A. h. occidentalis. However, these extremes intergrade circuitously through A. h. “‘treganzai,”’ A. h. “‘hyperon- ca”’ and A. h. “‘sanctilucae’’ with a similar pattern of geographic variation, except that A. h. occidentalis has a significantly longer culmen than all but one A. h. wardi (sensu latu) population—that in eastern Mexico, which has a sample size of only one! Tarsus length.—Nesting populations with the longest tarsi are 33.2% and 27.3% greater than those with the smallest in males and females, respectively. Means in males are shortest in A. h. fannini and then A. h. “hyperonca”’ X A. h. fannini, each of which has a significantly shorter tarsus than all other populations of the species (Table 1). The same is true with females, except that the means of those two populations are essentially identical. Elsewhere, males and females average smallest in A. h. herodias and A. h.“‘hyperonca”’ which differ signif- icantly from those with the longest tarsi in Florida A. h. wardi and A. h. occidentalis. However, these extremes intergrade circui- tously through A. h. “treganzai,”’ A. h. “sanctilucae”’ and Texas/eastern Mexican populations of A. h. wardi. Discussion and Conclusions Based on these findings, I recommend recognizing three subspecies among North American nesting populations of the Great Blue Heron, excluding the white-plumaged A. h. occidentalis of the Carribbean Basin. The first is Ardea herodias herodias L, with its moderately gray plumage and a nesting range as outlined above (see Results section). This the most highly migratory of the subspecies, with birds regularly moving southward into Central America and the Caribbean and as far as Belize, Panama, Colombia, Venezuela, Curacao, and the Do- minican Republic (also eastward to Ber- muda). In addition, lesser numbers move elsewhere, including northward to Hudson Bay, northern Quebec, Anticosti Island, and Newfoundland (plus as a vagrant to Green- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 246 el ds (S991) SLI-OSI 6 el dS (ZT 107) L7C-OL1 L6 ds (9'°CS1) P91-Cr1 CL dS (O'291) O81-Lr1 c6l LL ds (6°8L1) €0C-491 v6 ds (6'E81) 861-L91 6L dS (L°881) OOC-ELI L383 ds (9°€61) TITF81 (CO) ds (0'661) TIT-S8I SCC 9VI dS (8°COT) ETT—081 Ls ds (8°6L1) 881-691 snsieL 6I Ic 9C 6C vl 91 8C u (O'S) as (SEL) SVI-VEl cS dS (L'6S1) L9I—-0S1 c9 ds (8'PZ1) €€1-911 CL dS (9°CE1) 61-071 9SI 99 ds (6LY1) O9I-€EI 038 dS (S9rI1) I9I-€€1 cs ds (L'6v1) 8S1-6E1 Oe dS (1°6S1) p91-rS1 (69) ds (0'091) S9I-8rI 99] o8 ds (9'VS1) TLI-OF1 19 ds (l'rrl) SSI-871 uowynD CE Of 09 ds (O'LLI) 881-191 so[euloy CL ds (C781) S6I-ILI ys ds (9681) 861-781 v9 dS (€061) 802-081 col V9 ds (CL81) 10C-vLI 89 ds (6'8L1) 881-S9I 9°83 dS (L'6L1) 161-091 crv ds (Q'181) 681-9LI (vb) dS (8'C81) 981-8ZLI 86] 9S ds (L781) S6I-ILI 8S ds (V'LLI) C61-S9T So[vIN LEAL VE 6 6C VC 61 Nn N C6 dS (8°CSr) O8h-lEr OOL ds (108) S6r7-CLr 801 GS (O'VLy) 887-8Sr CCI GS (6 78) 80S—-09P OOS VII ds (8'16r) 9IS-PLY orl ds (O'9Lr) S6r-rEr 9 II ds (9°S8r) TOS-09P L6 ds (L681) 867-SLY LC! dS (C'S8h) 00S-99F 81S sel dS (O'€6r) EES-OLV 9CI dS (O'OLb) 96t-lbr pioyo Sur 6C le 8C 61 u SDIPOAIY S1]DIUAp1IIO AJIIS nsuas ‘uauupf +. DoUOsAdKY,, K quiuupf zpouosadky adK{, BIUIOJI[VD UloyINOS pur [eUSD pouosadKy, , ‘ ¢, JDZUDSAJ, , vIuIOjeD eleg IDINJNIUDS, , ‘ OdIXdJJ JO ISVOD FIND sexo Ipipm adK J, EPO 1pipM |SDIPO1AY ee uonejndog De “UONPIANp piepur}s pue ‘(uvour) aSuri “Iaquinu YIM ‘eOLIOWIW YON Wo (svIposay Yapsy) SUOCIOH IN[_ WosiH ynpve uosves Busou ATISOUL JO SIOJOLUT][MUW UL SJUSWSINSvII— | IGP 247 VOLUME 117, NUMBER 3 “BUTTOIR) YINOG “eUTpOIeD YON “erursirA ‘pueyArepy “ejOyeq LION O91 ds (6°C61) 9@Z-L9OT ee ds (81ST) LSI-9F1 TL ds (9°CST) E9I-ChI CL dS (1991) P8I-CSI EL ds (9°891) S8I-ScI OO! ds (8°OL1) Z8I-CSI C8 9 Tl ds (V'OLI) Z8I-€STI vel ds (8°€61) 9@Z-181 CI 8C CC 06 GS (OSI) TLI-CrI VL ds (V LIT) 9@I-V01 67 dS (6°STL) ZEI-8TI cL dS (CEL) TSI-€TI L9 ds (€6EL) 9SI-€TI Le ds (T°8€1) OVI-TEl LSI VL dS (9%) ESI-€El 9°9 ds (OPI) LSI-8EI vl Ic O& Lv TL dS (8°ZL1) S8I-O91 vs ds (8981) S6I-r8I 9S ds (0'081) O61-L9I LS ds (S'8L1) 061-891 69 ds (L’SLI) C6I-091 Sr ds (O'ELT) O8I-F9T ILI €¢ ds (CLLT) C8I-ILI 79 dS (V8L1) 761-691 cl CC 9¢ OV PUNO WITTTLAA POULT O1 YOU “eysePY [eIseOD pu spue]sT sNO[IeYD useNd « “UOISUTYSe AA [BISBOD Pur eIQUINJOD YsHiIg uoyINog x “OOTXO MON “BUOZITY “eTUIOFITED “ye ‘uose1Q ‘oyepy , SLI ds (F'99r) O1S—Sbr OTI ds (8°L9) 687-6Sr SII ds (S€9b) S6P—-Sbr € Ol ds (9°89r) S8h-0Sr Lel ds (€6Sr) l6r—-CEr (A ORS (T 19h) O8b-8Er cov COI ds (9'OLP) E8h—-Srr C9Ol ds (T9OLv) SOS—CSb al 6C Lv (TI6I) JesfoyieqO Wor sjuouaInsvoyy z “UBSTYOI] “UISUOOSIAA “kULIPUT IOR MON ‘siesnyoessey] “sure ‘noyo0UUOD |, SIpUap1II0 MIDIS nsuas ‘Tuiuupf pp2uosadky % quluupl BIUIOFTED YING ‘[eyuaD . DIU0sadKy, , ¢. JDZUDSAL, , BIUIO}eD vleg << @VINJHIUDS OOrXafJ JO ISVOD JNH ” SBXOL, epuloly 1plDM snsie u uowynD u [RL u Proyo Sur, u uone[ndog ‘ponunuog— | 21qRL 248 land), and west to California, Arizona, New Mexico, and Colorado. A. h. adoxa and A. h. olgista of Oberholser (1912) and A. h. lessoni Wagler were based on migrant A. h. herodias. Oberholser restricted the type lo- cality of A. lessoni to the Valley of Mexico. However, there are no records of the species ever having nested in the Valley, and Payne (1979) was correct in just citing Mexico as the type locality. Hellmayr and Conover (1948) erroneously placed A. h. olgista in the synonomy of A. h. hyperonca. However, the wing chord of the type (433 mm) is far too short for that population and is even short for A. h. herodias! It is also darker, as in the nominate population. The second subspecies recognized here is the pale A. h. wardi, which includes A. h. “terganzai,’’ A. h.“‘hyperonca,”’ and A. h. “santilucae.’’ Payne (1979) wrote in a foot- note (p.198) ““The type of wardi was taken on 5 January 1881. It is not known whether this was a local breeding bird or a wintering bird from a more northern population.” There is no doubt that it was from the local population. The type of A. h. wardi is a very large bird (Oberholser 1912, Table 1), larger in all measurements than any male A. h. herodias, and it has longer tail, culmen, and tarsus measurements than any other male A. h. wardi. Size is largest in the southeast (Table 1) and smallest in the Great Basin region (A. h. “‘treganzai’’), and only extremes can be identified based on measurements (Dickerman 1992, 2002). Oberholser described A. h. “hyperonca”’ as the color of A. h. herodias, but larger. The type is from northern California and is somewhat intermediate towards A. h. fan- nini and indeed inseparable from A. h. her- odias in color, but it is larger than the larg- est male of A. h. fannini or of A. h. herodias (Table 1). However, specimens from central California in the California Academy of Science and the Museum of Vertebrate Zo- ology labeled A. h. hyperonca are insepa- rable in color from a topotype of A. h. tre- ganzai, from early nesting season speci- mens from southern New Mexico and south PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Texas, or from a midwinter specimen of A. h. wardi from Florida. Contra Payne’s comment on clinal vari- ation in size in the east (1979), there are not enough nesting colony or even nesting season specimens yet available to fully doc- ument a cline. As mentioned earlier, there are no nesting season adult males from Del- aware, Virginia, West Virginia or Ken- tucky; only single males from Maryland, Tennessee, South Carolina, and Alabama: and only two from North Carolina. Indeed the best clinal variation in size is on the west coast, with an increase in culmen and tarsal length from A. h. fannini in the north, through an intermediate population in southern British Colombia and Washington, to the long-billed, long- legged A. h. “‘hy- peronca”’ population of California (Dick- erman 2004). A. h. wardi may be separated from A. h. herodias as follows: 1. Neck of A. h. herodias darker, colder ““vinaceous” gray vs. ““warmer,” paler neck of A. h. wardi.. 2. Chestnut of mid-ventral neck stripe is more extensive and darker in A. h. her- odias, and usually extends to the area behind and below the eye, thus faintly outlining the white of the throat; in A. h. wardi the area behind and below the eye is always white; chestnut of mid-neck is reduced and paler. 3. Dorsum and wings are darker in A. h. herodias and paler in A. h. wardi. The third subspecies recognized here is A. h. fannini, which differs from A. h. her- odias in being darker gray in color and in having the exposed culmen and tarsus sig- nificantly shorter and tail longer (plus wing in males, Dickerman 2004). A. h. fannini differs in being notably much darker gray than A. h. wardi (sensu latu), and in having the exposed culmen and tarsus significantly shorter. In addition, males have significant- ly shorter wings than all A. h. wardi pop- ulations except A. h. “‘treganzai”’ on the in- terior western U.S. A. h. fannini seems to VOLUME 117, NUMBER 3 differ from other Great Blue Herons in that it perforce fishes much of the time from rocks rather than wading, as do the other longer-legged subspecies. It appears to be largely resident within its nesting range (contra A.O.U. 1957), with the only extra- limital specimen being an adult taken at Wainwright on the Arctic coast of Alaska (Brock 1959). I know of no specimens of A. h. fannini (as here defined) from south of the Queen Charlotte Islands. A. h. fannini intergrades southward with A. h. “‘hyperon- ca’ and perhaps A. h. “‘treganzai”’ (both here = A. h. wardi) in southwestern British Columbia (including Vancouver Island) and western Washington (Dickerman 2004). For example, that population is paler gray as in A. h. wardi (sensu latu), but it is closer to A. h. fannini in the shorter exposed culmen, tarsus, and male wing chord. Acknowledgments The author has measured or compared specimens of Great Blue Herons in over 30 museums, sometimes more than once! He wishes to express his appreciation for the many courtesies he has received at the fol- lowing institutions: American Museum of Natural History, New York; Academy of Natural Sciences, Philadelphia; California Academy of Sciences, San Francisco; Car- negie Museum of Natural History, Pitts- burgh; Coleccion Ornitologico Phelps, Ca- racas; Colorado State University Coopera- tive Wildlife Research Vertebrate Collec- tion, Fort Collins; Cornell University Museum of Vertebrates, Ithaca; Cowan Ver- tebrate Museum, University of British Co- lumbia; Delaware Museum of Natural His- tory, Greenville; Denver Museum of Natu- ral History, Denver; Donald R. Dickey Col- lection, University California, Los Angeles; James Ford Bell Museum of Natural His- tory, University of Minnesota; James R. Slater Museum of Natural History, Univer- sity of Puget Sound; Museum of Natural History of Los Angeles County, Los An- geles; Museum of Comparative Zoology, 249 Harvard; Museum of Natural Science, Lou- isiana State University; Museum of South- western Biology, University of New Mex- ico; Museum of Vertebrate Zoology, Uni- versity of California, Berkeley; National Museum of Canada, Ottawa; National Mu- seum Natural History, Washington, D.C; North Carolina State Museum of Natural Sciences, Raleigh; Peabody Museum of Natural History, Yale; Royal British Co- lumbia Museum, Victoria; Sam Noble Mu- seum of Natural History, University of Oklahoma; San Diego Museum of Naturai History, San Diego; Texas Cooperative Wildlife Collection, Texas A&M; The Field Museum, Chicago; Thomas Burke Memo- rial Washington State Museum, University of Washington; University of Alaska Mu- seum, Fairbanks; University of Kansas Mu- seum of Natural History, Lawrence; Uni- versity of Nebraska State Museum, Lin- coln; Western Foundation Vertebrate Zool- ogy; Camarillo; Virginia Tech Museum Natural History, Blacksburg; Zoology Mu- seum, University of Wisconsin; and the pri- vate collection of the late Allan R. Phillips. He especially wishes to thank the cura- tors of collections who kindly shipped to New York specimens that permitted final color comparisons at the American Muse- um of Natural History. These include: Cal- ifornia Academy of Sciences, San Francis- co; Carnegie Museum of Natural History, Pittsburgh; Denver Museum of Natural His- tory; Los Angeles County Museum of Nat- ural History; Museum of Southwestern Bi- ology, Albuquerque; Museum of Vertebrate Zoology, Berkeley; National Museum of Natural History, Washington, D.C.; Utah Museum of Natural History, Salt Lake City; and the Western Foundation of Vertebrate Zoology, [then in Los Angeles]. Christine Blake of the AMNH graciously received and repacked all specimens. John P. Hub- bard suffered through several revisions of this manuscript and improved it greatly. Literature Cited American Ornithologists’ Union. 1931. Check-list of North American birds, 4th edition. Lancaster, Pennsylvania, 526 pp. 250 . 1957. Check-list of North American birds, 5th edition. Lord Baltimore Press Inc., Baltimore, Maryland, 691 pp. Bond, J. 1935. The status of the Great Blue Heron in the West Indies—Auk 52:76-77. Dickerman, R. W. 1992. Northeastern records of Ardea herodias wardi from the southeastern United States —Kingbird 42:10-13. . 2002. An adult Ardea herodias wardi from the northeast.—Kingbird 52:35-37. . 2004. Characteristics and distribution of Ar- dea herodias fannini with comments on the ef- fects of washing on the holotype. Northwestern Naturalist 85:130—-133. Friedmann, H., L. Griscom, & R. T. Moore. 1950. Dis- tributional check-list of the birds of Mexico, part 1. Pacific Coast Avifauna 29. Hancock, J., & H. Elliott. 1978. The herons of the world. Harper and Row Publ. New York. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hellmayr, C. E. and H. Conover. 1948. Catalogue of birds of the Americas. part 1. No. 2. Zool. Se- ries, Field Mus. Nat. Hist. 8:434 pp. Grinnell, J.. & A. H. Miller. 1944. The distribution of the birds of California. Pacific Coast Avifauna 27: 608 pp. Palmer, R. S. 1962. Handbook of North American birds, vol. 1. Loons through Flamingos. Yale University Press, New Haven, Connecticut. 567 Pp. Payne, R. B. 1979. Ardeidae. In E. Mayr and G. W. Cottrell, eds., Check-list of birds of the world, vol. 1, 2nd edition. Mus. Comp. Zool., Cam- bridge, Massachusetts. Peters, J. L. 1931. Check-list of birds of the world, vol. 1. Harvard Press, Cambridge. Massachu- setts. 345 pp. Oberholser, H. C. 1912. A revision of the forms of the Great Blue Heron (A. herodias Linnaeus).— Proc. U.S. Nat. Mus. 43:531—559. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):251—265. 2004. A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Forét des Mikea of southwestern Madagascar Steven M. Goodman and Voahangy Soarimalala (SMG) Department of Zoology, Field Museum of Natural History, 1400 Roosevelt Road, Chicago, Illinois 60605, U.S.A., e-mail: goodman@fmnh.org and WWF-Madagascar, BP 738, Antananarivo (101), Madagascar, e-mail: sgoodman@wwf.mg; (VS) Département de Biologie Animale, Université d’ Antananarivo, BP 906, Antananarivo (101), Madagascar and Ecology Training Program, WWE-Madagascar, BP 738, Antananarivo (101), Madagascar, e-mail: etp@wwf.mg Abstract.—A new species of Microgale, M. jenkinsae (Lipotyphla: Tenre- cidae), is described based on two specimens taken during an early 2003 bio- logical survey of the Forét des Mikea in southwestern Madagascar. It is distin- guished from other congeners by numerous pelage, cranial, and dental char- acters. M. jenkinsae is the fourth known species in this genus confirmed to occur in the dry western and southern forests of the island. The Forét des Mikea, the only site M. jenkinsae is known from, is the last remaining block of a distinctive forest habitat and is under considerable threat from human habitat degradation. Action needs to be taken to protect this unique region. Résumé.—Une nouvelle espéce de Microgale, M. jenkinsae (Lipotyphla: Tenrecidae), est décrite a partir de deux spécimens récoltés au cours d’un in- ventaire biologique mené au début de l’année 2003 dans la forét des Mikea située au sud-ouest de Madagascar. On le distingue de ses autres congénéres par divers caractéres de pelage, craniens et dentaires. M. jenkinsae est la qua- trieme espéce connue de ce genre dont la présence est confirmée dans les foréts seches de l’ouest et du sud de Vile. La forét des Mikea, le seul site d’ou M. jenkinsae a été rapporté, est le dernier bloc d’un habitat forestier distinctif qui est cependant extrémement menacé par la dégradation de |’habitat perpétrée par ’ homme. Des actions doivent étre prises pour protéger cette région unique. On the island of Madagascar there is an endemic family of Lipotyphla, known as the Tenrecidae, that represents one of the most remarkable adaptive radiations found in living mammals (Olson & Goodman 2003). As currently circumscribed, Micro- gale (shrew tenrecs) a tenrecid genus in the subfamily Oryzorictinae, comprises 18 spe- cies (Jenkins 2003). On the basis of biolog- ical inventories and associated museum studies conducted over the past few de- cades, seven species of Microgale new to science have been named (Jenkins 1988, 1992, 1993; Jenkins et al. 1996, 1997; Goodman & Jenkins 1998: Jenkins & Goodman 1999), although one of these, M. pulla, has since been synonymized (Jenkins et al. 1997). Subsequent to the publication of MacPhee’s (1987) taxonomic revision of the genus Microgale, there has been a re- newed interest in the small mammals of Madagascar. With the advent of pit-fall de- vices to trap these animals, there has been a massive increase in available shrew tenrec specimens. This has lead to a series of pub- lications refining some of MacPhee’s taxo- nomic conclusions and a greater under- standing of intra-specific, particularly age related, and inter-specific variation amongst these animals. As witnessed by the recent 45:00 252 43:00 43:30 44:00 44:30 21:00 T : = + 21:30 } IN Morombe |, angcky | ) Lac lhotry —“\W——_— 22:00 + “ Befandriana Atsimo Vorehy Ankazoabo Atsimo it 2:30 | Collection site Vohibasiaf” | Bes Ankiloaka Praesens Tsifota a Manombo sa ae ~~ ng omSakarahe 23:00 | See ee a Fee fe & lfaty ie _ a eee Toliara\@ ___ Tropic of Capricorn | 23:30 St. Augustin LS Oni = a = Betioky 24:00 | | \ Lac Tsimanampetsotsa 24:30 = 0) 50 100 150 EE _ ee _ Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON N Antsiranana i) Vohemar Meena ve ) \ Antananarivo Morondava Belo-sur-Mer ‘g Morombe| 200 kilometers Map showing the collection locality of Microgale jenkinsae in the Forét des Mikea, the forested zone between Morombe and Manombo, in southwestern Madagascar (expanded inset). description of six new valid shrew tenrec taxa (an increase of 33%), Microgale tax- onomy is in flux as a result of ongoing bi- ological inventories and molecular and morphological studies. Of the currently recognized 18 species of Microgale, 15 are restricted to the eastern and northern moister portions of Madagas- car where they occur in either forests or marshes. Of the remaining three species, two have been collected over the past few decades in the dry western forests. These include M. brevicaudata, which occurs from the northern foothills of the Marojejy Massif in the northeast, a zone of humid forest but probably with a marked dry sea- son, north to Vohemar and the region of Antsiranana at the north end of Madagascar, and then south along the west portion of the island to at least the Onilahy River near To- liara (Goodman et al. unpublished; Fig. 1). The second species, M. nasoloi, is known from two inland isolated forests in south- western Madagascar in the vicinity of Sa- karaha—the Analavelona Massif and the Forét de Vohibasia (Jenkins & Goodman 1999; Fig. 1). M. longicaudata is the third species falling into this group and has been collected from both eastern humid forests and western dry forests. However, M. lon- gicaudata, as currently defined, includes several cryptic species and will soon be re- vised (Olson et al. in press). MacPhee’s (1987, Fig. 13) map of col- lecting localities for Microgale included 31 sites in the eastern humid forest where a total of nine species were trapped, two sites in the western dry deciduous each with sin- VOLUME 117, NUMBER 3 gle species (one based on owl pellets), and three sites in the southern spiny-bush each with two species (all based on owl pellets). Although considerable advances have been made concerning the species richness and distribution of shrew tenrecs since Mac- Phee’s important revision, these data indi- cate a greater diversity of this group in the more mesic portions of the island. Recent biological inventories of the western and southern forests of Madagascar have largely upheld this view. However, during a 2003 survey of the Forét des Mikea, the region between Morombe and Manombo (Fig. 1), we captured a Microgale that represents a previously undescribed species of shrew tenrec. Materials and Methods Our small mammal collection made in the Forét des Mikea contains two speci- mens of Microgale, and in order to deter- mine their taxonomic identity, we have con- sulted material housed in several natural history museums, which include: BMNH— The Natural History Museum, London (for- merly British Museum of Natural History); FMNH—Field Museum of Natural Histo- ry, Chicago; MNHN—Muséum National d’ Histoire Naturelle, Paris; and UADBA— Université d’ Antananarivo, Département de Biologie Animale. Five external measurements in millime- ters were taken from our two specimens be- fore preparation and included: total length, head and body length, tail length, hind foot length (not including claw), and ear length. Mass was measured with the use of a spring balance and recorded in grams. An additional six cranial and two dental measurements were taken using a digital calipers accurate to the nearest 0.1 mm. These measurements, and their definitions, are: breadth of braincase: the greatest dis- tance measured across the hamular process- es of the squamosals to the mastoid bullae; greatest length of skull: the distance be- tween the tips of the nasals and the poste- 253 rior most portion of the cranium; interor- bital breadth: the minimum distance across the frontal bones between the orbital fossae; length of mandibular tooth row: the maximum distance from distal surface of the third molar to anterior surface of the first incisor; length of nasal: the maximum distance from the posterior extension of the nasals to their anterior tip; length of palate: the shortest distance between the tip of the postpalatal spine and anterior surface of the first upper incisor; length of maxillary tooth row: the maximum distance from dis- tal surface of the third molar to anterior sur- face of the first incisor; and zygomatic breadth: the maximum span between the zygomatic processes of the maxillae. Tooth abbreviations include: I = incisor, d = deciduous, C = canine, PM = pre- molar, and M = molar. Upper case tooth abbreviations with superscript are used for upper teeth and lower case abbreviations with subscript for lower teeth. Cranial and dental nomenclature follows Hershkovitz (1977) and MacPhee (1987). After comparison of the two specimens collected in the Forét des Mikea to all de- scribed forms of Microgale, these individ- uals could not be allocated to any known form and are therefore described as a new species. Microgale jenkinsae, new species Fig. 2, 3, Tables 1, 2 Holotype.—FMNH 176215, sub-adult male, collected on 18 February 2003 by Steven M. Goodman and Voahangy Soari- malala, field number SMG 13489. The specimen was preserved as a round study skin, with associated skull and partial post- cranial skeleton. Tissue samples were pre- served in EDTA. The skin is in good con- dition with a small hole in the left thigh. The skull and partial postcranial skeleton are intact. Dental age is sub-adult with P still erupting and matches MacPhee’s (1987) eruption pattern stage 1. The basis- penoid-basioccipital sutures are unfused. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5 Fig. 2. Photograph of the holotype of Microgale jenkinsae (FMNH 176215), a sub-adult male collected on 18 February 2003 in the Forét des Mikea, 9.5 km west Ankiloaka, 22°46.7’S, 43°31.4’E. (Photograph taken by S. M. Goodman.) VOLUME 117, NUMBER 3 Table 1.—External measurements (in millimeters) and weight (in grams) of Microgale jenkinsae and other species of small Microgale. Measurements presented as mean + standard deviation (minimum—maximum, n). For samples of two or fewer specimens only the measurements are presented. Head and body length Tail length Hind foot length Ear length Weight Total length Species 62 719 15 18 4.9 143 M. jenkinsae (Holotype FMNH 176215) M. jenkinsae 5.3 18 14 81 59 147 (FMNH 176154) M. nasoloi 14.0 16 13 53 81 141 (Holotype FMNH 156187) M. pusilla 3.5 = 0.40 3.14.2,n =7 11.4 + 0.98 10-13, n =7 11.4 + 0.53 11-12, n 69.9 = 4.18 65-77, n = 7 51.4 = 2.70 127.4 + 6.16 119-136, n = 7 = 7 7 SID 2 ANY 51-64, n = 11 47-56, n 3.2 + 0.56 2.14.1, n = 11 8.6 = 0.45 9.6 + 0.50 9-10, n = 11 58.4 + 4.39 53-66, n = 11 117.7 + 5.83 110-128, n = 11 M. parvula 11 WD 2= Bw 12-16, n = 13 n 8-9, 8.9 = 1.59 6.3-12, n= 11 12.4 + 0.96 41.9 + 5.92 115.7 + 6.68 68.9 = 0.35 63-74, n = 13 35-41, n = 13 107-129, n = 13 M. brevicaudata 11-14, n = 13 9.4 + 2.97 6.8-15, n = 10 16.1 + 0.67 15-17, n = 12 15.9 = 0.90 85.8 + 3.98 80-93, n = 12 159.3 + 6.34 70.8 + 6.71 64-85, n = 10 150-169, n = 11 M. fotsifotsy 15-17, n = 12 255 External measurements are: total length 143 mm, head and body length 62 mm, tail length 79 mm, hind foot length (without claw) 15 mm, and ear length 18 mm. The animal weighed 4.9 gm (Table 1). Type locality.—Madagascar: Province de Toliara, Forét des Mikea, 9.5 km west An- kiloaka, 22°46.7'S, 43°31.4’E, elevation about 80 m above sea level (Fig. 1). The site is about 17 km inland from the Moz- ambique Channel. Habitat.—The holotype was obtained in partially disturbed dry transitional decidu- ous forest growing on red sands. It was cap- tured in a pitfall trap placed in relatively dense understory composed primarily of Xerophyta (Velloziaceae), small bushes, and succulent Euphorbiaceae. Diagnosis.—A relatively small member of the genus Microgale with a head and body length of 59-62 mm, tail length of 79-81 mm, and greatest skull length of 18.7-18.8 mm. Deciduous PM? is simple, caniniform, and single-rooted. The color of the dorsal pelage is mixed agouti and the venter is gray with white-tipped fur. The ears are notably long (18 mm) for a shrew tenrec of this size. Paratype.—FMNH 176154 (SMG 13492), sub-adult female from the same lo- cality as the holotype, collected 19 Febru- ary 2003, and prepared as fluid preserved specimen with extracted skull. The dental eruption pattern fits MacPhee’s (1987) stage 1. Tissues saved in EDTA. Distribution.—Microgale jenkinsae is known only from the type locality in the Forét des Mikea, southwestern Madagascar. Description.—A small species of Micro- gale having a tail longer than the head and body (Fig. 2). The dorsal fur is relatively dense and soft. Pelage from the level of the ears to the base of the tail (including the flanks), is a mixture of completely black and tannish-brown hairs, or those that are tannish-brown along most of their length and black-tipped, imparting an agouti ap- pearance. The agouti pattern runs anteriorly from the level of the ears to the eyes. An- 256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Cranial and dental measurements (in millimeters) and weight (in grams) of Microgale jenkinsae and other species of small Microgale. Measurements presented as mean =~ standard deviation (minimum — maximum, n). For samples of two or less specimens only the measurements are presented. Greatest length Species of skull M. jenkinsae 18.8 (Holotype FMNH 176215) M. jenkinsae 18.7 (FMNH 176154) M. nasoloi DBw) (Holotype FMNH 156187) M. pusilla 16.6 = 0.71 15.7-17.5, n = 7 M. parvula 16.5 + 0.47 15.5-17.0, = 12 M. brevicaudata 20.7 + 0.79 19.0—21.9, n = 13 M. fotsifotsy 21.1 + 0.76 20-22, n= 12 terior and lateral to this band, the pelage is distinctly paler in coloration, with the ma- jority of hairs being pale tan to silvery- white. Individual hairs along the dorsum measure 4—5 mm. Guard hairs are medium gray in color. The ventral pelage, with the exception of the portion surrounding the gular to mental regions, is gray based with off-white tips. The difference between the ventral and dorsal color pattern is pro- nounced, but grade into each other laterally instead of forming a well-demarcated line. Upper surfaces of fore feet and hind feet are covered with short silver-white fur, which on the hind feet extends slightly be- yond the claws as ungual tufts. The color of mystacial and rhinarial vibrissae vary from either completely beige-white or black, to black at the base and gradually becoming beige-white at the tips. Mystacial vibrissae reaching up to 20 mm and rhinar- ial vibrissae about 5 mm in length. Pinnae are notably long (18 mm) for a small Mi- crogale, dark brown in color, and covered internally and externally with fine, silvery- gray fur. The hind foot is relatively long (14-15 mm) for a small species of Microgale (Ta- ble 1). The first digit of the hind foot is less than one-third the length of the second dig- Zy gomatic Interorbital breadth minimum 6.9 4.0 6.9 4.0 8.3 >. I 6.1 + 0.24 3.4 + 0.17 5.6-6.3, n = 8 3.1-3.7, n = 9 Spall 22 O23 3.7 = 0:19 4.7-5.4, n = 12 3.3-4.0, n = 12 8.0 = 0.51 4.8 = 0.27 7.2-8.8, n = 13 4.3-5.2,n = 14 8.2 = 0.32 5.0 + 0.18 7.7-8.7, n = 11 4.6-5.2, n = 12 it. The second and third digits are subequal in length, with the fourth digit slightly lon- ger. The fifth digit is about two-thirds the length of the fourth. There are five plantar tubercles and, based on FMNH 176154 (SMG 13492), these are located at the base of digit 1 and digit 5, in intermediate po- sitions between the base of digits 2 and 3 and digits 3 and 4, and notably reduced as distal hypothenar and proximal thenar pads. The skin of the tail is dark brown dor- sally and tannish-brown ventrally, and forming a relatively well-demarcated line laterally separating these two surfaces. The tail is clothed with very fine silvery-white fur, which becomes slightly denser at the tip. In FMNH 176154 (SMG 13492) the last 10 mm of the tail is mottled dark-white. The skull is relatively short (Table 2), slightly flattened dorsolaterally, with a con- stricted interorbital region. The rostrum is relatively short and tapers anteriorly. The anterior portion of frontals consist of two slightly concave plates divided at the mid- dorsal line and the posterior portion is slightly-domed. The braincase has a slightly bulbous parietal and interparietal, a rounded supraoccipital and occipital, and a weakly defined occipital crest. Dentally the holo- type is a sub-adult with the erupting crown VOLUME 117, NUMBER 3 Table 2.—Extended. Braincase Length Length Length of Length of width of nasal of palate maxillary toothrow mandibular toothrow 8.4 8.9 8.1 8.2 8.0 8.2 8.0 Wall 8.2 8.1 9.2 10.3 9.8 10.1 9.7 6.9 = 0.11 7.0 + 0.37 1.3) = O34! 7.4 + 0.40 7.0 + 0.36 6.87.1, n = 7 6.4-7.5, n= 8 6.8-7.8, = 8 6.7-7.7, n=8 O54, 0 = 7/ 6.8 = 0.18 Wed) = O22 Wo3) 22 O23 TA + 0.28 7.2 + 0.19 6.5—7.0, n = 12 6.9-7.7, n = 12 6.9-7.6 ,n = 11 6.7-7.6, n= 12 2O= Os S12 8.7 = 0.27 9.3 + 0.62 9.3} 2£ O.S7/ 9.3) 22 0.5 8.8 + 0.35 8.2-9.1, n = 13 8.4-10.5, n = 14 8.2-10.2, n = 14 8.3-10.2, n = 14 8.1-9.4, n= 14 9.3 + 0.26 9.4 + 0.45 OFS 0139) 10.0 + 0.43 9.6 + 0.47 8.9-9.7, n = 12 8.9-10.3, n = 12 9.1-10.2, n= 12 9.3-10.7, n = 12 9.0-10.5, n = 12 of I present and all antemolars are decid- uous, fitting stage 1 in MacPhee’s (1987) tooth eruption pattern. The upper toothrows from dI' to dPM? slightly converge anteri- orly. The lingual margins of dPM? and M! to M2 are roughly parallel. Palatal foramina are present. Pterygoids are relatively short and broad, and the pterygoid processes winged-shape and curved mid-ventrally. The glenoid fossa is shallow and narrowly curved. The mandibles are slender, the cor- onoid processes are relatively narrow at their bases and pointed dorsally, and the an- gular processes are short and narrow, and the dorsal surface is not expanded (Fig. 3). The dentition is not markedly robust (Fig. 3). There is a gap between the dI' and dI* and between the dpm, and dpm,. The first upper incisor (dI') is small, bicuspid (bidentate), and the distostyle moderately well developed; the dl? with approximately the same crown height as dl’, the tricuspid (tridentate) has the anterior accessory cusp more developed than distostyle; the dl? is one-half crown the height of dI*? and reach- ing just beyond the level of the distostyle of the dI’, the bicuspid with small distos- tyle; the dC! robust with crown height reaching that of the dI’, with small acces- sory anterior cusp and pronounced distos- tyle; the dP? small, equal in crown height to distostyle of the dC'; the dP® is large, slightly greater in crown height than the dC', lingual ledge with well-developed pro- tocone, and the parastyle, mesiostyle, an- terior ectostyle, and distostyle present; the dP* is large, longer in crown length than M! to M?, with elongated paracone, the lingual ledge with a protocone more developed than M' to M?, anterior ectostyle approxi- mately same length as paracone, and the parastyle, mesiostyle, and distostyle pre- sent; the M! and M? large, parastyle, me- siostyle, anterior ectostyle, and distostyle present, and centro-buccal cleft slightly more prominent in M? than M!; the M? is reduced in size and compressed anteriopos- teriorly. The first lower incisor (di,) is large, slightly shorter in crown length to the di,, the posterior accessory cusp well-devel- oped; the di, is large, the posterior acces- sory cuspid well-developed; the di, is small, about one-half the crown length of lower (deciduous) canine; the dc is large, poste- rior accessory cuspid present, no anterior accessory cuspid; the dpm, is small, slightly shorter that posterior accessory cuspid of lower canine, poorly developed anterior ac- cessory cuspid and posterior accessory cus- pid, and single-rooted; the dpm, is moder- 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Views of the cranium and mandible of the holotype of Microgale jenkinsae (FMNH 176215): upper left, dorsal view; upper right, ventral view; lower center lateral view of cranium and mandible. (Photograph taken by J. Weinstein, image number Z94379_05d.) VOLUME 117, NUMBER 3 ate in size, slightly longer in crown height than the pm,, moderately developed ante- rior accessory cuspid and posterior acces- sory cuspid; the dpm, is large, equal in crown height (formed by prominent proto- conid) to m,, the anterior accessory cuspid and posterior accessory cuspid present; the m,and m, are large, the m, slightly subequal in crown height to the m,, both with well- developed protoconid, anterior accessory cuspid, and posterior accessory cuspid, and slightly elongated anterobuccal cingulum; the m, is large and equal to m, in crown height, and with a well-developed proto- conid, anterior accessory cuspid, and prom- inent posterior accessory cuspid, and slight- ly elongated anterobuccal cingulum. Given that the individuals of M. jenkinsae are stage 1 sub-adults, no information can be provided on the adult dentition or antemolar replacement pattern of this species. Comparisons.—The fact that our two specimens of Microgale jenkinsae are stage 1 sub-adults complicates comparisons to a certain degree. However, sub-adult mem- bers of this genus, at this stage of dental eruption, exhibit the pelage coloration of adults and, in general are similar to adults in external measurements (MacPhee 1987; Jenkins et al. 1996, 1997). M. jenkinsae is readily distinguished ex- ternally from other relatively small mem- bers of this genus by pelage coloration and measurements. The contrasting agouti dor- sum and grizzled-gray venter is unique among small shrew-tenrecs. The pelage pigmentation in M. nasoloi is a relatively uniform gray; M. fotsifotsy has less gray in the dorsum than M. jenkinsae and is notably darker; M. parvula has a dark brown dor- sum and dark grayish-brown ventrum; M. brevicaudata is medium-brown dorsally and dull grayish-brown ventrally; and in M. longicaudata and M. pusilla the dorsum is a mixed light brown and medium brown and ventrum gray broadly edged with dark tan-brown. Further, there is no overlap in tail measurements between M. jenkinsae and any of these taxa, with the exception of 259 M. fotsifotsy, but M. jenkinsae can be dif- ferentiated from it based on pelage charac- teristics, a non-white-tipped tail, upper sur- faces of the feet clothed with short silver- white fur, and several external measure- ments (Table 1). The presence of a single-rooted second lower premolar separates M. jenkinsae from all other named small members of the genus Microgale, with the exception of M. pusilla. In the latter species the root form is iden- tical in individuals with deciduous and per- manent antemolar dentitions. M. pusilla is notably smaller than M. jenkinsae in all ex- ternal, cranial, and dental measurements (Tables 1 and 2), and is separable based on pelage characters. A generic revision of all members of Mi- crogale is currently in preparation and in- cludes molecular characters (Olson and Goodman, in prep.). The results of this study will be presented elsewhere and will address aspects of the phylogenetic position and sister-taxa relationships of M. jenkin- sae. Etymology.—This new species of Micro- gale is named after Paulina D. Jenkins of The Natural History Museum, London, for her important contributions to Tenrecidae systematics. Discussion Ecology.—Microgale jenkinsae is cur- rently known only from the Forét des Mi- kea, between Morombe and Manombo (Fig. 1), in the southwestern portion of Mada- gascar, a zone of transitional dry deciduous and spiny bush habitat (Seddon et al. 2000; Goodman and Soarimalala in press). This region receives, on average, about 400—500 mm of rainfall per year (Chaperon et al. 1993), with probably more rainfall in the inland higher ground than along the coastal plain. Differences in forest types within the Forét des Mikea tend to follow this pattern, with more deciduous forest on the slightly higher ground away from the coast and spiny bush along the coastal plain. 260 The climax vegetation of the dry decid- uous forest has been characterized as being dominated by the genera Dalbergia, Com- miphora, and Hildegardia (Humbert 1965). The formation has a canopy 10 to 15 m high, sometimes reaching 20 m, with a open medium stratum and diffuse undergrowth. All trees and most of the shrubs shed their leaves in the dry season. The vertebrate communities inhabiting the Forét des Mikea are similar to other arid portions of the island, although there are apparently several strict and regional en- demic vertebrates (e.g., the reptiles Furcifer belalandaensis and Paroedura vahiny, the birds Uratelornis chimaera and Monias benschi). The known small mammal com- munity consists of six Lipotyphla (Tenrec ecaudatus, Setifer setosus, Echinops telfai- ri, Geogale aurita, Microgale jenkinsae, and Suncus madagascariensis), one intro- duced murine rodent (Rattus rattus), and two endemic Nesomyinae (Macrotarsomys sp. and Eliurus myoxinus) (Carleton and Schmidt 1990; Soarimalala and Goodman 2004; Goodman and Soarimalala in press). Trapping.—During the survey of the Forét des Mikea, six sites were visited and systematically trapped using standard live and pit-fall traps (for more details on tech- niques see Goodman & Carleton 1996; Goodman et al. 1996). Pit-fall devices have been particularly useful for sampling Li- potyphla difficult to trap by other methods. For example, the only known specimens of another western Microgale, M. nasoloi, were captured with this technique (Jenkins and Goodman 1999). Three pit-fall lines, each composed of 11 12-liter buckets placed 10 m apart, were installed at each of the six survey sites. There was considerable variation between sites in trap success and species diversity. At site 1, 20 individuals (Tenrec, Setifer, Echinops, M. jenkinsae, and Geogale) were caught in 198 bucket nights between 14 and 19 February 2003. At site 2, 10 individuals (Echinops and Geogale) were obtained in 198 bucket nights between 21 and 27 February 2003. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON At site 3, 5 individuals (Tenrec, Echinops, Rattus rattus, and Geogale) were taken in 132 bucket nights between 2 and 5 March 2003. At site 4, only 4 individuals (Tenrec, Geogale, and Suncus madagascariensis) were trapped in 165 bucket nights between 8 and 12 March 2003. At site 5, nothing was captured in 198 bucket nights between 14 and 19 March 2003. At site 6, 7 indi- viduals (Geogale) were captured in 132 bucket nights between 22 and 25 March 2003. This level of faunal heterogeneity (as reflected by trapping-success) may be relat- ed to microhabitat differences between the sites, but further research is needed to test this hypothesis. Natural history.—Little definitive infor- mation can be gleaned on the natural his- tory of Microgale jenkinsae on the basis of two specimens, and the following extrapo- lations are tentative. Using foot structure and the context in which this species was trapped, it is terrestrial. Both specimens were taken in a portion of the Forét des Mikea dominated by more dry deciduous forest than by spiny bush habitat. Several of the other sites inventoried during the Forét des Mikea survey tended to be dom- inated by spiny bush habitat. The two M. Jenkinsae were obtained in different pitfall lines installed in portions of the forest hav- ing a relatively dense understory—one line was dominated by Gramineae associated with the regeneration of an old forest ex- ploitation track and the other line in a mi- crohabitat with a dense growth of Xerophy- ta (Velloziaceae) mixed with other low- growing plants and some succulent Eu- phorbiaceae. An analysis of soil samples taken at each of the 18 pitfall lines installed during the Forét des Mikea survey indicate that at site 1, where the two specimens of M. jenkinsae were collected, the average percent carbon in the soil was higher (1.9%, range 0.98— 2.3%) than four of the other five sites (all less than 0.9% carbon). The outlier is site 2 which had a slightly higher soil carbon content (2.4%, range 0.08—4.5%) than site VOLUME 117, NUMBER 3 1. Given that shrew tenrecs are believed to be primarily insectivorous, one may expect that their distribution would be correlated with soils relatively rich in organic materi- al, which in turn would support a higher density and diversity of invertebrates. Both specimens are dentally sub-adults, and, thus, it is not unexpected that they did not show any signs of reproductive activity. However, shrew tenrecs with deciduous an- temolar dentitions can be reproductive (e.g., Jenkins et al. 1996). Our survey was con- ducted during the rainy season, a period of the year that normally coincides with breed- ing activity in small mammals in this area of the island (Ganzhorn et al. 1996; Ran- drianjafy 2003). Paleoecological implications.—Remains of Microgale pusilla have been reported in disintegrated owl pellets of unknown age, but by extrapolation almost certainly Hol- ocene, from the sites of Lelia and Anjo- himpaty in southwestern Madagascar (MacPhee 1986, 1987), a zone of xero- phytic spiny bush habitat on an exposed limestone substrate. M. pusilla is consid- ered to be an inhabitant of the more mesic portions of the island, including the eastern humid forest and central highlands. More recently bone remains of M. pusilla have been identified in owl pellets collected in the capital city of Antananarivo, at least 80 km from the nearest intact natural forest block, and it is assumed that this species may live in surrounding marshlands and rice fields (Goodman et al. 1997a). At sev- eral sites in the central highlands it has been captured in marshlands within close vicinity to natural forest (Goodman et al. 2000a, Soarimalala et al. 2001). Thus, its occur- rence in Owl pellets in southwestern Mad- agascar can be interpreted in at least two ways: this species is a generalist and is able to live in a variety of ecological conditions from humid forests to marshlands to xero- phytic bush—however, on the basis of re- cent inventories of the drier portions of the island there is no evidence of its occurrence in this latter habitat—or the undated owl 261 pellets collected in the southwest are from a past geological period when this region of Madagascar was distinctly more mesic. The specimens of M. pusilla described by MacPhee (1986, 1987) from Lelia and An- johimpaty were deposited in the Service de Paléontologie collection at the Université d’Antananarivo. A detailed search of that collection, however, did not uncover these specimens. Nevertheless, a comparison of our material of M. jenkinsae to the illustra- tions and description of these specimens (MacPhee 1986) indicates considerable similarity in size, morphology, and dental structure. Most important in this regard is that dpm, (in M. jenkinsae and M. pusilla) and pm, (in M. pusilla) are simple in cor- onal structure and single-rooted, characters used to separate M. pusilla from all other small members of this genus before our rec- ognition of M. jenkinsae. Further, on the ba- sis of a scale provided with the line drawing of the Anjohimpaty mandible (MacPhee 1986, Fig. 5), the approximate lower tooth- row length is 18.2 mm, which is within the range of M. jenkinsae, but notably larger than M. pusilla (Table 2). We strongly sus- pect that these specimens, reported as M. pusilla, may be referable to M. jenkinsae. Recent biological surveys of the Parc National de Tsimanampetsotsa (Fig. 1), for- merly under the statute of a Réserve Na- turelle Intégrale, did not find any species of Microgale living in this protected area (Goodman et al. 2002), which is relatively close to Lelia and Anjohimpaty. Our small mammal surveys in the Forét des Mikea at six different sites, with a minimum of 132 pit-fall nights per site, yielded a total of 1023 pit-fall nights, yet only two individ- uals of M. jenkinsae were captured, both at the first site. This would indicate that this species is either rare or difficult to capture and presumably occupies specific micro- habitats. The important point here is that before significant paleoenvironmental infer- ences can be made associated with the pres- ence of certain taxa known only as subfos- sils, it is critical that detailed biological in- 262 ventories be conducted in the general re- gion of the paleontological site to thoroughly document the extant fauna. Conservation.—Historically, most field efforts associated with the exploration and documentation of Madagascar’s unique fau- na have been in the humid forests on the eastern, central, and northern portions of the island. Further, there is a preponderance of reserves and parks protecting this biome as compared to the drier western and south- ern regions of the island (ANGAP 2001). On the basis of several recent biogeograph- ic analyses of small mammals and birds, species turnover along the nearly 1200 km long eastern humid forests of the island is relatively low (Goodman et al. 1997b, 2000b). A number of endemic species in this biome have broad distributions, many extending the complete length of this hab- itat. More recent biological surveys of Madagascar’s western deciduous forests and southern spiny bush lands have re- vealed a previously unrecognized biota, in- cluding numerous terrestrial vertebrates. The growing realization is that levels of plant and animal species turnover along a latitudinal transect of western Madagascar is notably higher than the east, and this is probably related to a greater geological complexity and associated botanical com- munities in the west (Du Puy & Moat 2003). The recent surveys of the Forét des Mi- kea, which has no official protection, and forested regions to the north and south of this zone, are a case in point. Two unde- scribed species of mammals (Microgale jJenkinsae and Macrotarsomys nov. sp.) have been discovered in the Forét des Mi- kea that are unknown from any other region in the west. Further to the north, in the vi- cinity of the Bemaraha Plateau, there are at least two species of rodents that appear to be endemic to the region (Carleton et al. 2001). The recent discovery and description of Microgale nasoloi from unique forest formations in southwestern central Mada- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gascar seems to indicate another regional endemic with a very limited distribution. The drier western and southern forests of Madagascar have been subjected to consid- erable anthropogenic degradation, perhaps greater than in the humid east (see Smith 1997, Dufils 2003). In areas such as the Forét des Mikea, which was estimated in 1999 to contain forest cover in excess of 3700 km?, habitat loss rates have increased over the past few decades associated with pressures in the form of selective logging, cattle pasture, hunting, and clearing for ag- ricultural crops (Seddon et al. 2000). Given the levels of habitat heterogeneity and mi- croendemism in the west, action needs to be taken to protect the remaining large blocks of natural habitat in this region. On the basis of recent exploration of the Forét des Mikea this area should be given priority amongst the zones in need of rapid protec- tion. Acknowledgments We are grateful to the Direction des Eaux et Foréts for issuing permits to conduct fau- nal surveys in the Forét des Mikea. For ac- cess to specimens in their care we are in- debted to Géraldine Veron, Muséum Na- tional d’Histoire Naturelle, Paris; Paulina D. Jenkins, The Natural History Museum, London; and Prof. Daniel Rakotondravony, Université d’ Antananarivo, Département de Biologie Animale, Antananarivo. This field project was supported by WWF-Madagas- car, Fonds Frangaise pour |’ Environnement Mondiale (l’Agence Frangaise de Dével- oppement), and the Volkswagen Founda- tion. We are grateful to Link Olson for comments on an earlier draft of this paper and his important aid in numerous ways. Two anonymous reviewers also helped to improve this paper. 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Goodman, ed., A floral and faunal inventory of the eastern slopes of the Réserve Naturelle Intégrale d’Andringitra, Madagascar: with reference to elevational variation.—Fieldiana: Zoology, new series 85. , C. J. Raxworthy, & R. A. Nussbaum. 1997. A new species of Microgale (Insectivora, Ten- recidae), with comments on the status of four other taxa of shrew tenrecs.—Bulletin of the Natural History Museum, London (Zoology) 63:1-12. MacPhee, R.D.E. 1986. Environment, extinction, and Holocene vertebrate localities in southern Mad- agascar.—National Geographic Research 2: 441-455. . 1987. The shrew tenrecs of Madagascar: sys- tematic revision and Holocene distribution of Microgale (Tenrecidae, Insectivora).—Ameri- can Museum Novitates 2889:1—45. Olson, L. E., & S. M. Goodman. 2003. Phylogeny and biogeography of tenrecs. Pp. 1235-1242 in S. M. Goodman and J. P. Benstead, eds., The nat- ural history of Madagascar. The University of Chicago Press, Chicago. , & A. D. Yoder. in press. Illumination of cryptic species boundaries in long-tailed shrew tenrecs (Mammalia: Tenrecidae; Micro- gale), with new insights into geographic varia- tion and distributional constraints. Biological Journal of the Linnean Society. Randrianjafy, R. V. 2003. Contribution a l’étude de biologie de conservation de la communauté mi- cromammalienne d’Ankarafantsika. These de Doctorat de 3éme _ cycle, d’ Antananarivo, Antananarivo. Seddon, N., J. Tobias, J. W. Yount, J. R. Ramanam- pamonjy, S. Butchart, & H. Randrianizahana. 2000. Conservation issues and priorities in the Mikea Forest of south-west Madagascar.—Oryx 34:287-304. Smith, A. P. 1997. Deforestation, fragmentation, and reserve design in western Madagascar. Pp. 415— 441 in W. F Laurance and R.O.W. Bierregaard Jr., eds., management, and conservation of fragmented communities. The University of Chicago Press, Chicago. Université Tropical forest remnants: ecology, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Soarimalala, V. R. L., & S. M. Goodman. 2004. Les Rodentia, Lipotyphla et Carnivora de la forét des Mikea. Pp. 69-80 in A. P. Raselimanana and S. M. Goodman, eds., Inventaire floristique et faunistique de la forét de Mikea: Paysage écologique et diversité biologique d’une pré- occupation majeure pour la conservation.--Re- cherches pour le Développement, Série Scienc- es biologiques, No. 21. Soarimalala, V.,S. M. Goodman, H. Ramiarinjanahary, L. L. Fenohery, & W. Rakoronirina. 2001. Les micro-mammiféres non-volants du Pare Nation- al de Ranomafana et du couloir forestier qui la relie au Pare National d’ Andringitra. Pp. 197— 229 in S. M. Goodman and VY. R. Razafindrat- sita, eds., Inventaire biologique du Pare Nation- al de Ranomafana et du couloir forestier qui la relie au Pare National d’ Andringitra. Recherch- es pour le Développement, Série Sciences Biol- ogiques, Centre d’ Information et de Documen- tation Scientifique et Technique, Antananarivo, no. 17. Appendix | List of specimens of Microgale spp. examined during the course of this study. Microgale parvula.—Province d’ Antsiranana, Parc National de Marojejy [formerly Réserve Naturelle In- tégrale de Marojejy], along tributary of Manantenina River, 8 km NW Manantenina, 14°26.2'S, 49°46.5’E, 450 m (FMNH 159681); Parc National de Marojejy [formerly Réserve Naturelle Intégrale de Marojejy], along tributary of Manantenina River, 10 km NW Manantenina, 14°26.0'S, 49°45.7’E, 775 m (FMNH 159682, 159683, 159684); Parc National de Marojejy [formerly Réserve Naturelle Intégrale de Marojejy], 11 km NW Manantenina, Antranohofa, 14°26.2’S, 49°44.5'E, 1225 m (FMNH 159685); Pare National de Marojejy [formerly Réserve Naturelle Intégrale de Ma- rojejy],10.5 km NW Manantenina, along tributary at head of Andranomifototra River, 14°26.4’S, 49°44.5’E, 1625 m (FMNH 159686). Province de Fianarantsoa, approx. 45 km S. Ambalavao, east bank Iantara River, along Ambalamanenjana-Ambatoboay trail, edge of Pare National d’Andringitra [formerly Réserve Natu- relle Intégrale], 22°13'20"S, 47°01'29"E, 720 m (FMNH 151621); Pare National d’Andringitra [for- merly Réserve Naturelle Intégrale d’Andringitra], approx. 43 km S. Ambalavao, junction of Sahanivo- raky and Sahavatoy Rivers, 22°13’40"S, 47°00'13”E, 810 m (FMNH 151622); Pare National d’Andringitra [formerly Réserve Naturelle Intégrale d’ Andringitra], approx. 38 km S. Ambalavao, on ridge east of Volot- sangana River, 22°11'39”"S, 46°58'16"E, 1625 m (FMNH 151623, 151723, 151794, 151801). Microgale nasoloi.—Province de Toliara, Forét de Vohibasia, 59 km northeast Sakaraha, 780 m, VOLUME 117, NUMBER 3 22°27.5'S, 44°50.5'E (FMNH 156187); Forét d’ Analavelona, Antanimena, 12 .5 km NW Andranoh- eza, 22°40.7'S, 44°11.5'E, 1050 m (FMNH 161576). Microgale fotsifotsy.—Province de Fianarantsoa, Pare National d’Andringitra, 8.5 km SE Antanifotsy, Campement Andohan’ Ambola, PP ANOB2TBaS: 46°56.758'E, 1960 m (FMNH 165694, 165778, 165779) ; 2 km W. Andrambovato, along Tatamaly River, 21°30.7'S, 47°24.6'E, 1075 m (FMNH 170749); Forét de Vinantelo, at foot of Mt. Ambodivohitra, 15.5 km SE Vohitrafeno, 21°46.6'S, 47°20.8’E, 1100 m (FMNH 170750); approx. 40 km S. Ambalavao, along Volotsangana River, 22°13’22"S, 46°58'18”E, 1210 m (FMNH 151646, 151647); Province de Mahajanga, western side of Anjanaharibe-Sud, 13.5 km SW Befin- gotra, 14°47.0'S, 49°26.5’E, 1200 m (FMNH 167428). Province de Toliara, Pare National d’ Andohahela [for- merly Réserve Naturelle Intégrale d’ Andohahela], par- 265 cel I, 8 km NW Eminiminy, 24°37.55'S, 46°45.92’E, 440 m (FMNH 156569); Parc National d’ Andohahela [formerly Réserve Naturelle Intégrale d’ Andohahela], parcel I, 13.5 km NW Eminiminy, 24°35.04’S, 46°44.08'E, 1200 m (FMNH 156424); Pare National d’Andohahela [formerly Réserve Naturelle Intégrale d’Andohahela], parcel I, 15.0 km NW Eminiminy, 24°35.15’S, 46°43.85’E, 1500 m (FMNH 156570). Microgale pusilla.—Province d’ Antananarivo, 13 km NE Antananarivo, in Tyto alba pellets (FMNH 151606, 151607); 10 km SE Tsinjoarivo, Forét de Ma- hatsinjo, Andasivodihazo, 19°40.7’S, 47°46.2’E, 1550 m (FMNH 166123, 166124, 166125) ; Réserve Spé- ciale d’Ambohitantely, 24 km NE Ankazobe, 18°10.1'S, 47°16.6’E, 1450 m (FMNH 165489). Prov- ince de Fianarantsoa, Manambolo Forest, Ambavafa- tra, along Andohabatotany River, 17.5 km SE Sendri- soa, 22°8'58"S, 47°1'25”, 1300 m (FMNH 167612, 167619, 167621). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):266—270. 2004. Designation of the type species of Musaraneus Pomel, 1848 (Mammalia: Soricomorpha: Soricidae) Neal Woodman USGS Patuxent Wildlife Research Center, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013 Abstract.—The genus name Musaraneus often is attributed to Brisson (1762), however, most of Brisson’s names are unavailable. Pomel (1848) sub- sequently made the name Musaraneus available, but did not designate a type species. The 18 species that Pomel listed under Musaraneus currently are dis- tributed among five modern genera, two of which (Cryptotis Pomel, 1848 and Diplomesodon Brandt, 1852) are predated by Musaraneus. Because Cryptotis and Diplomesodon potentially could be considered junior synonyms of Musar- aneus, I propose Sorex leucodon Hermann, 1780 (= Crocidura leucodon) as the type species for Musaraneus, thereby establishing Musaraneus as a junior synonym of Crocidura Wagler, 1832. The generic name Musaraneus Pomel, 1848 derives from mus araneus (“‘spider mouse’’), one of the terms commonly used alongside sorex and mus caecus by classical Latin writers (e.g., Plinius n.d.; Columella n.d.; Serenus n.d.) to refer to small mam- mals now generally interpreted as shrews (family Soricidae). The classical name mus araneus has a long history of use in early zoological literature. It was adopted and used widely by Renaissance natural histo- rians and made the transition from a Latin common name to being incorporated into more formal taxonomies. The vernacular mus araneus generally was applied to the small mammal called locally by a variety of names that included ‘“‘muzeraigne,”’ “spitzmus, shrew, erd shrew,’ or “shrew-mouse’”’ (Gesner 1551, 1560, 1602: Marggraf 1648; Jonston 1657; Topsell 1658; Ray 1693). A number of early tax- onomists attempted to establish the name as Mus Araneus or Musaraneus within heir- archical classifications (Charleton 1668; Klein 1751; Brisson 1756, 1762). It is of interest that Gesner (1551, 1560, 1602) and subsequent writers (e.g., Topsell 1658, Charleton 1668) interpreted sorex as dis- 99 ce 29 ce tinct from mus araneus, in some cases as a broader category that might include mus ar- aneus (e.g., Klein 1751), or as a separate set of animals, typified by mus avellana- rum, the “haselmus” or “‘hasel-mouse” (Gesner 1560, Topsell 1658), or by the “rat”? (Charleton 1668). Gesner’s (1551, 1560, 1602) print of mus araneus is an il- lustration of a soricid (Fig. 1A), possibly a white-toothed shrew of the genus Croci- dura, whereas his picture of a sorex is iden- tifiable as a garden dormouse (Eliomys quercinus—Fig. 1B). His illustrations were copied and republished by subsequent writ- ers (e.g., Topsell 1658) and likely influ- enced later interpretations of the names. In contrast, Linné (1746, 1748, 1758) explic- itly and consistently applied Sorex to those mammals that previous authors had called mus araneus or Musaraneus, and Sorex Linné, 1758 is the name that survived in the taxonomic literature. Musaraneus con- tinues to be reflected in modern words for shrew in a number of romance languages, e.g., musarana (Spanish), musaraigne (French), musaranho (Portuguese), musar- agno (Italian). It also survives, in part, in VOLUME 117, NUMBER 3 Fig. 1. Gesner’s (1602) illustrations of (A) mus araneus and (B) mus avellanarum from Historiae Animalium. Photographs courtesy of the Smithsonian Institution Libraries, Joseph KR Cullman 3" Library of Natural History, Washington D.C. Reproduced with permission. the scientific name for the European com- mon shrew, Sorex araneus Linné, 1758. As a genus-level name, Musaraneus is often attributed to Brisson (1762; see Pomel 1848, Sherborn 1902, Palmer 1904, Mc- Kenna and Bell 1997, Kretzoi and Kretzoi 2000). Because Brisson (1762) did not con- sistently apply binomial nomenclature in his work, however, most of his names are unavailable in accordance with Article 11.4 of the International Code of Zoological No- menclature (ICZN 1999; but see Hopwood 1947; ICZN 1998). In a subsequent classi- fication of insectivores, Pomel (1848) re- described Brisson’s Musaraneus as one of four genera (with Talposorex Pomel, Sorex Linneus, and Galemys Pomel) within the tribe Soriciens in his family Spalacogalae. Pomel (1848) made the name Musaraneus available, and therefore, he is the author of this name, as noted by Sherborn (1928). Hopwood (1947) recorded a number of oth- er generic names used by Brisson (1762) that similarly were made available by later authors (see also ICZN 1998). Pomel’s (1848) Musaraneus comprised 18 species distributed among three “‘sec- tions”’ (subgenera): Cryptotis, a new taxon with a single North American species; Myo- sorex Gray, 1838, an existing taxon com- prising three African species; and Croci- dura Wagler, 1832, an existing taxon con- taining 14 Old World species. Based on the list of included species, the genus Musar- aneus included representatives from five modern genera. In addition to those genera representing Pomel’s (1848) three sections (Cryptotis, Myosorex, Crocidura), one spe- cies (Musaraneus puchellus) represents Di- plomesodon Brandt, 1852, and three others (M. crassicaudatus, M. vulgaris, M. Bach- mani) represent Sorex Linné, 1758. Pomel (1848) was uneven in designating species that he considered to be typical of the genera he described. In his classification of insectivores, Pomel noted a “typical spe- cies” for his newly-described Talposorex, but not for his names Galemys or Musara- neus. For these latter genera, he provided lists of species divided among several sec- tions (subgenera). Pomel (1848) wrote the latter name as, ““Genre Musaraneus Briss, 268 Pom.,” clearly indicating the influence of Brisson (1762). It is curious, however, that his version of Musaraneus did not explic- itly include any of the three “‘species”’ (Mu- saraneus, Musaraneus aquaticus, M. bras- iliensis) that Brisson (1762) had allocated to his genus Musaraneus. Pomel (1848) may have considered Brisson’s (1762) uni- nomial “‘species’” Musaraneus to be repre- sented by what Pomel called Musaraneus (Crocidura) vulgaris, which he equated with araneus of authors. The type species of Musaraneus Pomel, 1848 is important to modern taxonomists because Musaraneus Pomel, 1848 predates Cryptotis Pomel, 1848 and Diplomesodon Brandt, 1852, and Musaraneus could be in- terpreted as a senior synonym of one of these genera. Kretzoi and Kretzoi (2000: 241) indicated that the type species for Mu- saraneus Pomel is ““Crocidura (M.) priscus Pomel”’ (sic). There are a number of im- portant and confusing errors in their ac- count for this name, however. Both the original description of the genus and the designation of Crocidura priscus as the type species are credited by them to “Po- mel 1853,” which is referenced as “Arch. Sci. Phys. Nat., Bibl. Univ. Genéve, 9: 249,’ but this reference is a conflation of several different publications. Pomel (1853a, 1853b) are parts of his “Catalogue des Vertébrés Fossiles,” which was pub- lished in at least three sections in Annales Scientifiques, Littéraires et Industrielles de L’Auvergne: the first in October and No- vember of 1852 (Pomel 1852), the second in March and April of 1853 (Pomel 1853a), and the last in May and June of 1853 (Po- mel 1853b). Insectivores, including the de- scription of the fossil species Musaraneus (Crocidura) priscus, appear in the first part of this work, and the correct citation for that name is Pomel (1852:351). Pomel’s original description of Musaraneus is in an earlier publication (Pomel 1848) that was pub- lished in Volume 9 of Archives des Sciences Physiques et Naturelles, Genéve. In order for a species to be designated a type species PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON by a subsequent author, it must have been included in the genus by the original author (ICZN 1999: art. 69.2). However, Pomel (1848) did not include the name priscus among the species he listed under Musar- aneus, as he had not yet described the spe- cies. Therefore, the selection by Kretzoi and Kretzoi (2000) is invalid. Pomel’s (1848:249) original description of Musaraneus reads: Trois intermédiaires en haut, deux en bas, estomac oblong avec poche bien marquée sous le boyau py- lorique. My translation of this description is: Three upper intermediary teeth, two lowers, stom- ach oblong with well-marked pouch below the py- loric constriction. I interpret Pomel’s upper ““intermédiaires” to represent the simplified upper dentition between the large, hooked first incisor and the roughly molariform fourth premolar (P4) that commonly are referred to as “‘un- icuspids”’ (Choate 1970). The lower “inter- médiaires”’ are the teeth designated the low- er unicuspid and lower fourth premolar (p4). Among the five modern genera Pomel (1848) included within Musaraneus, all but Myosorex have a single lower unicuspid and p4; Myosorex typically has two lower unicuspids in addition to p4. In the upper dentition, Sorex has five unicuspids, Cryp- totis and Myosorex each have four, and Di- plomesodon has two. Only Crocidura has three upper unicuspids in accord with Po- mel’s (1848) description. Although it is not required that the type species match the original description for a genus, it is highly desirable. Among the modern species of Crocidura that Pomel (1848) included in Musaraneus, the majority are African, one is from Japan, and one, Crocidura leucodon, is widespread in continental Europe, including France, where Pomel lived. Among the recommen- dations for selecting a type species for sub- sequent designation are that the species be common and that it be well known to the VOLUME 117, NUMBER 3 original author (ICZN 1999: Recommen- dations 69.A1, 69.A7). Therefore, I select Sorex leucodon Hermann, 1780, as used in the name combination Musaraneus (Croci- dura) leucodon by Pomel (= Crocidura leucodon), as the type species of Musara- neus Pomel, 1848. By designating this tax- on as the type species, Musaraneus Pomel, 1848 becomes a junior synonym of Croci- dura Wagler, 1832, thereby stabilizing the generic names Cryptotis and Diplomesodon in accordance with their long-established usage. Acknowledgments My thanks to Alfred L. Gardner for orig- inally pointing out the potential problems engendered in the choice of a type species for Musaraneus. The Department of Special Collections, University of Kansas Libraries; Dale Miller, Leslie Overstreet and Daria A. Wingreen in the Joseph EF Cullman 3” Li- brary of Natural History, National Museum of Natural History; and Kirsten van der Veen in the Dibner Library of the History of Science and Technology, National Mu- seum of American History graciously pro- vided access to, and/or photocopies of, im- portant pre-Linnean manuscripts. Sandy Feinstein, Alfred L. Gardner, Robert M. Timm, and an anonymous reviewer provid- ed valuable comments on previous versions of my manuscript. Literature Cited Brandt, J. F 1852. Zoologischer Anhang. Die von Leh- mann gesammelten oder auf seinen Reisen beo- bachteten Wirbelthiere des Orenburger Gouver- nements, ferner der Uralischen, Kaspischen, Kirgisischen und Uralischen Steppen, ebenso wie Buchara’s und Samarkand’s. Beitrége zur Kenntniss des Russischen Reiches und der an- granzenden Lander Asiens 17:279-342. [not seen] Brisson, M. J. 1756. Regnum animale in classes ix. Jean—Baptiste Bauche, Paris. . 1762. Regnum animale in classes ix. Theo- durom Haak, Leiden. Charleton, W. 1668. Onomasticon zoicon. Jacob Al- lestry, London. 269 Choate, J. R. 1970. Systematics and zoogeography of Middle American shrews of the genus Crypto- tis.—University of Kansas Publications, Muse- um of Natural History 19:195—317. Columella, L. J. M. n.d. De Re Rustica. (H. B. Ash, E. S. Forster, and E. H. Heffner, translators, 1960, 1968). Loeb Classical Library, Harvard University Press, Cambridge, Massachusetts. Gesner, C. 1551. Historiae animalium Lib. I. de quad- rupedibus uiuiparis. C. Froschoverus, Zurich. . 1560. Icones animalium quadrupedum vivi- parorum et oviparorum, quea in historiae ani- malium Conradi Gesneri. C. Froschoverus, Zu- rich. . 1602. Historiae animalium. Liber primus de quadrupedibus viviparous, 2nd edition. Biblio- polio Cambieriano, Frankfurt. Gray, J. E. 1838. Revision of the genus Sorex, Linn.— Proceedings of the Zoological Society of Lon- don 1837:123-126. Hermann, J. 1780. In W. E. A. Zimmermann. 1780. 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Etudes sur les carnassiers insecti- vores. (Extrait) Seconde partie, Classification des insectivores.—Archives des Sciences Phy- siques et Naturelles, Genéve, 9:244—251. . 1852. Catalogue méthodique et descriptif des vertébrés fossiles découverts dans le bassin hy- drographique supérieur de la Loire et surtout dans la vallée de son affluent principal, Vallier—Annales Scientifiques, Littéraires et Industrielles de L Auvergne, 25:337—380. 1853a. Catalogue des vertébrés fossiles (suite.)—Annales Scientifiques, Littéraires et Industrielles de L Auvergne, 26:81—176. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1853b. Catalogue des vertébrés fossiles (suite et fin.). Remarques générales sur les caractéres des diverses faunes du Valey & de La Limange, comparées entr’elles et avec celles de différen- tes régions—Annales Scientifiques, Littéraires et Industrielles de L Auvergne, 26:177—229. Ray, J. 1693. Synopsis methodica animalium quadru- pedum et serpenti generis. S. Smith & B. Wal- ford, London. Serenus Sammonicus, Quintus. 7d. Liber medicinalis. Http://www.forumromanum.org/literature/serenusx. html. [Accessed 18 November 2003] Sherborn, C. D. 1902. Index mammalium. Section 1. C. J. Clay and Sons, London. . 1928. Index mammalium. Section 2, part 17, pp. 4195-4450. Trustees of the British Muse- um, London. Topsell, E. 1658. The history of four-footed beasts, vol. 1. E. Cotes, London. [Reprinted 1967 by Da Capo Press, New York]. Wagler, J. 1832. Mittheilungen iiber einige merkwiir- dige Thiere. Isis von Oken 25:275—282. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):271-302. 2004. The mammals of Palawan Island, Philippines Jacob A. Esselstyn, Peter Widmann, and Lawrence R. Heaney (JAE) Palawan Council for Sustainable Development, P.O. Box 45, Puerto Princesa City, Palawan, Philippines (present address: Natural History Museum, 1345 Jayhawk Blvd., Lawrence, KS 66045, U.S.A.) (PW) Katala Foundation, PRO. Box 390, Puerto Princesa City, Palawan, Philippines; (LRH) Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605 U.S.A. Abstract.—The mammal fauna of Palawan Island, Philippines is here doc- umented to include 58 native species plus four non-native species, with native species in the families Soricidae (2 species), Tupaiidae (1), Pteropodidae (6), Emballonuridae (2), Megadermatidae (1), Rhinolophidae (8), Vespertilionidae (15), Molossidae (2), Cercopithecidae (1), Manidae (1), Sciuridae (4), Muridae (6), Hystricidae (1), Felidae (1), Mustelidae (2), Herpestidae (1), Viverridae (3), and Suidae (1). Eight of these species, all microchiropteran bats, are here reported from Palawan Island for the first time (Rhinolophus arcuatus, R. ma- crotis, Miniopterus australis, M. schreibersi, and M. tristis), and three (Rhin- olophus cf. borneensis, R. creaghi, and Murina cf. tubinaris) are also the first reports from the Philippine Islands. One species previously reported from Pa- lawan (Hipposideros bicolor) is removed from the list of species based on re- identificaiton as H. ater, and one subspecies (Rhinolophus anderseni aequalis Allen 1922) is placed as a junior synonym of R. acuminatus. Thirteen species (22% of the total, and 54% of the 24 native non-flying species) are endemic to the Palawan faunal region; 12 of these are non-flying species most closely related to species on the Sunda Shelf of Southeast Asia, and only one, the only bat among them (Acerodon leucotis), is most closely related to a species en- demic to the oceanic portion of the Philippines. Of the 28 insectivorous bats, 18 species are somewhat to highly widespread in Indo-Australia, 2 are shared only with the Sunda Shelf and Indochina, 1 with the Sunda Shelf alone, 3 occur on the Sunda Shelf and the oceanic Philippines, 1 occurs in Palawan, Sulawesi, and the oceanic Philippines, 2 occur only on Palawan and in the oceanic Philippines, and | occurs on Borneo, Sulawesi, and throughout the Philippines. Though the insectivorous bats tend to be widely distributed, these data, particularly the distributions of the non-volant species, strongly reinforce the perception of Palawan Island (and associated smaller islands) as a biogeo- graphic unit of the Sunda Shelf, with only limited similarity to other portions of the Philippine Islands. The Philippine archipelago is remarkable for the large number of indigenous land mammal species (ca. 175), and especially for the number of endemic species (ca. 112). Given its relatively small land area, the Philippines has perhaps the greatest concentration of endemic mammals in the world (Heaney et al. 1998, Heaney & Re- galado 1998, Mittermeier et al. 1997). These species, especially the endemics, are not distributed homogeneously over the country; rather, there is a large number of discrete biogeographic units, and these cor- respond to the limits of the islands that ex- 272 isted during periods of low sea level during the late Pleistocene (Heaney 1986, 1991a, 1991b, 2000). With a single exception, cur- rent geological evidence indicates that none of these ““Pleistocene islands” has had dry- land connections to the Asian mainland or to other areas. Rather, each arose as a de novo oceanic island, some from a combi- nation of oceanic crust and volcanic mate- rials, and some as uplifted areas of conti- nental rock that had been submerged for long periods, and all of these have remained isolated by sea channels (Hall 1998, 2002; Heaney 1985, 1986, 1991a). The sole ex- ception is the Palawan faunal region, which generally has been considered to be a por- tion of the Sunda Shelf, both geologically and biogeographically, with many species shared with Borneo (Dickerson 1928, Ev- erett 1889, Heaney 1986). Although Pala- wan was initially also a de novo oceanic island, its biogeographic affinity to the Sun- da Shelf has been thought to be due to the presence of a shallow shelf between Borneo and Palawan with an intervening depth of ca. 145 m (Heaney 1986, 1991a). Previous- ly, evidence indicated that sea levels dropped to about 165 m below present lev- els during the penultimate glacial episode (Gascoyne et al. 1979), which would have resulted in a dry-land connection of Pala- wan to Borneo at about 165,000 BP (Hea- ney 1985, 1986, 1991a). However, recent evidence suggests that sea level dropped only to about 135 m (Rohling et al. 1998) or perhaps as little as 115 m below present levels (Siddall et al. 2003, Voris 2000) dur- ing that glacial episode, leaving open the question of when Palawan was connected to Borneo, or if the gap simply became very narrow. The mammals of Palawan Island, the largest part of the faunal region at 11,785 km/?, and the associated smaller islands have been documented over the course of more than a century (Allen 1910, Allen 1922, Ev- erett 1889, Heaney et al. 1998, Hoogstraal 1951, Kuntz 1969, Reis & Garong 2001, Sanborn 1952, Taylor 1934, Timm & Bir- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ney 1980), but the fauna is still poorly known in many respects. Little information has been available for most species on ecol- ogy and distribution, including habitat re- quirements, and only a few studies have considered phylogenetic relationships (e.g., shrews: Heaney & Ruedi 1994; bats: Mus- ser et al. 1982; squirrels: Heaney 1979; mu- rid rodents: Musser 1979, Musser & New- comb 1983, Musser & Heaney 1992; pan- golin: Feiler 1998; leopard cats: Groves 1997). In particular, microchiropteran bats have been only superficially documented. The limitations of available data have thus limited understanding of the Southeast Asian fauna, and the Philippine fauna in particular, from both biogeographic and ecological perspectives, and hence limited conservation planning in a nation that is of- ten cited as one of the most in need of ef- fective conservation action (Mittermeier et al. 1999, Ong et al. 2002, Wildlife Conser- vation Society of the Philippines 1997). Two of us (Esselstyn and Widmann) re- cently conducted extensive surveys of the mammals of Palawan Island, focusing on the 15 sites described below. Esselstyn worked from December 1999 to November 2000 at Sites 1-11, emphasizing (though not exclusively) insectivorous bats, which are the mammals most poorly known in the Philippines (Heaney et al. 1998, Heaney & Mallari 2002), and Widmann conducted studies of bats, rodents, and larger mam- mals from 1997 to 2002 at Sites 12—15; Heaney visited briefly in April 2000. In this paper, we report information collected dur- ing these studies, emphasizing new data on bats, and we include additional unpublished records of mammals. We summarize infor- mation on all additional species that were not taken during this study but have been documented on the island, and re-examined some key specimens from prior studies. We include descriptions of the habitats where we conducted our surveys because of a gen- eral paucity of such information, and use all available information to evaluate conser- vation status of the mammals. We include VOLUME 117, NUMBER 3 measurements of the skulls of selected spe- cies of insectivorous bats that have been es- pecially poorly known. Methods At Sites 1-11, small non-volant mam- mals were captured using locally made live (cage) traps and Victor (snap) rat traps. Ap- proximately 90% of live traps used mea- sured 11 X 11 X 24 cm, and 10% measured 13 X 16 X 13 cm. Trap lines consisting of approximately 70 traps (50 live traps and 20 snap traps) were placed in areas of tra- versable terrain. Individual traps were placed in locations of likely capture (e.g., near holes, along fallen logs, near root but- tresses, etc.) along the line spaced at 5—15 m intervals. Most traps were set at ground level, but in forest habitats we placed 5— 15% of the traps in elevated locations up to 2 m above ground level on fallen logs, hor- izontal vines, etc. All but two trap lines were set for three nights; the exceptions at Sites 3 and 7 were set for five and two nights, respectively. At Sites 12—14, a mix- ture of live traps were used (see Site de- scriptions). Most trap lines were baited with fresh grilled coconut coated with peanut butter. Other trap lines were baited with live earthworms or bananas. All traps were checked in the early morning and late af- ternoon. Baits were changed at least once daily, usually in the afternoon, and as nec- essary in the early morning. Most live an- imals were released at the site of capture. We captured bats using a harp trap (ca. 2 x 2 m, 4 bank), mist nets (2 X 6 m, 16 mm mesh), butterfly nets, and hand capture at Sites 1-11; only nets were used at Sites 12—15. Harp trap and net locations were se- lected to be locations of likely capture (e.g., natural canopy breaks, over streams and trails, around fruiting trees, and near poten- tial roosting locations). The harp trap and mist nets were usually set in a location for three nights, and occasionally for only one or two nights. During surveys of caves, we primarily used the harp trap, and its loca- 273 tion was frequently changed. Mist nets were continuously monitored during peak activ- ity periods from 1800 to 2000 h (at Sites 12 and 13, until 2400 h) and then checked again in the early morning. The harp trap was checked periodically between 1800 and 2100 h and again in the early morning. In forested areas, we searched for bats in po- tential roosting locations (e.g., hollow trees, rock formations, new banana leaves, etc.). Most bats were released at the site of cap- ture. For many of the bats, we report the proportion of adult females that were preg- nant on certain dates. These were palpated externally to determine the presence or ab- sence of an embryo before being released. Forearm and cranial measurements were taken by L. R. Heaney and D. S. Balete at the Field Museum of Natural History (FMNH). All voucher specimens from Es- selstyn, which are cited below as specimens examined, were preserved in fluid (with skulls later removed and cleaned) and cat- aloged at the FMNH; half of the vouchers have been deposited at the National Muse- um of the Philippines (NMP). Additionally, data on several previously unreported spec- imens housed in the University of Michigan Museum of Zoology (UMMZ) and United States National Museum of Natural History (USNM) are included here. Site Descriptions See Fig. | for approximate locations of study sites. The province of Palawan in- cludes the main island called Palawan and many smaller, nearby islands. The island is politically divided into 13 municipalities, one of which is Puerto Princesa City; the municipalities and the city are subdivided into barangays, and barangays into sitios. Site I (10°03’00"N, 119°00'44"E) was in lowland primary forest located along the Tarabanan River in northern Puerto Prin- cesa Municipality, at elevations ranging be- tween ca. 100 and 200 m. Slope in the area was generally moderate, rising from the riv- er to the ridge-tops. Forest in the area was 274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Jose Rizal @ Sofronio Espanola Brookes Point Bataraza Fig. 1. Map of Palawan Island, showing the locations of the primary towns (solid circles) and research sites (solid triangles), and the position of Palawan in the Philippines (inset). nearly undisturbed, which is uncommon at lection of minor forest products and hunting this low elevation. We were aware of only of Sus barbatus and Macaca fascicularis. one small human-made clearing (ca. 0.5 ha) Canopy ranged from 20 to 30 m in height in the area; other disturbances included col- and was multi-layered. Canopy trees ranged VOLUME 117, NUMBER 3 in diameter from 40—80 cm and had light buttress development. Leaf litter was thin. We surveyed small volant and non-volant mammals for 13 days in January 2000. Four trap lines were run for three nights, yielding 3 Tupaia palawanensis, 21 Max- omys panglima, and 1 juvenile Viverra tar- galunga in 792 trap-nights. Three of the lines were baited with coconut and peanut butter, while one line was baited with live earthworms. Forty-eight net-nights pro- duced 1 Cynopterus brachyotis and 12 harp-nights produced 3 Rhinolophus acu- minatus, 3 R. arcuatus, and 1 R. creaghi, plus other means produced 11 Megaderma spasma and 1 R. acuminatus. Site 2 (9°42'14"N, 118°32’01”"E) was in lowland primary forest located mid-way up Mt. Salakot, between 300 and 700 m ele- vation. Slope was rolling to moderately steep. Several small streams dissected the area. The only major human-caused distur- bance in the area was an unused helicopter landing pad and an abandoned road; the road had a dense regrowth of ferns and small trees. Sus barbatus was hunted, and some almaciga trees (Agathis sp.) near the upper reaches of the site had fallen due to over-collection of resin. Agoho trees (Ca- suarina sp.) gradually became more com- mon as elevation increased. Canopy ranged from 15 to 30 m in height and was multi- layered. Canopy trees ranged in diameter from 35—60 cm with the largest emergents reaching 80—90 cm. Buttress systems were only slightly developed and stilt root sys- tems were present above 600 m, but rare. Leaf litter was slightly deeper than at lower elevations. We surveyed small volant and non-volant mammals at this site for 20 days during March and July 2000. Six trap lines (three coconut and peanut butter-baited lines, one banana-baited line, and two earthworm-baited lines) yielded 16 Tupaia palawanensis, 55 Maxomys panglima, and 1 Sundamys muelleri from 1272 trap nights. Fifty-six net-nights yielded 1 Cynopterus brachyotis, 1 Hipposideros diadema, 8 Rhinolophus arcuatus, and 1 R. creaghi. AHS Ten harp-nights yielded 1 Hipposideros diadema, 16 Rhinolophus arcuatus, 10 R. creaghi, 2 R. virgo, and 2 Kerivoula hard- wickil. Site 3 (10°07'28"N, 118°59'36"E) was in primary montane and mossy forest located near the peak of Cleopatra’s Needle (max- imum elevation 1603 m), at elevations ranging from 1300 to 1600 m. Slope was moderate to extreme. Other than a trail to the peak occasionally traveled by tourists, there was little human-caused disturbance in the area. No above-ground sources of water were found near the site, but mist was frequently present: during our stay of two weeks during dry season, the area was al- most continuously shrouded by a dense fog. Vegetation type was montane forest up to ca. 1500 m. At this elevation, the vegetation began a transition to mossy forest. In mon- tane forest, the canopy reached a height of approximately 10 m. Trees, rocks, fallen logs, and other stable surfaces were covered with a thin layer of moss; epiphytic ferns and orchids were abundant. Many trees had an adventitious root system, but maintained straight boles. The canopy was more open here than at lower elevations. Above 1500 m, trees were shorter (2—4 m in height) and took on a shrub form above 1550 m. Moss growth was heavier at this elevation, and vegetation became extremely dense at the upper reaches. Pitcher plants (Nepenthes) began to appear at about 1500 m and were abundant by 1550 m. We surveyed small volant and non-volant mammals at this site for 12 days during February and March 2000. Three trap lines yielded 1 Tupaia pa- lawanensis, 33 Maxomys panglima, and 3 Rattus tiomanicus in 740 trap-nights. Two lines were run for three nights each and one line was run for five nights; all three lines were baited with coconut and peanut butter. Forty-eight net-nights produced 2 Rhinolo- phus arcuatus, and 12 harp-nights produced no captures; 2 Pipistrellus javanicus were captured by hand. Site 4 (9°33'45"N, 118°27'54’"B) is a cave known locally as ““Ma-ngit’’. It is located 276 along the Iraan River near Sitio Pamolkoan, Barake, Aborlan, at ca. 430 m elevation. The cave is in a small valley, restricted by mountains to the east and west. A small, seasonal stream flows through the cave. At least six entrances to the cave were evident, and many small tunnels connected medium to small caverns, which ranged from well- lit to completely dark. Very little distur- bance was evident in or around the cave due to its isolated location (ca. 10 hour hike from the nearest road). The cave was sur- rounded by a large expanse of primary low- land forest, but some agricultural areas were present within ca. 5 km. Disturbances to local vegetation included collection of rattan (Calamus spp.). We surveyed bats in Ma-ngit Cave for six days during December 1999. We captured 819 bats belonging to seven species (9 Eonycteris spelaea, 43 Hipposideros diadema, 367 Rhinolophus creaghi, 4 R. virgo, 9 Miniopterus australis, 386 M. schreibersi, and 1 M. tristis) inside the cave. Site 5 (10°05'00"N, 118°51’06"E) is a complex area of limestone karst containing probably greater than 100 caves in Baran- gays Tagabinet and Cabayugan, Puerto Princesa; elevation is ca. 50 m. Caves in the area ranged from tiny cracks too small to enter, to large complexes of multiple cav- erns with multiple entrances. Local terrain was generally flat except for the sometimes- massive limestone outcrops, which form high-rising cliffs throughout the area. Caves probably are present all over these complex formations, but only the very few found near ground level were accessible. We cap- tured bats in and around five different caves/cave complexes. These represented some of the most accessible caves in the area. Disturbance at the caves was moder- ate, with vandalism and guano excavation evident at most caves. Most of the caves were surrounded immediately by agricul- tural development. Both primary and sec- ondary lowland forests were present in the surrounding hills. We surveyed bats at these caves for 14 days between March and May PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2000. A total of 575 bats belonging to 10 species (18 Cynopterus brachyotis, 1 Me- gaderma spasma, 100 Hipposideros ater, 239 H. diadema, 14 Rhinolophus arcuatus, 33 R. creaghi, 10 R. macrotis, 86 R. virgo, 15 Miniopterus australis, and 59 M. schrei- bersi) were captured. Site 6 (9°28'25"N, 118°30'21"E) was a mostly abandoned agricultural area located in Barake, Aborlan Municipality, at eleva- tion ranging from 40—80 m. A small stream flowed through the area and topography was flat to rolling. Vegetation was a mosaic of grassland (primarily Imperata cylindri- ca) with sparse trees (mostly Vitex sp.), cashew plantations, dense brush, and very small (<1 ha) areas of secondary growth. Frequent fires appeared to maintain this area as a grassland. We surveyed small vo- lant and non-volant mammals for four days during June 2000. A single trap-line baited with live earthworms yielded 17 Rattus ex- ulans in 186 trap-nights. We captured 7 Cy- nopterus brachyotis in 6 net nights, and 1 Kerivoula whiteheadi in 3 harp-nights. Site 7 (9°29'15"N, 118°29'24”E) was in a narrow band of secondary forest in Barake, Aborlan Municipality, located between dis- turbed habitat at lower elevation (Site 6) and primary and good secondary forest at higher elevation. Elevation ranged from 80—140 m; slope was rolling to moderately steep, and two small streams dissected the area. Canopy height varied from 5—20 m. Woody vines and lianas were common and vegetation was quite dense in areas. Wild bananas (Musa spp.) were abundant, but patchy in distribution; leaf litter depth was highly variable. We surveyed this site for small volant and non-volant mammals for four days during June 2000. A single trap line baited with live earthworms yielding 120 trap-nights produced 1 Rattus exulans. Six net-nights produced 27 Cynopterus brachyotis and 1 Macroglossus minimus, and 3 harp-nights yielded 1 Hipposideros diadema and | Kerivoula pellucida. Site 8 (9°59'47"N, 118°56'43"E) is a large cave complex located in a limestone VOLUME 117, NUMBER 3 karst formation on top of the first ridge up from San Rafael, Puerto Princess, at an el- evation of ca. 250 m. The cave complex is known locally as ““Taraw’’. The cave sys- tem appeared to be quite large; we were unable to explore much of it due to a lack of climbing equipment and expertise. Some caverns exceeded 20 m in height, while oth- ers were quite small. We found no perma- nent water in or around the cave, but evi- dence of storm flow was present. Evidence of vandalism and guano collection was pre- sent. A mixture of habitats surrounded the cave complex: between the cave and the community of San Rafael, the vegetation was dominated by brushland and agricul- tural developments, while on the other side of the cave secondary and primary forest dominated, with mixed areas of slash-and- burn fields. We surveyed bats at this cave for five days during July 2000. We captured 2775 bats representing 10 species (7 Hip- posideros ater, 43 H. diadema, 90 Rhino- lophus acuminatus, 240 R. arcuatus, 239 R. creaghi, 25 R. macrotis, 151 R. virgo, 1257 Miniopterus australis, 711 M. schreibersi, 4 immature M. sp., and 8 Myotis macrotar- SUS). Site 9 (9°39'40"N, 118°27'48"F) is a small cave located in Sitio Labtay, Napsan, Puerto Princesa City. The cave, which con- sisted of a single chamber 1—2 m wide, 3— 6 m high, and ca. 30 m long, was in a nar- row canyon along the Panagurian River at ca. 280 m. Vegetation in the area consisted of good-quality secondary forest, second- growth forest, and agricultural develop- ments. We trapped bats with a harp trap, mist nets, and a butterfly net in the cave and surrounding forest for five days during August 2000. We captured 73 bats (4 Cy- nopterus brachyotis, 68 Hipposideros dia- dema, and 1 Rhinolophus arcuatus). Site 10 (10°44'00”, 119°34’23”) is a cave located near sea level in Sitio Sader, Ban- tulan, Taytay Municipality. The cave con- sisted of a single chamber (ca. 3—8 m wide, 3-7 m high, and 40 m long) with a large (>2.5 m diameter) entrance at each end. 277 Minor damage had been done by both trea- sure hunters and guano collectors. The sur- rounding vegetation was dominated by ag- ricultural areas with some strips and patches of residual forest. Large expanses of sec- ondary and logged-over forest were found nearby in the vicinity of Lake Manguao. We captured 115 bats belonging to 4 species (48 Eonycteris spelaea, 1 Rousettus am- plexicaudatus, 64 Hipposideros diadema, and 2 Miniopterus tristis) using a harp trap and butterfly net at one of the entrances to the cave during three days in October 2000. Site 11 (10°46'34"N, 119°31'52”E) was located around the perimeter of Lake Man- guao, in Barangays Poblacion and Bantu- lan, Taytay Municipality. The area was dominated by secondary and logged-over forest, with disturbance from slash and burn agriculture being found throughout the area. The area retained ca. 60% forest cover. Slope was generally moderate to steep, and elevation ranged from ca. 40-250 m. We trapped for small volant and non-volant mammals in forest, agricultural habitats, and two small caves near the lake for 14 days during October and November 2000. We totalled 471 trap-nights in three lines baited with coconut and peanut butter, and captured 3 Tupaia palawanensis, 23 Max- omys panglima, and 1 Rattus exulans. For- ty-two net-nights yielded 53 Cynopterus brachyotis, 2 Macroglossus minimus, and 1 Rousettus amplexicaudatus, and 13 harp- nights produced 1 Megaderma spasma, | Rhinolophus acuminatus, 2 Kerivoula hard- wickii, and 5 Tylonycteris pachypus. Addi- tionally, we captured 4 Megaderma spasma and 4 R. acuminatus by hand. Site 12 (9°27'48'N, 118°32'16"E) was lo- cated at the “rainforestation”’ site in Sitio Kandis, Aborlan Municipality, in forest/ grassland mosaic at ca. 40 m above sea lev- el, about seven km away from the next good secondary forest in the foothills of the Victoria Range. Charcoal making, logging, rattan collection, grazing and burning were common until 1994 when such activities were made illegal. The terrain was flat to 278 rolling, dissected by two creeks. The site consisted of 5.5 ha Imperata cylindrica grassland interspersed with single shrubs and trees, predominantly Antidesma ghae- sembilla, which forms the fire climax in more open situations, with Vitex pubescens, Guioa pleuropteris, Tarenna stenantha, Fa- graea fragrans, Lantana camara, and Mus- saenda philippica in protected areas not af- fected by fire in the last ten to fifteen years. About 4.5 ha consisted of regenerating for- est, close to two seasonal creeks, dominated by Garcinia benthami, G. parviflora, Can- arium asperum, Polyscias nodosa, and Bar- ringtonia curranii. Undergrowth was mod- erate to very dense. Canopy height was on average eight meters, with some taller emergents such as Nephelium sp. and Dip- terocarpus gracilis. The most conspicuous vine was Gnetum latifolium. Macrophytic epiphytes were virtually absent. Leaf litter layer was usually not closed, except in very dry years. Surveys were conducted regular- ly from 1997-2000 with 10 medium-sized Sherman live traps, 10 commercial live rat traps, and 55 wire mesh traps (measure- ments equal to medium Shermans). Baits were roasted coconut with peanut butter, but mostly fruits available in the area. A total of 3514 trap-nights yielded 169 small mammals (28 Tupaia palawanensis, 8 Sun- dasciurus juvencus, 11 Rattus exulans, 16 Rattus tiomanicus, 104 Maxomys panglima, and 2 Sundamys muelleri). Mist nets (2 X 6 m) were set along trails in forest, in grass- land, gaps in the shrub cover, and rarely in the canopy (ca. 8 m high). Capture sites were often near or in fruiting shrubs or trees, since the main focus of the study was on frugivores. From 1997 to 2000, a total of 482 net-nights yielded 1257 bats (1 Ac- erodon leucotis, 829 Cynopterus brachy- otis, 394 Macroglossus minimus, 5 Eonyc- teris spelaea, 4 Rousettus amplexicaudatus, 18 Megaderma spasma, \ Hipposideros diadema, 4 Scotophilus kuhlii, and 1 Mu- rina cf. tubinaris); all but the M. cf. tubi- naris were released. Site 13 (09°13'N, 118°26'E) was on Rasa PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Island, Narra Municipality, a small (8.3 km?) shallow coral island, 1.8 km offshore in the Sulu Sea. Approximately two-thirds of the island was covered with mangrove and one-third with coastal forest over lime- stone. About five percent of the latter had been converted into coconut plantation. Se- lective logging was done until the early 1990s, resulting in the complete loss of ma- ture Intsia bijuga. The mangrove consisted of nine species of the genera Rhizophora, Sonneratia, Avicennia, Bruguiera, Aegicer- as, and Ceriops. Canopy height was vari- able, usually between 8 and 15 m. Emer gent trees (e.g., Garuga floribunda and Pterocymbium taluto) ranged up to 42 m. Leaf litter layer was not closed, except un- der very dry conditions. Barren coral rocks and crevices were ubiquitous. Buttresses were a common feature of all emergent for- est trees. Under open conditions, an herbal layer consisting of /mpatiens sp. was pre- sent. Vines were abundant, including climb- ing bamboo Dinochloa sp., often forming dense tangles. Macrophytic orchids were present, but relatively scarce. Traps and nets were set along a trail within the coastal for- est. Traps were baited with roasted coconut with peanut butter, and a few with crickets. Most traps caught hermit crabs. Nets were set in the understory, which was very open from January to April 2002 and only pro- vided very few fruits due to an extended dry spell. Trapping totaled 104 trap-nights, and produced 3 Rattus tanezumi and 2 R. tiomanicus. Netting totaled 28 net nights, and yielded 2 Cynopterus brachyotis, 5 Ma- croglossus minimus, and 5 Megaderma spasma; all bats were released, and the rats preserved as vouchers. Site 14 (9°17'N, 118°27’E) was in fresh- water swamp forest in Narra Municipality, in about 5 ha of remnant forest along Tar- itien River. The habitat was dominated by two woody species, Nauclea orientalis and Pandanus sp., at lower elevations, which are flooded for at least six months. The herb layer was not extensive and was dominated by Acrostichum sp. The higher portions VOLUME 117, NUMBER 3 were dominated by pioneering species of early to medium successional stages, such as Trema orientalis, Vitex pubescens, and Commersonia bartramia. Even during ex- treme dry spells like that in the first half of 2002, there were isolated open water bodies left, which connected to several creeks dur- ing the rainy season. The swamp forest is bordered by ricefields and grassland. Twen- ty trap-nights yielded 1 Rattus exulans and 1 R. tiomanicus. Twelve net-nights yielded 27 Cynopterus brachyotis, 5 Macroglossus minimus, and 1 Megaderma spasma; all bats were released and the rats preserved as vouchers. Site 15 (10°12'N, 118°55’E) was along the “‘jungle trail” near the Central Park Sta- tion in Puerto Princesa (formerly St. Paul) Subterranean River National Park (PPSRNP). Primary lowland forest on steep slopes ascended from sea level to about 40 m. Five net-nights on 15 September 1996 yielded 1 Cynopterus brachyotis, 2 Rhino- lophus arcuatus, 3 Rhinolophus virgo, and 2 Rhinolophus sp.; all were released. Ad- ditionally, all three authors made visual ob- servations at various times. Accounts of Species Order Insectivora Family Soricidae—Shrews Crocidura palawanensis.—We never en- countered this poorly known species. It is endemic to the Palawan faunal region and has been taken in old-growth rain forest and shrubby second growth (Heaney & Ruedi 1994); the holotype came from “‘deep forest near the sea at ... Brooke’s Point’”’ (Taylor 1934), a second from near sea level in Ba- buyan, Puerto Princesa (Hoogstraal 1951, Sanborn 1952), and a third from 3600— 4350 ft (ca. 1100—1300 m) on Mt. Mantal- ingajan (USNM); two additional specimens are from Balabac (Heaney & Ruedi 1994). IUCN (2002) lists this species as Vulnera- ble, but current definitions suggest that Data Deficient would be more appropriate. Crocidura sp.—Reis & Garong (2001) 279 reported a single humerus of a shrew, sub- stantially smaller in size than C. palawa- nensis, from undated sediments in a small rock-shelter cave near Tabon Cave on Lip- uun Point, near Malunut Bay, in Quezon Municipality (near the location of the town of Quezon as shown in Fig. 1). They de- scribed the specimen as being similar in size to C. monticola from Borneo. We ten- tatively include it in our tallies of native species of Palawan, but we recommend that it be sought by trapping with small snap- traps baited with live earthworms and pit- fall traps. Suncus murinus.—This introduced com- mensal is abundant in urban and agricultur- al areas (Rabor 1986); in forest, it is rarely present, but occasionally is common (Hea- ney et al. 1989, Heaney & Tabaranza 1997). It is found throughout Asia and Indo-Aus- tralia, including the Philippines (Heaney et al. 1998). We observed this species fre- quently in houses in Puerto Princesa City and the State Polytechnic College of Pala- wan in Aborlan Municipality. Order Scandentia Family Tupaiidae—Tree Shrews Tupaia palawanensis.—TVhis common species is endemic to the Palawan faunal region (Wilson 1993); it is related to T. glis, which is widespread on the Sunda Shelf (Corbet & Hill 1992). It is widespread on Palawan (Taylor 1934), and is usually com- mon in secondary and primary lowland for- est, though local densities may be highly variable between apparently similar habitats (Dans 1993, Hoogstraal 1951, Sanborn 1952). It is rare in montane forest, and com- mon but patchy in agricultural areas. We captured and/or observed this species in co- conut and cashew plantations, brushy areas with a few small trees (Sites 6, 11, and 12), secondary and logged-over forest (Sites 7 and 11), and primary forest (Sites 1, 2, 3, and 15) from near sea level to 1400 m. IUCN (2002) lists this species as Vulnera- ble, but we concur with Heaney et al. 280 (1998) that the species should be delisted due to the variety of habitats used and its apparent abundance. Specimens examined: Deasitew lan) SiteyZn Gy): Order Chiroptera Family Pteropodidae—Fruit Bats We follow Ingle & Heaney (1992) and Heaney et al. (1998) in regarding reports of Haplonycteris fischeri (Kock 1969) and Ptenochirus minor (Yoshiyuki 1979) from Palawan as erroneous, probably originating in mislabeled specimens. Pteropus hypo- melanus is known from Cuyo Island, at the northeast edge of the Palawan faunal region (Heaney et al. 1998), as well as in the oce- anic Philippines and on islets around Bor- neo, and should be sought on Palawan. Acerodon leucotis.—This poorly known species is endemic to the Palawan faunal region. Hoogstraal (1951) found the species in an area with patches of ““much disturbed remnants of original forest and dense sec- ond growth forest’? on Busuanga Island, and two specimens were taken at Santiago, Iwahig (in Puerto Princesa) on Palawan (Sanborn 1952). A specimen from Bat Is- land (Barangay Tagburos, in Honda Bay), taken by P. O. Glass in 1978, is housed in the UMMZ. Heaney sighted large numbers of medium-sized, pale-furred flying foxes at Site 15 in April 2000, in the clearing of the old park headquarters near the center of the park (“Central Park’’), that were probably this species. Widmann captured one at Site 12 at a height of 5 m, and saw others feed- ing in the canopy at ca. 8 m height. The IUCN (2002) lists this species as Vulnera- ble; we regard it as Data Deficient. Cynopterus brachyotis.—Found through- out Southeast Asia; in the Philippines, it is common to abundant in secondary forest and agricultural areas, and rare in primary forest (Heaney et al. 1998); Sanborn (1952) reported many from Palawan. We netted this species frequently in secondary forest and agricultural areas at Sites 6, 7, 11, and 12, in freshwater swamp forest at Site 14, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and in coastal forest on Rasa Island (Site 13). We also captured this species in pri- mary forest in a tree fall gap (Site 1), over a stream (Site 2), and at a place with no visible disturbance (Site 15). We found them roosting in various-sized groups in three caves at Sites 5 (there appeared be less than 50 in each of two caves) and 9 (ca. 300 individuals); although this species occasionally roosts in caves on Borneo (Payne et al. 1985), there are no previous records of such roosts in the Philippines. On several occasions we captured them car- rying whole green figs (Ficus sp.) during flight; two of these individuals were return- ing to a cave at Site 5 between 1900— 2000 h. Out of 20 adult females caught at Site 14 on 20 March 2002, 15 were pregnant and 5 were carrying a single suckling young. On 1 and 2 April 2000, we captured three adult females at Site 5; all were carrying a single suckling infant during flight. On 17 May 2000 we captured eight adult females at Site 5; one was pregnant, one was carrying a suckling infant during flight, and two were emaciated and may have recently weaned their young. On this date, we also captured a male with enlarged mammaries (see Francis et al. 1994). On 30 October 2000, among 25 adult females, none were pregnant but one was lactating. Specimens examined: 4, Site 1 (1), Site 5 (3). Eonycteris spelaea.—This widespread Southeast Asian species 1s common in ag- ricultural areas in the Philippines, where all known roosts are in caves (Heaney et al. 1998, Rickart et al. 1993). Sanborn (1952) reported a large series from a cave above Tanabog, Palawan. We netted five individ- uals at Site 12, but most of our records came from caves in lowland forest. We found this species roosting 1n caves at Sites 4 and 10; at Site 4, the roosting population appeared to exceed 2000. At Site 10, there was an extremely large population (proba- bly >50,000) of small pteropodids roosting inside the cave. We captured 49 pteropodids at the entrance to the cave, 48 of which VOLUME 117, NUMBER 3 were E. spelaea and one was a Rousettus amplexicaudatus. On 19 December 1999, all three adult females we captured at Site 4 were pregnant. On 21 October 2000 at Site 10, we captured 11 adult female E. spe- laea, four of which were carrying an infant during flight and five of which were preg- nant. This species is heavily hunted in some areas of the Philippines (Rickart et al. 1993, Utzurrum 1992), but we observed no evi- dence of that being the case on Palawan. Specimens examined: 5, Site 4 (3), Site 10 (2). Macroglossus minimus.—In the Philip- pines, this widespread Australasian species is common in secondary forest and agri- cultural areas and uncommon in primary forest up to more than 2000 m (Heaney et al. 1998, 1999). We captured this species in secondary lowland forest (Sites 7 and 12) and agricultural clearings (Site 11), usually near wild or domestic banana plants (Musa spp.), and in freshwater swamp forest in Narra Municipality and Rasa Island (Sites 13 and 14). Specimens examined: 3, Site 7 GD, Sit i), Pteropus vampyrus.—In the Philippines, this widespread Southeast Asian species oc- curs in primary lowland forest and adjacent agricultural areas (Heaney et al. 1998; Ra- bor 1955, 1986; Rickart et al. 1993; San- born 1953; Taylor 1934). Widmann esti- mated 400 individuals on Malinau Island, Aborlan Municipality in 1998, 570 on Rasa Island on 12 November 1999, and a small colony (ca. 40 individuals) at Lagan on Du- maran Island on 27 October 2001, based on departure counts. Flying foxes commonly sighted in Puerto Princesa City around mango and guyabano (= sour sop) trees are probably this species. In 1998, we found two individuals of this species that ap- peared to have been electrocuted on power lines, one at the Provincial Agriculture Cen- ter in Irawan, Puerto Princesa, and the other at the State Polytechnic College, Aborlan. We believe this species to be common over- all, but under moderate pressure due to 281 hunting and perhaps to electrocution on power lines. Rousettus amplexicaudatus.—Within the Philippines, this widespread Southeast Asian species is commonly found in agri- cultural habitats up to 500 m and rarely in primary lowland forest (Heaney et al. 1998). All known roosting sites are in caves (Heaney et al. 1989, 1991, 1998, 1999; Hei- deman & Heaney 1989; Rickart et al. 1993). According to Payne et al. (1985), R. amplexicaudatus often roosts in association with Eonycteris spelaea. We netted one in- dividual from a cave at Site 10 containing a large population of E. spelaea, one in an agricultural clearing in Site 11, and four in forest-grassland mosaic at Site 12; all of these sites are in heavily disturbed areas be- low 60 m. Specimens examined: 2, Site 10 (1), Site 11 (1). Family Emballonuridae—Sheath-tailed Bats There are no known records of Saccolai- mus saccolaimus from Palawan, but its widespread distribution from India to New Guinea, including the oceanic Philippines (Heaney et al. 1998), suggests that it may be present and should be sought. Emballonura alecto.—The Philippine sheath-tailed bat is known from Borneo, the Philippines, and Sulawesi (Heaney et al. 1998); we never encountered this species on Palawan, but Taylor (1934:200) captured five individuals “under an overhanging rock along Iwahig River, near the base of Thumb Peak’. Taphozous melanopogon.—The bearded tomb bat is widespread in southern Asia (Heaney et al. 1998). In the Philippines, it is common in urban areas and lowland ar- eas with limestone caves and rare in forest (Rickart et al. 1993, Sanborn 1952). There is a previous record from the vicinity of Puerto Princesa (Allen 1922), and A. C. Al- cala collected 6 specimens from Sitio Mal- abusog, Tinitian, Roxas Municipality in 282 1984 which are deposited in the UMMZ. We never encountered this species. Family Megadermatidae—False Vampire and Ghost Bats Megaderma spasma.—This widespread southern Asian species is common in pri- mary lowland forest and disturbed forest in the Philippines (Heaney et al. 1991, 1998, 1999; Rickart et al. 1993). We captured this species from sea level to ca. 500 m in sec- ondary forest (Site 7), primary forest (Sites 1 and 2), in a bamboo thicket (Site 11), and in or near caves (Sites 5 and 11). It was the most common insectivorous bat netted in forest-grassland-mosaic (Site 12), in swamp forest (Site 14), and in coastal forest (Site 13). At Site 1, we found this species roost- ing in small groups (<10) in four hollow trees distributed throughout the area. At Site 11 we found ca. 12 individuals roosting in a small cave (ca. 0.5—3 m wide, 0.3—1.5 m high, and 10 m long) along with Rhino- lophus acuminatus. We also found two in- dividuals roosting in a small cave (also Site 11) that had been severely disturbed by treasure hunters three years earlier. Cranial measurements of three individuals (Table 1) are slightly smaller than those of specimens from Leyte and Biliran (Rickart et al. 1993) and southern Luzon (Heaney et al. 1999). Specimens examined: 3, Site 1 (3). Family Rhinolophidae—Horseshoe and Roundleaf Bats Several poorly known but apparently widespread species in this family occur on the Sunda Shelf and in the oceanic Philip- pines and should be sought on Palawan; these include Hipposideros cervinus and H. lekaguli (Balete et al. 1995, Heaney et al. 1998, Ingle & Heaney 1992). Hipposideros ater.—Occurs from India to Australia (Heaney et al. 1998). Known from lowland and montane forest and caves (Heaney et al. 1991, 1998; Payne et al. 1985, Rickart et al. 1993). We found this species to be uncommon to abundant in PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON three caves in disturbed lowland forest at 50 to 250 m elevation at Sites 5 (17% of 575 captures) and 8 (<1% of captures). During March to April 2000, none of the 26 females we captured at Site 5 were preg- nant or lactating, but on 19 and 20 May 2000, 25 of 30 adult females were pregnant. We have re-examined a specimen from Pa- lawan in the UMMZ identified by Allen (1922) as H. bicolor, and a series from the Tigoplan River, Palawan in FMNH reported by Sanborn (1952), and now consider them to be H. ater; thus, we now know of no records of H. bicolor from Palawan. Cranial measurements of 5 individuals (Table 1) are smaller than those of H. bicolor (Heaney et al. 1999, Ingle & Heaney 1992) but match those of H. ater (Ingle & Heaney 1992, Rickart et al. 1993). Specimens examined: 5, Site 5 (4), Site 8 (1). Hipposideros diadema.—The diadem roundleaf bat is widespread from Myanmar to the Solomon Islands, with many previous records from Palawan (Allen 1922, Heaney et al. 1998). In the Philippines, it is com- mon in disturbed forest, agricultural areas (Ingle 1992, Rickart et al. 1993), and pri- mary forest (Heaney et al. 1998, Rickart et al. 1993). Reis & Garong (2001) reported specimens from sediments in a rock-shelter near Tabon Cave, Quezon Municipality dat- ed to 11,130 BP. We captured this species from sea level to 600 m in disturbed grass- land-forest mosaic (Sites 6 and 12), second- ary forest (Site 7), primary forest (Site 2), and at nearly all caves we visited (Sites 4, 5, 8, 9, and 10), and we observed large numbers (probably thousands) in the un- derground river cave at PPSRNP. All of the roosts we identified held groups of H. dia- dema numbering greater than 200. Thirteen adult females captured in December 1999 included none that were pregnant or lactat- ing. At Site 5 in March to April 2000, none of the 54 adult females were pregnant or lactating, but between 15 and 20 May 2000 26 of 43 were pregnant and one was car- rying a suckling infant during flight. One of 21, one of 12, and none of 13 adult females 283 VOLUME 117, NUMBER 3 (61t-8' Or) (€9'S—-Er'b) (OS E-E'E) (OL Y-LO'Y) (9E9-SI'9) (68b-—18'y) (IS8-Sr'8) (818-898) (8h 9I-ZL’ST) Cir aus Ore CLY EGS L8V 878 CL 8 OT 9L € J (CIP-T6E) (LES-S8') (99'E-CE) (S8P-6S) (€79-F09) (h6'P-L'b) (CL'8-Iv'8) (S6'8-8L'8) (97 9I-66'S1) COV (Z) 80°S Loe ILV 919 v8 v7 co8 L388 (C) E191 € ul os41a snydojouyy (S$ 8r-6 9r) (SLE-IV'€) (S6'7-6'y) (8L°9-8r'9) (CZ P-LI'b) (996-66) (OL8-19'8) (8081-87 LI) LLY (1) Le9 8S e €6'1 €9°9 OCT 876 99°83 89°LI G J (8'Sp-I'Sh) (9L'S-79'S) (EvE-—9E'€) (lOr-H8'r) (€S9-6H9) (Crb-Eeb) (hh 6-ZE6) (9'8-7Z9'8) (8ELI-I7 LI) SHOAIDUL CoV (Z) L's 6£ € L830 Ig°9 LEV Le 6 €9°8 (Z) ELI € ui snydojouryy (97S-S' 1S) (869-949) (90'S-SS'h) (ZS'9-LED) (198-68) (65°9-L8'S) (8LOI-IS 01) (LZ TI-E8'01) (€0'1Z-L'07) VCS (Z) L8°9 E87 cr9 crs So9 c9'01 COT (7) L807 v J vVVS 89 ves c9'9 968 L9 ama LL II 9C CC I wi 148pa19 snydojoulyy SISUIOU 8° CL vss ely LOS c9°9 887 LS8 y9'8 98°91 I J -40oq snydojouryy (S9r-9'Sh) (8r'9-S0'9) (L7r-80'r) (€9'S-67'S) (LH L-S69) (8I'S-ZO'S) (9€ 6-498) (C96-I7'6) (16 8I-Ir'81) 0-97 le9 Cl y srs VIL 80'S 90'6 €S°6 89°81 € J (LLY-CSb) (LO9-159) (ICr-E0'ry) (L9°S-9F'S) (OS L-CSL) (IH'S-10'S) (CS 6-186) (98'6-CL'6) (97 6I-E'81) SNIDNI COV vL9 CLV (G55 VSL Ics cS'6 6L°6 82 81 C ul -1 snydojouryy (adAjoyou) syimbav 1uas 6c OV = IS y 009 BOL 619 98°6 a0) a I ul -lapup snydojouiyy (1'8t-9'9p) (S6'7-99'y) (OV'9-8L'S) (SLL-S'L) (179-S0'9) (8L6-€9'6) (C6'0I-S'°0I) LLY (1) 95°9 OLY 109 6S L LI9 89'6 OL OI (1) vr'6l V J SH! COV VIL LLY i) SOL €co9 186 68°01 Se 0c I ur -1uinov snydojouyy (VOr-l'0r) (Ly'S-S6'y) (SPE-IvV'e) (MrOr) (LO4-SLb) (HO r-19'y) (6E8-€7 8) (88L-9L'L) (LL YI-IE V1) € OV Ics eve COV 987 BL [e8 C8 L vs vl C J (66€-1'6£) (VOr-E8'y) (EEE-CTE) (ITP-CLE) (64-89) (9S +-€'h) (O€'8-97'8) ©GLPLL) (LE VI-I7 41) L6e L810 8CE one 81 oT 8o8 e8L 6c v1 € ul Jai sosapisoddi py evs = 6Le 90'L 06 LOL ce Ll LI vl = I J (0'9S-¥'SS) (yOP-SE) (81'L-S6'9) (16'8-L9'8) (EP L-EL) (87TI-97 IL) (SS rI-cS FI) Ls 9 LL¢ LOL 6L'8 MEL Lol vs v1 a G ul pusvds pusaposapy ysugy yysug] yipesiq AOIYIOO} qejour tsug] Y)PIM YIpIM wyisugy u xoS soroodg WdedI0F feqeyeg [eiered WHOJLUe[OIY 4Se] 0} [e1g10 projseyyy onew0skZ, SAISIOUIO[ApUOD, oururea ‘sourddiiyg ‘puvys] uemvjeg wor oeprydojouryy pue sepneunopeseayy Jo s}[npe Jo sjuouomMsvou! [eruRIO Jo sosuvi pue suvoyy— | s[qQuy, 284 from July, August, and October were preg- nant (Sites 8, 9, and 10, respectively). Spec- imens examined: 5, Site 4 (2), Site 5 (2), Site 9 (1). Rhinolophus acuminatus.—This poorly known species occurs from Thailand to Lombok and Palawan, but not elsewhere in the Philippines (Heaney et al. 1998; speci- mens from Negros reported by Csorba et al. (2003) as this species were mislabelled). It is found in lowland forest on Borneo (Payne et al. 1985) and in secondary low- land dipterocarp forest on Banggi (Md. Nor 1995). We captured this species in caves from ca. 60 to 250 m in caves (Sites 8 and 11), a bamboo thicket (Site 11), and pri- mary forest (Site 1). At Site 8, we captured 90 individuals out of 2775 captures. At Site 11 we found ca. 20 individuals roosting in a small cave (ca. 0.5—3 m wide, 0.3—1.5 m high and 10 m long) with ca. 12 Megad- erma spasma. At Site 1, we took two in- dividuals over a small stream just below a rock outcrop containing many fissures suit- able for roosting bats. We also captured a single individual in our temporary living quarters at Site 1 after we observed for sev- eral days a bat feeding inside our semi-en- closed tent for several minutes daily be- tween 0430 and 0600 h. Of 15 adult fe- males taken in July 2000 at Site 8, one was pregnant and one was lactating. Cranial measurements (Table 1) match those pre- viously available (Ingle & Heaney 1992) that are based on series from Balabac and Busuanga reported by Kuntz (1969) and housed in the USNM. We also refer two specimens collected by A. C. Alcala on 13 July 1984, at Malabusog, Roxas Munici- pality housed at UMMZ (162885 and 162886) to this species, and include them in Table 1. Sanborn (1952) reported a single speci- men from Palawan (housed in FMNH) that he regarded as the first record from Pala- wan. However, we have determined that a single specimen (UMMZ 53112) collected in the late 1800s by the Beal/Steere Expe- dition and subsequently named R. ander- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON seni aequalis (Allen 1922) was this species. Heaney compared this specimen, which is the holotype and only known specimen, to series of all species currently known from Palawan. It was unambiguously identified as R. acuminatus; as noted by Medway (1977:32), a dorsal connecting process with a prominent triangular point (as shown by Medway 1977 fig. 6a and by Ingle and Hea- ney 1992 fig. 13a) is present, the base of the sella is not expanded into a cup, the median groove of the horseshoe is not broadened, and papillae are not present. The ears (18 mm) are less than half of the length of head plus body, and the forearm is 46.4 mm. The skull (measurements in Ta- ble 1) is virtually identical to those in the series from Sites 1 & 8, including overall size and shape, nasal swellings, braincase breadth and inflation, toothrows, palatal bridge, foramina in the roof of the posterior portion of the nasal passage, and bullae. Be- cause R. acuminatus was named by Peters in 1871 (from Gadok, Java), we therefore recognize R. anderseni aequalis as its ju- nior synonym. We note that Cabrera (1909) described Rhinolophus anderseni “proba- bly from Luzon’’. Aside from the holotype of R. anderseni aequalis, no specimens have subsequently been referred to this spe- cies. Csorba et al. (2003) tentatively as- signed R. anderseni Cabrera as a junior synonym of R. arcuatus on the basis of the original description plus new drawings of the noseleaf and measurements of the skull, but without examining the holotype. We provisionally accept this, but point out the need for direct examination and compari- sons. IUCN (2002) lists R. acuminatus as Data Deficient. Specimens examined: 8, Site 1 (4), Site 8 (1), Malabusog (2), and the holotype of R. anderseni aequalis. Rhinolophus arcuatus.—Widespread from Sumatra to New Guinea (Heaney et al. 1998). Specimens from the Philippines currently identified as R. arcuatus may con- sist of two or more species (Heaney et al. 1991, 1999; Ingle & Heaney 1992: Rickart et al. 1993). Individuals referred to this VOLUME 117, NUMBER 3 “‘species”” have been found in agricultural areas, secondary forest, and primary low- land, montane, and mossy forest (Heaney et al. 1991, 1999; Ingle 1992; Rickart et al. 1993). We regularly captured this species at elevations from sea level to 1400 m in low- land primary forest (Sites 1 and 2), mon- tane forest (Site 3), and caves (Sites 5 and 8). We also captured a single individual in the understory of mature but disturbed for- est near a cave at Site 9, and we tentatively identified two individuals from Site 15 (which were released) as belonging to this species. Of 8 adult females taken between 15 and 20 May 2000 at Site 5, 5 were preg- nant and one was lactating. Of 22 adult fe- males taken in July 2000 at Site 2, one was lactating, and of 189 captured in July at Site 8, none were pregnant but 14 were lactat- ing. These are the first specimens of this species from the Palawan faunal region. Cranial measurements (Table 1) closely match those of specimens from Leyte (Rickart et al. 1993) and southern Luzon (Heaney et al. 1999). Specimens examined: 5, Site 1 (2), Site 2 (1), Site 3 (2). Rhinolophus cf. borneensis.—A_ single specimen taken by P. O. Glass on 31 Jan- uary 1978 at “Sabang, Buenavista” (in Barangay Cabayugan, near Ulugan Bay in Puerto Princesa Municipality, ca. 10°05’N, 118°49'E; UMMZ 161395) appears to be this species. It was previously known from Indochina, the Malay Peninsula, Java, and Borneo, as well as some smaller islands in the southern South China Sea (Corbet & Hill 1992, Csorba et al. 2003); this is the first record from Palawan and from the Philippines. Cranial measurements and fea- tures (Table 1) closely match specimens in FMNH from Sarawak, the Natuna Islands, and Sabah, and external features are similar, but because we have only a single speci- men, the identification is tentative. On Bor- neo, “the species roosts in caves, some- times in colonies of several hundred indi- viduals” (Payne et al. 1985). P. O. Glass (in litt.) noted that he captured the specimen in a mist net in a small banana grove in an 285 area of mixed agricultural/second growth forest within 1 km of mature forest. Spec- imen examined: 1, from Sabang. Rhinolophus creaghi.—This species was previously known from Borneo and Madura Islands, where it often roosts in caves (Cor- bet & Hill 1992, Csorba et al. 2003, Koop- man 1993, Medway 1977, Payne et al. 1985). This is the first record of this species from Palawan Island and the Philippines. On Palawan, we found it to be common in primary lowland forest from near sea level to at least 700 m. We captured one individ- ual at Site 1 and 11 individuals at Site 2. It roosts in caves, often in large numbers; we captured 368 (45% of captures) at Site 4, 239 (9% of captures) at Site 8, and 33 (6% of captures) at Site 5. Of 135 adult females captured in December 1999 at Site 4, 8 were pregnant. Of 16 captured at Site 5 in March to April, none were reproductively active; of 11 at Site 2 and 151 at Site 8 (July 2000), none were pregnant but one and 12 were lactating, respectively. Cranial measurements (Table 1) show this to be the largest member of the genus on Palawan; our specimens are not distinguishable from a small series from Borneo (FMNH 4707 1— 47075). Two previously unidentified speci- mens from Mt. Salicod, 2300 ft. (which may be the same mountain as Mt. Salakot, Site 2; P. O. Glass, in lit.), taken by P. O. Glass in 1978 and housed in the UMMZ, were taken earlier but were not reported; cranial measurements from these specimens are included in Table 1. This species is list- ed as Near-Threatened by IUCN (2002). Specimens examined: 7, Site 4 (3), Site 5 (1), Site 8 (1), Mt. Salicod (2). Rhinolophus macrotis.—This poorly known species ranges from India to Su- matra and the Philippines, where it is known from lowland forest with some re- cords from caves (Heaney et al. 1998, Ingle 1992). Our specimens represent the first re- cord from Palawan. We captured this spe- cies in or near three caves in disturbed low- land forest at SO—250 m at Sites 5 (10 cap- tures) and 8 (25 captures). The species ap- 286 pears to be uncommon. At Sites 5 and 8 it represented less than 5% and 1% of our captures, respectively. At Site 5 on 20 May 2000, we captured and examined two adult females; both were pregnant. Of 8 adult fe- males captured in July at Site 8, none were pregnant or lactating. Cranial measurements of 5 individuals (Table 1) fall within or near the range for the species in Ingle and Hea- ney (1992). Specimens examined: 5, Site 5 (4), Site 8 (1). Rhinolophus virgo.—This Philippine en- demic is widely distributed within the Phil- ippines (Heaney et al. 1998). It is known from secondary forest, primary lowland forest, reaching mossy forest on small, low- lying islands, and often roosts in caves (Heaney et al. 1991, Ingle 1992, Rickart et al. 1993); Sanborn (1952) reported a series from Tanabog, Palawan. This species ap- peared to be rare in the cave at Site 4 (0.5% of captures), common at Site 8 (151 cap- tures, about 6% of total) and abundant in some caves at Site 5 (15% of captures). We also captured this species in primary forest at Sites 2 and 15. Of 29 adult females cap- tured between 27 March and 4 April 2000, none were pregnant or lactating. Of 35 fe- males captured at Site 5 between 15 and 20 May 2000, 16 were pregnant and 5 were lactating. Of 78 females taken at Site 8 in July 2000, 6 were pregnant and 2 were lac- tating. Cranial measurements (Table 1) fall within the range (Ingle & Heaney 1992) or very near the range (Rickart et al. 1993) of previously available individuals. IUCN (2002) lists this species as Near-Threatened. Specimens examined: 6, Site 4 (2), Site 5 (4). Family Vespertilionidae—Vesper and Evening Bats This diverse family of bats is generally poorly documented, and more species should be sought on Palawan, including such widespread taxa as Harpiocephalus harpia, Murina cyclotis, Myotis ater (which Hill 1983 and Corbet & Hill 1992 have PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON shown to be distinct from Myotis muricola and to be present on Culion Island in Pa- lawan faunal region), Philetor brachypte- rus, and Pipistrellus tenuis. Glischropus tylopus.—This poorly known species is found from Myanmar to the Molucca Islands and Palawan (Heaney et al. 1998). In Peninsular Malaysia it roosts in rock crevices, bamboo, and in new ba- nana leaves (Payne et al. 1985). We never encountered this species, but it is repre- sented by a specimen in the USNM (Hol- lister 1913). Kerivoula hardwickii.—This species is widespread from India and southern China to the Lesser Sunda Islands and the Phil- ippines (Heaney et al. 1998). It was previ- ously known from lowland, montane, and ridge-top mossy forest from 500 to 1600 m in the Philippines (Heaney et al. 1999, Rickart et al. 1993). Everett (1889) includ- ed mention of this species from Palawan. A previous record from UMMZ (Heaney et al. 1998) has been re-identified as K. white- headi, as noted below. Payne et al. (1985) reported the species to “‘frequent the un- derstory of tall forest’”’ on Borneo, and Md. Nor (1995) caught one in primary lowland dipterocarp forest on Banggi Island “in the axil of a leaf on a rattan vine | m above ground’’, and netted them in the understory of primary forest on Balambangan Island. We captured two adult females, one of which was lactating, in a bamboo thicket at Site 11 (ca. 60 m asl), and two individuals in the understory (ca. 2—4 m above the ground) of primary lowland forest at ca. 650 m elevation at Site 2, all in harp-traps. Specimens examined: 3, Site 2 (1), Site 11 (2). Kerivoula_ pellucida.—This poorly known species is known from the Malay Peninsula, the Sunda Shelf, Jolo, and Pa- lawan (Heaney et al. 1998). Known only from lowland forest (Payne et al. 1985). Taylor (1934) reported two specimens from Palawan (no locality given) that he obtained from a group of seven that he found flying together in daylight: his map (Fig. 13), VOLUME 117, NUMBER 3 shows the locality in the vicinity of Broo- ke’s Point. On 21 June 2000, using a harp- trap we captured an adult female carrying a suckling infant in secondary lowland for- est (ca. 80 m) over a small stream at Site 7. Cranial measurements of the adult female (Table 2) are smaller than the one specimen from Davao del Norte, Mindanao available to Ingle and Heaney (1992), but they are otherwise very similar; additional speci- mens are badly needed to examine patterns of variation. Specimens examined: 2, Site 7 (2). Kerivoula whiteheadi.—This poorly known species is widely distributed from southern Thailand to Borneo and the Phil- ippines (on Luzon, Mindanao, and Pala- wan; Heaney et al. 1998). In the Philip- pines, it is known only from near sea level in disturbed forest and agricultural areas (Sanborn 1952). A single specimen in the UMMZ captured by P. O. Glass on 29 Sept. 1978 at Irawan, Puerto Princesa Municipal- ity (noted as 2 km N Irawan, at the base of Mt. Beaufort by P. O. Glass, in litt.) was erroneously reported by Heaney et al. (1998) under both K. hardwickii and this species. Additionally, two specimens taken “under banana fronds” by C. A. Ross on 8 April 1987 at Barangay Binwang, Quezon Municipality, are housed in the USNM. We captured a single individual of this species in a harp-trap near ground level in a cogon grassland (Umperata cylindrica) at Site 6. Cranial measurements of these four individ- uals (Table 2) are similar to those in Ingle & Heaney (1992) of a specimen from Min- danao, though some variation in size is pre- sent; more specimens are needed to assess geographic variation. Specimens examined: 4, Site 6 (1), Irawan, Puerto Princesa Mu- nicipality, 60 m (1), and Binwang, Quezon (2). Miniopterus australis.—This common species is found from India to Australia; it is widespread in the Philippines, but this is the first record from Palawan (Heaney et al. 1998). It is known to roost in caves in low- land areas of agriculture or second growth 287 (Heaney et al. 1991, Rickart et al. 1993, Sanborn 1952). We captured this species in varying numbers at several caves. In pri- mary and disturbed lowland forest at Sites 4 and 5 it was scarce, represented by less than 1% and less than 3% of captures, re- spectively. At Site 8 it was abundant (45% of 2775 captures). Cranial measurements of A individuals (Table 2) are similar to those reported by Ingle and Heaney (1992) and Rickart et al. (1993) from the Philippines, and by Corbet & Hill (1992) from through- out the species range. Specimens examined: 4, Site 4 (2), Site 5 (2). Miniopterus schreibersi.—This common species is found from Europe to the Solo- mon Islands and is widespread in the Phil- ippines, but this is the first record from Pa- lawan (Heaney et al. 1998). It is common in caves throughout the lowlands in agri- cultural areas and forest and known from both lowland and montane forest (Heaney et al. 1991, 1999; Rickart et al. 1993; San- born 1952). We captured this species in and around caves in disturbed and primary low- land forest at Sites 4, 5, and 8. At Sites 4 and 8 the species was abundant, represented by 47% and 26% of captures respectively. At Site 5 it was common at 10% of 575 captures. Of 214 adult females captured in December 1999 at Site 4, only one was pregnant. Of 36 captured between 15 and 20 May 2000 at Site 5, 15 were pregnant and none were lactating. Of 421 captured at Site 8 in July 2000, none were pregnant but 4 were lactating. Cranial measurements (Table 2) are similar to those reported by Ingle and Heaney (1992) and Rickart et al. (1993) from the Philippines, and by Corbet and Hill (1992) from throughout the species range. IUCN (2002) lists this species as Near-Threatened, but its abundance in heavily disturbed habitat in the Philippines (Heaney et al. 1998, Rickart et al. 1993) makes this inappropriate. Specimens ex- amined: 6, Site 4 (4), Site 5 (1), Site 8 (1). Miniopterus tristis.—This widespread species is found from the Philippines to the Solomon Islands; this is the first record PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 288 (9'7-8°CT) (SO'P-LO'E) (I8C-IL7) (HL T-6S7) (Sr'E-LE'E) (OB'E-LS'E) (969-659) (96L-9S°L) (HE OI-OL 01) DET 98 € CLT 89°C Ive pL cL9 CLL VC Ol G J snd VEC 16'€ 98°C 69°C cre Efe IL9 86'L 6c OL I ul -KYond sisajoKuojs I, OSE — EC te 8Le OLY 68 7 O9'L 106 cO'el I J OVE oe Cee C8 t 887 SES 06L Ie6 Oe el I we snoiupavl snjjassidig 8S 69°6 b8't 88'S Ors 169 89'6 CCCI sl 6l I J snjoidofns suoxp 187 898 CIV les OE Ll 889 IES) sl 88 LI I ut SNSADIOLIDUL SOK] Tee v9'9 AWE O6'€ ec Ss 80'S LL 116 c9' VI I ul SUDUIQHY “JO DULINGY, (€E€S-E'€S) (1E°8) (OF t-EE pr) (185-995) (16 L-SL'L) (9L:9-OL'9) (€6'6-79'6) (8LOI-VL 01) (PL 8I-IL'81) ees Ie8 Mew els €8L EL) 9L'6 OL 01 CL 81 C J CVS €18 COL 99°S CLL I¢‘9 C86 cL Ol OS 81 I ul SUSI sniajdonnpy (6€t-76E) (8P9-LE9D) (PS E-9TE) (9S P-CEH) (ST9-0'9) (CES-O6'p) (EE8-CI'8) (LS8-0E'8) (rerI-L9'rI) € CV ev9 6 € San v9 co's 618 Ge) 8L V1 ¢ J 1S1OG V LV ce9 coe Bor 809 Tes €L8 83°38 ee Sl I ul “lays Sniaidomlpy (QLE-T9E) (6L°S-95'S) (9D'E-16'7) (6LE-EL'E) TTS-9T'S) (Ob r-9Tb) (EE L-ETL) (8SL-67L) (SIEI-0671) Wie 99'S 00'€ OL'e 61's 6c PV 8CL 6e L 90° €1 € 5 L9€ 6L'S 90'€ O8'€ Ics OV OTL ICL SC El I we sypusny sniaidomp (yOE-r'67) (80°9-OL'S) (6r7Z-rE'?) (OEE-ZI'E) (C7S-LO'S) (LL9-95°9) (p9'L-rr'L) (87TI-S8'11) L'6C C8'S Iv'c Ble LVS aa 99'9 OOL 00°C! v J IpooayouiyM vjnoALay ele v69 8S L8¢ LOS 80°€ I¢L 978 Ic el I J ppionjjad pjnoaLiay (LEE-V'EE) (FO'L-16'9) (967-18'7) (78'E-S9'E) (HL'S-OS'S) (LO'P-LO'Y) (ES L-SO'L) (9L'8-79'8) (L8°EI-€9'ET) OEE 869 06°C We (65S LOY 6c L OL'8 cLel C J Tee 89 CLS ecg 6S cor PIL 878 6e el I UL = MYDIMpADY DINOALMAY sue] wsuay tppreiq MOIY}OOY qe[our wsue] (pin (pin yisuey u xaS saroadg UIvOI04 Teqered elered WIOJLIL[OI 4se] 0} [euqio proisey onewioshzZ SAISIOUL ourued -o[ApuoDd ee —————————— ‘sourddiryg ‘puels] uemeleg Woy oepruorniedso, jnpe JO SJUSWOINSveUI [eIULIO JO SasuvI pue sULIIAI—Z [GUL VOLUME 117, NUMBER 3 from Palawan (Heaney et al. 1998). The species is known to roost in caves and for- age in disturbed forest (Rickart et al. 1993, Sanborn 1952). We captured one specimen in a cave surrounded by old-growth forest at Site 4 and two in a cave surrounded by disturbed areas and secondary forest at Site 10. Miniopterus tristis appeared to be con- sistently less common than the other spe- cies of Miniopterus. Cranial measurements of 3 individuals (Table 2) are similar to those reported by Ingle and Heaney (1992) and Rickart et al. (1993) from the Philip- pines, and by Corbet and Hill (1992) from throughout the species range. Specimens examined: 3, Site 4 (1), Site 10 (2). Murina cf. tubinaris.—A single speci- men of a small tube-nosed bat (genus Mu- rina) was taken in a lowland grassland-for- est mosaic at Site 12 on 24 March 1997, and is housed in the Staatliches Museum fur Naturkunde in _ Stuttgart, Germany (#49238). The specimen (Table 2) is very similar to a series from Tonkin, Vietnam (FMNH 32203-32204, 46626-46627), though slightly larger. In our specimen, as in the Vietnamese series, the upper tooth- rows converge slightly, the anterior pre- molars are reduced, and the canines are short but longer than the premolars, as not- ed by Koopman and Danforth (1989) and Corbet and Hill (1992). The length of fore- arm (33 mm) falls at the center of the range given by Koopman and Danforth for M. tubinaris (28—35 mm), and at the high end given for M. suilla (26-33 mm). Koopman and Danforth (1989) considered M. florium, M. suilla, and M. tubinaris to be members of a species group, and perhaps to be con- specific, noting that few specimens are available. Corbet and Hill (1992) re-empha- sized the uncertainty in current taxonomy, but took a somewhat different view, refer- ring specimens from Borneo to M. suilla, rather than to M. tubinaris. While we agree entirely on the need for more specimens and further study, we follow Koopman & Danforth (1989) on referring Bornean spec- imens to M. tubinaris, and provisionally re- 289 fer the specimen from Palawan to this same species. Specimen examined: 1, Site 12 (1). Myotis horsfieldii.i—This common spe- cies is distributed from southeastern China to the Malay Peninsula, Sulawesi, and the Philippines (Heaney et al. 1998). On Bor- neo, the species “‘roosts in crevices or bell- holes in caves, usually not far from large streams or rivers” (Payne et al. 1985). In the Philippines, it has been recorded in low- land forest and agricultural areas, up to 800 m (Heaney et al. 1998). We never encoun- tered this species; two specimens in the UMMZ taken by A. C. Alcala in Sitio Mal- abusog, Tinitian, Roxas Municipality in 1984 were reported by Heaney et al. (1998). Myotis macrotarsus.—This species is known from Borneo and the Philippines (Heaney et al. 1998); it roosts in caves near sea level and forages in agricultural areas (Heaney and Utzurrum, unpubl. data). Md. Nor (1995) caught the species over a dry river bed and in the understory of primary lowland forest on Balambangan Island. In a cave in disturbed lowland forest at Site 8, this species was represented by <0.5% of 2775 captures. We also observed small numbers in the cave at PPSRNP. Cranial measurements (Table 2) of one individual show it to be slightly larger than those re- ported by Ingle & Heaney (1992). IUCN (2002) lists this species as Near-Threatened. Specimens examined: 1, Site 8 (1). Myotis rufopictus.—This poorly known Philippine endemic has been recorded from primary lowland and montane forest (Hea- ney et al. 1999, Mudar & Allen 1986). We never encountered this species; on Palawan, it is known from a single specimen in UMMZ reported by Allen (1922). We fol- low Ingle & Heaney (1992) in regarding this as one of several distinct species within the subgenus Chrysopteron, rather than rec- ognizing only a single species, Myotis for- mosus, within the subgenus (e.g., Corbet & Hill 1992). This species is not listed by IUCN (2002); we recommend listing as Data Deficient. Measurements in Table 2 of the specimen reported by Allen (1922) were 290 taken by Heaney. Specimen examined: 1, Puerto Princesa (1). Pipistrellus javanicus.—This species is distributed from Korea to Java and the Phil- ippines (Heaney et al. 1998). Taxonomic status is uncertain; P. imbricatus has been reported from Palawan (e.g., Allen 1922, Corbet & Hill 1992, Sanborn 1952), but In- gle and Heaney (1992) were unable to dis- tinguish more than one species of Pipis- trellus of this size in the Philippines; de- tailed study is needed. It is common in pri- mary montane forest and uncommon in lowland and mossy forest (Heaney et al. 1999, Ingle 1992, Sanborn 1952). We cap- tured two individuals from a roost in a hol- low tree in montane forest (ca. 1300 m) at Site 3. The opening in the tree appeared to have formed where a branch had been bro- ken off the tree and was quite small (ca. 1.5 < 5 cm). Cranial measurements (Table 2) fall within the range of specimens reported by Ingle and Heaney (1992), and are slight- ly smaller than a series from southern Lu- zon (Heaney et al. 1999). Specimens ex- amined: 2, Site 3 (2). Scotophilus kuhliiimTYhis common spe- cies is widespread from Pakistan to Taiwan and the Philippines (Heaney et al. 1998); it is abundant in urban and agricultural areas, and roosts in buildings and “tents”? made from modified palm leaves (Heaney et al. 1998; Rickart et al. 1989, 1993). Hollister (1913) and Taylor (1934) reported it from Puerto Princesa, Sanborn (1952) reported it from Brooke’s Point, and we found it to be abundant in buildings at the Provincial Ag- riculture Center, Irawan, Puerto Princesa, and in staff houses of the State Polytechnic College in Puerto Princesa, and in mixed urban/agricultural areas (Site 12). Tylonycteris pachypus.—This tiny bat is widespread from India to the Philippines (Heaney et al. 1998). In the Phillipines, it is known from bamboo stands in agricul- tural areas (Heaney & Alcala 1986); Hol- lister (1913) reported a specimen from Puerto Princesa. We captured several indi- viduals of this species in a bamboo thicket PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON at Site 11. Very near the capture site we observed what appeared to be more than a dozen individuals of this species foraging over a few remnant trees in a cleared area with houses that is immediately surrounded by logged-over and secondary forest. Spec- imens examined: 5, Site 11 (5). Tylonycteris robustula.—This species is also widespread from southern China to the Lesser Sunda Islands and the Philippines (including records from Calauit, Luzon, and Palawan); its habitat is apparently similar to that of 7. pachypus (Heaney & Alcala 1986, Heaney et al. 1998). We never encountered this species. Family Molossidae—Free-Tailed Bats This family is generally poorly known in Southeast Asia, partly because they typi- cally fly high above the canopy and are therefore rarely netted. At least one species (Chaerophon plicata) is widespread in the region and should be sought on Palawan. Cheiromeles torquatus.—This poorly known species is found from Sumatra to Java, Borneo, and Palawan, but not the rest of the Philippines (Heaney et al. 1998). It roosts in large caves and hollow trees and forages in open areas, over streams, and above forest canopy on Borneo (Payne et al. 1985). We never encountered this spe- cies, which was documented on Palawan by Sanborn (1952) based on a single specimen. IUCN (2002) lists this species as Near- Threatened. Mops sarasinorum.—This very poorly known species occurs in Sulawesi and the Philippines; the Palawan record is based on a single specimen in the Senckenberg Mu- seum, Frankfurt (Heaney et al. 1998). It prob- ably occurs in lowland forest (Heaney et al. 1998). We never encountered this species. It is listed by IUCN (2002) as Near-Threatened, but we recommend Data Deficient. Order Primates Family Cercopithecidae—Monkeys Macaca fascicularis.—This common monkey occurs from Myanmar to Timor VOLUME 117, NUMBER 3 and the Philippines (Fooden 1995, Heaney et al. 1998). It is known from agricultural areas near forest, second growth, secondary forest, and primary lowland and montane forest (Heaney et al. 1998, 1999; Rickart et al. 1993); Sanborn (1952) reported speci- mens from Iwahig, Puerto Princesa, and Brooke’s Point. Reis & Garong (2001) re- ported a specimen from sediments in a rock-shelter near Tabon Cave, Quezon Mu- nicipality dated to 11,130 BP. We common- ly observed this species at all of our sites (except Site 13), in secondary and primary forest (including mangrove, swamp forest, beach forest, and lowland forest) from sea level to 1000 m; at forest edge near agri- cultural areas and houses they seem to be less common and more shy. On Palawan, the species is under moderate hunting pres- sure for meat and the local pet trade, but appeared to have stable populations. In most areas, it was quite wary of humans, but in areas such as the PPSRNP (Site 15), the species did not associate humans with danger, and had become a regular thief of picnic baskets. It is listed by IUCN (2002) as Near-Threatened. Order Pholidota Family Manidae—Pangolins Manis culionensis.—This endemic spe- cies of the Palawan faunal region, with re- cords from Palawan and Culion Islands (Heaney et al. 1998), was formerly included within Manis javanica (Feiler 1998). It is known from primary and secondary low- land forest, possibly localized in distribu- tion (Allen 1910, Hoogstraal 1951, Sanborn 1952, Taylor 1934). We sighted several in lowland grassland/forest mosaic at Site 12. It is hunted for its skin, which is used to treat asthma. We have seen it for sale in Puerto Princesa and our guide at Site 11 said that it is hunted in logged-over lowland forest in that area. The species was de- scribed by local informants as fairly com- mon, but hunting pressure is moderately heavy. Manis javanica is listed by IUCN 291 (2002) as Near-Threatened; M. culionensis probably deserves the same status. Order Rodentia Family Sciuridae—Squirrels Hylopetes nigripes.—This large gliding squirrel is endemic to the Palawan faunal region; the number of museum specimens (Allen 1910, Sanborn 1952) suggests that it is common. Reis & Garong (2001) reported two specimens from sediments in a rock- shelter near Tabon Cave, Quezon Munici- pality dated to 11,130 BP. Taylor (1934) found the species in primary and secondary lowland forest where they nest in cavities in large trees. We observed an individual running up the side of a large hollow tree in primary forest at Site 1, and we frequent- ly heard and twice spotlighted them in ma- ture lowland forest at Site 15. We also heard the distinctive calls several times in selectively logged but largely intact forest near Barake, Aborlan Municipality, in the Victoria Range. According to local resi- dents, the species is common in mature for- est and is occasionally hunted as a source of food. IUCN (2002) lists this species as Near-Threatened; by current criteria, it should be listed as Data Deficient. Sundasciucus juvencus.—This tree squir- rel is endemic to central and northern Pa- lawan Island (Heaney et al. 1998). Hoogs- traal (1951) and Sanborn (1952) reported this species from primary and secondary lowland forest. We commonly observed this species in primary and secondary lowland forest at Sites 1, 2, 7, 11, 12, and in both secondary forest and grassland/degraded forest mosaic at Site 15. We also found it in a very small (<1 ha.) patch of secondary lowland forest surrounded by grassland and agricultural areas at Site 6. We observed the species on Dumaran Island, but not on Mal- inau or Rasa. The species is reportedly a common pest in coconut plantations. It is occasionally hunted as a source of food and for the local pet trade. It is listed by IUCN (2002) as Endangered, but this is strongly 292 contradicted by the available data, and we recommend de-listing. Sundasciurus rabori.—This_ poorly- known species, described from 5 specimens taken at 3600—4350 ft (ca. 1100—1300 m) on Mt. Mantalingajan, is endemic to Pala- wan Island (Heaney 1979). P. C. Gonzales deposited 2 specimens at the UMMZ that he collected on Mt. Gorangbato in Brooke’s Point Municipality in 1984; these are the only reported specimens aside from the original type series from Mt. Mantalingajan (Heaney 1979). Although we worked in some seemingly suitable habitats on Cleo- patra’s Needle (Site 3), we did not specifi- cally seek this species, and we never en- countered it. The IUCN (2002) lists S. ra- bori as Vulnerable, but based on current IUCN criteria, it should be considered Data Deficient. Sundasciunis steerii.—This species is en- demic to Balabac and southern Palawan Is- land (Heaney et al. 1998); Sanborn (1952) reported a large series from Brooke’s Point. Heaney et al. (1998) listed it as common in lowland forest and coconut and banana plantations. Because all of our study sites were in central and northern Palawan, we never encountered this species. It is listed by IUCN (2002) as Near-Threatened; since its habitat use is similar to the closely-re- lated S. juvencus, it is probably not threat- ened. Family Muridae—Mice Chiropodomys calamianensis.—This poorly known arboreal mouse is endemic to the Palawan faunal region; it is closely re- lated to species on the Sunda Shelf (Musser 1979). Reis & Garong (2001) reported a specimen from sediments in a rock-shelter near Tabon Cave, Quezon Municipality dat- ed to 11,130 BP. It is known from forest near sea level (Taylor 1934), coconut plan- tations, bamboo thickets, and buildings (Sanborn 1952); there are 12 specimens from Palawan in FMNH and NMP from the Hoogstraal expedition (Sanborn 1952). The PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON genus is apparently difficult to capture (Musser 1979); we never encountered this species. We recommend IUCN listing as Data Deficient. Haeromys pusillus.—This species is known only from Borneo, Palawan, and Ca- lauit Islands (Musser & Carleton 1993, Musser & Newcomb 1983). It is cited in Heaney et al. (1998) as ““Haeromys sp. A’ as potentially endemic to Palawan, but we follow Musser (pers. comm.) in treating it as conspecific with H. pusillus. We never encountered this species, but Musser & Carleton (1993) cited a specimen from Pa- lawan. A specimen of H. pusillus was taken in Sabah, Borneo, in a pit-fall trap near the edge of tall dipterocarp forest (Payne et al. 1985), and A. C. Alcala stated that he cap- tured the specimen from Calauit (in FMNH) by hand in a bamboo thicket (pers. comm.). IUCN (2002) listed this species as Vulnerable, but based on current criteria, it should be considered Data Deficient. Maxomys panglima.—This common rat is endemic to the Palawan faunal region; the genus is common on the Sunda Shelf, but is absent from oceanic portions of the Philippines (Musser et al. 1979). Sanborn (1952) reported large series from several lo- calities. We found it to be the most com- monly captured small mammal in agricul- tural/forest mosaic at Site 12 (62% of 169 captures), and was common to abundant in secondary forest (Site 11), primary lowland (Sites 1 and 2), and montane forest (Site 3) from near sea level to at least 1550 m. We captured a single juvenile in mossy forest at 1580 m at Site 3. Because we found this species to be common, although sometimes patchy, in all lowland and montane forested sites where we trapped extensively, and in mixed agricultural/second growth areas at Sites 11 and 12, we consider the IUCN (2002) listing as Near-Threatened to be un- justified. Specimens examined: 5, Site 1 (3), Site 3 (2). Mus musculus.—This introduced com- mensal has a nearly world-wide distribu- tion, although Southeast Asian populations VOLUME 117, NUMBER 3 are sometimes treated as a separate species, M. castaneus (Musser & Carleton 1993). It is common in human habitations in urban and rural areas (Heaney et al. 1998). We captured several in a residential area at Site 12, and it is most likely common in such places throughout Palawan. Palawanomys furvus.—This_ poorly known monotypic genus is endemic to Pa- lawan Island. It has been taken from a sin- gle locality on Mt. Mantalingajan and prob- ably occurs in high mountain forest (Mus- ser & Newcomb 1983). Our survey efforts on Cleopatra’s Needle (Site 3) failed to find this species; perhaps it is restricted to the more extensive mountain ranges of south- ern Palawan. The IUCN (2002) lists this species as Endangered; the lack of data and lack of damage to its presumed habitat (montane and mossy forest) suggest that it should be listed as Data Deficient. Rattus exulans.—This introduced com- mensal species is widespread from Bang- ladesh to Easter Island (Heaney et al. 1998). The first records from Palawan were named as a distinct species (luteiventris) by Allen (1910), but it is currently treated as a junior synonym of R. exulans (Musser & Carleton 1993). It is common in agricultural areas (Barbehenn et al. 1973, Rabor 1986) and sometimes present in disturbed forest and rare in primary forest (Barbehenn et al. 1973; Heaney et al. 1991, 1998). We found this species in grassland (Site 6), agricul- tural areas (Sites 6, 11, 12, and 14), and in secondary lowland forest (Site 7). The spe- cies appears to be absent from primary (e.g., Sites 1, 2, and 3) and logged-over for- est (e.g., Site 11) on Palawan. Specimens examined: 4, Site 6 (3), Site 11 (1). Rattus tanezumi.—This introduced com- mensal, formerly included within Rattus rattus (Musser & Carleton 1993), is wide- spread from Afghanistan to New Guinea and Micronesia (Heaney et al. 1998). It is often abundant in urban and agricultural ar- eas and common in disturbed forest up to 1800 m (Danielsen et al. 1994; Heaney et al. 1989, 1999; Rabor 1986; Sanborn 1952). 293 Hoogstraal (1951) and Sanborn (1952) found them to be common on Palawan in some agricultural and residential areas. We captured three at Site 13, and three in a res- idential area in Puerto Princesa; vouchers were deposited in the NMP and the collec- tion of the Palawan Council for Sustainable Development. Rattus tiomanicus.—This indigenous rat is found on the Malay Peninsula and the islands of the Sunda Shelf, including Pala- wan (Heaney et al. 1998). Payne et al. (1985) reported the species from secondary forest, agricultural areas and gardens, scrub, and grassland. We captured this species in grassland/forest mosaic at Site 12 (9% of captures), two in selectively logged forest at Site 13, one in a ricefield at Site 14, and three individuals from mossy forest and the transition zone between mossy and montane forest at Site 3. At Site 3, two individuals were taken at ca. 1580 m during the night and one at ca. 1540 m during the day. Spec- imens examined: 3, Site 3 (3). Sundamys muelleri.—This moderately large rat is found from southern Myanmar to the Sunda Shelf, including Palawan (Heaney et al. 1998); the genus is absent from the oceanic Philippines. Sanborn (1952) described a subspecies endemic to Palawan, S. m. balabagensis, from a single specimen taken at 3000 ft (ca. 900 m) “‘in thick forest near the top of Mt. Balabag”’; two additional specimens in the USNM are from Pinigisan, on the lower slopes of Mt. Mantalingajan at 2100—2500 ft (ca. 640— 760 m). Additional specimens from the Pa- lawan region are from Culion Island (San- born 1952), Balabac, and Busuanga (USNM; Heaney et al. 1998). On Borneo, the species occurs in forest, often near streams (Payne et al. 1985) at elevations usually below 3500 ft (ca. 1070 m; Med- way 1977). Md. Nor (1995) caught the spe- cies on Banggi, Balambangan, and Mol- leangan Islands “‘mostly in primary forest on low ground and near streams’’. We cap- tured one individual from a riparian zone in primary forest at ca. 700 m at Site 2, and 294 two in lowland grassland/forest mosaic at Site 12. Specimen examined: 1, Site 2 (1). Family Hystricidae—Porcupines Hystrix pumila.—The only porcupine found in the Philippines is endemic to the Palawan faunal region; other species occur widely on the Sunda Shelf and continental Asia. Reis & Garong (2001) reported 3 specimens from sediments in a rock-shelter near Tabon Cave, Quezon Municipality dat- ed to 11,130 BP. It is known to occur in secondary and primary lowland forest and to den in abandoned mine shafts (Hoogs- traal 1951, Sanborn 1952). We observed this species at dusk along the edge of sec- ondary forest at Site 11, once at night at Site 15 (where it was feeding on fruit of Terminalia catappa), and several times at night in grassland/forest mosaic at Site 12. Local guides reported them at Site 14, at the Iwahig Penal Colony in Puerto Princesa, in Rizal Municipality, and in Dumaran Mu- nicipality (on the mainland). This was re- ported as the most important game species for the Tagbanua ethnic community in Bar- ake, Aborlan Municipality (Lacema & Wid- mann 1999); they are often dug out of their subterranean dens. Not listed by IUCN (2002) but listing as Data Deficient or Near- Threatened seems justified. Order Carnivora Family Felidae—Cats Prionailurus bengalensis.—This small cat is widespread from Siberia to Pakistan and Bali, with reports from the Philippines on Busuanga, Cebu, Negros, Palawan, and Panay Islands only (Heaney et al. 1998, Taylor 1934). The population from the Pa- lawan faunal region was described recently as a distinct subspecies, P. b. heaneyi, by Groves (1997); it is well represented by museum specimens (Allen 1910, Sanborn 1952). Rabor (1986) reported the species from agricultural areas and forest from sea level to ca. 1500 m. We spotlighted one PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON along a river trail in Barake, Aborlan Mu- nicipality. Family Mustelidae—Weasels, Otters, and Badgers Amblonyx cinereus.—This otter occurs from India to Taiwan and the Sunda Shelf (Heaney et al. 1998). On Palawan, it is found in coastal rivers and bays (Hoogstraal 1951, Rabor 1986, Sanborn 1952). Payne et al. (1985) and Sanborn (1952) reported the species feeds on crustaceans, mollusks, and fish where there is permanent water and some tree cover. Rangers at PPSRNP re- ported to Heaney and Widmann that otters frequently visit along the beach and small streams, and local people reported them from the Iwahig River (Puerto Princesa), Aborlan River (Aborlan Municipality), Malatgao and Taritien Rivers (Narra Mu- nicipality), and adjacent mangrove and freshwater swamp forest. We received one report of an otter raiding a prawn pond. IUCN (2002) lists this species as Near- Threatened. Mydaus marchei.—This badger is en- demic to the Palawan faunal region; it is related to a species that occurs on the Sunda Shelf. It has been documented in mixed grassland and secondary forest (Hoogstraal 1951, Kruuk 2000, Rabor 1986, Taylor 1934), and Sanborn (1952) reported series from several localities. We occasionally smelled its strong odor in areas of mixed agriculture and secondary forest throughout Palawan; we sighted it often in residential and cultivated areas, grassland, and grass- land/forest mosaic at Site 12, and rarely in ricefields and freshwater swamp forest at Site 14. One individual living in a den on campus at the State Polytechnic College was easily followed and observed. Because it is widespread and moderately common on Palawan, and is rarely hunted (Grim- wood 1976, Kruuk 2000), we agree with Kruuk (2000) that the IUCN listing of this species as Vulnerable is not justified. VOLUME 117, NUMBER 3 Family Herpestidae—Mongooses Herpestes brachyurus.—The only mon- goose found in the Philippines is distributed from the Malaysian Peninsula to Borneo and Palawan (Heaney et al. 1998). The Pa- lawan population was named as a distinct species (H. palawanus Allen 1910), but currently is treated as a subspecies (Corbet & Hill 1992). Allen (1910) described them based on one specimen from Iwahig; San- born (1952) reported one specimen from Puerto Princesa and one from Brooke’s Point, and Rabor (1986) found the species most often near rivers. On Borneo, Payne et al. (1985) found the species to occur in primary and secondary lowland forest, plantations, and gardens. We never encoun- tered this species; but we received reports of them at Site 14. Family Viverridae—Civets Arctictis binturong.—The binturong is known from northern Myanmar to the Sun- da Shelf (Heaney et al. 1998). On Borneo, the species is arboreal and terrestrial, most- ly nocturnal, and occurs in old-growth and secondary forests, sometimes entering ag- ricultural areas near forest (Payne et al. 1985). The Palawan population, which ini- tially was named as a distinct species (A. whitei Allen 1910) from four specimens, is still represented by few specimens (Heaney et al. 1998). Rabor (1986) reported obser- vations from primary and secondary low- land forest up to 200 m. Our guide at Site 11 reported that a juvenile repeatedly en- tered remnant trees that were fruiting in a clearing surrounded by secondary forest. At Site 2, we observed A. binturong drinking water from a stream at ca. 400 m during mid-day. We spotlighted one in a fruiting Ficus tree at Site 15, and twice saw one in grassland/forest mosaic at Site 12 feeding on fruits of Guioa pleuropteris. Local peo- ple reported hunting them for food, and also catching them and selling them as pets. The IUCN (2002) listing of A. binturong whitei as Vulnerable seems justified. 295 Paradoxurus hermaphroditus.—This common species is found from Sri Lanka to the Lesser Sunda Islands and the Phil- ippines (Heaney et al. 1998). Recorded in agricultural areas and forest over a wide elevational range (Allen 1910; Heaney et al. 1991, 1999; Hoogstraal 1951; Rabor 1986); Sanborn (1952) reported large series from several localities. We often saw them feed- ing in fruiting trees and shrubs in grassland/ forest mosaic at Site 12, and we saw road- kills along the coastal highway. They are hunted, but the large number of museum specimens and sightings indicate that they traditionally have been and probably remain the most common carnivore on Palawan (e.g., Allen 1910, Sanborn 1952). Viverra tangalunga.—tThis civet is found from the Malay Peninsula to Sulawesi and the Philippines (Heaney et al. 1998). Known from primary and secondary low- land, montane, and mossy forest (Allen 1910, Heaney et al. 1999, Rickart et al. 1993). We captured and released a juvenile of this species in a cage trap in lowland primary forest at Site 1, and we observed two in forest-grassland mosaic at Site 12. Order Artiodactyla Tragulus napu and Axis calamianensis both occur in the Palawan faunal region, but we found no evidence of either species on Palawan Island. Family Suidae—Pigs Sus barbatus.—The bearded pig is found from the Malay Peninsula to Borneo and Palawan (Heaney et al. 1998). Rabor (1986) and Payne et al. (1985) reported the species from primary and secondary forest from sea level to the highest peaks; Sanborn (1952) reported a series from Iwahig. Groves (2001) has tentatively suggested that the population of this species from the Palawan faunal region, which has been recognized as a distinct subspecies, may warrant rec- ognition as a distinct species, Sus ahoeno- barbus. We regularly observed this species 296 or evidence of its occurrence in forest hab- itats (including fragmented forest) from sea level to montane forest at ca. 1500 m (Sites 1, 2, 3, 4, 7, and 11). We also observed evidence of the species entering cultivated areas near forest and damaging crops. Wild pigs are heavily hunted on Palawan with snares, low caliber rifles, and small, baited explosive devices known as “pig bombs”. The species appears to be locally common, but is in decline due to heavy hunting pres- sure (Caldecott et al. 1993, Oliver 1992). The IUCN (2002) lists S. barbatus ahoe- nobarbus as Vulnerable. Discussion Adequacy of sampling.—Small fruit bats on Palawan (Cynopterus, Eonycteris, Ma- croglossus, and Rousettus) appear to have been fairly completely sampled; no species have been added in over 50 years (exclud- ing the apparently erroneous reports of Haplonycteris fischeri and Ptenochirus mi- nor), despite extensive netting. It is inter- esting that our mist netting in primary for- est produced very few captures; for exam- ple, in primary lowland forest at 150 m el- evation (Site 1), we captured | fruit bat in 42 net-nights; in primary lowland forest at ca. 500 m (Site 2), we captured 1 fruit bat in 56 net-nights, and in primary montane forest at ca. 1400 m (Site 3), we captured no fruit bats in 48 net-nights. Although our sample size is limited, all of these values fall well below what would be typical on islands in the oceanic portion of the Phil- ippines (Heaney et al. 1989, 1999), sug- gesting that small fruit bats are not as abun- dant in primary forest on Palawan (e.g., Heideman & Heaney 1989, Heaney et al. 1989). Indeed, all of the small fruit bats currently known from Palawan predomi- nately occur in disturbed habitats, in con- trast to the oceanic Philippines, where sev- eral endemic genera (Alionycteris, Haplon- ycteris, Otopteropus, and Ptenochirus) are most common in old-growth forest. The ecology of large fruit bats (Acerodon PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Pteropus) has been very poorly studied on Palawan and elsewhere in the Philip- pines. This is the direct result of the diffi- culty in capturing these species by any means other than shooting. Despite the pau- city of ecological information on these spe- cies, their distribution among major island groups appears to be moderately well un- derstood because their activities and roosts are highly conspicuous, and no additional species have been found on Palawan in over 50 years. It seems unlikely that additional species will be found on Palawan, with the possible exception of P. hypomelanus. Insectivorous bats are clearly the least known of all Palawan mammals. Distribu- tions remain poorly documented, ecological information is scanty for most species, and the taxonomy is often uncertain. Our survey efforts and examination of previously col- lected insectivorous bats documented eight species (Rhinolophus arcuatus, R. ct. bor- neensis, R. creaghi, R. macrotis, Miniopte- rus australis, M. schreibersi, M. tristis, and Murina cf. tubinaris) on Palawan for the first time, and several more are noted in the text as very likely to be present. Our knowledge of small non-volant mammals (including Soricidae, Tupaiidae, Sciuridae, and Muridae) on Palawan is un- even. Some lowland species (e.g., Tupaia palawanensis, Sundasciurus juvencus, Maxomys panglima, Rattus tiomanicus, Sundamys muelleri, and the non-native mu- rines) are common and well known. Very little is known of several other species (e.g., Crocidura palawanensis, Crocidura sp., Sundasciurus rabori, Chiropodomys cala- mianensis, Haeromys pusillus, and Pala- wanomys furvus). Perhaps we failed to lo- cate these poorly known species because we did little trapping in trees or other places above the ground surface (C. calamianensis and H. pusillus), sampled only one site above 1000 m (S. rabori and P. furvus), and our trapping techniques were limited, i.e., we did not use pitfall traps (Crocidura spp.). The presence of so many poorly known species suggests that other species VOLUME 117, NUMBER 3 may await discovery, especially in high mountain habitats and high in the canopy. Musser & Newcomb (1983) suggested that unknown species are yet to be discovered on Palawan, citing the report of Hoogstraal (1951), which narrated their attempt to cap- ture a “very large rat with a white tail” described to them by native Palaw’an. Oth- er large islands in the Philippines have been shown to support diverse communities of endemic small mammals which are restrict- ed to montane areas (e.g., Heaney 2001, Heaney & Rickart 1990, Rickart 1993), and perhaps the same awaits discovery on Pa- lawan. Medium to large mammals (Cercopithe- cidae, Manidae, Hystricidae, the carnivores, and Suidae) are possibly the most thor- oughly inventoried subset of Palawan’s mammalian fauna. Because of their large size, they are easily observed compared to other mammals. Many of these species are also commonly hunted, so obtaining speci- mens is often easier than for non-game spe- cies. It is unlikely that other medium to large mammals await discovery on Pala- wan, though the ecology of all requires much additional study. Biogeography.—As noted above, the Pa- lawan faunal region is part of the Sunda Shelf and may have been connected to mainland Asia via Borneo (Everett 1889) during a Pleistocene episode of glacially- induced sea-level lowering (Heaney 1985, 1986, 1991a), though current data leave this uncertain. All other islands in the Philip- pines are oceanic and have probably never had a dry-land connection to any mainland area (Heaney 1985, 1986, 2001). Of the 58 native species currently known from Pala- wan Island (tentatively including the small Crocidura sp.), 13 species arc endemic to Palawan (and usually to some of the smaller islands that were included in Pleistocene Greater Palawan; Heaney 1986); 12 of these are non-volant, all of which have their closest relatives on the Sunda Shelf. Eight of Palawan’s 11 native rodents (73%) are endemic; all three non-endemics are mu- 297 rids. Only one endemic species is a bat (Ac- erodon leucotis), and only it has its closest relatives in the oceanic Philippines. This pattern of endemism is clearly consistent with the geological history of the Philip- pines and also highlights the importance of the greater vagility of bats over non-flying mammals. Of the 28 insectivorous bats, 18 species are somewhat to highly widespread in Indo-Australia (and some beyond), 2 are shared only with the Sunda Shelf and In- dochina (Rhinolophus acuminatus and Rhinolophus cf. borneensis), 1 with the Sunda Shelf only (Cheiromeles torquatus), 3 occur on the Sunda Shelf and the oceanic Philippines (Kerivoula pellucida, K. white- headi, Myotis macrotarsus), 1 occurs on Palawan, Sulawesi and the oceanic Philip- pines (Mops sarasinorum), 2 occur only on Palawan and in the oceanic Philippines (Rhinolophus virgo and Myotis rufopictus), and one occurs on Borneo, Sulawesi, and throughout the Philippines (Emballonura alecto). These data again demonstrate that the bats are more widely distributed and do not clearly reflect the geological history, as do the non-flying mammal species, which in the Philippines are usually restricted to a single area that was united by dry land dur- ing the late Pleistocene. From the highly diverse fruit bat fauna of Borneo (17 spe- cies; Payne et al. 1985), only five species extend to the northern continental land- bridge islands of Sabah (Md. Nor 1995). These are the same five non-endemic spe- cies that can be found just to the north on Palawan Island. We note that the combined totals of na- tive non-volant mammal species on Pala- wan that either are shared with Borneo and other portions of the Sunda Shelf or that are endemic to Palawan and have their closest relatives on Borneo or adjacent areas is 22 out of 24 (92%). The apparent exceptions are Hylopetes nigripes (related to H. albon- iger of Indochina) and Palawanomys furvus (an enigmatic genus of unclear phylogenet- ic position). If this analysis were extended to the entire Palawan faunal region, Axis 298 calamianensis (related to A. porcinus in In- dochina) would also be included here. Four species are widespread in Southeast Asia (including parts of Wallacea), and no spe- cies is shared with the oceanic Philippines except for those 4 species (Macaca fasci- cularis, Prionailurus bengalensis, Paradox- urus hermaphroditus, and Viverra tanga- lunga). If Sus barbatus, which occurs on Palawan, the Sunda Shelf, and in the mar- ginal islands of the Sulu Archipelago is counted as a widespread Southeast Asian species, the total rises to five species. These data, in sum, strongly reinforce the conclu- sion of Everett (1889), based on his anal- ysis of bathymetric features of the ocean floor and the pattern of relationships of the 18 species then known from Palawan and associated smaller islands, that the Palawan faunal region is an extension of the Sunda Shelf, probably due to a fairly recent dry- land connection, with only a small portion of its fauna shared with the oceanic Phil- ippines. Patterns of species richness relative to is- land area are also of interest. Insectivorous bats are so poorly known in the Philippines that it is possible only to say that the 28 species documented here compare favor- ably with most islands in the Philippines; that is, there is no evidence that Palawan is species-poor (Heaney et al. 2002). Fruit bats, on the other hand, are represented on Palawan by only 6 species, and this places their diversity far below that on much smaller islands in the oceanic Philippines; Maripipi, for example, has 10 species, but is only 22 km/?. It seems certain that Pala- wan has a depauperate fruit bat fauna, as well as probably having lower fruit bat den- sity, as noted above, compared to the oce- anic Philippines (Heaney 1991b). Species richness of non-flying mammals, on the other hand, at 24 is much above the species/ area curve documented in the oceanic Phil- ippines, though well below that of islands on the primary portion of the Sunda Shelf (Heaney 1984, 1986; Heaney et al. 2002). It is interesting that carnivores are notably PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON more diverse on Palawan than in the oce- anic Philippines, and murid rodents notably less diverse, for an island the size of Pala- wan. Conservation issues.—The most pressing issue facing terrestrial wildlife in Palawan is the rapid loss of forest cover, especially in the primary lowland forests that are tar- geted for logging. Palawan’s forests are less commercially valuable than the diptero- carp-dominated forests of the other islands, and, consequently, deforestation occurred later on Palawan. However, once forests were exhausted on Luzon, Mindanao, Ne- gros, etc., commercial logging operations began working in lowland forest on Pala- wan (Environmental Science for Social Change 1999) during the 1970’s and 1980’s at unsustainable levels (Quinnell & Balm- ford 1988, Kummer 1992). Since a logging ban was imposed in the early 1990s throughout the Province of Palawan, log- ging has declined, but the large commercial operations appear to have been replaced by small-scale, illegal commercial logging. We have seen that forest continues to disappear from the most accessible areas, and forest edges are gradually creeping higher and higher up the contours, in a manner similar to that experienced on Leyte (Rickart et al. 1993) and southern Luzon (Heaney et al. 1999). Lowland primary forest has been eliminated from many parts of Palawan and the destruction shows few signs of easing. Due to almost complete conversion of the coastal plain into ricefields, coconut, or oth- er plantations, distinctive ecosystems such as freshwater swamp forest and beach forest have virtually disappeared (Widmann 1998). Slash and burn agricultural practices have also been very damaging to forests, and Palawan has experienced a population explosion due to high birth and immigration rates. It should be noted that caves are crucial to maintaining the fauna, since approxi- mately 18 (32%) of Palawan’s mammals are bats that roost in caves. Caves have been the focus of much destruction in the Phil- VOLUME 117, NUMBER 3 ippines; common activities in caves on Pa- lawan include guano mining, general van- dalism, recreational exploration, and trea- sure hunting. Unlike many other parts of the Philippines (e.g., Heaney et al. 1991, 1999: Rickart et al. 1993), we never en- countered any evidence of cave-roosting bats being hunted on Palawan. We found guano mining to be common, usually near the mouth of caves. Recreational explora- tion of caves is steadily increasing; many caves are currently being developed or ad- vertised for this purpose, while many more proposals are in the planning stages. At PPSRNP (Site 15), hundreds of visitors may enter a one kilometer stretch of the cave daily. The bats are clearly disturbed by the activity, but the ultimate result of such disturbance is unknown. Many medium to large sized mammals are under significant hunting pressure on Palawan for their meat, the live animal trade, and medicinal use, as noted above, but few data are available on the impact. The recent and on-going shift from subsis- tence to market economies among members of the Tagbanua and other ethnic groups may contribute to the decline of some spe- cies (Lacerna & Widmann 1999), such as Sus barbatus, Pteropus vampyrus, and Hys- trix pumila for meat, Macaca fascicularis and Arctictis binturong as pets, and Manis culionensis for traditional Chinese medi- cine. Acknowledgments Funding for these studies was provided by the Palawan Council for Sustainable De- velopment, United States Peace Corps, Philippine Cockatoo Conservation Program through the Loro Parque Fundacion, Ten- eriffe, Spain, and the Barbara Brown and Ellen Thorne Smith Funds of the Field Mu- seum. Jun Saldajeno, Apollo Regalo, Ben- igno Maca, Erlito Porka, and Manual Lar- dizabal made significant contributions to field work, often under difficult living con- ditions. Adelwisa Sandalo, Lualhati Tabu- 299 gon, Ariel Carino, Leilani Berino, Dr. Ter- esita Salva, Dr. Edgardo Castillo, Dr. Pa- cencia Milan, Dr. Josef Margraf, Indira Lac- erna-Widmann, Siegfred Diaz, Deborah Villafuerte, and the wildlife wardens of Rasa Island provided valuable logistical and administrative support. JAE thanks the Ta- bugon family for their extraordinary hos- pitality during his stay on Palawan. We thank A. C. Alcala, P C. Gonzales, and P. O. Glass for depositing important speci- mens at UMMZ, and P. Myers for access to those specimens. H. Kafka, J. Mead, and R. Thorington provided access to specimens at the USNM, and E Dieterlen kindly loaned specimens from SMNH. We are grateful to P. O. Glass for additional details about spec- imens he collected. Ding Padilla, Clara Simpson, and Lisa Kanellos produced the map in Fig. 1. Eric Rickart gave sound ad- vice during the early stages of the project and Danilo Balete helped with the transport and identification of voucher specimens; Eric Rickart and Robert Timm provided constructive reviews of the manuscript. The Department of Environmental and Natural Resources, through the Protected Areas and Wildlife Bureau and the Provincial Envi- ronment and Natural Resources Office, pro- vided permits and encouragement. LRH thanks the Geological Sciences Department, Northwestern University, for use of office space during a sabbatical leave. Literature Cited Allen, G. M. 1922. Bats from Palawan, Philippine Is- lands.—Occasional Papers of the Museum of Zoology, University of Michigan 110:1—5. Allen, J. A. 1910. Mammals from Palawan Island, Philippine Islands.—Bulletin of the American Museum of Natural History 28:13-17. Balete, D. S., L. R. Heaney, & R. I. Crombie. 1985. 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Bishop Museum, Honolulu, Hawaii 96817, U.S.A., e-mail: (FK) fkraus@hawaii.edu, (AA) allison@hawaii.edu Abstract.—We describe a new species of natricine snake of the genus T7ro- pidonophis from the D’Entrecasteaux Islands, off the southeastern peninsula of New Guinea. The new species is large, with 15 unreduced scale rows, a high ventral and low subcaudal scale count, and a distinctive color pattern of dark mid-dorsal bands and offset lateral blotches on a yellow or brown ground color. The species is known from two specimens collected at 900—1090 m in primary lowland hill forest. Close relationships with other members of the genus are not apparent. The natricine genus Tropidonophis con- sists of 18 species of small to medium-sized snakes distributed from the Philippines (two endemic species) to the Bismarck Archi- pelago (two endemic species), with 12 spe- cies found on New Guinea and its offshore islands, four in the Moluccas, and one in Australia (Malnate and Underwood 1988). The genus is thought to be most closely re- lated to the Southeast Asian Xenochrophis on the basis of shared scalation, hemipenial, and osteological features (Malnate and Un- derwood 1988). Most Tropidonophis spe- cies are nondescript, frequently with a uni- formly dark dorsal ground color of brown or gray, sometimes with darker spots, short lines, or narrow bands. Some specimens of a few species have more conspicuous pat- terns of dark bands on a lighter ground col- or (cf. O'Shea 1996: 95). Tropidonophis species typically inhabit rainforest, occur from sea level to 2200 m (Malnate and Un- derwood 1988), and are reported to dwell frequently near permanent water sources (O’Shea 1996). During the course of conducting biolog- ical surveys in the D’Entrecasteaux Islands in 2002 we collected a strikingly colored specimen of Tropidonophis that is unas- signable to any currently recognized spe- cies. A search of museum collections re- vealed another specimen belonging to the same taxon. We take this opportunity to provide this species with a name. Materials and Methods Specimens were collected under appli- cable national and provincial permits, fixed in 10% buffered formalin, and transferred to 70% ethanol for storage. Measurements were made to the nearest mm in the field with a fiberglass tape; mass was measured to the nearest gram in the field with a Pe- sola scale. Diagnostic features and compar- isons to other species were based on data provided in the comprehensive study of Tropidonophis by Malnate and Underwood (1988) and by reference to specimens housed in the Bernice P. Bishop Museum, Honolulu (BPBM). Specimens are deposited in the BPBM and American Museum of Natural History (AMNH). Tropidonophis dolasii, new species Figs. 1, 2 Holotype.—BPBM 16539 (field tag FK 6118), adult female, collected by D. Sale- puna on E slope of Oya Tabu (Mt. Kilker- 304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. bar equals one cm. ran), 9.4555°S, 150.7857°E, 1090 m, Fer- gusson Island, Milne Bay Province, Papua New Guinea, on 23 August 2002. Paratype.—AMNH 73979, adult female, collected by L. Brass on E slope of Good- enough Island, 900 m, Milne Bay Province, Papua New Guinea, on 27 October 1953. Diagnosis.—A large species of Tropi- donophis with 15 dorsal scale rows at mid- body and one head length anterior to the vent, 161—162 ventrals, 63 subcaudals, 2 preoculars, 3 or 4 postoculars, 8 supralabi- als, 8 or 9 infralabials, no postocular dark stripe, and yellow or brown ground color (A) Lateral, and (B) dorsal view of head of holotype (BPBM 16539) of Tropidonophis dolasti. Scale with vaguely defined mid-dorsal black bands offset by lateral black blotches on scale rows 1—4. These dark bands and blotches are not solid, rather they are formed by a network of darkened scale margins. Description of holotype.—Adult female. Dorsal scale rows 15 (reduction to 15 rows occurs at the level of the 15" ventral); all rows except first keeled; first row weakly keeled on those scales posterior to approx- imately 15 ventrals anterior to vent; keels on dorsal scales more weakly developed an- teriorly and laterally and more strongly de- VOLUME 117, NUMBER 3 veloped posteriorly and dorsally; paired apical pits obvious on those dorsal scales retaining the horny epidermal layer; all dor- sal scales, except on first row, notched at the posterior tip. Rostral twice as wide as high; internasals longer than wide; prefron- tals wider than long, as are frontal, supra- oculars, and parietals; lateral extension of parietal contacts middle postocular, exclud- ing upper postocular from contact with an- terior temporals on each side. Nasals divid- ed by large nares; loreal higher than long; preoculars 2; postoculars 3 (right) and 4 (left); anterior temporals 2, upper a narrow sliver approximately 20% the size of lower, lower excluded from contact with posto- culars on right side, with point contact to second and third postoculars on left; pos- terior temporals 3, the most anterior lying on posterior slope of lower anterior tem- poral and in contact posteriorly with only the middle posterior temporal (Fig. 1). Su- pralabials 8, 4 and 5 contact eye; infra- labials 9 (right) and 8 (left), four contact anterior chin shields. Posterior chinshields separated along their entire length by 1 + 1 + 2 intergenials; lateral gulars separated from posterior chinshields. Pits present in the loreal, preoculars, postoculars, anterior temporals, posterior temporals, parietals, and supralabials; absent from the rostral, nasals, internasals, prefrontals, frontal, su- praoculars, infralabials, and chin shields; many small tubercles present on all head shields. Ventrals 161; anal divided; subcaudals 63, excluding tip; subcaudal pits unobserv- able because horny epidermal layer missing for all subcaudals; subcaudals/(ventrals + subcaudals) = 0.28. Dorsal scales on tail reduced to six rows at level of subcaudal 18, reduced to four rows at level of sub- caudal 41, and reduced to two rows at level of subcaudal 61. Total length 1145 mm; snout-vent length 905 mm; tail length 240 mm (21% of total length); mass 285 g. Maxillary teeth on left side 29, the last three enlarged. 305 Dorsal ground color in preservative yel- low, varying from deep orange-yellow an- teriorly to pale straw-yellow posteriorly. In- terstitial skin bright orange anteriorly, be- coming gray posteriorly. Head mustard yel- low, darker than neck and anterior body, without dark postocular stripe; supralabials and infralabials with black posterior mar- gins (Fig. 1A). Dorsum bears pattern of ~ 48 bands; each band spans middle 7—9 scale rows and is 1-2 scales long; mid-dorsal bands staggered against equal number of lateral blotches, each 3—4 scales high and 1—2 scales long. All bands and blotches formed by black outlining of affected scales; scale centers (and usually the pos- terior margins) retain ground color, impart- ing vague and indefinite appearance to bands and blotches (Fig. 2). First (reduced) dorsal band appears at level of ventral 15, first nearly complete dorsal band at level of ventral 28, and first trace of lateral blotch at level of ventral 30. Series of black dashes on first dorsal scale row up to level of ven- tral 25, each dash extending for 1—5 scales (Figs. 1A, 2). Tail with ~ 20 dark bands; bands increasingly reduced and poorly de- fined posteriorly. Venter yellow, fading from deep orange-yellow on chin to light straw yellow on tail, with gray flecks lat- erally at origin of dorsal banding and grad- ually filling in mid-ventrally; the last third of venter evenly, though not heavily, freck- led with gray. Color in life (from field notes).— “‘Dor- sum mustard yellow with vague mid-dorsal and lateral blotches created by black outlin- ing along the margins of affected scales. Dorsum becoming more orange anteriorly and top of head orange-brown. Sides turn- ing to yellow. Venter bright yellow with a tendency to orange-yellow on chin and throat. Black flecks scattered on venter be- ginning ca. 4-way down body and increas- ing in frequency posteriorly.” Variation.—The paratype is smaller (to- tal length ~810 mm, snout—vent length ~756 mm, tail length 54+ mm) than the holotype, has a broken tail, and is eviscer- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Holotype (BPBM 16539) of Tropidonophis dolasii in life. ated anteriorly. It differs from the holotype in having prefrontals longer than wide; postoculars 4 on right side, 3 on left; an- terior temporals two on each side, the upper (anteriormost) % as large as the lower; post- erior temporals two on each side; four in- fralabials in contact with anterior chin shields on left side, five on right; posterior chinshields meeting anteriorly and separat- ed posteriorly by 1 + 2 intergenials; pits on head scales not observable because of loss of horny epidermal layers; ventrals 162; subcaudals 15 before tail broken; maxillary teeth on left side 32, the last 4 enlarged. Dorsal ground color brown, no darker posteriorly than anteriorly; dorsum with ~51 bands, all bands and lateral blotches formed by dark brown, not black, margin- ing and more solidly filled in than for ho- lotype. Venter pale yellow anteriorly, changing to brown posteriorly. Barring on lips dark brown, less distinct than in holo- type due to general suffusion of brown pig- ment on the head. Comparisons to other species.— Tropi- donophis dolasii is distinguished from T. negrosensis in lacking a posterior reduction in dorsal scale rows; from T. dahlii, T. den- drophiops, T. doriae, and T. hypomelas in having 15 (vs. 17) dorsal scale rows; from T. mairii in having 2 (vs. 1) preocular; from T. truncatus in having 3 or 4 (vs. 2, rarely 3) postoculars; from T. halmahericus, T. mcdowelli, and T. punctiventris in having 8 (vs. 9) supralabials; from 7. aenigmaticus, T. novaeguineae, and T. picturatus in hay- ing a larger number of ventral scales (161— 162 vs. 140-152, 128-143, and 117—140, respectively); from 7. elongatus, T. mon- tanus, T. multiscutellatus, and T. parkeri in having fewer subcaudal scales (63 vs. 85— 108, 71-89, 74-103, and 80-100, respec- VOLUME 117, NUMBER 3 tively); and from T. statisticus in its larger size (~810—1145 mm vs. maximum of 870 mm), dorsal pattern of dark bands, offset with lateral blotches, on a yellow or brown ground (vs. uniform gray or brown with series of dorsal spots), and strongly barred labials (vs. unbarred). Only five other species of Tropidonophis attain a size greater than one meter: 7. dah- lii, T. doriae, T. elongatus, T. halmahericus, and T. montanus. The first is restricted to New Britain and the last three to the west- ern half of New Guinea or the Moluccas. Only 7. doriae approaches the geographic range of 7. dolasii, being found on the ad- jacent mainland of Milne Bay Province (Malnate and Underwood, 1988; O’Shea, 1996), but this species has 17 dorsal scale rows, no more than 153 ventrals in females, and no fewer than 71 subcaudals in fe- males. The conspicuous yellow dorsal and ven- tral coloring of 7. dolasii (brown dorsally in the long-preserved paratype) and its pat- tern of lateral blotches combined with mid- dorsal bands are apparently unique among Tropidonophis. In other dorsally banded Papuan species (e.g., 7. doriae, T. hypo- melas), the bands are typically solid, in- stead of being formed by a network of dark- ened scale margins, and extend across the entire dorsum, instead of lying just on the mid-dorsal scale rows. Ecological notes.—The holotype was collected in small-crowned lowland hill for- est (Paijymans, 1975, 1976) on steep terrain at 1090 m. The collection site faces east but receives little direct sunlight because sur- rounding ridges and frequent clouds block the sun throughout much of the day. Near- est water source was a small (~30-cm wide) trickle among rocks in a narrow ra- vine approximately 50—100 m elevation be- low the collecting site. At the time of col- lection, the region had been in a month- long drought, although moisture was still present under logs and some rocks. Tem- perature varied from 15.8—21.0°C during the two weeks of our stay. The specimen 307 was active in mid-morning and attempted to escape. Other snakes occurring in the same area were Aspidomorphus lineaticol- lis, Boiga irregularis, and Tropidonophis aenigmaticus. The paratype was noted to come from “transition oak-rain forest”’ and had an un- identified Rana in its stomach. Etymology.—The name is a patronym honoring Dolasi Salepuna of Ulua, Fergus- son Island, who was an invaluable field as- sistant and captured the holotype. Distribution.—The species is known only from the uplands of eastern Fergusson Island and eastern Goodenough Island (Fig. 3). It likely occurs throughout the higher elevations of the D’Entrecasteaux Islands. Remarks In their revision of Tropidonophis, Mal- nate and Underwood (1988) placed consid- erable importance in their key on the num- bers of anterior and posterior temporal scales, even while documenting that these characters show considerable intraspecific variation. We have not emphasized these scales for diagnosing 7. dolasii because we are uncertain of their modal distribution, given our few specimens and the consid- erable variation these characters exhibit in the genus. In considering the holotype, it is especially uncertain |) whether the third, small scale in the anterior-posterior series should properly be considered an anterior or posterior temporal, and 2) whether the first and the third small scales in the series are normally present or are aberrant divi- sions unique to that specimen. We have re- ferred to the third, small scale as a posterior temporal based on the definition provided by Malnate and Underwood (1988: 75) that those scales meeting either the posterior slope of the supralabial apex or the anterior temporals constitute the posterior tempo- rals. However, comparison with the para- type, whose temporals appear more normal in size and placement than those of the ho- lotype, shows the region occupied by this PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Elevations above 1000 m are not demarcated. = a Magnetic Nonth D’Entrecasteaux | Goodenough Islands Tis | a %o 8 Fergusson a Normanby Coral Sea Fig. 3. Milne Bay Map of southeastern New Guinea showing type locality for Tropidonophis dolasii (star) on Oya Tabu (Mt. Kilkerran), Fergusson Island, and approximate locality for paratype (dot) on Goodenough Island, D’Entrecasteaux Islands. small scale in the holotype to be part of the parietal in the paratype, suggesting this scale is not homologous with the other tem- poral scales. Further comparison of the two specimens shows the small anterior tem- poral of the holotype to be of similar place- ment but much smaller size than the cor- responding scale in the paratype (~20% the size of the larger anterior temporal in the holotype vs. ~66% in the paratype). Given these observations, it seems likely that the temporal scalation seen in the holotype is aberrant. If one assumes that the temporal scala- tion seen in the paratype is normal for the species, then, among New Guinean Tropi- donophis, having two anterior temporals would serve as a further character helping to distinguish 7. dolasii from T. statisticus, T. m. mairii, T. mcdowelli, and T. truncatus. Similarly, two posterior temporals would be a further character diagnosing our species from 7. dahlii, T. hypomelas, and T. pic- turatus. The nearest relatives of Tropidonophis dolasii are not immediately evident. It shares with eight other species (7. aenig- maticus, T. elongatus, T. montanus, T. mul- tiscutellatus, T. novaeguineae, T. parkeri, T. picturatus, and T. statisticus) the com- VOLUME 117, NUMBER 3 mon scale conditions of 15 unreduced dor- sal scale rows, two preoculars, three or more postoculars, and eight supralabials. Of these eight, only 7. elongatus and T. mon- tanus attain an equivalent size (the remain- der never exceed 950 mm and individuals usually are much smaller). Only T. elon- gatus, T. multiscutellatus, and T. novaegut- neae sometimes have dorsal bands, al- though the bands are solid and unlike the margined construction seen in 7. dolasii and are superimposed on a brown or gray, instead of yellow, ground color. Of the eight species, all except T. novaeguineae typical- ly bear a postocular dark stripe, not seen in T. dolasii. Ventral scale counts of Tropi- donophis dolasii overlap only with those in T. elongatus, T. montanus, T. parkeri, and T. statisticus; subcaudal counts overlap only with 7. aenigmaticus and T. pictura- tus; and ventrals + subcaudals overlap only with 7. aenigmaticus, T. multiscutellatus, and T. statisticus. Given this chaotic pattern of character-state similarities and our small sample size, attempts to identify the sister taxon of 7. dolasii would be premature. Acknowledgments We thank E Malesa, D. Salepuna, and J. Tekwae for field assistance; D. Mitchell and Conservation International for logistical as- sistance in Milne Bay Province; D. Libai, E Malesa, and B. Salepuna for logistical as- sistance on Fergusson Island; C. Kishinami for specimen curation; L. Ford and C. Rax- worthy for loan of the paratype; B. Evans for preparing the map; A. Kodani for pre- paring the line drawings; the PNG National Museum and Art Gallery for providing in- country collaborative assistance; and the PNG Department of Environment and Con- servation, PNG National Research Institute, and Milne Bay Provincial Government for permission to work in Milne Bay Province. This research was supported by National Science Foundation grant DEB 0103794. 309 Literature Cited Malnate, E. V., & G. Underwood. 1988. Australasian natricine snakes of the genus Tropidonophis.— Proceedings of the Academy of Natural Scienc- es of Philadelphia 140:59—201. O’Shea, M. 1996. A guide to the snakes of Papua New Guinea. Independent Group Pty. Ltd., Singa- pore, 239 pp. Paijmans, K. 1975. Vegetation of Papua New Guin- ea.—CSIRO Land Research Series 35:1—25 + 4 maps. . 1976. New Guinea Vegetation. Australian Na- tional University Press, Canberra, 212 pp. Appendix Specimens examined Tropidonophis aenigmaticus: BPBM 16534, E slope Oya Tabu, Fergusson Island, 9.4556°S, 150.7896°E, 1050 m, Milne Bay Prov., Papua New Guinea; BPBM 16535, Ulua, Fergusson Island, 9.4520°S, 150.8251°E, 0-10 m, Milne Bay Proy., Papua New Guinea; BPBM 16536, S slope Oya Waka, Fergusson Island, 9.4562°S, 150.5596°E, 980 m, Milne Bay Prov., Papua New Guinea; BPBM 16537, 17241, Saidowai, Normanby Island, 9.9637°S, 150.9546°E, 0-10 m, Milne Bay Proy., Papua New Guinea; BPBM 16538, 1.4 km NE Saidowai, Normanby Island, 9.9530°S, 150.9607°E, 40-80 m, Milne Bay Proy., Papua New Guinea; BPBM 17243, Sibonai, Normanby Island, 10.13578°S, 150.9708°E, 0-40 m, Milne Bay Prov., Papua New Guinea; BPBM 17242, 17244, S end Sewa Bay, Nor- manby Island, 10.0340°S, 150.9822°E, 60 m, Milne Bay Prov., Papua New Guinea. Tropidonophis doriae: BPBM 13135, E branch Avi Avi River, 5.5km S, 5.6km W of Tekadu Airstrip, 7.735°S, 146.496°E, 120 m, Gulf Prov., Papua New Guinea. Tropidonophis hypomelas: BPBM 12022, Weitin River Valley, 10 km N, 8.5 km W of river mouth, New Ireland, 4.533°S, 152.95°E, 250 m, New Ireland Prov., Papua New Guinea; BPBM 12163, Weitin River Val- ley, 8 km N, 7 km W of river mouth, New Ireland, 4.554°S, 152.964°E, 150 m, New Ireland Prov., Papua New Guinea. Tropidonophis mairii: BPBM 3104, 3299-3300, Balimo, 8.00°S, 142.55°E, 10 m, Western Prov., Papua New Guinea. Tropidonophis multiscutellatus: BPBM 5030, Biak Island, 1.02°S, 136.27°E, Papua, Indonesia; BPBM 3783, May River, 400 m, East Sepik Prov., Papua New Guinea; BPBM 17232-33, W Alotau, 10.3092°S, 150.3471°E, 5-10 m, Milne Bay Prov., Papua New Guinea; BPBM 5506, Wau, 7.343°S, 146.713°E, Mo- robe Prov., Papua New Guinea. Tropidonophis picturatus: BPBM 4133, Garaina, 7.883°S, 147.142°E, 750 m, Morobe Prov., Papua New Guinea. 310 Tropidonophis statisticus: BPBM_ 17293-95, 17299300, vic. Bunisi Village, 10.0171°S, 149.6002°E, 1420-1490 m, Milne Bay Proy., Papua New Guinea; BPBM 17296—98, Siyomu Village, 10.0145°S, 149.5970°E, 1300 m, Milne Bay Prov., Papua New Guinea; BPBM 4146, 5125, vic. Wau, 7.343°S, 146.713°E, 1600-1650 m, Morobe Prov., Papua New Guinea; BPBM 4128, 5459, 6239—40, Mt. Kaindi, 7.348°S, 146.667°E, 1800-2250 m, Morobe Prov., Papua New Guinea; BPBM 5458, 6337, Edie Creek, 7.358°S, 146.658°E, 2000-2200 m, Morobe PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Prov., Papua New Guinea; BPBM 6484, Bulldog Rd., 9 km SE Wau, 2200 m, Morobe Prov., Papua New Guinea; BPBM 3734, Sarawaget Range, 1920 m, Mo- robe Proy., Papua New Guinea; BPBM 5497, Kililo, Sarawaget Range, 2100 m, Morobe Prov., Papua New Guinea; BPBM 5498, SW Kabwum, Sarawaget Range, 2300 m, Morobe Prov., Papua New Guinea; BPBM 2272, Banz, 5.50°S, 144.35°E, 1680 m, Western High- lands Prov., Papua New Guinea; BPBM 2895, 16 km NW Banz, Western Highlands Prov., Papua New Guin- ea. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):311-316. 2004. A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras James R. McCranie and Franklin E. Castaneda (JRM) 10770 SW 164" Street, Miami, Florida 33157-2933, U.S.A., e-mail: jmccrani @bellsouth.net; (FEC) Posgrado en Biologia, Universidad de Costa Rica, San José, Costa Rica, e-mail: castanek @ yahoo.com Abstract.—A new species of Omoadiphas is described from the Cerro Tex- iguat Wildlife Refuge in the Cordillera Nombre de Dios of northern Honduras. The new species differs from the congeneric O. aurula in number of subcaudal, supralabial, infralabial, and postocular scales, in color and pattern, and in hay- ing the posterior nasal scale in contact with the prefrontal scale. Even though the type-locality is declared a wildlife refuge by the Honduran government, rapid deforestation of the area does not bode well for the continued existence of the species at its type (and only known) locality. Resumen.—Se describe una nueva especie de Omoadiphas del Refugio de Vida Silvestre Texiguat, ubicado en la Cordillera Nombre de Dios en el norte de Honduras. La nueva especie difiere de su congenerico O. aurula en el numero de escamas subcaudales, supralabiales, infralabiales y postoculares, en color y patron y en que tiene la escama nasal posterior en contacto con la escama prefrontal. Aunque la localidad tipo ha sido declarada como un Refugio de Vida Silvestre por el gobierno de Honduras, la rapida deforestaci6n que se observa en el area es una amenaza para la nueva especie. The Cordillera Nombre de Dios of north- ern Honduras is an area of extremely high endemism among amphibians and reptiles. The Cerro Texiguat Wildlife Refuge, in the western portion of this mountain range, is known to harbor 18 Honduran endemic spe- cies of amphibians and reptiles, eight of which have their type-locality within the re- serve (McCranie, pers. observ.). In Septem- ber 2003, we collected a specimen of snake in this reserve that represents an unde- scribed species of the recently described ge- nus Omoadiphas Kohler, Wilson, & Mc- Cranie and another endemic for the refuge. Herein we describe this species. Methods We follow the format of the description of the holotype in Kohler et al. (2001) in describing this new taxon. The Dowling (1951) method was used in counting ventral scales. Head and scale measurements were made to the nearest 0.1 mm with dial cali- pers held under a dissecting microscope. Snout-vent length and tail length measure- ments were made to the nearest mm along- side a ruler. Measurements are abbreviated to: snout-vent length (SVL); total length (TL); head length (HL); and head width (HW). Scale dimensions were made at the longest or widest points along the longitu- dinal or breathwise dimensions of the body, respectively. Color (capitalized) and codes (in parentheses) in life follow those of Smi- the (1975-1981). The term “‘goo-eaters”’ is used in the sense given it by Cadle & Greene (1993) and Fernandes (1995). Com- parative statements about other snake gen- 312 era are taken from KGhler et al. (2001) and references cited therein. Systematics Omoadiphas texiguatensis, new species Figs. 1-3 Holotype.—USNM 559599 (National Museum of Natural History), an apparently subadult female from approximately 2.5 airline km NNE of La Fortuna, 15°25’49’N, 87°18'32"W, 1690 m elev., Cerro Texiguat Wildlife Refuge, Departamento de Yoro, Honduras, collected 3 September 2003 by Franklin E. Castaneda & James R. Mc- Cranie. Original number LDW 13565. Diagnosis.—Omoadiphas texiguatensis can be distinguished from the holotype of O. aurula (SMF 78865; subadult female), the only known specimen of the only other known species in the genus, in having 47 subcaudal scales (24 in O. aurula), six su- pralabials (seven), seven infralabials (eight), one postocular (two), the posterior nasal contacting the prefrontal (posterior nasal separated from prefrontal by loreal), a dorsal pattern of a dark stripe on scale row three on each side (dark stripe only on vertebral row), and dark brown to nearly black ventral surfaces in preservative (pale yellow). The affinities of the two species of Omoadiphas appear to lie with a group of six other genera of snakes (see Kohler et al. 2001) that are part of a larger group re- ferred to as “‘goo-eaters.’’ Omoadiphas tex- iguatensis differs from the species of these six other genera in the following ways: from Adelphicos in having 17 dorsal scale rows (15), 172 ventral scales (120—147), and no anterior temporal (anterior temporal present); from all Atractus in having a di- vided cloacal scute (undivided) and from select species of Atractus in lacking an an- terior temporal (anterior temporal present in some Atractus); from Chapinophis in hav- ing 172 ventral scales (178-196), 47 sub- caudal scales (29—40), no anterior temporal (anterior temporal present), no scale row re- duction anteriorly on body (scale row re- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON duction present), and a striped body pattern (stripes absent); from Chersodromus in having 172 ventral scales (124-142), 47 subcaudal scales (32—43), a divided cloacal scute (undivided), and a striped body pat- tern (stripes absent); from all Geophis in having a divided cloacal scute (undivided) and a striped dorsal pattern (stripes absent) and from select species of Geophis in lack- ing an anterior temporal (anterior temporal present in some Geophis); and from Ninia in having 172 ventral scales (122-157), no anterior temporal (anterior temporal pre- sent), smooth dorsal scales (keeled), a di- vided cloacal scute (undivided), and a striped body pattern (stripes absent). Description of holotype.—An apparently subadult female; TL 169 mm; SVL 143 mm; tail length 26 mm (15.4% of TL); HL 8.0 mm from front face of rostral to pos- terior end of mandible; HW 3.9 mm at broadest point (level of angle of mouth); head barely distinct from neck; snout broadly rounded in dorsal view; eye length 0.8 mm; snout length 1.9 mm, about 2.4 times as long as eye length; pupil circular; rostral about 2.0 times wider than high (0.6 mm X 0.3 mm); internasals about 2.0 times wider than long (0.4 mm X 0.2 mm); pre- frontals much larger than internasals, about as wide as long (0.9 mm X 0.9 mm), bor- dering orbit above loreal and anterior to su- praocular; median prefrontal suture (1.0 mm) 0.4 times as long as frontal; frontal broadly rounded anteriorly, strongly V- shaped posteriorly, about 1.6 times longer than wide (2.3 mm X 1.4 mm), much long- er than distance from its anterior edge to tip of snout (1.6 mm); parietals about 2.1 times longer than wide (3.4 mm X 1.6 mm), me- dian suture (1.9 mm) shorter than frontal length; supraoculars longer than wide (0.6 mm X 0.4 mm), bordering orbit, contacting postocular, separated from loreal by pre- frontal. Nasal divided, anterior nasal contacting rostral, internasal, and first supralabial, posterior nasal contacting internasal, pre- frontal, loreal, and first and second supra- VOLUME 117, NUMBER 3 313 Fig. 1. Omoadiphas texiguatensis. labials, nostril located in posterior portion of anterior nasal; loreal single, about 3.0 times longer than high (0.9 mm X 0.3 mm), lower edge contacting second and third su- pralabials, upper edge contacting prefrontal, loreal bordering orbit (no preocular): post- Drawing of dorsal (A) and lateral (B) surfaces of the head of the holotype (USNM 559599) of ocular single, about 2.0 times higher than long (0.6 mm X 0.3 mm); no anterior tem- poral, posterior temporal single, about 1.7 times longer than high (1.0 mm X 0.6 mm); supralabials 6—6, third and fourth bordering orbit, fifth contacting postocular, parietal, 314 Fig. 2. Schematic drawing of the midbody dorsal pattern of the holotype (USNM 559599) of Omoadi- phas texiguatensis. and posterior temporal, sixth contacting posterior temporal; mental about 3.0 times wider than long (0.6 mm X 0.2 mm), sep- arated from chinshields by first pair of in- fralabials, which contact each other along ventral midline; chinshields about 1.3 times longer than wide (1.5 mm X 1.2 mm), not extending to border of lip, separated from first ventral by two gular scales and four preventral scales; infralabials 7—7, first four contacting single pair of enlarged chin- shields (their suture length 1.2 mm); a few tiny scale organs (tubercles) present dorsal- ly and ventrally on head; dorsal scales dis- posed in 17-17-17 longitudinal rows, smooth throughout, lacking apical pits and supra-anal tubercles; dorsal scales in 10 rows at level of tenth subcaudal; ventrals 172; cloacal scute divided; subcaudals 47, paired; tail spine pointed. Color in life: Dorsal surfaces of head and neck Chestnut (32) with Sepia (119) spots; dorsal surface of body Prout’s Brown (121A) with Sepia (119) spots; Sepia (119) dorsolateral stripe present on scale row three on each side, lateral area below stripe Vandyke Brown (121); dorsal surface of tail PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Prout’s Brown (121A); ventral and subcau- dal surfaces Vandyke Brown (121); iris Vandyke Brown (121). Color in alcohol (about two weeks after preservation): Dorsal surface of head me- dium brown; dorsal surface of body dark brown with indistinct darker brown spots present on anterior one-third; darker brown longitudinal stripe present on scale row three on each side of body; a vague, slightly darker brown vertebral stripe present; dor- sal surface of tail darker brown than that of body; ventral surface of head pale brown, that of body dark brown anteriorly, becom- ing even darker brown posteriorly; subcau- dal surface very dark brown, almost black. Distribution and natural history notes.— Omoadiphas texiguatensis is known only from within the limits of the Cerro Texiguat Wildlife Refuge (Refugio de Vida Silvestre Texiguat). The holotype was crawling in leaf litter next to a rotten log. Only the snake’s tail was exposed when first sighted; its body was under the leaves. It was found at 1000 h in moderately disturbed cloud forest (Lower Montane Wet Forest forma- tion of Holdridge 1967) at 1690 m elev. A hard rain occurred from about 1850 to 1875 h the previous day, but the weather was clear and sunny when the snake was cap- tured. Etymology.—tThe specific name texiguat- ensis is formed from Texiguat and the Latin suffix —ensis (denoting place, locality, or country). The name refers to the Cerro Tex- iguat Wildlife Refuge where the holotype was collected. We use this specific name in an effort to stress the importance of this wildlife refuge to the conservation status of many Honduran endemic species of am- phibians and reptiles (but see Discussion). Discussion The genus Omoadiphas is now known from two apparently subadult females placed in two species, making it one of the most poorly known snake genera in the Neotropics. Kohler et al. (2001) concluded VOLUME 117, NUMBER 3 315 i Fig. 3. Dorsal (A) and ventral (B) views of the holotype (USNM 559599) of Omoadiphas texiguatensis, total length 169 mm. 316 that its relationships appear to lie with six other Neotropical genera that are part of a larger group called “‘goo-eaters” by Cadle and Greene (1993) and Fernandes (1995). The discovery of O. texiguatensis appears to support this relationship as well as sup- porting the distinctiveness of the genus. Omoadiphas texiguatensis is truly a dif- ficult snake to find. After collecting the ho- lotype, we spent much of the following three days in the area raking through leaves, overturning and ripping apart rotten logs, and overturning rocks in an unsuccessful at- tempt to find more specimens. We also walked through the area for several hours on two nights searching for active snakes. In addition, this was McCranie’s fourth col- lecting trip to the area. As noted by Wilson et al. (2001). and McCranie & Wilson (2002), most of the protected areas in Honduras exist on paper only. Such is the case for the Cerro Texi- guat Wildlife Refuge. There are no facilities or personnel of any sort or even signage to denote the presence of a protected area. In- deed, people living in San Francisco (the closest village to the type-locality of O. tex- iguatensis) and in the area between that vil- lage and the type-locality that we queried are unaware that the area is a wildlife ref- uge. In addition, crop fields and cleared ar- eas now dominate the area around the type- locality. We did not encounter any pristine forest in September 2003 within an hour or two walk in any direction from where the holotype of O. texiguatensis was collected. This is in sharp contrast to the condition of the area during McCranie’s first visit in 1991 when pristine cloud forest dominated the region. Clearly, the rapid rate of defor- estation in the area does not bode well for the continued existence of O. texiguatensis or any of the other species of amphibians and reptiles found in this region of unusu- ally high endemism. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments We thank C. Gonzalez, M. Moreno, and H. Portillo of COHDEFOR, Tegucigalpa, for issuing collecting and exportation per- mits. Sefor G. Enamorado, his son Bairon, and daughters Marina and Norma of San Francisco, Yoro, provided us with a ride and good company up the tortuous “road” to the type-locality and then back to San Francisco. We also thank EF D. Castaneda for loaning us the vehicle in which we reached San Francisco. An earlier draft of the manuscript was improved upon by L. D. Wilson. Figs. 1 and 2 were drawn by S. Mohammadi. Literature Cited Cadle, J. E., & H. W. Greene. 1993. Phylogenetic pat- terns, biogeography, and the ecological struc- ture of Neotropical snake assemblages. Pp. 281-293 in R. E. Ricklefs and D. Schluter, eds., Species diversity in ecological communities: historical and geographical perspectives. Uni- versity of Chicago Press, Chicago, 414 pp. Dowling, H. G. 1951. A proposed standard system of counting ventrals in snakes.—British Journal of Herpetology 1:97—99. Fernandes, R. 1995. Phylogeny of the dipsadine snakes. Unpublished Ph.D. dissertation, Univer- sity of Texas at Arlington, 115 pp. Holdridge, L. R. 1967. Life zone ecology, Revised edi- tion. Tropical Science Center, San José, Costa Rica, 206 pp. Kohler, G., L. D. Wilson, & J. R. McCranie. 2001. A new genus and species of colubrid snake from the Sierra de Omoa of northwestern Honduras (Reptilia, Squamata, Colubridae).—Sencken- bergiana Biologica 81:269-276. McCranie, J. R. & L. D. Wilson. 2002. The amphibi- ans of Honduras.—Society for the Study of Amphibians and Reptiles, Contributions to Her- petology 19:1—625 + pls. 1—20. Smithe, EF B. 1975-1981. Naturalist’s color guide. Part I. Color guide. The American Museum of Nat- ural History, New York, 182 color swatches. Wilson, L. D., J. R. McCranie, & M. R. Espinal. 2001. The ecogeography of the Honduran herpetofau- na and the design of biotic reserves. Pp. 109— 158 in J. D. Johnson, R. G. Webb andy O. Flo- res-Villela, eds., Mesoamerican herpetology: systematics, zoogeography, and conservation. Centennial Museum, University of Texas at El Paso, Special Publication 1:1—200. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):317-329. 2004. A new species of Kolpotocheirodon (Teleostei: Characidae: Cheirodontinae: Compsurini) from Bahia, northeastern Brazil, with a new diagnosis of the genus Luiz R. Malabarba, Flavio C. T. Lima, and Stanley H. Weitzman* (LRM) Museu de Ciéncias e Tecnologia, PUCRS, Av. Ipiranga 6681, 90619-900, Porto Alegre, RS, Brazil. and Depto. Zoologia, IB, Universidade Federal do Rio Grande do Sul, Av. Bento Gongalves 9500, 91501-970, Porto Alegre, RS, Brazil, e-mail: malabarb@pucrs.br; (FCTL) Museu de Zoologia da USP, Caixa Postal 42594, 04299-970 Sao Paulo, SP, Brazil, e-mail: fctlima@usp.br; (SHW) Division of Fishes, WG 12, Department of Systematic Biology, MRC-0159, Smithsonian Institution, RO. Box 37012, Washington D.C. 20013-7012, U.S.A, e-mail: weitzman.stan@nmnh.si.edu Abstract.—Kolpotocheirodon figueiredoi, a new species of the characid sub- family Cheirodontinae, tribe Compsurini, is described from the upper rio Pa- raguacu basin, Bahia, Brazil. A new diagnosis for the genus is proposed, based mostly on scanning electron microscope (SEM) analyses of the caudal organ of the new species and that of the single previously known species, Kolpoto- cheirodon theloura. The genus is diagnosed now in part by the presence of a previously undescribed, sexually dimorphic and apparently glandular, structure found in the lower caudal-fin lobe of males. The basal relative position of Kolpotocheirodon within the Compsurini, in which all species are inseminating, is further supported by the presence of aquasperm in both species rather than the apomorphic elongate sperm nuclei present in the remaining members of the tribe. Resumo.—Kolpotocheirodon figueiredoi é descrito para a por¢ao superior da bacia do rio Paraguacgu, Bahia, Brasil. Propoem-se uma nova diagnose para © género, baseada principalmente na analise de Microscopia Eletr6nica de Var- redura do 6rgao caudal da nova espécie e de Kolpotocheirodon theloura, a unica espécie conhecida anteriormente. O género é diagnosticado pela presenga de uma estrutura aparentemente glandular e previamente nao descrita do lobo ventral da nadadeira caudal dos machos. A posigao relativamente basal de Kolpotocheirodon em Compsurini, uma tribo de peixes com inseminagao de Cheirodontinae, é corroborada pela presen¢a de espermatozoides aproximada- mente esféricos (aquasperm) nas duas espécies, ao invés da presenga de es- permatozoides de nticleo alongado, como observado nos demais membros da tribo. The genus Kolpotocheirodon was recent- ly described by Malabarba & Weitzman (2000), from a single species, K. theloura, from the uppermost tributaries of the rio Sao Francisco and rio Parana central Brazil. The genus is a member of the tribe Comp- surini, subfamily Cheirodontinae (see Mal- abarba et al., 1998) and was diagnosed pri- marily by the presence of a unique special- ized caudal organ at the proximal region of the ventral caudal-fin lobe of males. This organ consists of hypertrophied elongate dermal flaps attached along the fin rays and a series of relatively flat tabs and papillae 318 attached along the exposed border of those flaps. These structures were unknown in other inseminating or externally fertilizing species of characids. At the time the research of Malabarba & Weitzman (2000) was conducted, FE C. T. Lima and colleagues were collecting in the rio Pratinha, a tributary of the rio Paragua- cu, Iraquara, Bahia, Brazil and there they discovered a new cheirodontine species that has a caudal organ similar to that present in K. theloura. This new compsurin species is herein described and ecological data and field observations from the type locality are presented. Data from the description of the new spe- cies, examination of a new collection of better-preserved specimens of K. theloura than originally available, and scanning elec- tron microscopy (SEM) observations of the caudal-fin structures of these two species al- low a reanalysis of the characters diagnos- ing Kolpotocheirodon and redescription of the autapomorphies that distinguish its type species. Methods and Materials The systematic methods for making counts and measurements for all specimens studied here are the same as those described and used by Malabarba & Weitzman (1999) and are not re-described here. However, un- like the convention for fin rays wherein the count for the rays for the holotype is given first followed by the range and mean sep- arately for the unbranched and the branched rays, the counts of jaw teeth do not report a single value followed by an indication of variation. Instead, only the range of the counts, for example, maxilla with 2 or 3 teeth, is provided. This is because we are confident only in counts taken from cleared and stained specimens. SEM photographs were taken from specimens fixed in for- malin and preserved in 70% ethanol. Before metalization with gold, the fins were passed through 99% ethanol, then acetone, and treated with a critical point dryer. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Institutional abbreviations are as listed in Leviton et al. (1985). Character polarity for the diagnoses of the two Kolpotocheirodon species and a revised analysis of Kolpoto- cheirodon monophyly is here established by use of parsimony through a re-analysis of the cheirodontine clade Compsurini that was first diagnosed by Malabarba et al. (1998). This new analysis also includes spe- cies of the genera Saccoderma, Compsura, Macropsobrycon, and the species Acinoch- eirodon melanogramma (“identified”’ as ““New Genus and Species B” in Malabarba et al, 1998), and Kolpotocheirodon theloura (then “identified” as “‘New Genus and Spe- cies A’). Kolpotocheirodon Malabarba & Weitzman Kolpotocheirodon Malabarba & Weitzman, 2000:270 (type species: Kolpotocheiro- don theloura Malabarba & Weitzman, 2000:271 by monotypy and original des- ignation). Comments preliminary to the diagnosis.— The genus Kolpotocheirodon was diag- nosed in Malabarba et al. (1998) (as New Genus and Species A) and in Malabarba & Weitzman (2000) by the presence of three apomorphic features that occur in its type species. These characters, as described by Malabarba and Weitzman (2000), are a spe- cialized part of a caudal organ located at the proximal region of the ventral caudal- fin lobe of mature males and consist of hy- pertrophied elongate dermal flaps attached along the fin rays together with a series of relatively flat tabs and papillae attached along the exposed border of these flaps (= character 36 in Malabarba 1998); hooks on the anal-fin rays of mature males distributed along the most posterior unbranched and five anterior branched anal-fin rays (= char- acter 30 in Malabarba, 1998); and the twelfth and thirteenth caudal-fin rays are dorsally concave along their basal halves and have ventrally expanded segments (= character 34, state 2 in Malabarba 1998). VOLUME 117, NUMBER 3 Fig. 1. Diagnosis.—By using SEM the special- ized caudal-fin organ described in the pre- vious diagnosis of Kolpotocheirodon 1s now found to be more complex than for- merly known. A new caudal organ, previ- ously undescribed, corresponds to a second- ary sexually dimorphic organ found exclu- sively in the ventral lobe of the caudal fin of males of both Kolpotocheirodon species. This “‘pineapple-like’’ organ is easily rec- ognized by its peculiar shape, somewhat cone shaped or papilla-like, but completely covered by smaller papillae-like bodies or knobs (see Figs. 1, 2). These are distributed among the large papillae of the caudal fin of males of K. theloura (see Fig. 3), but form the entire caudal-fin organ in K. fi- gueiredoi (see Fig. 1). This organ is found only in adult males of both species, sug- gesting that it may have a reproductive SEM of caudal organ in Kolpotocheirodon figueiredoi, male (MZUSP 55219, 25.5 mm SL), from rio Pratinha, Iraquara, Bahia, Brazil. (A) lower caudal-fin lobe; (B) detailed image showing the smooth border of the flap attached along the basal portion of the nineteenth caudal-fin; (C) and (D) detailed images of the pineapple organs of the ventral lobe of the caudal fin. function, possibly pheromone in nature. This pineapple organ has not been found in other cheirodontines or other characids, and its presence supports a hypothesis of close relationship between the two Kolpotochei- rodon species. Both Kolpotocheirodon species have a conspicuous small black spot at the mid- length of the first branched anal-fin ray of males (Figs. 5, 7 and 8), absent in females (Fig. 6). Such a spot is absent in all other known cheirodontines. It is here considered derived and a synapomorphy for the genus. Males of Kolpotocheirodon figueiredoi and K. theloura have the ventral body sur- face in the area covering the pelvic bone with a dark brown mark, nearly in the shape of an isosceles triangle. This pigment ap- pears to externally delineate an area corre- sponding to the muscles inserted on the pel- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. caudal-fin ray of males in Kolpotocheirodon theloura. MNRJ 18081, SL 26.2 mm, from lagoa Perta Pé, rio Sao Francisco drainage, Palmital, Minas Gerais, Brazil. (A) bar = 50 wm; (B) bar = 20 pm. vic bone (Fig. 8). Such a spot is absent in all other cheirodontines, and constitutes a synapomorphy for the two species. Malabarba & Weitzman (2000) described the presence of well-developed hooks along the last unbranched and five anterior branched anal-fin rays of males as derived, and diagnostic for Kolpotocheirodon (= character 30 in Malabarba 1998). The new specimens available of K. theloura, MNRJ 18081, have the last unbranched and five to seven anterior branched anal-fin rays of males bearing hooks (5 branched rays in 7 specimens, 6 in 23 specimens, and 7 in 3 specimens). Males of K. figueiredoi have the last unbranched and five to six anterior branched anal-fin rays of males bearing hooks (5 branched rays in 6 specimens, 6 in 6 specimens; 8 in one specimen). The anal-fin region bearing hooks also contains modified soft tissues, absent in the remain- ing portion of the fin. Although showing more variability than previously described, the condition found in both Kolpotochei- rodon species is different from that found in other compsurins, which have hooks along a larger number of anal-fin rays. We found that only in Saccoderma species among compsurins are anal-fin hooks re- stricted to the anterior anal-fin rays, in the last unbranched and four anterior branched rays. By parsimony both conditions are considered derived and autapomorphic for Detailed SEM images of the pineapple organs found between the papillae of the ventral lobe of the each genus. Note: Menezes et al. (in press) and Weitzman et al. (in press) have de- scribed and discussed glandular soft tissue in the anal fins of sexually active male char- acids of many kinds including glandulocau- dines, and some non-glandulocaudines. This tissue is most often associated with anal-fin hooks, but in one species a glan- dular organ was found. Kolpotocheirodon theloura Malabarba & Weitzman Fig. 7 Kolpotocheirodon theloura Malabarba & Weitzman, 2000:271—281 (description; relationships); 272, fig. 1 (holotype); 273-4, fig. 2-3 (paratypes); 275, fig. 4 (caudal-fin hooks); 276, fig. 5 (ventral caudal-fin lobe); 277, fig. 6 (anal-fin hooks); 278, fig. 7 (premaxillary and maxillary teeth), fig. 8 (pelvic-fin hooks). Material examined: All specimens listed in Malabarba & Weitzman (1999), plus MNRJ 18081, 135 spms. (10 examined, SL 24.3—27.4 mm), Brazil, Minas Gerais, Palmital, lagoa Perta Pé, rio Sao Francis- co drainage. Diagnosis.—Kolpotocheirodon theloura is diagnosed from the new Kolpotocheiro- don species and other characid fishes by the following autapomorphies: As described in Malabarba & Weitzman (2000), K. theloura VOLUME 117, NUMBER 3 Fig. 3. SEM images of caudal organ in Kolpotocheirodon theloura, male, MNRJ 18081, SL 26.2 mm, from lagoa Perta Pé, rio Sao Francisco drainage, Palmital, Minas Gerais, Brazil. (A) lower caudal-fin lobe, bar = 600 zm ; (B) detailed image of the flap attached along the basal portion of the nineteenth caudal-fin ray with a series of relatively flat tabs along its exposed border, bar = 100 pm; (C) detailed image of the flaps attached to the eighteenth through thirteenth or fourteenth fin-rays with a single series of papillae along its exposed border, bar = 200 pm. has hypertrophied elongate dermal flaps at- tached along the fin rays of the ventral cau- dal-fin lobe of males (= character 36 in Malabarba 1998). The flap attached along the basal portion of the nineteenth caudal- fin ray has a series of relatively flat tabs along its exposed border (Fig. 3). The flaps attached to the eighteenth through thirteenth or fourteenth fin-rays are relatively short, narrow and bear papillae in a single series along its exposed border (Fig. 3A, C). These modified flaps of the caudal organ are not exclusive to males in K. theloura, being also found in females, although less developed (Fig. 4). Modified flaps are also observed in the dorsal fin of males of K. 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. theloura (Fig. 9). These modified flaps are independent of the sexually dimorphic pineapple-like organs described as a syna- pomorphy for Kolpotocheirodon and are absent in K. figueiredoi. The modified flaps constitute an autapomorphy of K. theloura. As described in Malabarba & Weitzman (2000), the twelfth and thirteenth caudal-fin rays of K. theloura are curved, dorsally concave along their basal halves, and with ventrally expanded segments (= character 34, state 2 in Malabarba, 1998). This fea- ture was not observed in K. figueiredoi and is considered autapomorphic for the type species, K. theloura. Kolpotocheirodon theloura has 3—5 very small vertical bars crossing lateral body Detailed SEM images of the flaps bearing papillae along the basal portion of the ventral lobe of the caudal-fin ray in Kolpotocheirodon theloura, female, MNRJ 18081, SL 26.5 mm, from lagoa Perta Pé, rio Sao Francisco drainage, Palmital, Minas Gerais, Brazil. (A) bar = 500 pm; (B)bar = 200 pm. stripe between pseudotympanum and area ventral to dorsal fin (Fig. 7). Such bars are absent in the new Kolpotocheirodon species and in other compsurins and represent an autapomorphy for K. theloura. Kolpotocheirodon figueiredoi new species Figs. 5, 6 Holotype.-MZUSP 70037, 1 male, 30.5 mm SL, Brazil, Bahia, Iraquara, rio Pratin- ha, Fazenda Pratinha (12°21'13’S; 41°32'57”"W), 17-21 Dec 1998; P. Gerhard, E C. T. Lima, FE Di Dario and L. S. Rocha. Paratypes.—All specimens collected with the holotype: MCP 22345, 3 males, Fig. 5. Kolpotocheirodon figueiredoi, new species, holotype, male, MZUSP 70037, SL 30.5 mm; rio Pratin- ha, Iraquara, Bahia, Brazil. VOLUME 117, NUMBER 3 323 Fig. 6. tinha, Iraquara, Bahia, Brazil. 25.1—-30.5 mm SL, 2 females, 24.0—24.8 mm SL. MZUSP 55219, (6) 14 males, 24.2-28.2 mm SL, 8 females, 24.0—31.0 mm SL; (1 male 28.2 mm SL and 1 female 26.9 mm SL Alizarin red s and Alcian blue stained specimens cleared with trypsin; 1 male 26.2 mm SL and 1 female 26.4 mm SL sectioned for histology; 1 male 25.5 mm SL sectioned for TEM study). Diagnosis.—Kolpotocheirodon figueire- doi lacks all autapomorphies described above for K. theloura, but has no unambig- uous autapomorphies for its diagnosis. The Kolpotocheirodon figueiredoi, new species, paratype, female, MZUSP 55219, SL 30.0 mm; rio Pra- following characters have alternative states between K. figueiredoi and K. theloura, but these also occur alternatively among other compsurin species. Nevertheless they are most parsimoniously accepted either as au- tapomorphic for K. figueiredoi or apo- morphic for K. theloura. Whereas males of K. figueiredoi have no hooks on the caudal-fin rays, while males of K. theloura have the twelfth to the four- teenth or fifteenth principal caudal-fin rays bearing 4—6 retrorse hooks on each side in a row along their dorsal divisions (Mala- Fig. 7. Gerais, Brazil. Kolpotocheirodon theloura, male, MNRJ 18081, SL 25.0 mm; lagoa Perta Pé, rio Palmital, Minas 324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. covering the pelvic bone showing a dark brown mark, nearly isosceles triangle shape, apparently externally delineating the area corresponding to the muscles inserted in the pelvic bone. (B) and (C) Left lateral view of the dorsal (B) and anal fins (C), showing the dark spots of those fins. barba & Weitzman, 2000: fig. 4). The pres- ence of hooks on the caudal fin is known for several compsurins, including Acinoch- eirodon melanogramma (hooks on caudal- fin rays 13-14, rarely on ray 15), Saccod- erma hastata (hooks on caudal-fin rays 13— 18), “Odontostilbe’ dialeptura (hooks on caudal-fin rays 12-16), and Macropsobry- con uruguayanae (hooks on caudal-fin rays 12-14, plus several spinelets along the proximal half of caudal-fin rays 14 to 18). However, hooks are absent in Compsura heterura, Compsura gorgonae, and ”’ Od- ontostilbe”’ mitoptera. Malabarba & Weitz- man (1999, 2000) pointed out that although these hooks are present on the ventral lobe of the caudal fin in all these species, they do not all occur on the same caudal-fin rays in all species and are of different shapes. A previous analysis of character distribution Kolpotocheirodon. figueiredoi male, MCP23455, SL 30.5 mm. (A) Ventral body surface in the area (Malabarba et al., 1998) indicated the pres- ence of caudal-fin hooks as a synapomor- phy for the compsurin cheirodontines, and its absence a secondary reversal in some of its species. The inclusion of a new species bearing no hooks in the most basal genus of the tribe allows either the recognition of the presence of hooks as a synapomorphy for the tribe Compsurini with a reversal in K. figueiredoi, or the recognition of inde- pendent acquisitions of hooks in K. thel- oura and in the clade including the remain- ing compsurins. The first hypothesis is pre- ferred, since it better conforms with the pu- tative homology of caudal-fin hooks among compsurins (de Pinna 1991). Males of K. figueiredoi have a conspic- uous small black spot in the soft tissue be- tween midlength of first and second, and second and third branched dorsal-fin rays VOLUME 117, NUMBER 3 Fig. 9. rodon theloura, male, MNRJ 18081, SL 26.2 mm, from lagoa Perta Pé, rio Sao Francisco drainage, Palmital, Minas Gerais, Brazil. (A) bar = 500 wm; and (B) bar = 100 pm. Detailed SEM images of the flaps bearing papillae along the anterior dorsal-fin ray in Kolpotochei- (Figs. 5, 8). This is absent (Fig. 7) in K. theloura (= character 65 in Malabarba 1998). Among compsurins, a similar spot is observed in species of Compsura, Macrop- sobrycon and Acinocheirodon, but is absent in species of Saccoderma, This spot was previously proposed as a synapomorphy for a clade including the last four genera cited above. Again, the inclusion of a new spe- cies in the most basal genus of the tribe allows both the recognition of the presence of the dorsal black spot as a synapomorphy for the tribe Compsurini with a reversal in K. theloura, or the recognition of indepen- dent acquisitions of the dorsal black spot in K. figueiredoi and in the clade including re- maining compsurins. The first hypothesis is preferred because it better conforms to the putative homology of the dorsal black spot among compsurins. A further character distinguishing K. fi- gueiredoi is its caudal-peduncle/caudal-fin Table 1—Morphometrics of Kolpotocheirodon figueiredoi, new species. Standard length is expressed in mm; measurements through head length are percentages of standard length; the last four entries are percentages of head length. Range includes the holotype, MZUSP 70037, and paratypes MCP 22345, MZUSP 55219. Halewee Males Females male n Low High x SD n Low High xX SD Standard length (mm) 30.5 13 25.3 303 270 10 23.8 31.0 26.4 Snout to anal-fin origin 9.7) 13 57.3 63.0 60.0 1.30 1©@ © @5,3 ©3.7/ I35 Snout to dorsal-fin origin 50.5 13 45.7 52.9 495 1.80 1@ 4B S24 ayy 135) Snout to pelvic-fin origin 42.0 13 42.0 46.7 445 1.51 I@ = 4S5).3} E38} E33} INS IN/ Dorsal-fin base length 13}, Ih 13 Wi IS Id i120) 10 Wi 143 132 0.72 Anal-fin base length 25.9 13 24.5 29.0 26.8 1.27 10 235 26.8 25.4 1.08 Caudal peduncle length IS). 13 25 id. 4 O20) 10 Hiei 54 3s 25) Caudal peduncle depth 13.8 13 13.6 16.1 14.8 0.76 10 Wi 133 125 O93 Depth at dorsal-fin origin Slail 13 Sil B5./ B35) iis 10) 336) 39:4 35:4 Ese Dorsal-fin height 29.8 12 26.1 29.8 28.0 1.36 Q WZ WMS RA O83 Pelvic-fin length 19.0 13 167 19.3 IS2 O71 10 IS4b 16.3 146 O79) Pectoral-fin length 19.3 13 175 20.1 18.8 0.77 8 IS. IO2 WA tao Bony head length 23.3 13 Was) AP) DANS (O59) 1@ 23.0 250 Aj O72 Snout length 21.1 13 20323) 8 Oa 10 IQAE WZ.3) Biles 1335) Upper jaw length 31.0 13 M54 Zl@ 27.7 Way 10 2a SO2 ZI 2,13) Horizontal eye diameter 38.0 13 33.3 385 362 i139 I@ 34h} BV Bs 1s Least interorbital width 29.6 13 ADM Bill ASQ Weil I@ 254 2S Wei 1.32 326 spot (Figs. 5, 6) that is more or less rect- angular in shape and horizontally elongate. It never reaches the dorsal and ventral bor- ders of the caudal peduncle. The same spot is vertically elongate, lozenge-shaped and reaches the dorsal and ventral borders of the caudal peduncle in K. theloura (Fig. 7). Description.—Morphometric data sum- marized in Table |. Body elongate and compressed, greatest depth at dorsal-fin or- igin. Predorsal profile slightly convex. Pro- file of body along dorsal-fin base poster- oventrally inclined, nearly straight from base of posterior dorsal-fin ray to adipose fin. Ventral body profile convex from tip of lower jaw to pelvic-fin origin. Muscles cov- ering pelvic bone strongly prominent in ventral body profile, especially in males. Area between pelvic- and anal-fin origins slightly concave in females and notably concave in males, with a pair of concavi- ties, separated from each other by a small median keel visible only when pelvic fins moved out of way. These concavities lodge pelvic fins when later retracted. Ventral pro- file along anal-fin base slightly concave in females. In males same profile, slightly convex in region of anterior lobe and slight- ly convex along remaining posterior fin portion. Dorsal and ventral profiles of cau- dal peduncle nearly straight in females. Dorsal and especially ventral surfaces of caudal peduncle of males convex, with an internal translucent cavity, covered by cau- dal peduncle scales. Head small. Snout shorter than eye di- ameter. Mouth terminal. Maxilla short, po- sitioned at an angle of approximately 45 de- grees relative to long body axis. Posterior tip of maxilla reaching vertical that passes through anterior border of eye. Premaxilla with 4 teeth, each having 9 small evenly spaced cusps all about equal in size. Cutting edge arched. Maxilla with 2 or 3 teeth similar in form to those of pre- maxilla, with 7—9 cusps. Cutting edge slightly arched to almost straight. Dentary bone with 4 or 5 large teeth each with 7 cusps; followed by 2 or 3 smaller teeth with PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 7 or fewer cusps. Teeth posterior to second tooth asymmetrical with most lateral cusps situated towards tooth base and most me- dial cusp more distally located. Cusps small and regular and approximately equal in size. Cutting edge slightly arched to almost straight. Dorsal-fin rays, 11, 9, m = 22 (ii, 8 in one specimen). First unbranched ray less than half-length of second. Dorsal-fin origin ap- proximately at midlength of body. Adipose- fin origin nearly at vertical through inser- tion of posteriormost anal-fin ray. Anal-fin rays, ii, 18, (iii-iv X = 3.5, 16— 19, X = 17.5, n = 22). Anal-fin origin slightly posterior to vertical passing through base of posteriormost dorsal-fin ray in fe- males and at a vertical passing through base of two last dorsal-fin rays in males. Distal border of anal fin concave in females, with anterior 5—6 branched rays very long, form- ing prominent anterior lobe. Distal border of anal fin of males convex in the anterior lobe, decreasing in length gradually and forming a posterior concave border. Anal- fin rays of males with slender, elongate re- trorse hooks on longest unbranched ray, and anterior first 5 or 6 branched rays (scattered hooks present in branched rays 7 and 8 in one specimen). Hooks inserted at postero- lateral border of fin rays, bent over lateral surface of fin ray and anteriorly directed. Hooks located on posterior ray branches, less numerous on anterior ray branches. One, rarely two, bilateral pair of bony hooks per ray segment. Pectoral-fin rays, 1, 9 G, 8-9, X = 8.6, n = 22). Distal ends of longest rays not reaching pelvic-fin origin in females; reach- ing or not in males. Pelvic-fin rays, 1, 7 (1,7 in all specimens, n = 22). Pelvic-fin origin anterior to vertical passing through dorsal- fin origin. Distal tip of pelvic fin passing anal-fin origin in males, but not in females. Male pelvic fins bearing elongate ventro- medial retrorse hooks along branched rays DW ts, Principal caudal-fin rays 10/10 (10/9, but 10/10 and 9/9, in one specimen each, n = VOLUME 117, NUMBER 3 21). Lower caudal-fin lobe of males cov- ered with series of papillae from 12" or 13" ray to 18" or 19" principal caudal-fin rays. Papillae most numerous near caudal-fin base, extending in some specimens to near tip of lower caudal-fin lobe. Hooks or hy- pertrophied dorsal fin-ray flaps absent. Dor- sal fin-rays 9-10, and ventral procurrent caudal-fin rays 8-10, in two cleared and stained specimens. Scales cycloid, moderately large. Lateral- line pores incomplete, perforated scales 7, (5-9, X = 7.4, n = 20). Scales in lateral series 34, (332-36, X = 33.7, n = 19). Scale rows between dorsal-fin origin and lateral line 5, (5-6, X = 5.2, n = 20). Scale rows between lateral line and pelvic-fin origin 4, (4-5, X = 4.1, n = 20). Predorsal scales, when in regular series 11 (10-12, X = 10.8, n = 18). : Supraneurals, 4; precaudal vertebrae, 16; caudal vertebrae, 17—18 Gn two cleared and stained specimens). Color in alcohol.—(See Figs. 5, 6, 8). Head dark brownish dorsally with a silvery color in opercle and infraorbital area, where guanine not destroyed by fixative. Body pale brownish yellow; dorsolateral scales delineated in their borders with dark chro- matophores. Black lateral body stripe evi- dent, pale anterior to dorsal-fin origin, pro- gressively wider and conspicuous posteri- orly in larger specimens. Humeral spot ab- sent. A conspicuous caudal spot centered at posterior termination of caudal peduncle, rectangle-shaped and extending to base of middle caudal-fin rays; caudal spot never reaching ventral and dorsal borders of cau- dal peduncle. Dorsal fin of males with a conspicuous small black spot in soft tissue between approximately mid length of first and second, and second and third branched dorsal-fin rays; dorsal fin of females with- out distinct marks. Anal fin of males with a concentration of dark chromatophores along midlength of first branched anal-fin ray, forming a small and conspicuous spot in adult male specimens; absent in females. An inconspicuous dark line present along 327 anal-fin base in both sexes, plus a small dark line on body nearly parallel to longi- tudinal lateral body stripe in males and par- allel to anal-fin base in females. Pectoral and pelvic fins hyaline. Ventral body sur- face in area covering pelvic bone of males with a dark brown mark nearly shaped like an isosceles triangle, apparently delineating external area corresponding to muscles originating from pelvic bone. Ventral mid- line between pelvic-fin insertion and anal- fin origin of males with a pair of thin lateral black lines, seen only when pelvic fin ex- tended. A dark mark present on lower in- ternal border of orbits, visible only when eyes depressed. Color in life.—Described from color slides of a male taken in the field by Pedro Gerhard. Head dark brownish dorsally; op- ercle and infraorbital area silvery. Body light brownish yellow; dorsolateral scales slightly delineated with dark chromato- phores; belly silvery. Lateral body stripe evident, silvery, pale anterior to dorsal-fin origin. Humeral area unpigmented, but a dark area visible due to presence of a pseu- dotympanum. As described in preserved specimens, a conspicuous caudal spot cen- tered at posterior termination of caudal pe- duncle, rectangle-shaped, and extending to base of middle caudal-fin rays; never reach- ing ventral and dorsal borders of caudal pe- duncle. Caudal spot in colorful specimens bordered dorsally and ventrally by two yel- low spots. Small black spot on dorsal fin of males, located approximately at mid length of first and second, and second and third branched dorsal-fin rays, bordered dorsally by yellow pigmentation. Anal-fin spot of males located along mid length of first branched anal-fin ray, anteriorly bordered with yellow pigmentation. A small dark line along anal-fin base and a small dark line on body nearly parallel to longitudinal lateral body stripe visible above anterior lobe of anal fin. Pectoral and pelvic fins hyaline. Presence of marks on ventral body surface not visible in available photos. Sexual dimorphism.—Males are easily 328 recognized by their color pattern, display- ing two conspicuous small black spots on the dorsal and anal fins (see Fig. 5), and a triangular dark brown mark on the ventral body surface of the pelvic bones (see Fig. 8), absent in females (See Fig. 6). Sexes are also differentiated by the relative position of the pelvic and anal fins, both located more anteriorly in males than in females; by the larger caudal peduncle depth in males, having an expanded portion in its ventral and dorsal profiles; and by the larger pelvic-fin lengths of males (see Table 1 for all these measurements). Distribution.—Known only from the type locality, the rio Pratinha, Bahia, Brazil. The rio Pratinha is a tributary of the rio Santo Antonio, itself a tributary of the rio Paraguacu, a coastal drainage of eastern Brazil. Habitat and natural history notes.—For a complete description of the site of collec- tion of K. figueiredoi, the rio Pratinha, see Lima & Gerhard (2001: 112-113). In the rio Pratinha, K. figueiredoi was observed and collected only in those portions with a moderate water current. The species was most commonly collected in a riffle at a narrow stretch of rio Pratinha. Specimens were observed at midwater, swimming against the current, probably feeding on food items drifting downstream. During one occasion, one individual of this species was seen picking with its mouth on a large boul- der in a cave entrance. The ecological pref- erences of K. figueiredoi may be remark- able, given the fact that at least some other species of the Cheirodontinae, for example Cheirodon interruptus (Jenyns) and Chei- rodon tbicuhiensis Eigenmann, prefer lentic waters such as lagoons or pools in slow- moving water courses in coastal streams of Rio Grande do Sul, Brazil, personal obser- vation. Etymology.—We take great pleasure in naming this species in honor of José Lima de Figueiredo, a Brazilian ichthyologist at the Museu de Zoologia da Universidade de Sao Paulo. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Discussion.—The two Kolpotocheirodon species are included in the tribe Compsurini (Malabarba et al. 1998) by sharing two un- ambiguous synapomorphies with the mem- bers of that tribe: they are inseminating (Character 70 in Malabarba et al. 1998), and the anal-fin hooks are positioned along the posterolateral border of the anal-fin rays and bent more or less anteriorly over the lateral surface of the anal-fin ray to which each 1s attached (Character 26 in Malabarba et al. 1998). The presence of hooks and their distribution in the caudal fin were also previously employed for the diagnosis of the tribe, but are absent in K. figueiredoi. Alternative hypotheses explaining this are discussed above under the diagnosis of this species. Kolpotocheirodon theloura was the only species of the Compsurini known to have aquasperm (a nearly spherical or spherical sperm nucleus), a condition also found in the new Kolpotocheirodon figuei- redoi. All other species of the Compsurini so far investigated have elongate sperm nu- clei (see Burns et al. 1997:434, fig. 1B—H & 1998:242, fig. 11). Acknowledgments SEM pictures were made at Centro de Microscopia e Microanalises—PUCRS. We thank Marco Aurélio Azevedo and John Burns for the histological analyses of the gonads and caudal organ that provide im- portant data concerning the generic and species diagnoses presented herein. We thank P. Gerhard for the photos of live spec- imens. We also thank Pedro Gerhard, Fabio Di Dario and L. S. Rocha for their help dur- ing field work and Silvio Arruda and Rai- mundo Oliveira for logistic support. Finan- cial support was provided by CNPq (proc. 464545/00-5) and FAPESP (proc. O1/ 14449-2), Literature Cited Burns, J. R., S. H. Weitzman, K. R. Lange, & L. R. Malababa. 1998. Sperm ultrastructure in char- acid fishes. Pp. 235-244 in L. R. Malabarba, R. VOLUME 117, NUMBER 3 E. Reis, R. P. Vari, Z. M. S. Lucena and C. A. S. Lucena, eds., Phylogeny and classification of neotropical fishes. Porto Alegre, Edipucrs, 603 Pp. , S. H. Weitzmann, and L. R. Malabarba. 1997. Insemination in eight species of cheirodontine fishes (Teleostei: Characidae: Cheirodonti- nae).—Copeia 1997(2):433—438. Leviton, A. E., R. H. Gibbs Jr., E. Heal, & C. E. Daw- son. 1985. Standards in herpetology and ichthy- ology, part I. Standard symbolic codes for in- stitutional resource collections in herpetology and ichthyology.—Copeia 1985:802-832. Lima, FE C. T., & P. Gerhard. 2001. A new Hyphes- sobrycon (Characiformes: Characidae) from Chapada Diamantina, Bahia, Brazil, with notes on its natural history.—Ichthyol. Expl. Fresh- waters 12(2):105—114. Malabarba, L. R. 1998. Monophyly of the Cheirodon- tinae, characters and major clades (Ostariophy- si: Characidae). Pp. 193—233 in L. R. Malabar- ba, R. E. Reis, R. P. Vari, Z. M. S. Lucena and C. A. S. Lucena, eds., Phylogeny and classifi- cation of neotropical fishes. Porto Alegre, Edi- pucrs, 603 pp. , ©. H. Weitzman, AND J. R. Burns. 1998. Compsurini. Pp. 216-220 in Monophyly of the Cheirodontinae, characters and major clades (Ostariophysi: Characidae). Pp. 193-233 in L. R. Malabarba, R. E. Reis, R. P Vari, Z. M. S. Lucena and C. A. S. Lucena, eds., Phylogeny 329 and classification of neotropical fishes. Porto Alegre, Edipucrs, 603 pp. , & S. H. Weitzman. 1999. A new genus and species of South American fishes (Teleostei: Characidae:Cheirodontinae) with a derived cau- dal fin, including comments about inseminating cheirodontines. Proceedings of the Biological Society of Washington 112(2):410—432. , & . 2000. A new genus and species of inseminating fish (Teleostei: Characidae: Chei- rodontinae: Compsurini) from South America with uniquely derived dermal papillae on caudal fin.—Proceedings of the Biological Society of Washington 113(1):269—283. Menezes, N. A., S. H. Weitzman, & J. R. Burns. 2003. A systematic review of Planaltina (Teleostei: Characiformes: Characidae: Glandulocaudinae: Diapomini) with a description of two new spe- cies from the upper rio Parana, Brazil. Proceed- ings of the Biological Society of Washington 116:557—600. de Pinna, M. 1991. Concepts and tests of homology in the cladistic paradigm.—Cladistics 1991(7): 367-394. Weitzman, S. H., N. A. Menezes, J. R. Burns, and H.- G. Evers. 2004. A new genus and species of inseminating characid fish from the upper rio Xingu and rio Tapajos, Brazil, (Teleostei: Char- aciformes: Characidae) with comments on re- lationships among inseminating characids. Neo- tropical Ichthyology (in press). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):330—338. 2004. Astyanax biotae, a new species of stream fish from the Rio Paranapanema basin, upper Rio Parana system, southeastern Brazil (Ostariophysi: Characiformes: Characidae) Ricardo M. C. Castro and Richard P. Vari (RMMC) Laborat6rio de Ictiologia de Ribeirao Preto, Departamento de Biologia da Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto, Universidade de Sao Paulo, Avenida Bandeirantes 3900, 14040-901, Ribeirao Preto, SP. Brazil, e-mail: rmcastro @ffclrp.usp.br; (RPV) Vertebrate Zoology Section, Division of Fishes, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A., e-mail: vari.richard@nmnh.si.edu Abstract.—Astyanax biotae, a new species of characid, is described from a first-order stream in the Rio Paranapanema basin, upper Rio Parana system, in the interior of the state of Parana, southeastern Brazil. The species differs from its congeners in that region in a combination of morphometric and pigmentary features. Resumo.—Astyanax biotae, uma nova éspecie de caracideo € descrita de um riacho de primeira ordem da bacia do Rio Paranapanema, sistema do Alto Rio Parana, interior do Estado do Parana, sudeste do Brasil. A espécie descrita difere das demais espécies do género Astyanax ocorrentes na mesma regiao por uma combinagao de caracteres morfométricos e pigmentares. Astyanax Baird & Girard includes nearly 90 nominal species of neotropical characid fishes distributed from the southwestern United States to Argentina (Lima et al. 2003:106). The numerous nominal species assigned to Astyanax, in conjunction with the lack of a comprehensive treatment of the genus subsequent to Eigenmann (1921, 1927), often makes the identification of spe- cies problematic. Furthermore, Astyanax as now delimited is likely non-monophyletic, and various species encompassed in the ge- nus as traditionally defined (.e., characids with two rows of teeth in the upper jaw and with the inner tooth row consisting of five teeth) have been generically reassigned in recent years (e.g., Zanata 1997). This uncertainty applies even in regions such as the upper Rio Parana that until re- cently had been thought to be well known ichthyologically. Evidence from a series of fish groups (Britski & Langeani 1988; Me- nezes 1988; Vari 1988; Weitzman et al. 1988; Langeani 1990; Menezes 1996a, 1996b; Castro & Casatti 1997) demon- strates that the Rio Parana system upstream from the now submerged Sete Quedas Falls is an area of endemism (see Castro et al. 2003), a phenomenon likely correlated with the formidable barrier to fish migration pre- sented, until recently, by those falls. The numerous streams and headwaters that con- tribute to the large rivers of this system are inhabited primarily by fish species of small body sizes (mostly less than 12 cm in stan- dard length). Such small-sized species con- stitute at least 50% of the described fresh- water fish species of South America and typically demonstrate a high degree of geo- graphic endemism (Castro 1999). Such spe- cies are highly dependent on riparian veg- etation for food, shelter, and reproduction (see Bohlke et al. 1978; Lowe-McConnell 1987), but those habitats are threatened by a number of anthropogenic activities, most notably deforestation and the extensive use VOLUME 117, NUMBER 3 of fertilizers and pesticides in intensive ag- ricultural practices (see Lowe-McConnell 1975, 1987; Menezes et al. 1990; Sabino & Castro 1990; Aratyo Lima et al. 1995; Cas- tro & Menezes 1998). The lacunae in our understanding of the fish diversity within the upper Rio Parana basin and the possibility of extirpation of as-yet unrecognized species is clearly dem- onstrated by the species of Astyanax in that basin. In their comprehensive overview of the then-known species of Astyanax in the upper Rio Parana basin, Garuti and Britski (2000) recognized seven species of the ge- nus within that river system. Nonetheless, recent collecting efforts in that basin re- vealed at least two undescribed species of Astyanax, one of which is known only from a narrow first-order stream running through a narrow gallery forest that is a remnant of the originally widespread subtropical me- sophytic forest of that region. This species, which may be in danger of extinction, is described herein. Material and Methods Measurements are given as proportions of standard length (SL) except for subunits of the head that are presented as proportions of head length. Lateral-line scale counts in- clude all pored scales along that series, in- cluding scales posterior to the hypural joint. In fin-ray counts, lower-case Roman nu- merals indicate unbranched rays, and Ara- bic numerals indicate branched rays. The last anal-fin rays that are joined at the base were counted as one element. Counts for the holotype are indicated in square brack- ets in the text. Measurements were made following the methods outlined in Fink & Weitzman (1974:1—2) with the addition of head height measured at the vertical at the base of the supraoccipital spine. Cleared and stained specimens were prepared fol- lowing a modification of the method out- lined by Taylor & Van Dyke (1985). Ver- tebral counts include the four vertebrae as- sociated with the Weberian apparatus. 33) Stomach contents were analyzed on eight specimens (37.5 to 52.5 mm SL) using the methods of frequency of occurrence and percent composition described by Bowen (1992) and Hynes (1950), respectively. The food items were grouped in broad taxonom- ic or ecological categories reflecting their origins, with aquatic insects and algae con- sidered autochthonous and terrestrial in- sects, arachnids, and vascular plants allo- chthonous. The following institutional abbreviations are used: LIRP—Laborat6rio de Ictiologia de Ribeirao Preto, Departamento de Biolo- gia da Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto, Universidade de Sao Paulo, Ribeirado Preto, Brazil; MZUSP—Museu de Zoologia da Univer- sidade de Sao Paulo, Sao Paulo, Brazil; and USNM—National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C., U.S.A. Astyanax biotae, new species Fig. 1, Table 1 Astyanax sp. 2. Castro et al., 2003:13, 20, 21, fig. 6.6 [Brazil, Parana, Rio Parana- panema basin; ecology]. Holotype.—LIRP 4009, 49.8 mm SL; Brazil, Parana State, upper Rio Parana sys- tem, Rio Paranapanema basin, Municipio de Diamante do Norte, Fazenda Agua Mole, Cérrego Agua Mole (22°38’31.7'S, 52°48'59.0"W); collected by Ricardo M. C. Castro, Hertz EK Santos, Ricardo C. Benine, Katiane M. Ferreira, and Flavio C. T. Lima, 7 August 2000 (station PPAO29). Paratypes.—LIRP 2734, 15 specimens, 27.5—52.3 mm SL; LIRP 4021, 2 cleared and stained specimens, 51.3—52.5 mm SL; USNM 373492, 15 specimens, 31.2—52.2 mm SL; MZUSP 79807, 10 specimens, 32.4—45.6 mm SL; LIRP 4276, 34 speci- mens, 33.0—47.4 mm SL; collected with holotype. Diagnosis.—Astyanax biotae is readily distinguished from all congeners in the up- per Rio Parana basin in having the terminus 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. system, Rio Paranapanema basin, Municipio de Diamante do Norte, Fazenda Agua Mole, Corrego Agua Mole (22°38'31.7"S, 2°48'59.0"W). of the base of the dorsal fin situated along the vertical through the base of the first or second branched anal-fin ray, versus through the origin of the anal fin (A. fas- clatus, A. trierythropterus) or in the area of the vent (A. altiparanae, A. cf. eigeman- Astyanax biotae, new species, holotype, LIRP 4009, 49.8 mm SL. Brazil, Parana, upper Rio Paranda — niorum, A. paranahybae, A. scabripinnis, and A. schubarti). Furthermore, A. biotae has a distinct overall reticulate pattern formed by dark pigmentation on the ex- posed portion of the scales versus the lack of such a pigmentation pattern in all of the Table 1.—Morphometric values for holotype and 30 paratypes of Astyanax biota. Standard length is expressed in millimeters; measurements 1-15 as percentages of standard length; 16—21 as percentages of head length. Standard length 1. Greatest body depth 2. Snout to dorsal-fin origin 3. Length of base of dorsal fin 4. Posterior terminus of dorsal fin to adipose fin 5. Posterior terminus of dorsal fin to caudal-fin base 6. Snout to origin of pelvic fin 7. Snout to anus 8. Snout to origin of anal fin 9. Length of base of anal fin 10. Length of caudal peduncle 11. Length of longest dorsal-fin ray 12. Length of first pectoral-fin ray 13. Length of first pelvic-fin ray 14. Least depth of caudal peduncle 15. Head length 16. Head height 17. Snout length 18. Gape width 19. Orbital diameter 20. Postorbital head length 21. Interorbital width Holotype Paratypes Mean SD 49.8 27.5—52.3 44.20 6.11 34.7 34.7-41.8 38.68 1.83 54.5 50.4—56.9 53.86 32 13.3 12.3-15.1 IBS) 0.69 24.5 19.3—24.5 2290) 1.30 38.4 35.6-41.9 37.39 1.30 49.6 45.7-49.8 48.33 0.98 60.0 54.6-61.3 58.35 1.63 65.1 61.4-66.8 64.07 1.44 31.6 29.1-39.6 32.09 1.95 10.4 9.4-12.8 11.31 1128) AY <3) 26.8—30.8 28.21 1.19 21.3 19.2—24.4 22.01 1.25 16.5 16.0-19.2 17.44 0.82 12.0 10.9-13.7 12.40 0.55 Ql 25.4-28.7 27.23 0.86 94.2 94.2—113.5 102.15 4.40 26.8 23.5-29.3 25.86 1.53 29.0 26.3-34.8 30.59 1.88 31.9 31.9-40.0 34.85 2.17 42.8 35.143.6 37/2 2.13 37.0 34.8-40.9 38.12 1.70 VOLUME 117, NUMBER 3 other species of Astyanax that occur in the upper Rio Parana basin. Astyanax biotae and A. paranahybae can also be distin- guished by the difference in their relative body heights (approximately 35—42% of SL versus 25%, respectively). Description.—Morphometrics of holo- type and paratypes presented in Table 1. Body relatively deep, less so in individuals of less than 30 mm SL; greatest body depth located along vertical through insertion of pelvic fin. Dorsal profile of head distinctly convex from margin of upper lip to vertical through posterior nostril, straight to very slightly convex from that point to tip of su- praoccipital spine. Dorsal profile of body slightly to moderately convex from rear of head to origin of dorsal fin, straight and posteroventrally slanted along base of dor- sal fin, straight to slightly convex from pos- terior terminus of base of dorsal fin to ad- ipose fin, and slightly concave along caudal peduncle. Slight middorsal ridge present along predorsal region of body. Body trans- versely rounded overall dorsally, but some- what flattened middorsally between poste- rior terminus of base of dorsal fin and adi- pose fin. Ventral profile of head strongly convex anteriorly and then slightly convex as far as vertical through posterior margin of eye. Ventral profile of body convex to insertion of pelvic fin, nearly straight but slightly posteroventrally aligned from that point to origin of anal fin, straight to slight- ly convex and posterodorsally slanted along base of anal fin, straight to slightly concave along caudal peduncle. Head obtusely rounded anteriorly in lat- eral profile; mouth terminal, albeit very slightly upturned. Upper jaw with maxilla distinctly posteroventrally angled and ex- tending under orbit as far as vertical through anterior margin of pupil. Nostrils of each side very close together; anterior opening circular, posterior crescent-shaped. Eye relatively large and without distinct ad- ipose eyelid. Median fronto-parietal fonta- nel extending from mesethmoid to supra- occipital spine; width of fontanel approxi- 685) mately one-fourth of interorbital distance. Infraorbital series complete with third infra- orbital by far the largest. All infraorbitals carrying laterosensory canal segments proximate to inner margin of orbital rim. Supraorbital absent. Branchiostegal rays four. Gill-rakers long and _ setiform; 6+1+11 rakers on outermost gill-arch of 52.5 mm SL cleared and stained specimen. Description of dentition based on two cleared and stained specimens. Teeth on premaxilla in two rows, with teeth of inner row larger. Inner row with five teeth. Sym- physeal tooth of inner series quadricuspid and more elongate than other teeth. Second tooth more massive and pentacuspid. Re- maining teeth pentacuspid, with third and fourth teeth somewhat smaller than second tooth, and fifth tooth distinctly smaller than all other teeth in series. Outer row of teeth on premaxilla consisting of four tricuspid teeth arranged in regular series with size of teeth gradually decreasing laterally. Fourth tooth of outer tooth row separated from third tooth by distance twice that separating other teeth of series. Maxilla with single tri- cuspid or pentacuspid tooth. Dentary with eight to 10 teeth. Anterior five dentary teeth pentacuspid and arranged in single row. First four dentary teeth massive and fol- lowed by much smaller fifth tooth. Anterior five dentary teeth followed by gap and then three to five very small, elongate, conical teeth. Scales cycloid, relatively large, and firm- ly implanted. Lateral line decurved anteri- orly and then nearly straight along midla- teral line, completely pored from supra- cleithrum to base of caudal fin and followed by apparently unossified tubular extension running along membrane between middle rays of caudal fin. Lateral line scales 32 to 35 [34]; scales in transverse series from or- igin of dorsal fin to lateral line 6 or 7 [6]; scales in transverse series from insertion of pelvic fin to lateral line 4 or 5 [4]; scales in transverse series from origin of anal fin to lateral line 4 or 5 [5]; scales along mid- dorsal line between tip of supraoccipital 334 process and origin of dorsal fin 10 to 14 [11]; scales along mid-dorsal line between posterior termination of base of dorsal fin and adipose fin 8 to 11 [9]; horizontal scale rows around caudal peduncle 13 to 15 [14]. Vertebrae 32(3), 33 (17), or 34 (7) [33]. Dorsal-fin rays 11,9 [11,9]; anal-fin rays 11 to iv,22 to 26 [111,24]; total number of anal- fin rays 24 to 30 [28]; pectoral-fin rays 1,10 to 12 [1,12]; pelvic-fin rays typically 1,7, with i,6 in three specimens, and 1,4 in both fins in one apparently anomalous individual [1,7]; some specimens with anteriorly di- rected hooks on dorsal surface of pelvic-fin rays in adpressed fin; principal caudal-fin rays 10/9 [10/9]. Dorsal-fin margin distally rounded to slightly truncate; first unbranched ray ap- proximately 40% length of second un- branched ray. Dorsal-fin origin situated at vertical approximately at middle of SL. Or- igin of adipose fin located slightly anterior of vertical through posterior terminus of base of anal fin. Pectoral fin relatively well developed, profile distinctly acute in ad- pressed fin. Tip of pectoral fin extending to, or falling slightly short of, vertical through insertion of pelvic fin. Profile of expanded pelvic fin pointed, with lateral rays longest. Insertion of pelvic fin located distinctly an- terior to vertical through origin of dorsal fin. Tip of adpressed pelvic fin extending to origin of anal fin. Distal margin of anal fin ranging from somewhat concave to straight, with third unbranched and first and second branched rays longest and subequal or first through third branched rays longest; sub- sequent branched rays gradually decreasing in length. Caudal fin forked with lobes rounded. Color in life.—Description based on col- or transparencies of live holotype (see also Castro et al., 2003:fig 6.6). Overall colora- tion silvery-brownish with silvery high- lights on scales, particularly in abdominal region. Basal region of exposed portions of scales darker, particularly along regions slightly dorsal of midlateral line. Iris, an- teroventral portions of infraorbital region, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lower jaw, and ventral regions of head sil- very. Iris with green highlights. Dark pig- mentation as in preserved specimens. Coloration in alcohol.—Overall ground color of specimens fixed in formalin yel- lowish-brown on body, with guanine still present on ventral portion of head and on abdomen. Snout and dorsal portion of head relatively dark. Middorsal and immediately adjoining portions of body dark. Distinct, ventrally attenuated humeral spot extending from approximately two scales ventral of dorsal midline to about one scale dorsal of horizontal through insertion of pectoral fin. Scales of lateral surface of body posterior of humeral mark with dark pigmentation field on exposed portion of each scale. Dark spots forming irregular, discontinuous dark stripe along midlateral surface of body. Caudal peduncle with distinct, anteriorly-at- tenuating, dark mark. Dorsal, anal, and caudal fins with inter- radial membranes covered with small dark chromatophores. Dorsal fin with dark pig- mentation on interradials more prominent on distal one-half of central rays of fins, particularly in larger individuals. Dark pig- mentation on caudal fin particularly well developed along middle rays of fin. Anal fin with dark pigmentation distinctly more developed on distal half of fin in some in- dividuals; otherwise pigmentation of uni- form intensity across fin. Adipose fin often freckled with small dark spots. Pectoral and pelvic fins with small dark spots along fin- ray margins and on membranes. Etymology.—The species name, biotae, 1s in recognition of the important pioneering role of the ““-BIOTA/FAPESP—The Virtual Biodiversity Institute Program” (www. biota.org.br/) in the inventory, conserva- tion, and sustainable use of the biodiversity resources of the State of Sao Paulo, Brazil. This special research program of the Fun- dacgao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP) supported the collect- ing efforts that yielded all known speci- mens of the species. VOLUME 117, NUMBER 3 Fig. 2. Common name.—Brazil, Parana, Dia- mante do Norte: ““Lambari’” a name also generically applied to all other species of Astyanax and other small characids in southeastern Brazil. Distribution.—Known only from the type locality in the region called the Pontal do Paranapanema. Ecology.-The sample of Astyanax biota was collected during the winter dry season in the Corrego Agua Mole (see Castro et al. 2003:fig. 5.6), a first-order stream running through a narrow, not very dense gallery forest within an extensive cattle grazing area, at an elevation of approximately 300 m above sea level. This location lies within what was originally an extensive subtropi- cal mesophytic forest in southern and southeastern Brazil (Huek & Seibert 1981). The width of the stream varied between 0.7—1.0 m and the depth between 0.17—0.40 m, with a current speed of approximately 0.2 m.s''. The marginal vegetation was dominated by grasses of the family Cyper- acea (Fimbristylis sp.) and ferns (Pterydo- phyta) of the family Polypodiaceae. Water temperature was 18.6°C; pH 8.7; dissolved oxygen 10.6 mg.l-'; conductivity 17 335 Son, ae SR ae Map of the upper Rio Parana basin showing type locality for Astyanax biotae (star) and major river systems in the basin; A = Rio Paranapanema; B = Rio Parana, C = Rio Uruguay, D = Rio Tieté. S.cm~!; and horizontal water transparency 0.4 m. Collecting efforts along an 100 m long stretch of the stream yielded seven fish spe- cies in addition to Astyanax biotae: Calli- chthys callichthys, Corydoras aeneus, Crenicichla britskii, Gymnotus cf. inaequil- abiatus, Gymnotus cf. sylvius, Rhamdia quelen, and Phalloceros caudimaculatus. Astyanax biotae was the most abundant species in the sample (approximately 70% of the 110 specimens in the sample) and the second largest contributor to the fish bio- mass (approximately 31% of the total col- lected fish biomass) after Rhamdia quelen (approximately 53%). These values clearly indicate the ecological importance of Asty- anax biotae at this site. Although our food analysis results are derived from a single collecting event, the stomach content analysis of eight individ- uals (37.5 to 52.5 mm SL; one with an empty stomach) clearly demonstrates that Astyanax biotae feeds primarily on arthro- pods (approximately 80% of the diet com- position), with debris and seeds of vascular plants (approximately 15%) and filamen- tous algae (approximately 6%) significantly 336 less important in the diet. Aquatic insects (mostly aquatic larvae of the Chironomidae followed in order by aquatic larvae of the aquatic Coleoptera, aquatic larvae of the Plecoptera and Trichoptera (equal amounts of each), nymphs of the Ephemeroptera, na- iads of the Odonata, and a single adult of the aquatic Hemiptera) and terrestrial in- sects (primarily worker ants, Formicidae; followed by worker termites, Isoptera, and adult terrestrial Coleoptera) account for ap- proximately 30% of the ingested arthro- pods, followed by distinctly lower numbers of arachnids (mostly spiders, Aranae, and a pseudoscorpion). Overall, approximately 55% of the items in the stomachs of A. bio- tae were allochthonous and 45% were au- tochthonous, a clear indication of the im- portance of the riparian vegetation as a food source for this species of Astyanax. One of the examined specimens, a 52.5 mm SL fe- male (USNM 373492) with a greatly dis- tended abdomen was found to contain ap- proximately 350 roundish, well-developed, deep yellow oocytes 0.7—0.8 mm in diameter. Comparative material examined.—Asty- anax altiparanae, LIRP 35, 126 specimens, 43.0-80.1 mm SL; USNM 373491, 10 specimens, 41.1—79.9 mm SL. Astyanax cf. eigenmanniorum, LIRP 3401, 10. speci- mens, 55.0—70.8 mm SL; USNM 373495, 10 specimens, 48.5—68.3 mm SL. Astyanax fasciatus, LIRP 32, 28 specimens, 42.0— 93.5 mm SL; USNM 373493, 10 speci- mens, 45.7—83.8 mm SL. Astyanax schu- barti, MZUSP 4263, holotype, 82.9 mm SL; MZUSP 4264, 1 paratype, 90.4 mm SL. Astyanax scabripinnis, LIRP 124, 562 specimens, 19.1-75.0 mm SL; USNM 373494, 10 specimens, 36.5—74.8 mm SL. Astyanax trierythriopterus, LIRP 2017, 138 specimens, 26.3—41.2 mm SL; USNM 373496, 10 specimens, 27.8—41.1 mm SL. Acknowledgments The specimens of Astyanax biotae that served as the basis for this description were collected during a collaborative LIRP- MZUSP expedition supported by FAPESP PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Fundagao de Amparo a Pesquisa do Estado de Sao Paulo) within the “BIOTA/FA- PESP—The Virtual Biodiversity Institute Program” (www.biota.org.br/) through the Thematic Project “Fish diversity of the headwaters and streams of the upper Parana River system in the State of Sao Paulo, Bra- zil’”’ (FAPESP grant No. 98/05072-8, Ri- cardo M. C. Castro (LIRP) principal inves- tigator). Research associated with this pro- ject was supported by that grant, the Neo- tropical Lowland Research Program of the International Environmental Sciences Pro- gram of the Smithsonian Institution, and the PRONEX Project ““Conhecimento, conser- vacao e utilizag¢ao racional da diversidade da fauna de peixes do Brasil” (FINEP/ CNPq grant No. 661058/1997-2). The first author is a Conselho Nacional de Desen- volvimento Cientifico e Tecnoldgico do Brasil researcher (grant No. 301309/91-4). The success of the collecting effort was as- sured by the assistance of H. FE Santos, K. M. Ferreira and R. C. Benine (all of LIRP), and FE C. T. Lima (MZUSP). H. E Santos (LIRP) assisted with the preparation of the clear and stained specimens, the stomach extractions for diet analysis and the prepa- ration of the photograph of the holotype. A. L. A. Melo (LIRP) processed the speci- mens. A. C. Ribeiro (LIRP) produced the distribution map and K. M. Ferreira (LIRP) helped with the identification of the stom- ach contents. This paper was greatly im- proved by the suggestions and criticisms of S. H. Weitzman and C. J. Ferraris, Jr. Literature Cited Aratjo-lima, C. A. R. M., A. A. Agostinho, & N. E Fabré. 1995. Trophic aspects of fish communi- ties in Brazilian rivers and reservoirs. Pp. 105— 136 in J. G. Tundisi, C. E. M. Bicudo and T. M. Tundisi, eds., Limnology in Brazil. Acade- mia Brasileira de Ciéncias and Sociedade Bras- ileira de Limnologia, Rio de Janeiro, 376 pp. Bohlke, J., S. H. Weitzman, & N. A. Menezes. 1978. Estado atual da sistematica de peixes de agua doce da América do Sul.—Acta Amazonica 8: 657-677. Bowen, S. H. 1992. Quantitative description of the diet. Pp. 325-336 in L. A. Nielsen and D. L. VOLUME 117, NUMBER 3 Johnson, eds., Fisheries techniques. American Fisheries Society, Blacksburg, 468 pp. Britski, H. A. & E Langeani. 1988. Pimelodus par- anaensis, sp. N., um novo Pimelodidae (Pisces, Siluriformes) do Alto Parana, Brasil.—Revista Brasileira de Zoologia 5:409—417. Castro, R. M. C. 1999. Evolugao da ictiofauna de ria- chos sul-americanos: padr6es gerais e possiveis processos causais. Pp.139—155 in E. P. Cara- maschi, R. Mazzoni, C. R. S. FE Bizerril and P. R. Peres-Neto, eds., Ecologia de peixes de ria- chos: estado atual e perspectivas. Oecologia Brasiliensis VI, Rio de Janeiro, 260 pp. Castro, R. M. C., & L. Casatti. 1997. The fish fauna from a small forest stream of the upper Parana River Basin, southeastern Brazil.—tIchthyolog- ical Explorations of Freshwaters 7:337—352. , L. Casatti, H. E Santos, K. M. Ferreira, A. C. Ribeiro, R. C. Benine, G. Z. P. Dardis, A. L. A. Melo, R. Stopiglia, T. A. Abreu, E A. Bock- mann, M. Carvalho, E Z. Gibran, & E C. T. Lima. 2003. Estrutura e composigao da ictio- fauna de riachos do Rio Paranapanema, sudeste e sul do Brasil.—Biota Neotropica 3(1):1—31 {http://www/biotaneotropica.org.br/v3n1/pt/]. , & N. A. Menezes. 1998. Estudo diagnéstico da diversidade de peixes do Estado de Sao Pau- lo. Pp. 1-13 in R. M. C. Castro, ed., C. A. Joly and C. E. M. Bicudo, orgs., Biodiversidade do Estado de Sao Paulo, Brasil: sintese do conhe- cimento ao final do século XX, vol. 6. Verte- brados. WinnerGraph—FAPESP, Sao Paulo, 71 pp. Eigenmann, C. H. 1921. The American Characidae.— Memoirs of the Museum of Comparative Zo- ology 43(4):1-102. . 1927. The American Characidae.—Memoirs of the Museum of Comparative Zoology 43(4): 311-428. Fink, W. L., & S. H. Weitzman. 1974. The so-called Cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Characi- dae).—Smithsonian Contributions to Zoology 172:1—46. Garuti, V. & H. A. Britski. 2000. Descrigaéo de uma espécie nova de Astyanax (Teleostei: Characi- dae) da bacia do Alto Parana e considerag6es sobre as demais espécies do género.—Comun- icagdes do Museu de Ciéncias da PUCRS, Porto Alegre, Série Zoologia 13:65—88. Hynes, H. B. N. 1950. The food of fish-water stick- lebacks (Gasterosteus aculeatus and Pygosteus pungitius), with a review of methods used in studies of food fishes.—Journal of Animal Ecology 19:36—57. Huek, K. & P. Siebert. 1981. Vegetationskarte von Siti- damerica. Band Ia. Fischer, Sttutgart, 90 pp. Langeani, F 1990. Reviséo do género Neoplecostomus 337 Eigenmann & Eigenmann, 1888, com a descri- ¢ao de quatro novas espécies do sudeste brasi- leiro (Ostariophysi, Siluriformes, Loricari- idae).—Comunicagdes do Museu de Ciéncias da-PUCRS, Porto Alegre, Série Zoologia 3:3— Zi, Lima, F C. T. et al., 2003. Genera incertae sedis in Characidae. Pp. 106-169 in R. E. Reis, S. O. Kullander and C. J. Ferraris, Jr., orgs., Check list of the freshwater fishes of South and Central America. Edipucrs, Porto Alegre, Brazil, 729 Pp. Lowe-McConnell, R. H. 1975. Fish communities in tropical freshwaters: their distribution, ecology and evolution. Longman Publishers, New York, 337 pp. . 1987. Ecological studies in tropical fish com- munities. Cambridge University Press, Cam- bridge, 382 pp. Menezes, N. A. 1988. Implication of the distribution patterns of the species of Oligosarcus (Teleos- tei, Characidae) from central and southern South America. Pp. 295-304 in P. E. Vanzolini and W. R. Heyer, eds., Proceedings of a work- shop on neotropical distribution patterns. Aca- demia Brasileira de Ciéncias, Rio de Janeiro, 488 pp. 1996a. Conservac¢ao da diversidade da ictiofau- na da Bacia Parana-Paraguai-Uruguai. Anais XV Congresso Panamericano de Ciéncias Ve- terindrias, Campo Grande, MS, 4 pp. 1996b. Methods for assessing freshwater fish diversity. Pp. 289-312 in C. E. M. Bicudo and N. A. Menezes, eds., Biodiversity in Brazil. CNPq, Sao Paulo, 326 pp. , Castro, R. M. C., S. H. Weitzman, & M. J. Weitzman. 1990. Peixes de riacho da Floresta Costeira Atlantica Brasileira: um conjunto pou- co conhecido e ameagado de vertebrados. Pp. 290-295 in S. Watanabe, coordinator, Il Sim- posio de ecossistemas da Costa Sul e Sudeste Brasileira: Estrutura, fun¢ao e manejo. Acade- mia de Ciéncias do Estado de Sao Paulo, vol. 1. 448 pp. Sabino, J., & R. M. C. Castro. 1990. Alimentagao, per- {odo de atividade e distribuigao espacial dos peixes de um riacho da floresta Atlantica (sud- este do Brasil).—Revista Brasileira de Biologia 50:23-36. Taylor, W.R., & G. Van Dyke. 1985. Revised proce- dures for staining and clearing small fishes and other vertebrates for bone and cartilage study.— Cybium 9(2):107-119. Vari, R. P. 1988. The Curimatidae, a lowland neotrop- ical fish family (Pisces: Characiformes); distri- bution, endemism, and phylogenetic biogeog- raphy. Pp. 343-377 in P. E. Vanzolini and W. R. Heyer, eds., Proceedings of a workshop on 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON neotropical distribution patterns. Academia and W. R. Heyer, eds., Proceedings of a work- Brasileira de Ciéncias, Rio de Janeiro, 488 pp. shop on neotropical distribution patterns. Aca- Weitzman, S. H., N. A. Menezes, & M. J. Weitzman. demia Brasileira de Ciéncias, Rio de Janeiro, 1988. Phylogenetic biogeography of the Glan- 488 pp. dulocaudinae (Teleostei: Characiformes, Char- Zanata, A.M. 1997. Jupiaba, um novo género de Te- acidae) with comments on distribution of the tragonopterinae com osso pélvico em forma de other freshwater fishes in eastern and south- espinho (Characidae, Characiformes).—Iherin- eastern Brazil. Pp. 379—427 in P. E. Vanzolini gia, Série Zoologia, 83:88—106. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):339-345. 2004. Tetragonopterus lemniscatus (Characiformes: Characidae), a new species from the Corantijn River basin in Suriname Ricardo C. Benine, Gabriela Zanon Pelicao, and Richard P. Vari (RCB) Laboratorio de Ictiologia de Ribeirao Preto, Departamento de Biologia, FFCLRP- Universidade de Sao Paulo, Av. Bandeirantes, 3900, 14040-901, Ribeirao Preto, Sao Paulo, Brazil, e-mail: rbenine@hotmail.com; (GZPD) Laboratério de Ictiologia de Ribeirao Preto, Departamento de Biologia, FFCLRP- Universidade de Sao Paulo, Av. Bandeirantes, 3900, 14040-901, Ribeirao Preto, Sao Paulo, Brazil, e-mail: gzpd@usp.br; (RPV) Division of Fishes, Smithsonian Institution, RO. Box 37012, National Museum of Natural History, WG-14, MRC 159, Washington, D.C. 20013-7012, U.S.A., e-mail: vari.richard@nmnh.si.edu Abstract.—Tetragonopterus lemniscatus, a new species of characid characi- form, is described from the Corantijn River basin in western Suriname. The species is readily distinguished from its congeners (7. argenteus, T. chalceus) by the presence of dark, longitudinal stripes positioned between adjacent scale rows of the lateral surface of the body. Resumo.—Tetragonopterus lemniscatus, uma nova espécie de caraciforme caracideo, é descrita de bacia do Rio Corantijn, oeste de Suriname. Esta espécie é€ prontamente distinguida de seus congéneres pela presenga de um padrao estriado de coloragao ao longo do corpo, formado por faixas escuras presentes entre as fileiras de escamas adjacentes. The Neotropical characid characiform genus Tetragonopterus is characterized ex- ternally by a relatively deep body with a transversely-flattened prepelvic region that is bordered laterally, particularly proximate to the pelvic-fin insertion, by distinctly-an- gled scales, a pronounced ventral curvature of the anterior portion of the lateral line, an anal fin with a long base, and a complete outer row of teeth on the premaxilla. Recent authors (e.g., Géry 1977:450; Reis 2003: 212) have recognized only two species of Tetragonopterus, T. argenteus and T. chal- ceus, but the examination of samples of the genus that originated in the Corantijn River basin of western Suriname revealed a third species of the genus, which we describe herein. Material and Methods Measurements are given in terms of stan- dard length (SL). Lateral-line scale counts include all pored scales along that series, in- cluding scales posterior to the hypural joint. In fin-ray counts, lower-case Roman numer- als indicate unbranched rays, and Arabic nu- merals indicate branched rays. The last anal- fin rays that are joined at the base were counted as one element. Morphometric and meristic data were taken following the pro- cedures outlined in Fink & Weitzman (1974). Individual meristic values in the de- scription are followed by their frequency in parentheses, with values for the holotype in- dicated in square brackets. Gill rakers counts were taken from specimens that were cleared and counterstained following the method of Taylor & Van Dyke (1985). Vertebral counts were taken via radiographs and include the four vertebra of the Weberian apparatus and the terminal centrum. Institutional abbreviations follow Leviton et al. (1985) with the addition of LIRP, La- 340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Tetragonopterus lemniscatus, new species, holotype, USNM 225366, 47.5 mm SL; Suriname, Nick- erie District, tributary to Sisa Creek. boratorio de Ictiologia de Ribeirao Preto, Tetragonopterus lemniscatus, Departamento de Biologia da RPRFCLRP, new species Universidade de Sao Paulo, Ribeirao Preto, Fig. 1, Table 1 SP, Brazil; and NZCS, National Zoological Collection of Suriname, Paramaribo, Suri- Holotype.—USNM 225366, adult male, name. 47.5 mm SL, Suriname, Nickerie District, Table 1.—Morphometric data for holotype and 11 paratypes of Tetragonopterus lemniscatus. Paratypes Holotype Range Mean Standard length (mm) 47.5 41.8-81.4 Percentages of standard length Greatest body depth Silat 46.2—53.9 50.0 Snout to dorsal-fin origin 52.1 51.3-53.6 52.4 Snout to pectoral-fin origin 28.9 27.6-29.9 29.1 Snout to pelvic-fin insertion 50.9 47.5-52.2 49.6 Snout to anal-fin insertion 67.9 64.6-69.4 67.2 Caudal peduncle depth 12.0 10.3—12.0 11.0 Caudal peduncle length 9.3 7.1-9.5 8.3 Pectoral-fin length 22.8 21.7-24.3 23.0 Pelvic-fin length 19.6 17.6—21.1 IS), 7/ Dorsal-fin length 32.8 32.6-44.3 37.2 Orbit to dorsal-fin origin 38.5 37.1-42.1 38.8 Head length 26.0 25.6-29.4 28.1 Head depth DOPED, 20.4—22.2 21.4 Percentages of head length Snout length 27.0 24.4—28.2 26.2 Upper jaw length 41.3 41.3-44.9 43.3 Horizontal orbital diameter 41.8 38. 147.0 42.9 Least interorbital width 38.6 29.6-38.6 32.9 VOLUME 117, NUMBER 3 tributary to Sisa Creek, north side of stream approximately 700 m downstream of cross- ing of road from Amotopo to Camp Geo- logie, approximately 3°42’'N, 57°42'W, R. P. Vari et al., 20 Sep 1980. Paratypes.—All collected in Suriname, Nickerie District. USNM 374750, 4 speci- mens, 42.0—46.6 mm SL. LIRP 4928, 2 specimens, 47.5—47.9 mm SL, cleared and counterstained, collected with holotype. USNM 225523, 2 specimens, 74.0—81.4 mm SL. LIRP 4929, 1 specimen, 79.8 mm SL, stream at km 212 of Amotopo to Camp Geology road, at Machine Park-Camp 212, approximately 3°50’N, 57°34’W, R. P. Vari et al., 15 Sep 1980. NZCS F7062, 1 spec- imen, 62.1 mm SL, formerly USNM 225320, stream entering Corantijn River, at approximately km 385, slightly N of Tiger Falls, approximately 4°00'N, 58°02’W, R. P. Vari et al., 16 Sep 1980. USNM 224367, 2 specimens, 48.4-60.1 mm SL, Kamp Kreek, 100 m N of turnoff to Camp Geol- ogy, approximately 4°49’N, 57°28’W, R. P. Vari et al., 13 Sep 1980. Diagnosis.—Tetragonopterus lemnisca- tus 1s distinguished from its two recognized congeners, 7. argenteus and T. chalceus, by the dark pigmentation on the lateral surface of the body (presence of dark, longitudinal stripes formed by pigmentation fields along the margins of the adjoining scale rows ver- sus the absence of dark stripes, respective- ly). Tetragonopterus lemniscatus further differs from T. argenteus in the number of median scales between the tip of the supra- occipital spine and the base of the first dor- sal-fin ray (8 versus 12—16, respectively). Description.—Morphometric data are summarized in Table 1. Overall body size moderate (41.8—81.4 mm in SL). Body pro- portionally deep. Greatest depth of body at origin of dorsal fin. Dorsal profile of head slightly concave above orbit. Each nostril located closer to anterior margin of orbit than to each other. Supraoccipital spine elongate, but tip of spine not extending be- yond vertical through posterior margin of opercle. 341 Dorsal profile of body convex from tip of supraoccipital spine to posterior terminus of base of dorsal fin; slightly convex from that point to end of base of adipose fin. Caudal peduncle profile concave both dor- sally and ventrally. Ventral profile of body convex from tip of lower jaw to beginning of caudal peduncle. Prepelvic region of body transversely flattened, with flattening more pronounced proximate to pelvic-fin insertion. Scales along lateral margins of flattened region immediately anterior to in- sertion of pelvic fin with distinct angle. Ob- tuse median keel extending from immedi- ately posterior of insertion of pelvic fin to urogenital opening. Mouth terminal. Premaxillary teeth in two rows. Outer premaxillary tooth row with 4 (5) or 5 (7) [5] tricuspid teeth with median cusps most developed. Inner row with 5 teeth with tetracuspid symphyseal tooth followed by two pentacuspid, and then two, rarely one, tricuspid teeth. Max- illa with 3 tricuspid teeth along anterodorsal portion of free anterior margin. Dentary with 4 (4) or 5(8) [5] pentacuspid teeth fol- lowed by series of small tricuspid teeth (Fig. 2). Dorsal-fin rays 11,9 (12) [41,9]. Distal mar- gin of dorsal fin straight. Adipose fin well- developed. Anal-fin rays iv,29 (3), iv,30 (5), or iv,31 (4) [iv,30]. Posterior unbranched and anterior branched anal-fin rays longest, with distal margin of remainder of fin mod- erately concave. Principal caudal-fin rays i,17,1 (12) [1,17,1]. Pectoral-fins rays 1,11 (2), 1,13 (7), or 1,14 (3) [1,13]. Tip of pec- toral fin extending beyond vertical through insertion of pelvic fin. Pelvic-fin rays 1,7 (12) [1,7]. Tip of pelvic fin reaching to base of first or second unbranched anal-fin ray in smaller individuals, barely falling short of base of first unbranched ray in larger specimens. Scales cycloid. Median scales anterior to origin of dorsal fin 8 (12) [8]. Lateral line distinctly ventrally curved anteriorly, with 33(3), 34(5), or 35(4) [33] pored scales. Rows of scales above lateral line to origin of dorsal fin 6 (11) or 7 (1) [6]. Rows of 342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Tetragonopterus lemniscatus, paratype, LIRP 4928, 47.5 mm SL. Premaxilla, maxilla, and lower jaw showing form of dentition; left side, lateral view. Scale bar = 1 mm. scales below lateral line to origin of anal fin 5 (11) or 6 (1) [5]. Scales around caudal peduncle 14 (11) [14]. Scale sheath formed of one row of scales overlaps basal portions of all but three or four posterior most anal- fin rays. Field of small scales covering base of caudal fin; scale field extending further distally on fins along its dorsal and ventral margins. Two cleared and stained specimens with 9 gill-rakers on upper limb and 13 gill-rak- ers on lower limb of first gill arch. Vertebrae 30 in all specimens including holotype. Coloration in life-—(Based on photo- graph of recently captured specimen from the Corantijn basin by third author). Overall coloration silvery, but somewhat purplish on portion of body dorsal to horizontal run- ning approximately through dorsal margin of orbit. Humeral spot faintly apparent. Dark stripes on lateral surface of body ap- parent, but slightly masked by overlying guanine. Infraorbital series, opercle, ventral VOLUME 117, NUMBER 3 Atlantic Ocean Silas Fig. 3. 56° Map of Suriname showing collecting sites of Tetragonopterus lemniscatus. Star = holotype locality French Guiana and dots = paratype localities (some symbols represent more than one locality or lot). portion of head, and most of body bright silver. Iris yellowish with indications of red dorsally. Fins dusky with yellowish cast. Color in alcohol.—Overall ground col- oration yellowish tan. Dorsal portion of head, jaws, nape, and portion of middorsal region of body anterior and posterior to base of dorsal fin distinctly darker. Posterior margins of scales with band of dark chro- matophores. Dark pigmentation particularly well-developed on dorsal and ventral por- tions of exposed regions of scales and form- ing undulating, narrow, horizontal stripes along regions of overlap of scale rows on lateral surface of body. Stripes extending on anterior portion of body from horizontal through base of insertion of pectoral fin to region about two scales ventral of origin of dorsal fin. Stripes ventral of horizontal through dorsal margin of orbit decurved ventrally anteriorly, with posterior portion of ventralmost stripes posterodorsally-an- gled in region over base of anal fin. Smaller individuals with 9 or 10 dark stripes appar- ent. Dorsalmost stripes becoming variably masked by overall darker pigmentation on dorsolateral region of body in larger spec- imens. Humeral region with indistinct, slightly posterodorsally-aligned bar in area above second and third scales of lateral line. Humeral spot becoming progressively less apparent in larger specimens. Caudal pe- duncle with large, rounded, dark spot con- tinuing posteriorly onto basal portions of middle caudal-fin rays. Short, irregular, hor- izontal stripes extending anteriorly from an- terior margin of spot in some larger indi- viduals. Median fins with small, dark chromato- phores overlying both membranes and rays 344 of rayed fins and lateral surface of adipose fin. Distal margin of caudal fin somewhat darker in some large specimens. Pectoral and pelvic fins hyaline or with few, small, dark chromatophores. Distribution.—Tetragonopterus lemnis- catus is only known from localities in the Corantijn River basin in western Suriname (hig33)): Habitat.—The holotype locality of Tetra- gonopterus lemniscatus was a black water rainforest stream with a limited amount of emergent vegetation and shadowed by overhanging trees. The stream had a mod- erate rate of water flow over a sand bottom with areas of detritus. Although all other population samples of the species were also collected in black water, some of the loca- tions were in full sun and at other collecting sites the current was swift. Some locations at which the species was collected had areas of clay, rock, or mud bottom. Etymology.—The species name, lemnis- catus, from the Latin for beribboned, is in reference to the series of dark stripes along the lateral surface of the body in this spe- cies. Remarks.—Tetragonopterus was first re- ported from Suriname by Kner (1859:38) who cited T. chalceus for that country. That citation may have been the basis for the in- clusion of the species in the Surinamese ichthyofauna by Eigenmann (1912:68; 1917:58) and for the report of the occur- rence of the species throughout the Guianas by Géry (1977:450). Ouboter and Mol (1993:146) reported T. chalceus from both the upper portion of the Corantijn River and from the Kabalebo River, the major right bank tributary to the Corantijn River. It is likely that the above citations, in particular that of Ouboter and Mol (1993), were based, at least in part, on 7. lemniscatus. Tetragonopterus has not been reported from elsewhere in Suriname, although 7. chal- ceus has been reported from a series of lo- calities across French Guiana including the Fleuve Maroni along the Surinamese- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON French Guiana border (Planquette et al., 1996:320). Comparative material.—Tetragonopterus chalceus: MNHN A9812 (holotype); MCP 15145 (4, 1 C&S); USNM 66293 (1); MZUSP 29820 (3) (1 C&S); MCP 14015 (1, C&S); MZUSP 40819 (2, 1 C&S). Te- tragonopterus argenteus: MNHN A-9807 (1); MZUSP 15570 (4, 2 C&S); MZUSP 5091 (2, 1 C&S); USNM 224789 (4). Acknowledgments Financial support was provided by FA- PESP (Proc. 00/1920-6, 98/05072-6, and 98/10337-0), PRONEX (Proc. 059/97), and the Neotropical Lowland Research Program of the Smithsonian Institution. We thank Oswaldo T. Oyakawa (MZUSP), Carlos A. Lucena and José Pezzi da Silva (MCP) for the loan of specimens. Sandra Raredon (USNM) prepared Fig. 1 and Alexandre C. Ribeiro (LIRP) Figs. 2 and 3. Patrice Pru- vost (MNHN) provided radiographs of specimens of Tetragonopterus deposited at that institution. Tatiana X. Abreu (LIRP) and Angela M. Zanata (MZUSP) examined and provided photographs of the holotype of 7. chalceus. Marcelo R. de Carvalho, Flavio A. Bockmann, and Ricardo M. C. Castro (LIRP) and Thomas B. Vari provid- ed valuable comments on earlier drafts of the manuscript. Literature Cited Eigenmann, C. H. 1912. The freshwater fishes of Brit- ish Guiana, including a study of the ecological grouping of species and the relation of the fauna of the plateau to that of the lowlands—Mem- oirs of the Carnegie Museum 5:xii + 578, 103 plates. 1917. The American Characidae.—Memories of the Museum of Comparative Zoology 53(1): 1-102. Fink, W. L. & S. H. Weitzman. 1974. The so-called cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Characi- dae).—Smithsonian Contributions to Zoology 172:1-42. Géry, J. 1977. Characoids of the World. TFH Publi- cations, Neptune City, New Jersey, U.S.A., 672 PP. VOLUME 117, NUMBER 3 34 Nn Kner, R. 1859. Zur Familie der Characinen, III. Folge Planquette, P, P. Keith, & P-Y. Le Bail. 1996. Atlas der ichthyologischen Beitrage——Denkschriften des poissons d’eau douce de Guyane, vol. 1. der Akademie der Wissenschaften, Wien 17: Muséum National d’histoire Naturelle and In- 137-182. stitut National de la Recherche Agronomique, Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. Paris, 429 pp. Dawson. 1985. Standards in herpetology and’ Reis, R. E. 2003. Subfamily Tetragonopterinae. Pp. ichthyology, part I. Standard symbolic codes for 212 in R. E. Reis, S. O. Kullander and C. J. institutional resource collections in herpetology Ferraris, Jr., orgs., Check list of the freshwater and ichthyology.—Copeia 1985(3):802—832. fishes of south and central America. Edipucrs, Ouboter, P. E., & J. H. A. Mol. 1993. The fish fauna Porto Alegre, Brazil, 729 pp. of Suriname. Chapter 8. Pp. 133-154 in P. E. Taylor, W. R. & G. C. Van Dyke. 1985. Revised pro- Ouboter, ed., Freshwater ecosystems of Surina- cedures for staining and clearing small fishes me. Kluwer Academic Publishers, The Nether- and other vertebrates for bone and cartilage.— lands. Cybium 9:107-119. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):346-362. 2004. Longipalpa saltatrix, a new genus and species of the meiofaunal family Nerillidae (Annelida: Polychaeta) from an anchihaline cave in Bermuda Katrine Worsaae, Wolfgang Sterrer, and Thomas M. Iliffe (KW) Zoological Museum, University of Copenhagen, Denmark, e-mail: kworsaae @zmuc.ku.dk; (WS) Bermuda Natural History Museum, Flatts FLBX, Bermuda, e-mail: museum.bamz @ibl.bm; (TMI) Texas A&M University at Galveston, Texas 77553-1675, U.S.A., e-mail: iliffet@tamug.edu Abstract.—A new genus and species of the meiofaunal family Nerillidae is described from an anchihaline cave in Bermuda. The description is based on studies of live animals with dissecting and light microscopes, as well as studies of fixed material with light and scanning electron microscopy. Longipalpa sal- tatrix, new species, differs from all other nerillids by possessing a pair of extremely long latero-ventral palps on the prostomium and a pair of ciliated pygidial lobes. It is further characterized by the combination of the following characters: three very short dorsal prostomial antennae, eight chaetigerous body segments, single parapodial cirri from segment three to eight, compound chae- tae, and hermaphroditism. With 48 species in 17 genera (300 wm— 2 mm in length), the Nerillidae is the largest meiofaunal family in the Polychaeta. The family has been a member of the now re- jected group “Archiannelida’ (e.g., Beau- champ 1910, Goodrich 1912). The Nerilli- dae are now believed to be more closely related to a macrofauna family among the Aciculata and possibly have evolved by progenesis (Westheide 1990, Westheide & Purschke 1996, Rouse & Fauchald 1997, Rouse & Pleijel 2001). Nerillids are nearly all marine and dis- tributed worlwide, from the intertidal to abyssal depths (3660 m—see Worsaae & Kristensen 2003). While most nerillids are members of the interstitial sand fauna, some have been found in mud, fine silt, organic debris, bacterial mats, green algae and mac- rophytes (Jouin & Swedmark 1965, Gelder 1974, Sterrer & Iliffe 1982, Saphonov & Tzetlin 1997, Miiller et al. 2001, Worsaae & Kristensen 2003). Several nerillids are known from caves: Leptonerilla prospera (Sterrer & Iliffe, 1982) was described from caves in Bermuda with fine silt; Mesone- rilla diatomeophaga Nunez, 1997 in Nunez et al. (1997) was described from a cave in Lanzarote with diatom carpets on lapilli; Nerilla marginalis TYilzer, 1970 was de- scribed from a marginal cave in Istra; and Troglochaetus beranecki Delachaux, 1921 has been reported from various freshwater caves, groundwater reservoirs and moun- tain rivers in Europe and Colorado, U.S.A. [see Morselli et al. (1995) for review]. Ne- rillids are known from all continents, except Antarctica, and the wide geographical dis- tribution as well as the diversity in habitats may well reflect an old evolutionary origin of the family. The anchihaline Bermudian caves are in- habited by a rich and diverse fauna, con- sisting primarily of crustaceans (Sket & Il- iffe 1980; Iliffe et al. 1983; Manning et al. 1986; Iliffe 1993, 1994, 2000). The most abundant stygobiont taxa are copepods and ostracods with 18 species each. Non-crus- taceans include two ciliates, two gastro- pods, and two annelids—the nerillid Lep- tonerilla prospera and the tubificid oligo- chaete Phallodriloides macmasterae (Er- VOLUME 117, NUMBER 3 séus, 1986). Although most of these species are endemic to Bermuda, many of them have cave-adapted congeners from the Ca- ribbean, Mediterranean and the Pacific. Sty- gobiont taxa with such highly anomalous distributions are believed to be Tethyan rel- icts. Materials and Methods The geology of Bermuda is particularly unusual in that the island consists of a mid- ocean volcanic seamount, capped with ma- rine and eolian limestone of Pleistocene age. The numerous inland caves of Ber- muda are totally within this limestone and often contain tidal, anchihaline pools that extend below sea level to a maximum depth of about 25 m. Surface waters in these pools are brackish, with salinity increasing with depth to approach fully marine levels at 3—5 m depths. The island and its caves have been profoundly affected by changes in sea level associated with Pleistocene gla- ciation. During the Ice Ages, sea level was as much as 100 m lower and the caves of Bermuda were all dry and air filled. Large speleothems (stalactites and stalagmites) formed at this time by rainwater percolating through the ground and dripping into the caves. As glacial periods ended, sea level rose and flooded a substantial portion of the caves such that they are only accessible with the use of specialized cave diving techniques (Iliffe 1993, 1994). The material was collected in Roadside Cave, a small anchihaline cave located in the Walsingham Tract of Bermuda (32°21'N, 64°43'W) on 15, 20 and 21 Jan 2002. A low entrance crawlway opens to a small dark chamber containing a narrow marine lake, which extends underneath a rock ledge and has a maximum depth of no more than 10 m. Surface salinity and tem- perature recorded on 28 Oct 1981 were 30%0 and 23°C, respectively. Tidal magni- tude in the pool is 57% of that in the open ocean, with a lag of 80 minutes. A number of other anchihaline stygobionts inhabit this 347 small pool, including the platycopioid co- pepods Antrisocopia prehensilis Fosshagen, 1985 in Fosshagen & Iliffe (1985) and Nan- ocopia minuta Fosshagen, 1988 in Fosshag- en & Iliffe (1988); the calanoid copepod Paracyclopia naessi Fosshagen, 1985 in Fosshagen & Iliffe (1985); the misophrioid copepod Speleophria bivexilla Boxshall & Iliffe, 1986; the bogidiellid amphipod Ber- mudagidiella bermudensis (Stock, Sket & Iliffe, 1987); the pseudoniphargid amphi- pod Pseudoniphargus grandimanus Stock, Holsinger, Sket & Iliffe, 1986; the halocy- prid ostracod Spelaeoecia bermudensis An- gel & Iliffe, 1987; the mictacean Mictocaris halope Bowman & Iliffe, 1985; and the gastropod Caecum troglodyta Moolenbeek & Faber, 1987 in Moolenbeek et al. (1987). Similar to the new species of polychaete here described, the copepods Antriscopia prehensile, Nanocopia minuta and Speleo- phria bivexilla are known only from this cave. Samples were collected with a conical plankton net with a diameter of 30 cm and a mesh size of 40 wm. Rocks and projec- tions below the water surface were covered with a thin layer of fine silt. Before the samples were taken, the surface layer was whirled up with hands, fins or loose stones from 0.5—6.5 meter’s depth and thereafter the net was dropped and dragged through the suspended material. More than 70 specimens were sorted out alive from the collected samples. Several of these were observed and video recorded alive with a Hitachi VK C-350 video cam- era mounted on a Wild M 420 Makroskop dissecting microscope. Fourteen animals were studied and photographed alive with an Olympus BX51 light microscope mount- ed with a digital camera (Olympus c-3030). Twelve of these were afterwards prepared as permanent whole mounts. Unless other- wise mentioned, measurements were made on live animals. Before fixation, the ani- mals were anesthetized in an isotonic so- lution of MgCl,, which was added under the cover slip for the whole mounts. The MgCl, 348 of the whole mounts was replaced by a fix- ative (2% formaldehyde or a trialdehyde so- lution) and then by a glycerol series from 2—100% (diluted in distilled water). When the glycerol was fully dehydrated after two days, the cover slip was sealed with Gly- ceel®: Twenty-six specimens were fixed for scanning electron microscopy (SEM) in a modified trialdehyde solution (Lake 1973) and postfixed in 1% OsO,, or fixed directly in 1% OsO,. The specimens were trans- ferred to distilled water, dehydrated through an acetone series, critical point dried, mounted on stubs, sputter coated with pal- ladium, and examined with a JEOL JSM- 6335F Field Emission scanning electron microscope. The study of live animals was carried out at the Bermuda Aquarium and. Zoo (BAMZ) and the study of fixed material was carried out at the Zoological Museum, University of Copenhagen (ZMUC). Types are deposited in the Zoological Museum, University of Copenhagen (ZMUC), Den- mark, and in the Smithsonian Institution, National Museum of Natural History (USNM), Washington D.C., U.S.A. Family Nerillidae Levinsen, 1883 Longipalpa, new genus Diagnosis.—Longipalpa is unique among nerillids by having two extremely long palps on the prostomium and two densely ciliated lobes on the dorsal side of the pygidium. It is further characterized by the combination of the following charac- ters: eight chaetigerous segments between prostomium and pygidium; prostomium with three very short simple dorsal anten- nae; compound serrated chaetae; single parapodial cirri from segment three to eight; two pygidial lobes; one anterior and one posterior group of non-motile cilia ar- ranged in distinct patterns and a pair of short bands of motile cilia on prostomium; tranverse discontinuous rows of ciliary tufts on dorsal and ventral surface; cuticular PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON plates in pharyngeal apparatus; two pairs of segmented nephridia from segment II-III and IJI-IV; hermaphroditic reproduction with one pair of spermioducts from seg- ment VI-VII and one pair of oviducts from segment VII-VIII. Type species.—Longipalpa saltatrix, new species, by present designation. Gender.—Feminine. Etymology.—From the Latin longus (=long) + English palp (=prostomial ap- pendage), in reference to the greater length of these appendages when compared to oth- er genera in the family. Similarity.—Longipalpa differs from the seventeen described nerillid genera by the two extremely long prostomial palps and two ciliated pygidial lobes. It furthermore differs from most genera by the very short length of the prostomial antennae, lack of parapodial cirri in segment 2 and possible lack of pygidial cirri. Four characters have been important in defining nerillid genera in recent years: number of body segments (7—9), compound or capillary chaetae, number of antennae (O—3), and number of cirri per parapodium (1-2) (Tzetlin & Larionov 1988, Tzetlin & Saphonoy 1992, Westheide & Purschke 1996, Miiller et al. 2001, Miiller 2002). Longipalpa resembles eight genera (Afro- nerilla, Akessoniella, Micronerilla, Nerilli- dium, Nerillidopsis, Thalassochaetus, Tro- chonerilla, Troglochaetus) by having eight segments. It thereby differs from four gen- era with seven segments (Aristonerilla, Bathychaetus, Paranerilla, Psammoriedlia) and five genera with nine segments (Lep- tonerilla, Meganerilla, Mesonerilla, Neril- la, Xenonerilla). It resembles seven genera with compound chaetae (Aristonerilla, Lep- tonerilla, Mesonerilla, Micronerilla, Neril- lidopsis, Paranerilla, Thalassochaetus) and six genera with three antennae (Aristoneril- la, Leptonerilla, Mesonerilla, Micronerilla, Nerilla, Trochonerilla), although only Tro- chonerilla possesses antennae of similar short length. Two genera (Leptonerilla, Mi- cronerilla) differ from Longipalpa by the VOLUME 117, NUMBER 3 presence of two cirri per parapodium (ver- sus One cirrus per parapodium in Longipal- pa). Six genera show resemblance to Longi- palpa in three out of the four “generic” characters mentioned above: Aristonerilla, Mesonerilla, Micronerilla, Nerillidopsis, Thalassochaetus, and Trochonerilla (see Table 1). Micronerilla may show the great- est resemblance with Longipalpa, but dif- fers by having two cirri per parapodium, pygidial cirri and two eyes. It furthermore diverges by the much longer antennae; many ciliary tufts on antennae, parapodial and pygidial cirri; parapodial cirri present on segment 2 (and sometimes on segment 1 as well) and absent on the last segment; gonochoristic reproduction and two pairs of spermioducts (Swedmark 1959, Jouin 1970, Saphonov & Tzetlin 1997, Miiller 2002). The other five genera likewise differ from Longipalpa in several important characters mentioned in Table 1. Leptonerilla prospera has previously been described from the caves of Bermuda (Sterrer & Iliffe 1982). The two Bermudian cave species have not been found in the same cave or Ccave-systems, although Road- side Cave and Walsingham Cave (type lo- cality for L. prospera) are separated by only 290 m. Their morphology is very different, and there is no reason to suspect that these two species should be closely related. Longipalpa saltatrix, new species Figs. 1-6, Table 2 Gen sp. A in Worsaae & Miiller (2004). Type material.—Holotype: ZMUC-POL 1675 (whole mount), 763 wm long, Road- side Cave, Bermuda (32°21'N, 64°43’W), 0.5—6.5 m depth, 20 Jan 2002. Paratypes: All paratypes with same locality as for ho- lotype, 0.5—6.5 m depth, collected 15, 20 and 21 Jan 2002. Nine specimens as whole mounts (ZMUC-POL 1676-1684) and 26 specimens on nine SEM-stubs (ZMUC- POL 1685-1693) are deposited in the Zoo- logical Museum, University of Copenhagen Table 1.—Comparison with relevant genera. Abbreviations: ( ), exceptions from remaining species of a genus; —, not applicable; segm, segment. Spermioducts Cirri per Pygidial parapodium Cirri Antennae Others segm Sex cirri segm Chaetae Eyes dimensions* Antennae Segm Genus VI-VII long palps, py- compound 3-8 hermaphroditic short Longipalpa gidial lobes ciliary tufts on VI-VII gonochoristic 2-7 compound long 7 Aristonerilla antennae IV-V, V-VI gonochoristic, 1-9 compound (O) 3 (2) long 9 Mesonerilla + VI-VII VI-VII + hermaphroditic gonochoristic ciliary tufts on 1-7 compound long 8 Micronerilla antennae VIL-VIl V-VI hermaphroditic 2-7 capillary + medium Nerillidopsis compound compound 2-7 2-7 8 8 Thalassochaetus VI-VII + palps present gonochoristic capillary short Trochonerilla VII-VIII 4 Short = antennae shorter than prostomium and palps, medium = antennae longer than prostomium but shorter than palps, long = antennae longer than prostomium and palps. 349 + Unpublished observations of Trochonerilla mobilis with palps (Worsaae, personal observation). 350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Wf j Wi4 sn yj en eg sd hg od 4 100 um Fig. 1. Reconstruction from light micrograph of live holotype of Longipalpa saltatrix, new species, dorsal view. Not all chaetae are drawn. Detailed information on nephridia, gonoducts and external dorsal ciliation is included from confocal laser scanning microscopy and scanning electron microscopy. Abbreviations: as, anterior field of sensory cilia; be, band of cilia; bm, bulbus muscle; ct, ciliary tuft; cp, cuticular plates; dg, dorsal glands; eg, eggs; en, enteronephridium; hg, hindgut; la, lateral antennae; mg, midgut; mo, mouth; no, nuchal organ; od, oviduct; pa, palp; pc, parapodial cirrus; pl, pygidial lobe; ps, posterior field of sensory cilia; sd, spermioduct; sg, salivary glands; sn, segmented nephridium; tb, tranverse ciliary band. VOLUME 117, NUMBER 3 ies) Nn — Table 2.—Meristics and morphometric characters of holotype and total type material (measurements on ju- veniles in parentheses). Abbreviations: excl., exclusive; incl., inclusive; L, length; min., minimum; max., max- imum; segm., segment; W, width. Holotype Total L excl. appendages, chaetae 763 max. W incl. parapodia 224 max. W excl. parapodia 192 prostomium L 72 W 81 max. L palps 696 L median antenna 43 max. L lateral antennae 56 trunk L segm. | 80 L segm. 2 123 L segm. 3 113 L segm. 4 109 L segm. 5 103 L segm. 6 67 L segm. 7 56 L segm. 8 35 L pygidium 8 max. L parapodia segm. 1 63 max. L other parapodia 48 max. L parapodial cirri 67 max. L pygidial lobes 42 chaetae max. no. chaetae segm. 1? max. no. chaetae notopodia* max. no. chaetae neuropodia? max. total L chaetae max. L shaft> L distal extension shaft? L blade? Min Max Average n 624 (471) 985 788 14 127 (116) 285 202 13 108 (99) 268 1 13 59 77 68 12 67 (66) 90 80 12 680 (660) 718 696 6 24 (17) 43 33 7 41 (31) 65 53 10 55 (46) 97 74 2 98 (77) 150 126 12 76 (71) 152 108 12 73 121 104 12 71 (69) 103 93 12 54 106 80 12 54 89 68 11 25 58 42 11 8 37 20 12 35 63 53 13 30 50 40 12 4] 73 60 13 30 50 4] 9 7 (6) 13 10 8 7 10 8 6 6 10 8 6 135 145 139 8 86 109 103 5 0) 2 1 5) 33 41 37 5) 4 Measured on fixed material by SEM. > Measured alive by LM and on fixed material by SEM. (ZMUC), Denmark. Two paratypes as whole mounts (USNM 1022181-1022182) are deposited in the Smithsonian Institution, National Museum of Natural History, Washington, D.C., U.S.A. Diagnosis.—Characters of the genus. Etymology.—From the Latin saltator (=dancer), in reference to the swimming skills of the species, which may swim in loops while waving and twisting the long palps. Description (see Table 2 for principle counts and measurements).—A relatively hyaline nerillid with brown pigmentation, especially along intestinal wall. The body consists of prostomium, eight chaetigerous segments and pygidium (Figs. 1, 2A, 4). Adults with eight segments and a total length of 624—985 wm (only 454-825 wm when fixed); juveniles with six to seven segments and a total length of about 500 wm. Maximum body width generally at segment five, up to 268/285 wm (excl./incl. parapodia); narrow restriction between seg- ment one and two, posterior to the esoph- agus. Prostomium up to 77 pm long, 90 ~m PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a re & F dee 22) BOOT — Fig. 2. Light micrographs of live holotype of Longipalpa saltatrix, new species. A, Whole specimen with two palps. B, Closer view of ciliation on palp and prostomium. C, Closer view of prostomium and segment one. Abbreviations: see Fig. 1; pp,, parapodium of segment one; pr, prostomium; I-VI, segments one to eight. VOLUME 117, NUMBER 3 Fig. 3. Light micrographs of live specimens of Longipalpa saltatrix, new species. A, Dorsal view of segment three to six showing midgut lining of glandular cells with vesicles. B, Ventral view of middle segments showing diffuse glandular pattern. C, Posterior part of animal. D, Parapodium. E, Segments seven-eight and pygidium. Abbreviations: see Fig. 1; arrowhead, extension of shaft; cb, chaetal blade; cs, chaetal shaft; dg, diffuse glandular pattern; fa, fascicle; gc, glandular cells; ov, ovoids; py, pygidium; VI-VIII, segments six to eight; ve, vesicles. 354 wide; first four segments longest, decreas- ing in length posteriorly, pygidium even shorter. Prostomium short, with two ventro-lat- eral palps and three dorsal antennae. Palps filiform and long, up to 718 wm (up to about 90% of body length in adults, and up to about 130% in juveniles), and with com- plex ciliation (see below) (Figs. 1, 2, 5A). Antennae short, filiform, with few distal cil- ia. Medium antenna up to 43 pm long, lat- eral antennae up to 65 pm long (Figs. 1, 4A, B, 5A). Nuchal organs paired, situated between palps and parapodia of segment one on a round elevated bulge on each lat- eral. side of the prostomium (Figs. 1, 5A, C). Parapodia of segment one very large (up to 63 pm long), up to twice the length of the following parapodia (Figs. 1, 2A, 4A, C). Parapodial cirri (with few distal cilia) be- tween dorsal and ventral chaetal bundles of parapodia of segment three to eight; length up to 73 wm, increasing towards the pos- terior segments (Fig. 4A). No trace of at- tachment of parapodial cirri on segment one and two, neither of scars from detached cir- ri, or rudimentary cirri. Appendages like cirri and palps, and even chaetae, were eas- ily lost during handling and fixing of the animals. Of the more than 70 specimens ob- served alive, none possessed parapodial cir- ri On segment one and two and scars were not found with SEM. Pygidial cirri were never observed, but it was difficult to ex- amine the pygidium thoroughly for scars of cirri with SEM. On one specimen, a pair of scar-like structures was found at the pygid- ium, which could be scars from lost pygid- ial cirri or just an artifact (Fig. 6G). All adult (but no juvenile) animals possessed two very special structures, here named py- gidial lobes due to their location on the dor- sal side of the pygidium. Each lobe is up to 50 wm long, with two projections and a dense ciliation (Figs. 1, 3E, 4A, 6G, H). All chaetae compound and relatively straight, shaft with minor pointed distal ex- tension, less than 2 wm long (Figs. 3D, 6B). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Chaetae very slightly serrated and generally with a hairy appearance (Fig. 6A—C). Seg- ment one uniramous, with up to thirteen chaetae in one chaetal fascicle; segments two to eight biramous, with dorsal and ven- tral fascicles comprising up to ten chaetae each. Similar numbers of chaetae in seg- ments two to six, somewhat fewer chaetae in the last two segments. No noticeable dif- ferences in number or length between dor- sal and ventral chaetae. Similar lengths in all segments of shaft, blade and total length of chaeta. Shaft up to 109 um, blade up to 41 pm, total length up to 145 pm (Figs. 3D, 4, 0A). Dorsal surface of prostomium with very specific ciliation characterized by three dif- ferent groups of cilia: a pair of short bands with motile cilia (>20 cilia, up to 20 pm long), one on each dorso-lateral surface next to the lateral antennae (Figs. 1, 5A); two transverse rows of non-motile cilia in front of antennae on the anterior most part of the prostomium (Fig. 5A, B), the last of which arranged in a distinct pattern; and a posterior group of twenty non-motile cilia (Fig. 5D, E), arranged in complex pattern near the origin of median antenna on pos- terior part of the prostomium. The patterns of the anterior field of cilia (probably sensory in function, see Discus- sion) and the posterior fields of cilia (prob- ably sensory) are characteristic of the spe- cies and are here given in detail: anterior field of cilia (S-15 pm long) with two transverse rows of cilia (Fig. 5B). Posterior row contains about 5 cilia, spaced 2—5 wm apart (cilia no. 1-5 in Fig. 5B). Anterior row with about 20 cilia (no. 6—25 in Fig. 5B). Cilia arranged in distinct pattern mir- rored halfway along the row. Moving to- wards the middle from the lateral sides, the first cilia are two groups of four cilia (no. 6-9 and 10-13), a single cilium next to them (no. 14 and 15), two groups of three cilia (no. 16—18 and 19—21), three cilia in the middle (no. 22—24), and one cilium (no. 25) in front of the middle cilium. ie) Nn Nn VOLUME 117, NUMBER 3 Fig. 4. Scanning electron micrographs of Longipalpa saltatrix, new species. A, Dorsal view of whole spec- imen with both palps lost. B, Lateral view of specimen with one palp lost. C, Ventral view of specimen with two palps lost. Abbreviations: see Figs. 1, 2; arrowhead, connection of chaetal shaft and blade; db;, dorsal ciliary band of segment three; ma, median antennae; mv, midventral ciliary band; vb,-vb,, ventral ciliary band on segments one to seven. 356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Scanning electron micrographs of Longipalpa saltatrix, new species. A, Dorsal view of prostomium, right palp, and segment one. B, Closer view of anterior field of twenty-five cilia. C, Closer lateral view of nuchal organ and extra ventral ciliary band. D, E, Close dorsal view of posterior field of twenty cilia from two speci- mens. Abbreviations: see Figs. 1, 2, 4; bc, band of cilia; sc, scar from lost palp; vc, ventral ciliation around mouth; xb, extra ventral ciliary band. VOLUME 117, NUMBER 3 357 Fig. 6. Scanning electron micrographs of Longipalpa saltatrix, new species. A, Chaetal bundle and para- podial cirrus. B, Closer view of chaetae with microvillar hairy appearance. C, Two hairy chaetae showing serration pattern (indicated by arrowheads). D, Left ventral side of prostomium and segment one. E, Right side of segment six with half dorsal ciliary band. E Left ventral side of segment three with half ventral ciliary band. G, Left dorsal side of posteriormost segments. H, Closer view of left pygidial lobe. Abbreviations: see Figs. 1- 5; ch, holes after lost chaetae; db, dorsal ciliary band of segment six; ex, extension of shaft; gr, dense ciliary groups; sc, scars from lost pygidial cirri or an artifact. 358 Posterior field of twenty cilia (3-15 pm long) covers an area about 12 wm wide and 8 wm long (Fig. 5D, E). Four cilia in a close square (no. 1—4 in Fig. 5D) surrounded by a common elevation of the cuticle, are found in the center of the field, posterior to the basal part of the antenna. Right next to these four cilia one cilium is found on each lateral side (no. 5—6), which as all the single situated cilia in the pattern, is surrounded by a cuticular collar. On each lateral side of the antenna is one cilium (no. 7—8). About 3 wm posterior of these two clusters are found, each with three cilia in a transverse line, surrounded by a common elevation of the cuticle (no. 9-14). Next to these, on the level of the 4 central cilia, are found 2 pairs of cilia next to each other on each lateral side (no. 15-18). The last pair of cilia (no. 19—20) is located a few micrometers poste- riorly with about 5 pm in between the cilia. Palps with complex ciliation containing transverse ciliary bandlets in a longitudinal row on the inner and outer lateral surfaces of the palp, respectively. More than 20 cilia, up to 20 wm long in each ciliary bandlet, positioned 5—20 pm apart in a row extend- ing to the tip. Farthest distance between bandlets on outer lateral surface of the palp. Longitudinal row of single ciliary tufts on both dorsal and ventral surface between the longitudinal rows of bandlets (Figs. 1, 5A). Less than 5 cilia, up to 7 wm long in each ciliary tuft, located in a row extending to the tip. Ciliary bandlets beat in metacronal waves, creating a water current leading par- ticles towards the base of the palp. Motility of ciliary tufts not clearly distinguishable due to intense beating of ciliary bandlets. However, we suspect these cilia to be non- motile due to their small number and short length. Dorsal surface of body segments not cil- iated, except for few ciliary tufts. Two to four tufts of motile cilia are situated in a transverse line across each segment between the parapodia on each side (Figs. 4A, B, 6E). Each tuft contains 20—200 motile cilia, up to ca. 25 4m long. Each pygidial lobe possess- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON es two large groups of cilia, one on each of the two projections of the lobe (Figs. 1, 4A, 6G, H). Each group contains more than 100 cilia, up to ca. 25 wm long. Ventral surface with dense ciliation around mouth on ventral side of prostomi- um, continuous with relatively narrow mid- ventral ciliary band extending to the anus on the dorsal side of the pygidium. Tran- verse rows of ventral ciliary tufts on each segments at the level of the parapodia: four pairs on segments one to three, three pairs on segments four to seven, two pairs on segment 8. Three additional pairs of ciliary tufts: one pair between prostomium and segment one, almost connecting ciliation around mouth with that of the nuchal or- gans; and two pairs between segment one and two (Figs. 4C, 6D). Pharynx with ventral opening between prostomium and segment one, and muscular bulb in segment one (Figs. 1, 2A, C). About six pairs of ventral brown glands (may have salivary function) open into buccal cavity on ventral side of pharynx (Figs. 1, 2A, C). Two additional dark brown, round cell- groups (probably glandular in function) dorsally of pharynx in prostomium. All groups contain several cells with relatively large round vesicles. A pair of triangular cuticular plates on ventral side of pharyn- geal bulb in anterior part of pharyngeal or- gan (Figs. 1, 2C). Large round glandular cells with many small vesicles and brown pigmentation line stomach wall (Figs. 1, 3A); large ciliated cells line hindgut (Figs. 1, 3C). Diffuse superficial glands create a unique pattern in the ventro-caudal epithe- lium (Fig. 3B). Studies by confocal scanning microscopy showed the distribution of nephridia and gonoducts in this and other species (Wor- saae & Miiller 2004). Longipalpa saltatrix is hermaphroditic with one pair of sper- mioducts in segments six to seven and one pair of oviducts in segments seven to eight (see Fig. 1 and Worsaae & Miiller 2004, fig. 2J). Two pairs of segmented nephridia are present, from segments two to three, and VOLUME 117, NUMBER 3 from segments three to four, respectively (see Fig. 1, and Worsaae & Miiller 2004, fig. 2G—I). Several enteronephridia line the hindgut (see Fig. 1, and Worsaae & Miiller 2004, fig. 2G, J). Fertile animals contain a maximum of two large eggs with diameters up to 170 wm, and an additional large num- ber (up to ca. 40 has been counted) of smaller ovoids with a diameter about 10— 20 wm. Distribution.—Presently known only from a certain anchihaline cave pool in Ber- muda. Motility.—The animals swim beautifully in the water column, describing loops and turns. Less frequently, they glide over the surface and if provoked make an escape re- action or quick turn by undulation of the body and fast curling up of the palps in a narrow spiral. When swimming, they are capable of bending the prostomium and body as well as waving, bending and curl- ing the long palps. The pygidial lobes are flapped between positions flat along the body to an almost right angle to the dorsal surface, thereby using the densely ciliated lobes as helms. The forward drift seems to be created mainly by the ventral ciliation, possibly with additional force from the cilia on palps and pygidial lobes. Remarks.—The description of Longipal- pa saltatrix not only adds a new genus to the family Nerillidae, but also expands the definition of the family. The extremely long palps of this species are not only unusual in their length but probably also in their function. The longitudinal rows of ciliary bandlets create a water current propelling particles towards the mouth opening, which has never been observed in other nerillids. It seems possible that the animals collect food particles by help of the palps, thereby increasing their feeding radius extensively. Foraging could happen when gliding over or through the substrate as well as when swimming through and above the sediment collecting particles in suspension. In several species of nerillids, the ciliation of the much shorter palps creates a water current 359 transporting particles away from the mouth (Worsaae, personal observations). This transport may indicate that other nerillid palps may also be functional in feeding be- havior, however, by transporting rejected particles away from the mouth and not by gathering them. The previous understand- ing of the nerrillid palps as being mainly sensory in function should probably be ex- panded to include a function in feeding be- havior. This view stands in contrast to the general comprehension of the ventral palps of the major taxon group Aciculata (see Rouse & Fauchald 1997) with which the family has the most apparent resemblance (see e.g., Schmidt 1848, Quatrefages 1866, Westheide 1990, Westheide & Purschke 1996, Rouse & Fauchald 1997, Rouse & Pleijel 2001). Aciculates are generally char- acterized by short sensory palps with no di- rect function in food collecting. If the neril- lids truly belong to the Aciculata, then the long palps of Longipalpa saltatrix may also influence the conception of the conserva- tiveness of the palps in this taxon group. The extremely long palps would proba- bly be disadvantageous in the interstitial habitat from which many nerillids are de- scribed. This disadvantage may be one of the reasons why these long palps have not been found in other nerillids. In the Ber- mudian caves with only a sparse layer of very fine silt on top of bare rocks, it may even be an advantage to be able to actively swim and perhaps also feed on suspended particles with the aid of the palps. However, Leptonerilla prospera which lives under rather similar conditions, except from more light in the Walsingham Caves in Bermuda, does not possess long feeding palps. Dif- ferences in habitat characteristics between cave and interstitial habitats include more space in caves, thus allowing for swimming as opposed to crawling, and different types of food, hence different feeding mecha- nisms. Many other anchihaline species are most commonly found within the water col- umn rather than on the sediments, implying that this is where food is primarily located. 360 The motile ciliary bands and the anterior field of cilia on the prostomium could easily be detected on live animals with light mi- croscopy (LM), whereas the posterior field of cilia could only be detected with SEM. The motile ciliary bands (be in Figs. 1, 5A) are most likely not mechanoreceptors be- cause of their long length, motility, and dense grouping. However, the anterior and posterior fields of cilia are probably sensory in function because of their non-motility, shorter length, and single appearance of cil- ia—each with a cuticular collar. Two fields of cilia (suggested to be sensory in func- tion) have also been described for the neril- lid Paranerilla limicola Jouin & Swed- mark, 1965 (Worsaae & Kristensen 2003). The anterior field of cilia in P. limicola con- sists of a little group of cilia and, except for the anterior position; it is very different from the two transverse rows of cilia ar- ranged in a pattern found in L. saltatrix. The posterior field in P. limicola is more similar with 14 cilia arranged in a distinct pattern. However, this pattern differs some from the pattern of 20 cilia found in L. sal- tatrix. It seems very possible that the two systematically significant prostomial fields of cilia (probably sensory in function) are a common feature of nerillids, which just demands SEM techniques to be described. A few of the unusual characteristics of L. saltatrix have previously been found in sin- gle occasions in otherwise very different nerillid species. Structures remarkably sim- ilar to the special triangular cuticular plates on the ventral part of the pharynx have been described for Thalassochaetus palpi- foliaceus Ax, 1954. A different, although also distinct pattern of diffuse superficial ventral glands are described for Nerillidium renaudae Jouin, 1970. Pygidial lobes have been described for the aberrant Nerillidium simplex Levi, 1953 (see also Jouin 1966, Swedmark 1959). These lobes are appar- ently not double-lobed or cilicated as in L. saltatrix; however, their position and square, non-cirriform appearance is very similar to L. saltatrix. The pygidial lobes PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON found in N. simplex have been interpreted as modified pygidial cirri (Levi 1953, Swedmark 1959), which may also count for the lobes of L. saltatrix. The superficial re- semblance with the lobes of N. simplex is probably a matter of convergence; however, the lobes may be functionally comparable. The two groups of cilia on each pygidial lobe of L. saltatrix show great resemblance in number and length with the two ciliary tufts found on each side of the body seg- ment dorsal to the parapodia. Furthermore, examined juveniles did not possess pygidial lobes, which would be expected if the lobes were modified pygidial cirri. These obser- vations could mean that the pygidial lobes are modified rudiments of a strongly re- duced ninth body segment. However, stud- ies with LM and SEM of L. saltatrix show that there are no remains of chaetae or para- podial muscles and cLSM studies show that there are no segmental nerves posterior to the eighth body segment. Further exami- nation of the development of L. saltatrix is needed to clarify the origin of the pygidial lobes. Acknowledgments We thank Dr. Martin V. Sgrensen for as- sistance with collecting of the material. The study was financially supported by the Ber- muda Zoological Society. This paper is contribution # 63 from the Bermuda Bio- diversity Project (BBP), Bermuda Aquari- um, Natural History Museum and Zoo. Literature Cited Angel, M. V., & T. M. Iliffe. 1987. 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A. e-mail: mhrocha@ciencias.unal.edu.co Abstract.—A new species of the genus Neostrengeria Pretzmann, 1965, N. lemaitrei from Magdalena Valley, Cundinamarca Department, is described. The genus is endemic to the Eastern Andes of Colombia, at altitudes ranging from 300 to 300 m above sea level. With the addition of N. lemaitrei the total number of species rises to 21. This new species, like all others in Neostrengeria, is distinguished primarily by the morphology of the first male gonopod, partic- ularly by the form of lateral and accessory lobes, and the shape of the apex. The genus Neostrengeria Pretzmann, 1965, comprises 21 species of freshwater crabs that inhabit mountain springs and streams on the slopes and high plain of the Eastern Andes in Colombia (2° to 9°40'N, 73° to 74°50'W), at altitudes ranging from 300 to 3000 m above sea level (Campos 1994). The taxonomy of Neostrengeria was re- viewed by Rodriguez (1982), with follow up studies by Campos (1992, 1994, 2000). Campos & Lemaitre (1998) presented a key for the identification of the species based on the morphology of the male first gono- pod. The distribution of the genus has been discussed by Campos & Rodriguez (1985), and Campos (1992, 1994). The present new species was found in the Magdalena Valley, at altitude of 720 m above sea level. The general carapace morphology of Neostrengeria species is very similar. The species are characterized primarily by the shape of the first male gonopod which has a distinct lateral lobe generally divided in two halves forming an accessory lobe. The form of the gonopod’s apex is also variable according to the species, and can be oval, oblong, or expanded into a projection. The terminology used for the different processes of the gonopod is that established by Smalley (1964), Rodriguez (1982) and Campos & Lemaitre (1998). The material is deposited in Museo de Historia Natural, In- stituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogota (ICN- MHN). The abbreviations cb and cl, re- ported as cl X cb, indicate carapace breadth and carapace length, respectively. Color no- menclature follows Smithe (1975). Family Pseudothelphusidae Rathbun, 1893 Tribe Hypolobocerini Pretzmann, 1971 Genus Neostrengeria Pretzmann, 1965 Neostrengeria lemaitrei, new species Fig. | Holotype.—Agua Blanca stream, Vereda Lamal, Inspecci6n Guadualito, Municipio Yacopi, Cundinamarca Department, Colom- bia, 720 m alt., 4 Nov 1995, leg. M. R. Campos: | male, 13.9 X 23.6 mm, ICN- MHN-CR 1991. Paratypes.—Same locality data as holo- type: 5 males, size range 8.1 X 12.9 mm, to 12.5 X 20.0 mm, 4 females, size range 9.4 X 14.7 mm, to 7.4 X 11.2 mm, ICN- MHN-CR 1533. Type locality.—Agua Blanca stream, Vereda Lamal, Inspeccion Guadualito, Mu- 364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Neostrengeria lemaitrei, new species, male holotype, ICN-MHN-CR 1991. A, left first gonopod, caudal view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view; E right carapace half, dorsal view; G, left opening of efferent branchial channel, external view; H, left third maxilliped, external view. 1, lateral lobe; 2, accessory lobe; 3, cephalic expansion; 4, mesocaudal projection of spermatic channel. VOLUME 117, NUMBER 3 nicipio Yacopi, Cundinamarca Depatment, Colombia, 720 m alt. Diagnosis.—Carapace without median groove; front lacking distinct upper border. Third maxilliped with exognath 0.67 times length of ischium. First male gonopod with lateral lobe semicircular distally, proximally narrow, with external margin concave; ac- cessory lobe elongated, semi-acute distally, forming excavated ridge on caudal surface; accessory lobe as long as lateral lobe. Apex outline oval with expansion projected ce- phalically into prominent, acute spine. Description of holotype.—Carapace (Fig. 1F) with cervical groove straight, shallow, ending some distance from lateral margin. Anterolateral margin lacking depression be- hind external orbital angle. Lateral margin with series of approximately 15 papilliform teeth. Postfrontal lobes oval, high, indicated anteriorly by 2 transverse depressions. Me- dian groove lacking. Front without distinct upper border, frontal area sloping down- wards, slightly bilobed in dorsal view, low- er margin visible in dorsal view, strongly sinuous in frontal view. Dorsal surface of carapace smooth, covered by small papillae, regions well demarcated. Third maxilliped with distal half of external margin of merus rounded, exognath 0.67 times length of is- chium (Fig. 1H). Orifice of efferent bran- chial channel open, irregularly ovate (Fig. 1G). First pereiopods heterochelous; palm of larger chela strongly swollen, fingers slight gaping when closed, smaller chela slight swollen, fingers not gaping when closed. Walking legs (pereiopods 2-5) slen- der, but not prominently elongated (total length 1.10 times the breath of carapace). First male gonopod wide in caudal view; mesial side forming convex expansion with deep subdistal notch; caudal margin wide with excavated surface, festooned (Fig. 1A, D); lateral lobe wide, semicircular distally, proximally narrow with external side con- cave, separated from accessory lobe by deep notch (Fig. 1A—D); accessory lobe elongated, semi-acute distally, forming ex- cavated ridge, covered with diminute papil- 365 lae and row of spinules on external border on caudal surface; accessory lobe as long as lateral lobe (Fig. 1A, C); apex outline oval in distal view with expansion projected cephalically into prominent, acute spine; mesial lobe subtriangular; mesocaudal pro- jection of spermatic channel with bifid tip; spermatic channel with conspicuous rows of spinules; proximal cephalic border with two setae (Fig. 1C, D, E); conspicuous se- tae along outline of prominent basal round- ed lobe, and a patch of setae on caudal sur- face (Fig. 1A). Color.—The holotype, preserved in al- cohol, is brown-olive (near 129, Dark Brownish Olive) on the dorsal side of the carapace. The dorsal and ventral surfaces of the chelae and the walking legs are brown (near 223, Raw Umber). The ventral sur- face of the carapace is beige (near 92, Pale Horn Color). Habitat.—The specimens were collected in shaded, moist banks of springs and streams. They were found in soft mud, un- der rocks. Etymology.—The species is named in honor of Colombian scientist Dr. Rafael Le- maitre, who has dedicated his life to study- ing Crustaceans. This species is not only a recognition of Rafael’s contributions to sci- ence, but to the stimulus he has provided to a new generation of up and coming Col- ombian scientists. Remarks.—A comparison of both de- scriptions and material of other species of the genus with that of this new species re- vealed that it is most similar to Neostren- geria gilberti Campos, 1992. The main dis- tinguishing feature between both species is the form of the first gonopod. The male first gonopod of N. gilberti has been described and illustrated by Campos (1992: 542, fig. 2). In this new species, the mesial side of the gonopod is convex expanded with deep subdistal notch, whereas in N. gilberti it is rounded basally, straight tapering distally without subdistal notch. The lateral lobe in N. gilberti is rounded distally with the prox- imal external side straight, whereas in UN. 366 Table 1.—Vertical distribution of the Neostrengeria species. Meters above sea Species level Neostrengeria appressa Campos, 1992 1125-1900 N. aspera Campos, 1992 1600 N. binderi Campos, 2000 470 N. botti Rodriguez & Tiirkay, 1978 1350-2600 N. boyacensis Rodriguez, 1980 2350-3000 N. charalensis Campos & Rodriguez, 1450-2150 1985 N. gilberti Campos, 1992 950-1250 N. guenteri (Pretzmann, 1965) 500-1575 N. lasallei Rodriguez, 1980 1110-2150 N. lemaitrei, new species 720 N. libradensis Rodriguez, 1980 1200 N. lindigiana (Rathbun, 1897) 1800-2350 N. lobulata Campos, 1992 1700-2350 N. macarenae Campos, 1992 300-500 N. macropa (H. Milne Edwards 1853) 2200-2900 N. monterrodendoensis Bott, 1967 1320-1500 N. niceforoi (Schmitt, 1969) 1000-1750 N. perijaensis Campos & Lemaitre, 1998 1270-1800 N. sketi Rodriguez, 1985 1800 N. tencalanensis Campos, 1992 1600-2400 N. tonensis Campos, 1992 1600-2400 lemaitrei it 1s distally semicircular, and proximally narrow with the external side concave. The apex outline in N. gilberti is oblong in distal view with a mesially di- rected semi-acute spine; the mesocaudal projection of spermatic channel is awl- shaped with a distal spinule on the inner side. In contrast, in N. lemaitrei, the apex outline is oval in distal view with the ex- pansion projected cephalically into a prom- inent, acute spine, and the mesocaudal pro- jection of spermatic channel has the tip bi- fid. Distribution of Neostrengeria species The distribution of the species of Neos- trengeria comprises both slopes and the high plain of the Eastern Cordillera of Co- lombia that encompasses the Magdalena, Orinoco and Catatumbo basins. It is limited to the north by Serrania de Perija, and to the south by Serrania de La Macarena (2° to 9°40'N, 73° to 74°50'W), (H. Milne Ed- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wards 1853; Rathbun 1897; Pretzmann 1965; Bott 1967; Schmitt 1969; Rodriguez & Tiirkay 1978; Rodriguez 1980, 1982, 1985; Campos & Rodriguez 1985; Campos 1992, 1994, 2000; Campos & Lemaitre 1998). Based on the collected material, the ver- tical distribution of the species of the genus Neostrengeria (Table 1) ranges from 300 m to 3000 m. Neostrengeria botti has the greatest altitude range of between 1350 and 2600 m. The species that exhibit a range of between 300 and 1000 m are N. binderi, N. macarenae and N. lemaitrei, new species. Most of the species are distributed between 1000 and 2400 m. The highest altitude, 3000 m, is reached by N. boyacensis. Acknowledgments I am especially grateful to the referees for providing useful comments of the paper. I also indebted to David H. Campos for crit- ically reading the manuscript. The illustra- tion was prepared by Juan C. Pinzon. Literature Cited Bott, R. 1967. Fluss-krabben aus dem westlichen Sii- damerika.—Senckenbergiana Biologie 48(5/6): 365-372. Campos, M. R. 1992. New species of fresh-water crabs of the genus Neostrengeria Pretzmann, 1965 (Crustacea: Decapoda: Pseudothelphusidae) from Colombia.—Proceedings of the Biological Society of Washington 105:540—554. . 1994. Diversidad en Colombia de los cangre- jos del género Neostrengeria.—Academica Co- lombiana de Ciencias Exactas Fisicas y Natur ales. Col. Jorge Alvarez Lleras No. 5:1—143. . 2000. Neostrengeria binderi, a new species of pseudothelphusid crab from the eastern Andes of Colombia (Crustacea: Decapoda: Brachyu- ra).—Proceedings of the Biological Society of Washington 113:401—405. Campos, M. R., & R. Lemaitre. 1998. A new fresh- water crab of the genus Neostrengeria Pretz- mann, 1965, from Colombia (Crustacea: Deca- poda: Brachyura: Pseudothelphusidae) with a key to the species of the genus.—Proceedings of the Biological Society of Washington 111: 899-907. Campos, M. R., & G. Rodriguez. 1985. A new species of Neostrengeria (Crustacea: Decapoda: Pseu- VOLUME 117, NUMBER 3 dothelphusidae) with notes on geographical dis- tribution of the genus.—Proceedings of the Bi- ological Society of Washington 98:718—727. Milne-Edwards, H. 1853. Observations sur les affini- tiés zoologiques et la classification naturelle des crustacés.—Annales des Sciences Naturelles, Zoologie 20:163—228. Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam- ilie Pseudothelphusidae.—Anzeiger der Oster- reichischen Akademie der Wissenschaften Mathematische Naturwissenschaftliche Klasse (1),1:1-10. . 1971. Fortschritte in der Klassifizierung der Pseudothelphusidae.—Anzeiger der Mathema- tisch Naturwissenschaftliche der Osterreichisch- en Akademie der Wissenschaften (1)179:14—24. Rathbun, M. 1893. Descriptions of new species of American freshwater crabs.—Proceedings of the United States Naitonal Museum 16(959): 6459-661, pl. 73-77. . 1897. Descriptions de nouvelles espéces de crabes d’eau douce appartenant aux collections du Muséum d’ Histoire naturelle de Paris.—Bul- letin du Muséum National d’ Histoire Naturelle, Paris 3(2):58-61. 367 Rodriguez, G. 1980. Description préliminaire de qu- elque espéces et genres nouveaux de crabes d’eau douce de 1’Amérique tropicale (Crusta- cea, Decapoda, Pseudothelphusidae).—Bulletin du Muséum National d’ Histoire Naturelle, Paris 4(3):889—-894. . 1982. Les crabes d’eau douce d’ Amérique. Famille des Pseudothelphusidae.—Faune Trop- icale 22:1—223. . 1985. A new cavernicolous crab (Crustacea, Decapoda, Pseudothelphusidae) from Colom- bia.—Bioloski vestnik, Ljubljana 33(2):73—80 Rodriguez, G., & M. Tiirkay. 1978. Der generische Status einiger Kolombianischer Stisswasser- krabben.—Senckenbergiana Biologica 59:297— 306. Schmitt, W. 1969. Colombian freshwater crab notes.— Proceedings of the Biological Society of Wash- ington 82:93-112. Smalley, A. 1964. A terminology for the gonopods of the American river crabs.—Systematic Zoology 13:28-31. Smithe, E B. 1975. Naturalist’s color guide: The American Museum of Natural History. New York, part 1: unnumbered pages. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):368—376. 2004. A new species of Agostocaris (Caridea: Agostocarididae) from Acklins Island, Bahamas Fernando Alvarez, José Luis Villalobos, and Thomas M. Iliffe (FDA, JLV) Colleccion Nacional de Crustaceos, Instituto de Biologia, Universidad Nacional Aut6noma de México, Apartado Postal 70-153, México 04510 D.E, México, e-mail: falvarez@servidor.unam.mix; (TMI) Department of Marine Biology, Texas A&M University at Galveston, Galveston, Texas 77553-1675, U.S.A. Abstract.—The new bresilioid shrimp Agostocaris acklinsensis is described from an anchialine cave in Acklins Island, Bahamas. This is the third species described in the genus. The new species is characterized by having small ex- opods on the third and fourth pereiopods, one spine on the ischium of the fifth pereiopod, and an outer ramus of the uropods with one distolateral spine. A key to the species of Agostocaris is provided. The family Agostocarididae Hart & Manning, 1986, was created to accommo- date Agostocaris williamsi, from Grand Ba- hama and Turks and Caicos, a species that appeared to be morphologically similar to some species in the Atyidae De Haan, 1849, and the Bresiliidae Calman, 1896, but had a distinct morphology of the propodus and dactylus of the first two pereiopods. Ken- sley (1988) described a second species from Cozumel, Mexico, Agostocaris bozanici, which exhibits the same unique pereiopodal morphology, placing it also in the Agosto- carididae. Holthuis (1993) synonymized the Agostocariidae with the Bresiliidae. How- ever, Martin & Davis (2001) have proposed to recognize the family Agostocarididae within the superfamily Bresilioidea Cal- man, 1896, where a hetereogeneous assem- blage of forms are included in five families. At best, as pointed out by Kensley (1988), the relationships of Agostocaris are unclear. The particular articulation of the proprodus of the first pair of legs, and the morphology of the chela of the second pair of legs, are unique characters not shared by any other genus in the Bresilioidea. With respect to the diagnosis of Agostocaris, with the new species described herein, the range of variation in taxonomically impor- tant characters increases, making it neces- sary to provide a new diagnosis for the ge- nus. Materials and Methods Specimens of the new Agostocaris de- scribed herein were collected during an ex- pedition to Crooked and Acklins Islands, Bahamas, in January 1999. The new spe- cies was captured in Jumby Hole Cave (22°29.275'N, 73°53.501’W), Snug Corner, Acklins Island, Bahamas, 11 January 1999 (Fig. 1). This cave is located about 250 m inland from the west side of the island fac- ing the shallow water Bight of Acklins. It is actually a complex of closely associated caves that were mined for guano in the past. More than 3 m of soil and guano were re- moved from pits within dry portions of the cave. One of these caves contains a 20 m diameter, shallow (30 to 50 cm deep) pool. Sediments in the pool consist of a thick lay- er of guano from a bat roost located directly above. Tidal range in the pool appeared to be about 30 cm. This pool is in total dark- ness but is close to 4 or more entrances on all sides. Salinity was measured at 32.5%o VOLUME 117, NUMBER 3 369 GULF OF MEXICO Crooked Island Bight of Acklins Acklins Island Liza Bay Cave Fig. 1. Map showing the location of the type locality of Agostocaris acklinsensis, Acklins Island in the Bahamas. 370 with a refractometer and water temperature was 25.5°C. Specimens of Agostocaris were observed walking across the surface of rocks and the guano bottom in 50 cm depth. They were collected by hand using glass vials. Other invertebrates collected from the cave pools included copepods, archiannelid and other polychaetes, mites and the shrimp Barbouria cubensis (von Martens, 1872) (Hippolytidae). The specimens representing the new spe- cies are deposited in the Coleccion Nacion- al de Crustaceos (CNCR), Instituto de Biol- ogia, Universidad Nacional Autonoma de México. Other abbreviations used are: cl, postorbital carapace length, and tl, total length. Results Agostocaris Hart & Manning, 1986 Diagnosis.—Rostrum well developed, with or without dorsal teeth. Carapace lack- ing spines and grooves. Eyes reduced, fused, without pigment or weakly pigment- ed. Antennal scale with lateral spine. First maxilliped with lash on exopod. Second maxilliped with terminal segments serial. Pleurobranchs on all pereiopods or on pe- reiopods 2—5. First and second pereiopods chelate, first pair heavier than second one. First pereiopod with propodus articulating with carpus at one third of its length. Sec- ond pereiopod with carpus undivided; dac- tylus digitiform, heavier and longer than propodus, both fingers without teeth or spines. Telson with 4—5 pairs of dorsal spines, posterior margin with variable num- ber of spines. Agostocaris acklinsensis, new species Figs. 2—4 Material examined.—Holotype, female, cl 7.3 mm, tl 21.5 mm; 11 January 1999; Jumby Hole Cave, Snug Corner, Acklins Is- land, Bahamas; collected by T. M. Iliffe; CNCR 19601. Paratypes, 8 females, cl 4.0— 8.0 mm, tl 13.6—21.7 mm; same locality, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON date and collector as holotype; CNCR 19602. Description.—Carapace globose, smooth, devoid of spines. Rostrum laterally com- pressed, triangular, ending in sharp tip, reaching distal end of first antennular seg- ment; without teeth in mature individuals, with three dorsal teeth with alternating setae in juveniles (Fig. 2a, b). Carapace without grooves, inferior margin of orbit and pter- ygostomian angle slightly produced (Fig. 2a), pterygostomian regions produced lat- erally (Fig. 2b). Abdomen smooth, somites 1—2 with rounded pleura, somites 3—5 with posterior angle of pleura subacute, sixth somite with posterior margin sinuous at insertions of telson and uropods. Telson 2.5 times as long as its basal width, tapering distally, distal width less than half of basal width; bearing four pairs of movable spines on dorsal surface, spines located on distal two thirds of dorsal surface; posterior margin rounded, bearing 9 spines, second pair from external one longest (Fig. 4g). Eyes pigmented, fused, forming part of a single plate, peduncle and cornea not dis- cernible, projected dorsally (Fig. 2c). An- tennule with first segment as long as seg- ments 2 and 3 combined; stylocerite acute, reaching distal margin of first segment (Fig. 4e). Antennal scale 1.8 times as long as wide, laterodistal tooth short not exceeding distal margin of blade (Fig. 4f), flagellum 1.25 times total length (Fig. 2a). Mandible with stout 2-segmented palp, incisor process with six distal teeth, molar process conical, sharp distal end (Fig. 2d). Both mandibles approximately symmetri- cal. First maxilla with distal lacinia oval shaped, bearing three rows of short, thick setae on mesial surface; proximal lacinia with single row of short, thick setae on dis- tomesial margin; palp bearing one distal, long setae and two subdistal short ones on internal margin (Fig. 2e). Second maxilla with scaphognathite approximately rectan- gular distally, subtriangular proximally; dis- tal margin with long, plumose setae; lateral VOLUME 117, NUMBER 3 37 Fig. 2. Agostocaris acklinsensis, new species, a female holotype, b-f female paratype: a, total lateral view: b, carapace, dorsal view; c, dorsal view of eyes, carapace removed; d, mandible: e, first maxilla; f, first maxil- liped. Scale bar represent: a—c, f, 1 mm; d—e, 0.5 mm. 372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Agostocaris acklinsensis, new species, female paratype: a, second maxilla; b, second maxilliped; c, third maxilliped; d, first pereiopod; e, detail of propodus and dactylus of first pereiopod; f, second pereiopod. Scale bars represent: a—d, f, 1 mm; e, 0.5 mm. VOLUME 117, NUMBER 3 373 Fig. 4. Agostocaris acklinsensis, new species, female paratype: a, third pereiopod; b, detail of proximal segments of third pereiopod; c, fourth pereiopod; d, fifth pereiopod; e, antennule; f, antenna; g; telson and uropods, left side omitted; h, first pleopod; i, second pleopod. Scale bars represent 1 mm. 374 margin with short plumose setae; internal margin with long simple setae, increasing in length distally, almost as long as sca- phognathite; palp digitiform, devoid of se- tae; distal endite trapezoidal, middle and proximal endites approximately rectangular, all three bearing simple setae on distal mar- gins (Fig. 3a). First maxilliped with triangular endite bearing marginal setae; palp digitiform, with apical tuft of setae; exopod elongated, bearing long, simple setae distally; caridean lobe broadly rounded, with submarginal row of short setae and long plumose setae along margin; epipod bilobed, both lobes trapezoidal, distal one smaller, devoid of se- tae (Fig. 2f). Second maxilliped with en- dopod pediform, 4-segmented, with contin- uous row of setae along margin; exopod slender, bearing long simple setae distally; epipod simple, flat, rounded (Fig. 3b). Third maxilliped with endopod 4-segmented, bearing setae on mesial margin; exopod as long as first segment of endopod, with dis- tal tuft of long setae; epipod digitiform, less than half the length of exopod; arthro- branches present (Fig. 3c). First pereiopod with ischium and merus of about same length and width, carpus wider proximally, propodus articulating with carpus at one third of its length, palm as long as fingers, cutting edges of both fin- gers with minute sharp teeth, dactylus with long setae arising from proximal half teeth (Fig. 3d); exopod as long as ischium and merus combined, with apical tuft of long setae; arthrobranch and pleurobranch pre- sent (Fig. 3d). Second pereiopod longer than first one, with merus slightly shorter than ischium, carpus becoming wider dis- tally and as long as merus, propodus with palm shorter than fixed finger, dactylus heavier and longer than fixed finger; exo- pod shorter than ischium and merus com- bined, bearing apical tuft of long setae; ar- throbranch and pleurobranch present (Fig. 3f). Third pereiopod with ischium with two spines, merus the longest segment, carpus and propodus of about the same length, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dactylus with corneous sharp tip and four smaller teeth on internal surface, arthro- branch and pleurobranch present, finger- like exopod arising from basis (Fig. 4a, b). Fourth pereiopod with ischium with two spines, merus the longest segment, carpus and propodus of about the same length, dactylus with corneous sharp tip and three smaller teeth on internal surface, arthro- branch and pleurobranch present, finger- like exopod arising from basis (Fig. 4c). Fifth pereiopod with ischium with one spine, propodus the longest segment, dac- tylus with corneous sharp tip and eight smaller teeth on internal surface, arthro- branch and pleurobranch present (Fig. 4d). First pleopod with exopod setose, endo- pod devoid of setae, one third the length of exopod (Fig. 4h). Second pleopod with en- dopod and exopod setose, appendix interna slender more than half the length of endo- pod (Fig. 41). Uropods with external ramus bearing one distolateral movable spine, distal margin broadly rounded, with long plumose setae on distal and internal margins. Internal ra- mus bearing marginal long plumose setae except on proximal third, distal margin sub- acute (Fig. 4g). Etymology.—The specific name is de- rived from “‘Acklins”, the name of the Ba- hamian island where the new species was captured. Key to the species of Agostocaris 1. First maxilliped with palp 2-segmented, ischium of fifth pereiopod devoid of spines, outer ramus of uropods devoid of distolateral spines, .. Agostocaris williamsi — First maxilliped with palp unsegmented, ischium of fifth pereiopod with spines, outer ramus of uropods with spines, out- er ramus of uropods with distolateral SPINES ce Paes see Sime aoe eee eee et 2 . Ischium of fifth pereiopod with two spines, outer ramus of uropod with two distolateral spines, telson with five pairs of dorsal spines i) VOLUME 117, NUMBER 3 — Ischium of fifth pereiopod with one spine, outer ramus of uropod with one distolateral spine, telson with four pairs of dorsal spines Agostocaris acklinsensis Remarks.—Agostocaris acklinsensis can be easily distinguished from the other two known species in the genus by the presence of: exopods on the third and fourth pereio- pods, a fifth pereiopod with one spine on the ischium and one distolateral movable spine on the outer ramus of the uropods. Other taxonomically important characters vary among the three species. A second maxilla with a palp devoid of setae and an unsegmented palp of the first maxilliped distinguish A. acklinsensis from A. william- si, whereas the number of dorsal spines on the telson, unpigmented eyes and two dis- tolateral spines on the outer ramus of the uropods seprate A. bozanici (Table 1). Noteworthy are the eyes of Agostocaris, which are composed of one single plate not differentiated into peduncle and cornea. This plate is projected outside the orbits creating the eye-like structures, which in the three species are pointed distally. Since all the species of Agostocaris are cave dwellers it is reasonable to suppose that the cornea was lost and later the peduncle was reduced, in such a way that the “‘eyes”’ we see now are part of the basal plate. This singular morphology merits further studies on its ontogeny and functionality. The placement of the genus Agostocaris is a matter of controversy. Holthuis (1993), by synonymizing Agostocarididae with the Bresiliidae, gave more weight to characters that are shared by many taxa in the Caridea (mandible with palp, carpus of second legs undivided, first two pairs of legs chelate, first pair of legs more robust than second one, Williams, 1984) with little resolution among families, than to exceptional auta- pomorphic characters such as the fused eyes and the particular morphology of the first two pereiopods of Agostocaris. We agree with Martin & Davis’ (2001) proposal of recognizing a superfamily Bre- Table 1.—Comparison of selected characters of the three species of Agostocaris. A. acklinsensis A. bozanici A. williamsi Weakly pigmented Without pigment Weakly pigmented Palp with setae Eyes Palp without setae Palp without setae Second maxilla Palp unsegmented Palp unsegmented Palp 2-segmented First maxilliped Basis with finger-like exopod Basis without exopod Basis without exopod Basis without exopod Basis without exopod Third pereiopod Basis with finger-like exopod Ischium with one spine Fourth pereiopod Fifth pereiopod Ischium with two spines Appendix interna less than half Ischium devoid of spines Appendix interna less than half Appendix interna two thirds length of Second pleopod length of endopod With four pairs of dorsal spines length of endopod With five pairs of dorsal spines endopod With four pairs of dorsal spines Telson Outer ramus without distolateral Outer ramus with two distolateral Outer ramus with one distolateral Uropods spine spines spines 375 376 silioidea, which includes five families, and concur with the opinion that this taxon still represents an artificial grouping. While it is beyond the scope of this paper to discuss the relationships among bresilioids, it is clear that Agostocarididae represents a dis- tinct family that can be easily separated from the other four bresilioid families. The Alvinocarididae Christoffersen, 1986, and Mirocarididae Vereshchaka, 1997, lack ex- opods on all pereiopods, whereas the Agos- tocarididae can have exopods on all five pe- reiopods. The Diascididae Rathbun, 1902, have well developed eyes with peduncle and cornea, a dorsoventrally flattened ros- trum and a disc-like dactylus of the first pe- reiopod, contrasting with the fused eyes, acuminate rostrum and typically shaped dactylus of pereiopod 1 of the Agostocari- didae. Finally the Bresiliidae, and the rest of the bresilioid families, can be separated from the Agostocarididae based on the car- pus-propodus articulation of the first pereio- pod which is normal in the former, being the distal end of the carpus articulated to the proximal end of the propodus; while in the latter the carpus is articulated to an area close to the middle portion of the propodus. In addition, the chela of the second pereio- pod in the Agostocarididae is unique in that the digitiform dactylus is longer than the fixed finger and lacks teeth or spines. Acknowledgments Collection of shrimp described herein was part of the January 1999 Anchialine Caves Expedition to the southern Bahamas led by Thomas Iliffe. Other members of the expedition included Texas A&M University graduate students Brett Dodson and Shelley Fetterolf. This expedition was funded by National Science Foundation, Biotic Sur- veys and Inventories Program award num- ber 9870219. We thank Neil Sealey (Media Publishing Ltd, Nassau, Bahamas), Dr. Nancy Elliott (Sienna College) and Dr. Wil- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON liam Keegan (Florida Museum of Natural History) for providing invaluable logistical information on Crooked and Acklins Is- lands. The drawings were prepared by Ro- lando Mendoza. Literature Cited Calman, W. T. 1896. On deep sea Crustacea from the south west of Ireland.—Transactions of the Royal Irish Academy 31:1—22. Christoffersen, M. L. 1986. Phylogenetic relationships between Oplophoridae, Atyidae, Pasiphaeidae, Alvinocarididae fam. n., Bresiliidae, Psalido- popidae and Disciadidae (Crustacea Caridea Atyoidea).—Boletim Zoologico, Universidade do Sao Paulo 10:273-281. De Haan, W. 1849 (1833-1850). Crustacea. Jn P. EK von Siebold, ed., Fauna Japonica sive descriptio an- imalium, quae in itinere per Japonium, Jussu et auspices superiorum, qui summum in India Ba- tava imperium tenent, suscepto, annis 1823- 1830 collegit, notis, observationibus et adum- brationibus illustravit. Lugduni-Batavorum, 243 Pp- Hart, C. W., Jr, & R. W. Manning. 1986. Two new shrimps (Procarididae and Agostocarididae, new family) from marine caves of the western north Atlantic_—Journal of Crustacean Biology 6:408-416. Holthuis, L. B. 1993. The Recent Genera of the Car- idean and Stenopodidean Shrimps (Crustacea, Decapoda). Nationaal Natuurhistorisch Muse- um, Leiden, 328 pp. Kensley, B. 1988. New species and records of cave shrimps from the Yucatan Peninsula (Decapoda: Agostocarididae and Hippolytidae).—Journal of Crustacean Biology 8:688—699. Martin, J. W., & G. E. Davis. 2001. An updated clas- sification of the recent Crustacea. Natural His- tory Museum of Los Angeles County, Science Series 39, 124 pp. Rathbun, M. J. 1902. Papers from the Hopkins Stan- ford Galapagos Expedition 1898-1899. VIII. Brachyura and Macrura.—Proceedings of the Washington Academy of Sciences 4:275—292. Vereshchaka, A. L. 1997. A new family for a deep-sea caridean shrimp from North Atlantic hydrother- mal vents.—Journal of the Marine Biological Association of the United Kingdom 77:425— 438. Williams, A. B. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, 550 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):377-384. 2004. A new species of caridean shrimp of the family Stylodactylidae from the eastern Pacific Ocean Mary K. Wicksten and Joel W. Martin (MKW) Department of Biology, Texas A&M University, College Station, Texas 77843-3258, U.S.A., e-mail: wicksten@mail.bio.tamu.edu (JWM) Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, U.S.A., e-mail: jmartin@nhm.org Abstract.—Four specimens of shrimp of the family Stylodactylidae were collected at two stations off Baja California, Mexico, and California, U.S.A. These are the first specimens of the family reported from the eastern Pacific. The shrimp are described as a new species, Bathystylodactylus echinus. The species can be recognized by the following features: rostrum straight, much longer than the carapace, bearing at least 23—27 dorsal and 18—25 ventral spines; eye small and without pigment, stylocerite slender and not reaching middle of first segment of antennular peduncle, carapace without prominent posterior dorsal hump, body set with minute spinules, posterior pereopods con- siderably longer than anterior two pair, slender and lacking fringe of setae. Shrimp of the family Stylodactylidae are recognized by their peculiar first and sec- ond pereopods, which end in elongate but nearly equal fingers with setae on the cut- ting edges. These pereopods and the max- illipeds are densely setose. Species of the family are widely distributed from tropical to temperate regions (e.g., Cleva 1990a), al- though most of the species described to date have come from the tropical Indo-Pacific (Chace 1983; Cleva 1990b, 1994, 1997; Okuno and Tachikawa 2000). Chace (1983) and Cleva (1994) reviewed the members of the family, described new species, and pro- vided keys. Hanamura and Takeda (1996) described an additional genus, Bathystylo- dactylus, for a new species (B. inflatus) from off Taiwan (and for the former Sty- lodactylus bathyalis from the Coral Sea), bringing to 5 the number of recognized genera in the family (Stylodactylus, Neos- tylodactylus, Parastylodactylus, Stylodac- tyloides, and Bathystylodactylus). There have been no previous reports of the family in the eastern Pacific Ocean. While sorting specimens in the Benthic Invertebrate Collection of Scripps Institu- tion of Oceanography, we found four spec- imens of shrimp of this family from three stations taken off California, U.S.A., and Baja California, Mexico. The specimens in- clude both males and females. We com- pared these specimens with specimens of Stylodactylus rectirostris in the collections of Texas A&M University (catalog number 2-7212, Oregon station 5916) and with published descriptions of other species in the family. The specimens represent an un- known species of Bathystylodactylus, de- scribed herein. Systematic Account Bathystylodactylus echinus, new species Figs. 1-5 Holotype: Male, carapace length (CL) 32.7. Basin off Magdalena Bay, Baja Cali- fornia, Mexico (24°35’N, 113°25’W), 3563-3621 m, 6-foot Sigsbee trawl, 24 June 1965, ship Horizon sta. MV65-I-38, Carl Hubbs, collector; Scripps Institution of 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Scale bar = 10.0 mm. Oceanography (SIO) catalog number C3188. Paratypes: Male, CL 41.4, same station as holotype, LACM CR 1965-349.1 (Nat- ural History Museum of Los Angeles County). Female, CL 29.7. Basin off Mag- dalena Bay, Baja California, Mexico (24°23'N, 113°17’W), 3427-3621 m, 45- foot otter trawl, 25 June 1965, ship Horizon 2 sta., MV65-I-39, C. Hubbs, SIO cat. no. 3203. Female CL 27.6 Patton Escarpment (32°25'N, 120°40'W), 3689-3630 m, 40- foot otter trawl, 7 Feb. 1981, ship New Ho- rizon sta. 133, collector S. Luke, SIO cat. no. C10324. Description: Rostrum (Figs. 1, 2B, C) nearly straight, nearly 2X length of cara- pace but broken in all specimens, with 23— 27 movable dorsal and 18—25 ventral spines; series of 7—9 minute spinules on carapace just posterior to rostrum proper, long setae along distal ventrolateral surface. Carapace (Fig. 2A) with hepatic depression well delineated. Antennal and branchioste- gal spines short but obvious, antennal spine Bathystylodactylus echinus, new species, male holotype, Scripps Institution of Oceanography C3188. located ventral to suborbital angle. Lateral surface of carapace inflated over branchial region, suprabranchial carina curved. Area posterior to eye and antennal origin slightly depressed. Anterior regions of carapace set with small, simple, movable spinules, pos- terior regions punctate or with few spinules. Abdomen (Fig. 1) with small spinules on dorsal and lateral surfaces, somites one and two rounded dorsally, somite three weakly carinate dorsally; somite four rounded to weakly carinate, with or without shallow depression interrupting dorsal carina; pleura of somites rounded, those of somites four and five (Fig. 5B) each with sharp poster- oventral spine; one specimen with minute spine on pleuron of somite three. Telson (Fig. 5C, E) 8X longer than wide, tapering to apex, with 11—13 pairs of dorsolateral spines located on weak ridges and numer- ous small spinules; two mesial spines flank- ing apex on either side. (Apex of telson pre- served in only one specimen; observed asymmetry may be due to injury.) VOLUME 117, NUMBER 3 37/9) Fig. 2. Bathystylodactylus echinus, new species, male holotype, carapace and rostrum. A, carapace and eye, lateral view. B, rostrum (attached at area of dashed lines in A and illustrated at same scale as A). C, higher magnification of region of rostrum shown in B and denoted by arrows. Scale bar = 10.0 mm A, B; 2.5 mm C. Eyes (Figs. 1, 2A, 5A) reduced, cornea without trace of pigment. Antennular peduncle (Fig. 5A) elongate. Stylocerite slender, not reaching middle of first segment. First and second segments subequal in length, third segment very short. Antennal scale (scaphocerite) more than 4X long as broad, outer margin slight- ly concave, with microscopic spinules, not reaching end of second segment of anten- nular peduncle, blade exceeding distolateral spine. Carpocerite covered by minute spi- nules, reaching second segment of anten- nular peduncle. Basicerite bearing strong lateral spine. Mandible (Fig. 3A) with molar process bearing teeth in the following configuration: 2 small, one large, 4 small and large blunt process; stout, 2-jointed palp present. First maxilla (Fig. 3B) with distal endite broad 380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Bathystylodactylus echinus, new species, male paratype (LACM CR 1965-349.1). A, mandible; B, first maxilla; C, second maxilla; D, inner surface of second maxilla slightly enlarged and showing palp:; E, first maxilliped. Scale bar = 5.0 mm A, B, D; 10.0 mm C, E. and with stiff mesial setae; proximal endite 2 more proximal endites; long palp ending curved inward and ending in brush of setae; in 5 setae, scaphognathite with anterior half palp ending in long setae and having tufts rounded, posterior half slender and curved of setae on lateral surface. Second maxilla mesially, bearing long setae. First maxilli- (Fig. 3C, D) with distal endite larger than ped (Fig. 3E) with long distal and short VOLUME 117, NUMBER 3 proximal endites; palp reaching 3/4 length of distal endite and ending in tuft of setae; exopod with lash, well developed caridean lobe and deeply bilobed epipod. Second maxilliped (Fig. 4A) much larger than inner mouthparts, with exopod having lash and reaching end of basal segments; podobranch and epipod present; basal seg- ments fringed with stiff curved setae; an- tepenultimate segment short, with long sim- ple setae on flexor margin at articulation with basal segments; penultimate segment with fringe of long setae on flexor margin; two terminal segments; that on flexor side longer than one on extensor side, both fringed with long setae. Third maxilliped (Fig. 4B) setose, with arthrobranch but without exopod, exceeding antennular pe- duncle by about length of distal segment. Ultimate segment longest, with dense setae on flexor side. Penultimate segment with long, pinnately branched setae. Antepenul- timate segment with both long and short se- tae. Pereopods all lacking exopods or epi- pods. First pereopod (Fig. 4B, C) with en- tire flexor surface fringed with long setae, merus longer than carpus, propodus about equal in length to carpus, ending in elon- gate chela (Fig. 4C); fingers simple, with long setae and shorter spine-like setae along cutting edges. Second pereopod similar to first. Third to fifth pereopods (Fig. 1) elon- gate, with few scattered setae; merus of third pereopod with 8—10 spines on flexor and lateral surfaces; merus of fourth pereo- pod with 15, merus of fifth pereopod 14; carpus shorter than merus; propodus broken and dactylus missing in all specimens. All pleopods densely setose. First pleo- pod shorter than second to fifth pleopods. Male second pleopod (Fig. 4D, E) with ap- pendix interna and appendix masculina, ap- pendix masculina reaching nearly % length of appendix interna, with apex notched and bearing small hooks. Lateral branch of uropod with spinules, margin nearly straight, two small teeth by 381 suture (Fig. 5D). Uropods shorter than tel- son. Etymology.—The specific name is de- rived from the Greek word for spiny. Remarks.—The new species can be as- signed to the genus Bathystylodactylus ac- cording to the features given by Hanamura and Takeda (1996). The new species bears a well-developed and two-jointed mandib- ular palp. Both sexes bear well-developed arthrobranchs on the four anterior pereo- podal somites. There is no supraorbital spine. The stylocerite falls far short of the mesiodistal margin of the basal segment. There are no fringes of setae on pereopods 3—5, as there are in Stylodactylus rectiros- tris and other species of Stylodactylus. Han- amura and Takeda (1996) mentioned that the third to fifth abdominal somites were ““weakly carinate”’ dorsally. In our speci- mens, only somite three is consistently weakly carinate. The posterior three pereo- pods definitely are longer than the anterior two in the new species, but due to breakage, their relative lengths to each other cannot be determined. Two species of Bathystlyodactylus have been described previously: B. bathyalis (Cleva, 1994), from the Coral Sea (as Sty- lodactylus bathyalis); and B. inflatus Han- amura and Takeda (1996), from off Taiwan (Hanamura and Takeda 1996). Bathystylo- dactylus echinus can be distinguished from the former by its curved rostrum and char- acteristic sharp spine on the ventral margin of abdominal pleuron three. Like Bathys- tylodactylus inflatus, B. echinus has a straight rostrum with numerous dorsal and ventral spines. The pleura of the fourth and fifth abdominal somites each bear a poster- oventral spine. However, in B. inflatus the carapace has a marked wide elevation near the posterodorsal margin. This is not pre- sent in B. echinus. The shape of the supra- branchial carina is more sinuous in B. infla- tus than in B. echinus. In B. inflatus, there are 11 spinules on the carapace posterior to the rostrum; in B. echinus, there are 8—9. In B. inflatus, there are 9—10 dorsal rostral 382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ~~ SSS SSS 5 SSS e e Pi tiy a Fi bia ) PER miei 1] \ us SNARK Fig. 4. Bathylstylodactylus echinus, new species, male paratype (LACM CR 1965-349.1) (A) and holotype (SIO C3188) (B-E). A, second maxilliped (paratype). B, right third maxilliped (upper appendage) and first pereopod (holotype). C, higher magnification of chela of first pereopod (tips of fingers broken). D, second pleopod (holotype). E, higher magnification of appendix interna and appendix masculina (arrow from D). Scale bar = 10.0 mm A, E, E; 7.5 mm B, C. VOLUME 117, NUMBER 3 383 Vv Oy _ Ze TD; 7 ZS > ZL LEE \ ) y] K — Fig. 5. Bathystylodactylus echinus, new species, male paratype (A) and holotype (B—E). An, antennule, antenna, and eye (e), right side, dorsal view, male paratype (LACM CR 1965-349.1). sc = scaphocerite; st = stylocerite. B, lateral view of abdominal somite 6 plus portions of the telson, uropods, and pleurae of somites 4 and 5, holotype. C, telson and right uropods, dorsal view, holotype. D, higher magnification of distolateral area of outer uropod (arrow from C). E, higher magnification of tip of telson (arrow from C). Scale bar = 10.0 mm A, B; 7.5 mm C; 3.75 mm D, E. 384 spines located proximally to the origin of the first ventral rostral spine; in B. echinus, there are no more than 4. The integument of B. inflatus was described as “thin” and the body consequently “soft.” In B. echi- nus, the integument appears to us to be typ- ical of a benthic caridean, and not membra- nous (as seen in midwater species of the Oplophoridae, for example). Cleva (1994) and Hanamura and Takeda (1966) described the body of Bathystylo- dactylus species as “‘pubescent.”’ Their il- lustrations show a very light coating of pile. In B. echinus, the spinules on the body are characteristic and easily seen, especially on the dorsal aspect of the carapace. These spi- nules conform in shape and structure to tac- tile or vibrational sensory structures seen in other crustaceans (Cohen and Dijkgraaf 1961). Bathystylodactylus echinus is the largest and deepest species known in its family. It was collected with the flatback lobster Wil- lemoesia inornata Faxon at stations MV65- I-38 and MV65-I-39, and with the galatheid crab Munidopsis antonii (A. Milne-Ed- wards) at station MV65-I-39. Acknowledgments We thank Larry Lovell, Scripps Institution of Oceanography, for allowing us to ex- amine the specimens and offering assis- tance and hospitality during a visit. We also thank an anonymous reviewer for alerting us to a potential synonymy. The study ben- efited from partial support from NSF grants DEB 9978193 to J. Martin and D. Jacobs (from the PEET Initiative of Systematic Bi- ology), DEB 0120635 to Cliff Cunningham et al. (from the Biocomplexity Genome-En- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON abled Research program), and DEB 0138674 to J. Martin et al. (for collection support). Literature Cited Chace, FE A. Jr. 1983. The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedi- tion, 1907-1910, part 1. Family Stylodactyli- dae.—Smithsonian Contributions to Zoology 381: 1-21. Cleva, R. 1990a. Sur les Stylodactylidae (Crustacea, Decapoda, Caridea) de Il’ Atlantique —Bulletin Muséum National d’Histoire Naturelle 4, sér. 12, sect. A, 1: 165-176. . 1990b. Crustacea Decapoda: les genres et les espéces indo-ouest pacifiques de Stylodactyli- dae. Pp. 71-136 in A. Crosnier, Résultats des Campagnes MUSORSTOM, vol. 6.—Mémoires du Museum National d’ Histoire Naturelle (A), 145. . 1994. Some Australian Stylodactylidae (Crus- tacea: Decapoda) with descriptions of two new species.—The Beagle, Records of the Museum and Art Galleries of the Northern Territory 11: 53-64. . 1997. Crustacea Decapoda: Stylodactylidae récoltés en Indonése, aux iles Wallis et Futuna et au Vanuatu (Campagnes KARUBAR, MU- SORSTOM 7 et 8). Données complémentaires sure les Stylodactylidae de Nouvelle-Calédonie. Pp. 385—407 in A. Crosnier & P. Bouchet, eds., Résultats des Campagnes MUSORSTOM, 16. Mémoires du Muséum National d’ Histoire Na- turelle, Paris 172. Cohen, M. J., & S. Dijkgraaf. 1961. Chapter 2. Mech- anoreception. Pp. 65—108 in T. H. Waterman, ed., The physiology of Crustacea, vol. II. Aca- demic Press, New York, 681 pp. Hanamura, Y., & M. Takeda. 1996. Establishment of a new genus Bathystylodactylus (Crustacea: De- capoda: Stylodactylidae), with description of a new species from northwestern Pacific.—Zoo- logical Science 13:929—934. Okuno, J., & H. Tachikawa. 2000. A new species of the genus Neostylodactylus Hayashi & Miyake, 1968 (Crustacea, Decapoda: Stylodactylidae) from southern Japan.—Proceedings of the Bio- logical Society of Washington 113:39—47. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):385-397. 2004. A new pedunculate barnacle (Cirripedia: Heteralepadidae) from the Northwest Atlantic L. Buhl-Mortensen and W. A. Newman (LB-M) Benthic Habitat Research Group, Institute of Marine Research, PB. 1870 Nordnes N-5817 Bergen, Norway, e-mail: lene.buhl-mortensen @imr.no; (WAN) Marine Biological Research Division, Scripps Institution of Oceanography, La Jolla, California 92093-0202, U.S.A. Abstract.—A species of Heteralepas has been discovered attached to a gor- gonian coral from 500 meters of depth off Nova Scotia (~42° N). A brief review of the previously described Heteralepas species is presented. Of the 29 previously described species (including 2 in synonymy), the new species is more similar to some from the Indo- West Pacific than to any of the 8 previously known species from the Atlantic. While the new species can be distinguished from Atlantic but not some Pacific species by some characters, it can be dis- tinguished from all species of the genus by small but marked differences in the configuration of the apertural region of the capitulum. Therefore it is pro- posed as a new species, Heteralepas cantelli, the most northern known member of the family. Introduction A specimen of Heteralepas was discov- ered during a survey of the coral-associated fauna at ~42°N on the continental shelf and slope off Nova Scotia, Canada, in 2002. This is not only several degrees of latitude farther north than any previously known species of the genus along the Atlantic sea- board, but at a higher latitude than any pre- viously known species of Heteralepas (Young 1999, Zevina 1982). It was collect- ed by a benthic trawl from ~500 m of depth, attached to the gorgonian Primnoa resedaeformis (Gunnerus, 1763). Thirty species of Heteralepas have been described, and 2 of these are presently in synonymy. Of the 28 recognized species (Table 1), 8 have been recorded from the Atlantic; 3 from the Western Atlantic, 4 from the Eastern Atlantic and 1 found in both areas; H. cornuta (Darwin, 1852), H. lankesteri (Gruvel, 1901), H. belli (Gruvel, 1901), and A. luridas (Zevina, 1975), and H. microstoma (Gruvel, 1901), H. meteo- rensis Carriol, 1998, H. alboplaculus Zev- ina & Kolbasov, 2000 and H. segonzaci Young, 2001 plus H. cornuta respectively (Zevina 1975, Young 2001). All these spe- cies are from relatively low latitudes and none compare favorably with the new form. The closest affinities of the new form are with species like A. japonica (Aurivillius, 1892) from the Indo-West Pacific. However, the new form can be distinguished from all previously described species by character- istics of the apertural regions, and therefore it is considered to represent a new species. Systematics Subclass Cirripedia Burmeister, 1834 Superorder Thoracica Darwin, 1854 Order Pedunculata Lamarck, 1818 Suborder Heteralepadomorpha Newman, 1987 Family Heteralepadidae Nilsson-Cantell, 1921 Pilsbry (1907b) revised Alepas and ex- tracted two distinct but related taxa from it, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 386 00S = CETT = O0E—08C = O0€ = O87 Ieosesepry] JON LIC Bog eulyD O0€ = OOV uedeg O00Z—00€ = 8CC == OSV eIUeUse ], [ephzcjuy ueder EL-SS purleoz MON 6CE-LLE ueder é spuv|s] 1eqosIn G — 8cr SIV spueys] ueieMmeyy VOC BISAR]eIAI 86 BISALRI vs BISAveyy S16-8EC SUeNS ye” €C9-69C = é — 00S I-16 = purleoz Mon ‘ueder puvleoZ MON ‘elensny ‘sourddiyyryq “sy uorun -dy 0} Isom pure JoyeM MOTTeEYS — JUNOUWIeSS IOD}9I/\ — JUNOWIvIS IODIO//\I — JUNOWIvaS IODI9//\ oytoed “a's = oyloed “A'S = BIUIOFE DA = JUNOUWILIS IO = -9J “VIlopey] “Sol0zVv BOHOWW °S 77 N = BNOOS PAON uvoqque) eqng [[ze1g “SOIpUuy 1soAA ee “AOU “ds 17/2JUD) “ET QZ 100% ‘SuNOX 19p2uU0sas “LZ 0007 “Aosegioy 2 vUIA[Z snjnonjdogy, “H 9 8661 TOLUeD sisuasoajau, “H +ST 7661 “ISplesi9ysg 2 eUIAIZ, vssaf E861 “USy sujiuis 786 “BUIAOZ, vaynf 786] “BUIAIZ, Dsodipp CL6[ PUIASZ sppliny P96| ‘ULUIMON DIOYdooDsKu 096[ “‘UPWIMeEN MmouljN LE61 ON wipy CCHL “YDOIG VIgnpy, L161 “Arqstid vjnjaa 606] “o[epueuuYy voLIMMgGoolU QLO6I “Argsttq snusko (82061 “A1qstid) xa4 (L061 20H) sinuay (L061 80H) s7pa0 (SO6I ‘S[epueuUY) DUDISKojDU “A 8 (SO6I “s[epueuuy) svsis YL Ke) (L061 “(eAnID) viuojsosIU “FY 2.9 (1061 “12ADID) 17729 “H xS (O06 “‘[PANID) Majsayun] “Hs (LE6] ‘OIN) BlODIUOUDsad “YT = (V68I ‘SNIALINY) vivIponb “Hy ¢ (1061 ‘TeANID) voIpul “= 0ZO'I-8r puvjeoZ MON 0} ueder == = = (Z68I ‘SnpIALNY) vo1u0dvl Fy Z JUNOUWIVIG ODIO] CLEbE sourddry1yg ‘eIlopey] O} Spuryst [Ize1g 0} euljores; OILZ-EL ‘eas uewiepuy eyitt@yn SIOYSJJO 2 VW AA YON ‘sorpuy iso (ZSQI ‘UIMIe) VInNUIOD “YY SS (uw) yidaq oytoeg 1S9A\-Opuy oyloeg uso\seq onuepy wloyseq onuepyy WI9}sa sedayeinjay eee ‘Aydeis0es 10/pue Asojoydiour [eussixa ur Ayre] IIs JO asnedeq “AOU “ds 1jajupd “FY YUM poreduroo ore yeu) saroods = ,, ‘(UOISSNOSIpP 998) AJUIeVIOOUN oTWIOUOXe) = { ‘([O0Z) SUNO,A pue (0007) ACseq[oy 2 CUIAOZ “(8661) [OLUVD “(€861) USA “(6L61) J9}SOq Wo pofiduios ejep yIdep pue Aj[eoO] WIOIJ ULID0 P]IOM oy) JO svdajpsajayy snuss ay) Jo so1loadgs— | 3IqGuL, VOLUME 117, NUMBER 3 Heteralepas and Paralepas, but he left them in the family Lepadidae. Nilsson-Can- tell (1921) noted that these two genera, in addition to lacking calcareous plates, dif- fered from the remaining Lepadidae in the nature of their trophi and cirri and therefore he proposed a new family, the Heteralepa- didae, for them. Species of Heteralepas are generally considered to have ctenopod or lasiopod cirri used for setose feeding, while those of Paralepas have acanthopod cirri, generally used to feed on the food or tissues of their hosts, including the eggs of hosts such as spiny lobsters. The two genera are further distinguished by the inner ramus of the posterior two pairs of cirri (cirri V & VI) being similar to the outer rami in Par- alepas, but conspicuously reduced in length and breadth in Heteralepas. However, as will be noted in the discussion, there is at least one species that is somewhat inter- mediate in these characters and it likely should be assigned a genus of its own. Heteralepas Pilsbry 1907b Heteralepas cantelli sp. nov. (Figs. 1—4) Type material.—The sole specimen (ho- lotype) is deposited in the National Muse- um of Washington, Washington, D.C. USNM 1019509. Etymology.—Named in honor of the Swedish cirripedologist, Carl August Nils- son-Cantell (cf. Newman 1990) who erect- ed the family Heteralepadidae. Material.—Known from a single speci- men collected in the Northeast Channel, south of Nova Scotia, Canada (41°55.9’N, 65°42.5'W), by a commercial bottom trawl- er on October 9, 2002 from 500 m depth. It was attached to the exposed skeleton of the gorgonian Primnoa resedaeformis. Diagnosis: Capitulum and peduncle rel- atively smooth, without tubercles, carinal ridge or indications of the insertions of the carapace adductor muscle; apertural region slightly recessed or depressed below gen- eral surface; aperture ~ % height of the ca- 387 pitulum, with crenulate lips restricted to up- per %. Description The fresh specimen was translucent yel- lowish pink. The capitulum is 3 cm high and 2 cm wide, globular or nearly ovoid in lateral aspect, slightly pointed apically, lat- erally compressed, frontal margin interrupt- ed by a depressed apertural region with slightly protuberant lips in the upper % of the aperture (Figs. 1A, B, 2A, B). The slightly recessed apertural region is outlined by a thin edge and in the region below it, where the carapace adductor muscle is found, there is a chin-like thickening. Oth- erwise the capitulum is smooth, without carinal crest or ridge, warts, bumps or pro- tuberances. Aperture % height of capitulum; crenate lips produced in upper 4%. Peduncle 1.2 cm in diameter, equal in length to ca- pitulum and marked with several folds and lines in the otherwise smooth cuticle, basal portion expanded into attachment disc. La- brum too damaged to describe; mandible (Fig. 3B) with 4 teeth including inferior an- gle, surface covered with numerous fine se- tae, lower margins of teeth 1—3 with a few fine pectinations (5 under the first and sec- ond, and 3 under the third tooth; Fig. 3B, al-a3). First maxilla (Fig. 3C) with cutting edge stepped (plane of superior cutting edge indented relative to plane of inferior cutting edge) rather than notched, with three major spines (one large flanked by two somewhat smaller ones) above and ap- proximately 14 spines below step, with soft setae in a group along the superior margin and spread out along the inferior margin, lateral surfaces clothed with numerous se- tae. Second maxilla (Fig. 3A) with a prox- imal cluster of long spine-like setae and a similar array of setae separated into two groups along the cutting edge. Cirrus I not separated from posterior pairs but modified as a maxilliped of rela- tively short, unequal, densely setose rami; cirri II-VI basically similar in structure, se- 388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lang, Il. ertural region; B, lateral aspect of entire animal; tation lasiopod (Fig. 4C). However, while cirri II-IV have long subequal rami nearly equal in length to the outer rami of cirri V and VI, the inner rami of V and VI are at- rophied (Fig. 4D, Table 2). The number of articles comprising the cirri is as follows: Cirrus: I Il Ill Inner ramus: 18 42 Outer ramus: Mal 5)II IV V VI 30) I) 2D" 2 S71 33 OO Caudal appendage (Fig. 4D) of 13 articles, slightly longer than pedicel of cirrus VI. Pe- nis (Fig. 4A, B) relatively long, slender, an- nulated, without specialized hooks or grap- Heteralepas cantelli sp. nov.: A, right-frontal aspect of capitulum, enlarged to show details of ap- ples but clothed with numerous, long soft setae distally. Discussion: The cirri of the new species are fully lasiopod (Fig. 4C) and the inner rami of the cirri V & VI are substantially reduced in length as well as breadth (Fig. 4D; Table 2). Therefore the new species is a Heteralepas in the strict sense. The num- ber of articles comprising the rami of the cirri and the form of the mouthparts, while sometimes useful in distinguishing species, are considered somewhat variable (Nilsson- Cantell 1921), as are elaborations of the ca- pitulum as well as its length relative to the peduncle (Young 2001). Therefore keys, VOLUME 117, NUMBER 3 1 cm 389 Fig. 2. region; B, lateral aspect: such as that presented by Zevina (1982) for the 19 species of Heteralepas recognized at the time, should be used with caution. Zevina (1982) did not include complete synonymies in her monograph, and at least two species once attributed to Heteralepas were assigned to Paralepas without amend- ing either genus. Therefore we review all species attributed to the genus and, as can be seen from Table 1, 28 species (including the new form) are presently recognized. In Heteralepas cantelli sp. nov.: A, left-frontal aspect of capitulum, enlarged to show details of apertural the process we encountered some problem- atic forms, and these are briefly discussed below before moving on to those that are strictly relevant to the new species. Heteralepas quadrata_ (Aurivillius, 1892). This shallow-water species [includ- ing 1) A. percnonicola as a junior synonym (Hiro 1937), 2) the forms attributed to the species by Rosell (1972), and 3) a little- known form from the Eastern Pacific (Zullo 1991] sat uncomfortably in Heteralepas un- 390 Wi A WW Wy | \\y A Fig. 3. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1mm Heteralepas cantelli sp. nov., mouth parts: A, second maxilla; B, mandible (al’-a3’, enlarged un- dersides of teeth al-a3); C, first maxilla. (A—C same scale). til Foster (1979) transferred it to Paralepas, a decision accepted by Zevina (1982). However, the species also sits uncomfort- ably in Paralepas because in some ways it is morphologically intermediate between the two genera. The characters largely in- volve the cirri, their setation being neither strictly lasiopod nor acanthopod, the rela- tively low number of articles of their rami, and the somewhat reduced inner rami of cirri V & VI, as well as the somewhat in- termediate armature of the mandible and first maxilla. This suggests that proposal of a new genus is in order, and such a study would be of interest to evolutionary biolo- gists as well as cirripedologists in light of the inferred relative primitiveness of the Heteralepadidae (Foster 1979), a view re- cently corroborated genetically and mor- phologically (Harris et al. 2000; Pérez-Lo- sada et al. 2004). While there are a number of samples from the Eastern Pacific attri- buted to this species in the Benthic Inver- tebrate Collection at Scripps Institution of Oceanography, an appropriate review of the situation would also require studying ma- terials from the Western Pacific. However, such a study is beyond the scope of the present paper. ?Heteralepas malaysiana (Annandale, 1905). From a telegraph cable at approxi- mately 54 m depth in the Gaspar Straits. Annandale (1909:84) accepted Pilsbry’s (1907b) revision of Alepas and transferred Alepas xenophorae Annandale, 1906 to Heteralepas (Paralepas) and described Heteralepas (Heteralepas) nicobarica sp. nov. In the same paper, in a list of species VOLUME 117, NUMBER 3 Amm (ww““7~ry, i en ji aie ) 7 391 Fig. 4. Heteralepas cantelli sp. nov., thoracic appendages: A, penis; B, enlargement of distal portion of penis; C, intermediate segments of outer ramus of cirrus VI; D, posterior of thorax supporting right caudal appendage and pedicel of cirrus VI with proximal portions of inner and outer rami in outline (narrow and wide respectively, boundaries of articles omitted). contained in the Indian Museum, Annan- dale (1909:130) included Heteralepas ma- layana (sic) under the subgenus Heterale- pas. This was presumably because Annan- dale (1905:81) had clearly stated that the posterior (= inner) ramus of cirrus V was ““... reduced to a mere thread, less than one-third as long as the anterior ramus” and that cirrus VI was “... in much the same condition”. However, subsequently, and without a word of explanation, he (An- nandale 1916:298) transferred Heteralepas malaysiana to the subgenus Paralepas. While Newman (1960) retained malaysiana in Heteralepas s.s., Zevina (1982) followed Annandale by returning it to Paralepas. This is puzzling, considering the habitat as well as the characteristics of cirrus V & VI given in the original description. In light of these considerations, and the fact that the ornamentation of the capitulum appears more similar to that of Heteralepas rex (Pilsbry, 1907a) from Hawaii and ZH. uti- nomit Newman, 1960 from Tasmania than it does to any species of Paralepas, we have tentatively returned the species to Het- eralepas. ?Heteralepas ovalis (Hoek, 1907): This species is represented by a single specimen taken along with Paralepas morula from an PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 392 (ZO6L [PANID) ,.sJoyooOID sop juoj10d soun-onbyjonb juop sarpuoie sasnounryo suonejnurss op, , “uOISSNOSIq 99S = (VI1UOdHl “FY 104) soinjons podeys-asea jeioads 0} siajoy = ‘(100Z) 8unoX Aq snowlAuouds posopisuod = .. sox ON sox sox Sox Bese uoneunoed a[qipuryy sued yinoyy el TI-OI ¢ v1 CI CI-v osepuodde [epneD 09 68 é 76 é 89-LE snuiv1 19]nQ CK TC-61 EG 97 97 LT-€1 snuiet 1ouUy seponty TA SnD 09 TO-VL é €6 é 69-9€ snuivt 19]nO SC 1Z-61 LT EC 6 6C-EI snuiet JouUy sopnTy A SMUD ON SOOKE SOK & Sox ON Jejnqn ON Sox ON j i, j Polwoiewiep UISIVUI IaMOT APYsits A|snonordsuo, ON Anysis é é ayepnusia, Anied Sox Sox ON LON ON apis WoO, Sule] y % % % A 6% % O/V oinyiedy ON Sox SOX SOX Sox SUIAIe A pouexsiy) uIsIeUL jeulie) ON Sox Sox & Sox ON yuosoid So9pD1eqn} SNOUTTIYD)». +. our OT €1-9'0 ¢0 Lv'0-ST 0 L0 ($0) 8 I-@0 d/o € SEE TE TL-1T'E ve (8 €) 9 TI-S0 (Wid) YsueT Ci € 1-60 660 €I-L'0 60 (60) 6 I-S'0 (WO) WIPIM gjounped € CUT 91 SI-CI Li (07) 9'€-6'0 (wo) ysueT G OCCT SC LIC CT (SI) €7-9'0 (WS) WPI unypnyide) opis [eulIes uonount Jejnounped-o] Jopim ApYyst]S JOpIM —sJOPIAA = Jopim ApysipsS — Joprm ApYsITS Surkre A -njides 0} unyniides Jo wpIAA gjounpad 2 un] ROM esl) Ieo[) ROM DLN SurAre A -nqides Us9MJog UOPIIBPWUOG “AOU ‘ds 1)]AIUDI ‘H LLOJSAYUD] ‘H 12]2q. ‘H *k ISUILOAJAU ‘H 4 DWUOISOADIUL ‘H poiuodpr ‘H (Q) wintnitdes Jo yYSrey 0} (y) oinjode Jo WYBIEy Jo ONeY = O/V (dq) BJouNped jo yISUZ] 01 (D) UN[NIIdeo Jo yISUZ] Jo OnRY = d/Q uo paseq “Aou ‘ds yjajuvo “FY pue Majsayun] “YY ‘1J2q “YY ‘sisuasoajaw “py ‘pwojsosonu “FY “‘voUodpl spdajvsajayy Suowe uostreduios Jesisojoydiop~y— Z 29". VOLUME 117, NUMBER 3 echinoid spine in Malaysian waters. Hiro (1936:223) noted that nothing is known of the internal parts, but from the original fig- ures it is evident that the capitulum to ap- erture ratio is approximately 3:1. This is suggestive of Paralepas, but for lack of more conclusive evidence we have left it in Heteralepas. Heteralepas cygnus Pilsbry, 1907b: The original description was based on a speci- men acquired from the ‘“Ward’s Natural Science Establishment, Monterey, Califor- nia’, and hence the specimen was presum- ably from California, but it has not been recorded from this region since. Further- more, Annandale (1909) indicated that there is a specimen in the Edinburgh Mu- seum, questionably from the West Indies. The description may be adequate to distin- guish it from similar albeit relatively undis- tinguished forms, but what ocean it came from remains uncertain. Heteralepas cornuta (Darwin, 1852): A species usually having more-or-less distinc- tive carinal protuberances on its capitulum, first reported from the Caribbean (presum- ably from 90 m or so). It has since turned up in the Gulf of Mexico (Gittings et al. 1986), off Madeira and other West African islands (cf. Haroun et al. 2003), and along the coast of Northwest Africa. Furthermore it has been found in the Indian Ocean, the Philippines and the Southeast Pacific, off Chile (4315 m!) (cf. Young 2001 for re- view). Young (2001) commented not only on its wide geographical range and the ex- traordinary depth of the Chilean record compared to other populations attributed to the species, but on differences in cirral se- tation of the Chilean form compared to the population he has studied from the eastern Atlantic. Thus H. cornuta may represent a number of similar species. In any event, like the previous species, it is sufficiently distinct from the new form to no longer concern us here. Heteralepas microstoma (Gruvel, 1901): Known from off Madeira, the Azores and Meteor Seamount immediately to the south. 393 While known to range from between 269— 623 m, it is most commonly found around 300 m (Young 2001). Zevina & Kolbasov (2000) illustrated and compared it to anoth- er recently described species, H. meteoren- sis Carriol, 1998, as well as to their new species, H. alboplaculus Zevina & Kolba- sov, 2000, which was also from Meteor Seamount. Having such similar forms sym- patric on Meteor Seamount, and then large- ly from the same depth, is troubling. Young (2001) synonymized H. meteorensis with H. microstoma, but he was apparently un- aware of the work of Zevina & Kolbasov (2000) who claimed that all three species can be distinguished from each other by mi- nute cuticular structures revealed by SEM. However, their photographs are not clear in this regard. As can be seen from our Table 2, there appears to be little in the way of macro-morphological differences among them, although the peduncle of H. meteo- rensis seems to be relatively longer and the aperture does not appear as tubular as in H. microstoma. On the other hand, Heterale- pas alboplaculus is described as having the capitulum and to some extent the peduncle covered by well-spaced tubercles contain- ing calcareous structures. Such calcareous structures are unprecedented in the family and could be the work of a pathogen. We hope that workers in the Atlantic will clar- ify this situation in the near future. In the meantime, while the specific status of H. meteorensis and H. alboplaculus is uncer- tain (Table 1), we have included the former as well as H. microstoma in Table 2 for comparative purposes. When it comes to determining the affin- ities of the new species, the logical place to begin is in the Atlantic. Of the 8 previously known species of Heteralepas noted in the introduction, 5 occur in the eastern Atlantic. These include H. cornuta, alboplaculus and segonzaci, and taking their capitular fea- tures at face value, they are distinct from the new form and therefore no longer con- cern us here. This leaves H. microstoma and meteorensis, which are very similar if 394 not synonymous, but as noted above both have been characterized in Table 2 for com- parative purposes. As for the western Atlantic species, H. cornuta, which ranges as far north as the Carolinas, was noted above as being dis- tinct from the new form. This leaves H. lur- idas, belli and lankesteri. The first, from 300-700 m of depth in the Caribbean, is known to range between 2 and 9.5 mm in height and the specimen illustrated in the original description is less than 6 mm high, so it is a small species. Its capitulum, with a somewhat tubular or flaring apertural re- gion, is otherwise undistinguished, and its cirral and caudal appendage counts are low- er than in the new species. So, assuming H. luridas is not based on juveniles, it too need no longer concern us. This leaves H. belli and lankesteri, and since in outward ap- pearance they are similar to the new form, they have been included in Table 2. As we shall see, so far none of the species includ- ed in Table 2 agree well with the new spe- cies in numerous detail; but what about spe- cies from the Indo-Pacific? Of the Indo-West Pacific species, H. ja- ponica and similar species such as H.. fulva from the Southeast Pacific are rather close to the new form. The former has been re- ported from between 18 and 1020 m depth from Japan to Singapore, Australia and New Zealand, the Nicobars in the Andaman Sea and Réunion Is. (Foster & Buckeridge 1995). Therefore, while not as wide—rang- ing as H. quadrata, it is wide-ranging com- pared to most species of the genus. Part of this range is due to synonymies, and that of Nilsson-Cantell’s (1927, 1938) for H. indica (Gruvel, 1901) has long been ac- cepted. This extended the range of the spe- cies to Singapore and into the Indian Ocean where it was reported from Nicobar Is. on floating wood. Furthermore, Foster (1979), in his report on New Zealand cirripeds, syn- onymized H. dubia Broch, 1922 from 55— 72 m in Disaster Bay, Australia, with H. japonica. However, Zevina (1982), without explanation, continued to recognize H. du- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bia as a distinct species, and subsequent au- thors have followed suit. Considerable variability in characters might be expected in such a wide-ranging species and for present purposes we are ac- cepting the opinion of these authors. How- ever, considering such variation in cirripeds as geographical rather than indicative of ge- netically distinct populations has generally proven wrong (Newman 1993). Thus cau- tion seems in order because the reported variations in the mandible of presumed H. Japonica from different populations, appear to go beyond the range of variability found within a species. Although Aurivillius (1894) did not illustrate the mandible of H. Japonica, his written description agrees with Nilsson-Cantell’s (1921:247, fig. 43b) and Pilsbry’s (1911:71, fig. 4A) illustra- tions, and also with that of Gruvel (1902: 284, Pl. 24, fig. 24) for H. indica. Thus the teeth of the mandible appear to be without pectinations, but on close inspection of Nilsson-Cantell’s illustration there might have been low pectinations of the lower margins of teeth 1—3, especially 2 and 3. However, since there is no such suggestion in the other illustrations, the evidence fa- vors the mandible being simple. The situation at the southern end of the range for H. japonica looks quite different with regard to the mandible. Foster (1979) synonymized H. dubia Broch from Austra- lia, and the population he was studying in New Zealand, with H. japonica. While Broch (1922:288, fig. 37B) gave no indi- cation of pectinations on the first tooth, he clearly illustrated them on the upper sides of teeth 2—4 as well as the lower sides of 2 and 3. Foster (1979:16, fig. 3J) illustrated the same for the upper sides, but limited pectinations on the underside to tooth 1. So, the populations attributed to this species from north and south of the equator appear to differ in the characteristics of the man- dible, and the new species, with its incon- spicuous pectinations on the lower sides of teeth 1-3 (Fig. 3, al-a3), differs from both of them. VOLUME 117, NUMBER 3 In view of the foregoing considerations we include H. dubia in Table 1 as a ques- tionable species rather than a synonym of H. japonica. Nonetheless, H. japonica still includes sufficient variability to make it an ideal Indo-Pacific representative similar to the new form from the Atlantic, and there- fore it is included in Table 2 for compara- tive purposes. Summary and Conclusions The essentially naked heteralepadids pre- sent a difficult problem to systematists since, being unarmored, they lack a number of distinct features customarily utilized in separating genera and species (Zullo & Newman 1964). Aside from the work of Nilsson-Cantell (1921, 1927), and to some extent, Young (2001), no studies have eval- uated the usefulness of morphological char- acters in distinguishing Heteralepas spe- cies. Thus it is difficult to establish a new species with a high degree of certainty. But, in spite of the latitude allowed by synony- my, the present form could not be assigned to any known species. As can be observed in Table 2, the At- lantic species most similar to the new spe- cies (H. microstoma, meteorensis, belli and lankesteri) are readily distinguished from it as well as from H. japonica from the Indo- West Pacific, by several characters. How- ever, the new species, H. cantelli, cannot be distinguished from H. japonica by the char- acters presented in the table. This is due in part to the variability attributed to H. ja- ponica, but there are notable differences be- tween these two species, not included in the table, that distinguish them. These include 1) the lack of any indication of a carinal thickening, crest, or protuberances along the carinal margin (but sometimes also found lacking in individuals of H. japoni- ca), 2) a marked crenation of the apertural margin largely restricted to the upper third rather than along its entire margin, and 3) a slightly depressed area around the entire apertural region, setting it off from the gen- 395 eral surface of the capitulum. The last two differences are sufficient not only to distin- guish the new form from H. japonica, but from all known heteralepadids. Acknowledgments We thank Paulo S. Young, Museu Na- cional/UFRJ, Rio de Janeiro, Brazil, who died tragically before the publication of this paper, for advice on the Atlantic species during preparation of the manuscript, and Pal B. Mortensen, Bedford Institute of Oceanography, Dartmouth, Canada and Vladimir E. Kostylev, Natural Resources Canada, Dartmouth, Canada, for helping with the translation of publications in Ger- man and Russian, respectively. While we would also like to thank two judicious ref- erees (John S. Buckeridge, EOS, Auckland University of Technology, as well as Paulo S. Young) for reviewing the manuscript, we are solely responsible for any errors that re- main. Literature Cited Annandale, N. 1905. Malaysian barnacles in the Indian Museum, with a list of Indian Pedunculata.— Memoirs of the Asiatic Society of Bengal 1(5): 73-84. . 1906. Preliminary report on the Indian stalked barnacles—Annals and Magazine of Natural History 17(7):389—400. . 1909. An account of the Indian Cirripedia Pe- dunculata, part 1—Family Lepadidae (s.s.). Memoirs of the Indian Museum 2:61—137. . 1916. Barnacles from deep-sea telegraph ca- bles in the Malay Archipelago.—Journal of the Straits Branch of the Royal Asiatic Society 74: 281-302 + pls. IV—VI. Aurivillius, C. W. S. 1892. Neue Cirripeden aus dem Atlantischen, Indischen und Stillen Ocean — Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, Stockholm 3:123—134. 1894. Studien iiber Cirripeden. Kongliga Svenska VetenskapsAkademiens Handlingar,— Uppsala 26(7):5—107 + 9 pls. 1-9. Broch, H. 1922. Studies on Pacific cirripeds. Pp. 215— 358 in Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-1916. X. Videnskabelige meddelelser fra Dansk Naturhistorisk Forening i Kébenhayn 73. Carriol, R. P. 1998. A new pedunculate cirriped (Thoracica, Heteralepas) 396 from the northeast Atlantic Ocean.—Zoosyste- ma 20(3):505—509. Darwin, C. R. 1852. A monograph on the sub-class Cirripedia, with figures of all the species. The Lepadidae; or, pedunculated cirripedes. Pp. 1— 400 + pls. 1-10. Ray Society, London (1851). Foster, B. A. 1979. The Marine Fauna of New Zealand; Barnacles (Cirripedia: Thoracica).—New Zea- land Oceanographic Institute Memoir 69:1—159 (1978). , & J. S. Buckeridge. 1995. Barnacles (Cirri- pedia, Thoracica) of seas off Réunion Island and the East Indies.—Bulletin du Muséum na- tional d’Histoire naturelle, Paris, 4° séries, 16(2—4):345-382. Gittings, S. R., G. D. Dennis, & H. W. Harry. 1986. Annotated guide to the barnacles of the northern Gulf of Mexico. Sea Grant College Program, Texas A & M University, College Station, 36 PP. Gruvel, A. 1900. On a new species of the genus Alepas (A. lankesteri), in the collection of the British Museum.—The Annals and Magazine of Natu- ral History. Ser. VI, 6:195—199 + pl. VII. . 1901. Diagnoses de quelques espéces nouvel- les de Cirrhipédes.—Bulletin, Muséum national d’ Histoire naturelle, Paris 7:256—263. . 1902. Sur quelques Lépadides nouveaux de la collection du British Museum.—Transactions of the Linnean Society, London, Second Series, 8: 277-294 + Pl. 24. Haroun, R., R. H. Pérez, & P. D. Santana. 2003. Cir- ripedia. Pp. 67—68 in L. M. Abad, J. L. M. Es- quivel, M. J. G. Sanahuja and I. I. Zamorna, eds., Lista de especies Marinas de Canarias (Al- gas, Hongos, Plantas y Animales), Consejeria de Politica Territorial y Medio Ambiente del Gobierno de Canarias, Tenerife, 200 pp. Harris, D. J., L. S. Maxson, L. E Braithwaite, & K. A. Crandall. 2000. Phylogeny of the thoracican barnacles based on 18S rDNA sequences.— Journal of Crustacean Biology 20(2):393-398. Hiro, E 1936. Descriptions of three new species of Cirripedia from Japan.—Bulletin of the Biogeo- graphical Society of Japan 6(23):221—230. . 1937. Studies on Cirripedian fauna of Japan. II. Cirripeds found in the vicinity of the Seto Marine Biological Laboratory.—Memoirs of the College of Science, Kyoto Imperial Univer- sity, Series B 12(3):385—478. Hoek, P. C. C. 1907. The cirripedia of the Siboga Ex- pidition,—Pedunculata. Siboga-Expeditie 31a: 1-127. Newman, W. A. 1960. Five pedunculate cirripeds from the Western Pacific, including two new forms.—Crustaceana 1(2):100-116. . 1990. Carl August Nilsson-Cantell, 28 De- cember 1893-14 January 1987.—Crustaceana 59(3):289-294. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1993. Darwin and cirripedology. Pp. 349—434 in J. Truesdale, ed., The history of carcinolo- gy.—Crustacean Issues 8. Balkema, Rotterdam. Nilsson-Cantell, C. A. 1921. Cirripeden Studien. Zur Kenntnis der Biologie, Anatomie und Syste- matik dieser Gruppe.—Zoologiska Bidrag, Uppsala 7:75—390. . 1927. Some barnacles in the British Museum (Nat. Hist.).—Proceedings of the Zoological Society of London 1927(3):743-790, figs. 1-19, pl. 1. . 1938. Cirripedes from the Indian Ocean in the collection of the Indian Museum, Calcutta— Memoirs of the Indian Museum 13(1):1—81 + pls. 1-3. Perez-Losada, M. J., J. T. Hgeg, & K. A. Crandall. 2004. Unraveling the evolutionary radiation of the thoracican barnacles using molecular and morphological evidence: a comparison of sey- eral divergence time estimation approaches.— Systematic Biology 53(2):244—264. Pilsbry, H. A. 1907a. Hawaiian Cirripedia—Bulletin of the Bureau of Fisheries 26:181—190 + pls. IV & V (1906). . 1907b. The barnacles (Cirripedia) contained in the collections of the U.S. National Muse- um.—Bulletin of the United States National Museum 60:1—122 + pls. 1-11. . 1911. Barnacles of Japan and Bering Sea.— Bulletin of the Bureau of Commercial Fisheries 29:61-84 + pls. VILI-XVII (1909). Ren, X. 1983. Five new species of suborder Lepado- morpha (Cirripedia Thoracica) from Chinese waters.—Oceanologia et Limnologia Sinica 14(1):74-87. Rosell, N. C. 1972. Some barnacles (Cirripedia Thor- acica) of Puerto Galera found in the vicinity of the U.P. Marine Biological Laboratory.—Na- tional and Applied Science Bulletin 24(4):104— 283. Young, P. S. 1999. The Cirripedia (Crustacea) collected by the “Fisheries Steamer Meteor” in the East- ern Atlantic—Arquivos do Museu Nacional, Rio de Janeiro 58:1—54 (1998). . 2001. Deep-sea Cirripedia Thoracica (Crus- tacea) from the northeast Atlantic collected by French expeditions.—Zoosystema 23(4):705— 756. Zevina, G. B. 1975. Cirripedia Thoracica of the Amer- ican Mediterranean.—Trudy Instituta Okeanol- ogii 100:233—258 (in Russian). . 1982. Barnacles of the suborder Lepadomor- pha of the world ocean. II. Pp. 1-222 in Fauna U.S.S.R., Zoological Institute, Russian Acade- my of Science, Leningrad, 133 (in Russian). , & G. A. Kolbasov. 2000. Barnacles of the genus Heteralepas (Thecostraca, Cirripedia, Thoracica) from the Canary Islands and the VOLUME 117, NUMBER 3 397 Azores. Description of mantle ultrastructure.— jacent regions in the tropical eastern Pacific. Pp. Zoologicheskii Zhurnal 79(11):1275-1283 (in 173-192 in M. J. James, ed., Galapagos marine Russian). invertebrates. Taxonomy, biogeography, and , & M. Y. Schreider. 1992. New species of Cir- evolution in Darwin’s Islands. Plenum Publish- ripedia (Crustacea) from the Indian Ocean.—Zool- ing Company, New York, New York. ogicheskii Zhurnal 71(10):39—46 Gn Russian). , & W. A. Newman. 1964. Thoracic Cirripedia Zullo, V. A. 1991. Zoogeography of the shallow-water from a southeast Pacific guyot—Pacific Sci- cirriped fauna of the Galapagos Islands and ad- ence 18(4):355-372. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):398—407. 2004. Two new species of seven-spined Bathyconchoecia from the North Atlantic and Indian oceans (Crustacea: Ostracoda: Halocypridae) Louis S. Kornicker and J. A. Rudjakov (LSK) Department of Zoology—IZ, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012, U.S.A., e-mail: kornicker.louis@nmnh.si.edu; (JAR) Museum of Comparative Zoology, Harvard University, 26 Oxford St., Cambridge, Massachusetts 02138-2902, U.S.A, e-mail: rudyakov @fas.harvard.edu Abstract.—A new species of halocyprid ostracode Bathyconchoecia omega from abyssal depths of the North Atlantic Ocean, off Newfoundland, Canada, is described and illustrated, and a new species Bathyconchoecia georgei is proposed for a specimen from the Indian Ocean previously referred to Bathy- conchoecia deeveyae Kornicker, 1969. The R/V Chain, operated by the Woods Hole Oceanographic Institution, collected in 1972 at a depth of 4400 m in the North Atlantic Ocean, off Newfoundland, Canada, a bottom sample containing a single A-1 male of Bathyconchoecia omega, new spe- cies. The A-1 male from off Newfoundland is considerably larger than previously de- scribed seven-spined species of the genus, and is the northernmost occurrence of the group. Additional ostracodes in the sample are mostly bottom-living Podocopida, Cla- docopida and Myodocopida, which sug- gests a bottom or near-bottom habitat for B. omega. However, a specimen of pelagic species of Conchoecia in the sample sug- gests that it contains some shallow water contaminants. Only three species of Bathyconchecia having seven spines on the carapace (four on right valve, three on left), have been de- scribed previously: B. deeveyae Kornicker, 1969, B. septemspinosa Angel, 1970, and B. longispinata Ellis, 1987. One of the specimens previously referred to B. deev- eyde is proposed as a new species herein. Thus, the number of 7-spined species of Bathyconchoecia is now five. Their distri- bution is shown in Fig. 1. Correction.—Kornicker (1981:1237) re- ported that the slide containing the append- ages of the holotype of B. deeveyae (USNM 123335) had been lost. It has been recov- ered. Bathyconchoecia omega, new species Figures 2—6 Holotype.—Unique specimen, A-1 male on slide and in alcohol, MCZ Harvard Uni- versity, MCZ50432. Type locality.—R/V Chain 106, 30 Aug 1972, Station 334, North Atlantic Ocean, off Newfoundland, Canada, 40°42.6'’N— 40°44'N, 46°13.8’W—46°14.6'W, epibenthic sled, depth 4400 m. Material.—Holotype. Description of A-I male (Figs. 2—6).— Carapace with linear dorsal margin except for slight bulge near middle just posterior to base of dorsal spine. Posterodorsal corner of each valve with gland on very slight bulge. Posterodorsal corner evenly rounded except for long spine on right valve; spine parallel with length of valve, but at slight upward angle (very tip of spine of specimen broken off; soft matter projects from broken tip). Base of spine projects slightly medial VOLUME 117, NUMBER 3 Fig.1. to slightly overlap posterior edge of left valve (Fig. 2B). Rostrum of each valve with anterior spine at slight angle to each other (Fig. 2B). Spine at midlength of dorsal mar- gin of each valve at slight outward and up- ward angle (Fig. 2A, B). Spine near ventral margin of each valve at about % length of valve at slight downward and outward an- gle. Anterior spines on rostra and posterior spine on right valve with surface ridges par- allel to lengths of valves; a few of the ridg- es of the rostral spines bear short stout spines. Other long spines with minute sur- face spines. Carapaces completely covered by distinct punctae and slightly curved ver- tical frills (not all shown in Fig. 2A). Frills generally on each side of 2 or 3 rows of punctae (Fig. 2C). Indistinct reticulations and ridges on anteroventral surface of valve ventral to incisure (Fig. 2A). Pigmentation: No black pigment spots on either carapace or body. Central adductor muscle attachments (Fig 2A): Indistinct, near center of valve 399 B. deeveyae B. georgei B. longispinata B. omega B. septemspinosa Distribution of species of seven-spined Bathyconchoecia. and consisting of 2 individual scars; stria- tions of muscle ends indistinctly visible from outside view of valve; scars not cov- ered by punctae. Carapace size (mm): Length including spines 3.79, length excluding spines 2.92, height excluding spines 1.68, width without spines 1.52. First antenna (Fig. 2E): Shaft short with indistinct segmentation. Brush-like struc- ture with about 315 filaments in about 9 rows, each with about 35 filaments. Dorsal bristle on segment following brush-like structure stout, spinous, about % length of brush filaments. Terminal segment with 4 bristles: 1 long stout bristle reaching well past brush filaments and with widely scat- tered marginal spines (not shown); 3 shorter than brush filaments. Limb with densely packed amber-colored cells. Second antenna (Fig. 3A—D): Protopod bare. Endopod: Ist article with 2 spinous dorsal bristles (1 long, 1 short) and few in- distinct medial spines near ventral margin; 400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Complete carapace from right side, length without spines 2.92 mm; B, Complete carapace, ventral view; C, Left valve, detail of ornamentation on outer surface; D, Posterodorsal corner of complete specimen (spine on right valve, glandular opening on left valve); E, Right Ist antenna, lateral view. VOLUME 117, NUMBER 3 401 Sen Rene Fig. 3. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Left 2nd antenna, medial view; B, Endopod right 2nd antenna, lateral view; C, Proximal part exopod left 2nd antenna, medial view; D, Distal part exopod right 2nd antenna, lateral view; E, Proximal part left Sth limb, lateral view; EF Right 5th limb drawn on body, lateral view. 402 2nd article with 1 minute bristle medial to 3rd article and 2 stout terminal bristles with few indistinct marginal spines (inner bristle stouter, both about same length as exopod bristles); 3rd article with 3 bristles: middle bristle longer and stouter than others, more than 1/2 length of bristles of 2nd article, with few marginal spines; outer bristle about % length of middle bristle, with many marginal spines; inner bristle similar in length to outer bristle, with marginal spines; base of 3rd article lateral to distal end of 2nd article; endopods of left and right limbs similar. Exopod: Ist article with short ven- tral spines and small medial terminal bris- tle; articles 2 to 8 with long natatory bristle; Oth article with 4 bristles (2 short lateral; 1 ventral of medium length and with short marginal spines; 1 long dorsal, with nata- tory hairs). Mandible (Fig. 4): Coxa (Fig. 4B-—E): Pars incisivus with 5 ventral teeth and slen- der distal tooth at ventral tip of triangular posterior section; anterior edge serrate (Fig. 4C). Proximal list with 10 teeth in 2 layers (Fig. 4D); distal list with 19 teeth in 3 lay- ers (Fig. 4E). Spined posterior part with 6 lobes with numerous spines and 7th lobe with short stout spinous bristle and minute spines along distal posterior edge of lobe near bristle (Fig. 4C). Anterior margin of coxa evenly rounded, without triangular process. Basis (Fig. 4A, E G): 2 long plu- mose bristles present on or near dorsal mar- gin and | long spinous medial bristle near midwidth some distance from dorsal margin (Fig. 4F); lateral surface with 3 long bare distal bristles and long spines (Fig. 4F); posterior margin spinous and with 2 short distal bristles (proximal bristle sclerotized and with ventral spines; distal bristle tube- formed) (Fig. 4G); anterior margin with long bare distal bristle (Fig. 4G); ventral margin with 5 short teeth with minute sec- ondary teeth (Fig. 4G); | short tooth with minute secondary teeth on posterior margin proximal to posterior ventral tooth. Poster- odorsal corner of basis with oval sclerite (Fig. 4A, B). Endopod (Fig. 4E H): Article PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1 with dorsal, ventral, lateral, and medial slender spines and 4 bristles (1 long, ter- minal, dorsal, spinous; 1 long, distal, ven- tral, bare; 2 medium length, medial with ba- ses close to ventral bristle, bare). Article 2 with 5 bristles (2 long and 1 shorter, ter- minal, dorsal, spinous; 2 medium length, distal ventral, bare) and few distal spines on ventral margin and medial and lateral sur- faces near ventral margin. Article 3 with long spinous terminal claw and 6 bristles (2 long, spinous, terminal (shorter of these with base on medial side), and 4 short, ven- tral, bare (1 of these with lateral base)) and medial spines. Maxilla (Fig. SA—C): Endite of precoxa with 2 tube-formed bristles, 3 claws, and 2 long spinous bristles. Coxa: dorsal margin with long stout dorsal bristle (Fig. 5B); proximal endite with 3 tube-formed bristles and total of 4 claws and claw-like bristles; distal endite with 2 tube-formed bristles and 4 claws. Basis with 2 long stout plumose bristles near dorsal margin, and short bare ventral bristle. Endopod: article 1 spinous with 4 dorsal bristles (3 proximal, | distal); medial surface with 4 distal bristles (3 long, 1 short); article 2 with 2 stout claws of un- equal length and 4 slender bristles. Fifth limb (Fig. 3E, F): Epipod with 3 groups of 4 stout plumose bristles; dorsal group with additional small 5th bristle (Fig. 3F). Precoxa with 3 ventral bristles (Fig. 3E). Coxa with 11 or 12 ventral bristles (not all shown). Basis with 6 bristles plus long terminal dorsal exopod bristle with minute widely separated marginal spines (not all shown). Endopod: article 1 with dorsal and medial spines and 3 bristles (not all shown). Article 2 with dorsal and medial spines and 4 bristles (3 near ventral margin and | lon- ger dorsal). Article 3 with 2 long terminal slender claws and 1 long ringed, terminal, slender ventral bristle. A muscle terminates at base of exopod bristle. Sixth limb (Fig. 6A): Epipod with 3 groups of 5, 5, and 6 (dorsal) long plumose bristles; dorsal group with additional short 7th bristle (Fig. 6A). Coxa with 1 spinous, VOLUME 117, NUMBER 3 403 Fig. 4. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Left mandible, junction of coxa and basis, lateral view. B—H, Right mandible, lateral views: B, Proximal part of coxa; C, Distal end of coxa; D, Detail of proximal tooth of coxa (detail from C); E, Detail of distal tooth of coxa (detail from C): E Basis and endopod; G, Distal end of basis; H, Endopod. 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Right maxilla, medial view (arrow indicates tube-formed bristles); B, Left maxilla, lateral view; C, Right maxilla, oblique dorsal view (not all bristles shown); D, Anterior of body from right side showing upper and lower lips (esophagus dashed); E, F Lower lip from left side, anterior of body to left; G, Upper lip, dorsal view. VOLUME 117, NUMBER 3 ~ LZ 4222 = LEE a M1 ANS \ A eV Ul] \ CRAP SesSone x 405 Fig. 6. Bathyconchoecia omega holotype, MCZ 50432, A-1 male: A, Right 6th limb, medial view; B, 7th limb; C, Left lamella of furca and unpaired bristle, lateral view; D, Right lamella of furca, lateral view; E, Posterior view of ventral end of body showing unpaired bristle and furca; E Posterior view of body showing copulatory organ on left side; G, Posterior of body from right side showing furca and copulatory organ; H, Copulatory organ from left side, anterior to upper left. 406 ventral, terminal bristle. Basis with spines, 4 spinous bristles and 1 long, terminal, dor- sal, exopod bristle with widely scattered minute spines (basis may consist of medial and lateral parts). Endopod: article 1 with 4 bristles; article 2 with 2 bristles; article 3 with 3 long terminal bristles (dorsal 2 claw- like). A muscle terminates at base of exo- pod bristle. Seventh limb (Fig. 6B): Broad thumb- like process with 2 long unequal bare bris- tles. Furea (Fig. 6C—G): Each lamella with 7 claws with teeth along posterior margins; | unpaired spinous bristle following claws on lamellae (Fig. 6E). Bellonci organ: Not developed. Lips (Fig. 5D—G): Upper lip with spinous posterior edge (Fig. 5D, G). Lower lip spi- nous (Fig. 5E, F). Copulatory organ (Fig. 6F—H): Organ with 2 separate branches on left side of body. Broad anterior branch with minute terminal teeth; narrow posterior branch with small tapered tip. Comparisons.—The length of the unique A-1 male from off Newfoundland (exclud- ing spines) is 2.92 mm, whereas A-1 instars of B. deeveyae and B. septemspinosa are shorter than 1.8 mm (Kornicker and Angel, 1975: table 1; Kornicker, 1981:1240). A length of 0.66 mm was reported for an A- 4 instar of B. deeveyae by Kornicker (1991: 30). The adult male of B. longispinata has a range of lengths of 1.95—2.11 mm (Ellis, 1987: Table Il), much shorter than the 2.92 mm length of the A-1 male referred herein to B. omega. The 2 mid-dorsal spines on the carapace of the later specimen are short- er than those of B. longispinata. Also, the fossae and frills of B. omega are on all parts of the valve, whereas, they cover only cer- tain areas on B. longispinata. The length of the adult male of B. georgei, new species, is 1.28 mm, much smaller than the length (2.92 mm) of the A-1 male of B. omega. The carapace of the former species is with- out the frills present on the carapace of B. omega. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The 2nd endopod articles of both man- dibles of the A-1 instar of B. omega bear 5 bristles compared to 4 on the A-1 mandi- bles of B. septemspinosa and B. deeveyae and the adult male mandible of B. longis- pinata, and 3 on the adult male mandible of B. georgei. Mandibles of a total of six A-1 and A-2 instars of B. septemspinosa examined by Kornicker and Angel (1975: Table 1) indicate that the number of bristles on the 2nd endopod article of the mandible of those instars do not vary from 4 bristles and, therefore, may be a reliable character to use to discriminate specimens of B. ome- ga, but reliability of the character in the lat- ter species is unknown. Bathyconchoecia georgei, new species Bathyconchoecia deeveyae Kornicker.— George, 1971: 141, figs. 1-9. Not Bathyconchoecia deeveyae Kornicker, 1969: 403, pl. 1, figs. 1-2. Etymology.—Species named in honor of Jacob George, National Institute of Ocean- ography, Cochin-18, India, who described the specimen upon which the new species is based. Holotype.—Unique specimen, adult male. Specimen is in a vial labeled 07.43, with serial number 0130, deposited in the archive room at the Indian Ocean Regional Centre, National Institute of Oceanography, Cochin — 14, India (there are no mounted slides). (Information about specimen sup- plied by Dr. Rosamma Stephen, Scientist, National Institute of Oceanography Region- al Center, Cochin, in correspondence with the junior author. Dr. Stephen did not ex- amine specimen in vial, but stated that she “could make out that there is a white spec- imen inside.’’) Type locality.—International Indian Ocean Expedition station Co. 62 (I. O. B. C.1969), in vertical haul from 200 to O m, off SW coast of India, 10°39'N, 75°22’E. Material.—None examined. Discussion of B. deeveyae Kornicker, 1969.—This species was described from an VOLUME 117, NUMBER 3 A-1 juvenile collected at a depth of 508— 523 m in a benthic trawl in the Peru-Chile Trench System, Pacific Ocean (Kornicker, 1969:403). A second specimen, an adult male, was collected in a vertical plankton haul from 200 to O m in the Indian Ocean off the SW coast of India (George, 1971: 141). A third specimen, an adult or A-1 fe- male, was collected at a depth of 520 m in an epibenthic sled from off Surinam, Atlan- tic Ocean (Kornicker, 1981:118). Ellis (1987:83) observed, “It is possible that these three specimens are not conspecific.” That observation prompted the present au- thors to reconsider the three specimens that had been referred to B. deeveyae, and led to our conclusion that the Indian Ocean specimen is not conspecific with the other two specimens of B. deeveyae from the At- lantic and Pacific Oceans. The Indian Ocean specimen was adequately described by George (1971: 141), so that only a brief diagnosis based on the adult male is pre- sented here. Diagnosis (adult male).—Carapace 1.28 mm long, excluding spines. Second endo- pod article of mandible with 3 bristles. Fur- ca with 8 claws on each lamella. Comparisons.—The carapace of the new species, B. georgei is much smaller than equivalent stages of B. septemspinosa, B. deeveyae, and B. longispinata (because only the adult male of B. georgei is known, the relative sizes of its instars is an extrap- olation). The 2nd endopod article of the mandible of the adult male B. georgei bears 3 bristles compared to 5 on the adult male of B. longispinata and 4 on the A-1 instars of both B. septemspinosa and B. deeveyae. The adult male B. georgei bears 8 claws on each lamella compared to 7 on the adult male B. longispinata. Acknowledgments We thank Elizabeth Harrison-Nelson for preparing the illustrations and text for pub- 407 lication, Molly Ryan for producing the spe- cies distribution map (Fig. 1), and Megan Bluhm for inking the illustrations from pen- ciled Camera Lucida drawings by the first author. We are greatly indebted to Dr. Ro- samma Stephen, National Institute of Oceanography, Cochin, India, for providing information about the present location of the type specimen of B. georgei. The junior author would like to thank Dr. Gonzalo Gi- ribet and Mrs. Ardis B. Johnston for their help and encouragement. Literature Cited Angel, M. V. 1970. Bathyconchoecia subrufa n. sp. and B. septemspinosa n. sp., two new halocy- prids (Ostracoda, Myodocopida) from the trop- ical North Atlantic and the description of the larval development of B. subrufa. —Crusta- ceana 19:181—199. Deevey, G. B. 1968. Pelagic ostracods of the Sargasso Sea off Bermuda: Descriptions of species, sea- sonal and vertical distribution. —Peabody Mu- seum of Natural History (Yale University) 26: 1-125. Ellis, C. J. 1987. Bathyconchoecia longispinata n. sp., a new species of halocyprid Ostracod with sey- en carapace spines.—Crustaceana, 53:83—93. George, J. 1971. On the occurrence of Bathyconchoe- cia deeveyae Kornicker (Ostracoda, Halocypri- didae) in the Indian Ocean. —Crustaceana 21: 141-144. Kornicker, L. S. 1969. Bathyconchoecia deeveyae, a highly ornamented new species of Ostracoda (Halocyprididae) from the Peru-Chile Trench system.—Proceedings of the Biological Society of Washington 82:403—408. . 1981. Range extension and supplementary de- scription of Bathyconchoecia deeveyae (Ostra- coda: Halocyprididae).—Proceedings of the Bi- ological Society of Washington 94:1237—1243. . 1991. Myodocopid Ostracoda of Hydrother- mal Vents in the Eastern Pacific Ocean. Smith- sonian Contributions to Zoology, 516, 46 pages, 25 figures, 2 tables. , & M. V. Angel. 1975. Morphology and on- togeny of Bathyconchoecia septemspinosa An- gel, 1970 (Ostracoda: Halocyprididae). — Smithsonian Contributions to Zoology 195: 1— 21. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):408—422. 2004. The hermaphroditic sea anemone Anthopleura atodai n. sp. (Anthozoa: Actiniaria: Actiniidae) from Japan, with a redescription of A. hermaphroditica Kensuke Yanagi and Marymegan Daly (KY) Costal Branch of Natural History Museum and Institute, Chiba, 123 Yoshio, Katsuura, Chiba Pref., 299-5242 Japan, e-mail: yanagi@chiba-muse.or.jp; (MD) Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence KS 66045 U.S.A., Current address: Dept. of Evolution, Ecology & Organismal Biology, Ohio State University, Columbus, Ohio 43210 U.S.A. e-mail: daly.66@osu.edu Abstract.—A new species of internally brooding sea anemone, Anthopleura atodai, is described from the middle to northern Pacific coasts of Honshu, Japan. This species attaches to mussels or in rock crevices of the higher tidal zone. This is the second hermaphroditic species and fourth internally brooding species of Anthopleura to be reported; it is distinguished from other members of Anthopleura by a combination of the following features: brooding its young, synchronously hermaphroditic, S-shaped basitrichs in filaments, 40 to 68 ten- tacles, verrucae in the proximal part of the column larger than those in the distal part, cobalt-blue spot at the distal end of each siphonoglyph. Anthopleura hermaphroditica, the species that most closely resembles A. atodai, is rede- scribed to clearly differentiate it from A. atodai and to resolve questions about its taxonomy and identity. Anthopleura Duchassaing and Michelot- ti, 1861, one of the largest genera in the Actiniaria, includes about 50 species (Carl- gren 1949; Dunn 1974, 1978, 1982a; Fautin 2003). In Japanese waters, six species of Anthopleura are known: Anthopleura asia- tica Uchida & Muramatsu, 1958; A. fusco- viridis Carlgren, 1949; A. kurogane Uchida, 1938; A. mcmurrichi Wassilieff, 1908; A. pacifica Uchida, 1938; A. uchidai England, 1992. Additionally, Atoda (1954) reported the post-larval development of an uniden- tified species of Anthopleura, which broods its young in the colenteron. Although Atoda (1954) mentioned that the species could be distinguished from other species of Antho- pleura by its coloration, it has never named; we formally describe it here as a new spe- cies, A. atodai. Internal brooding is widely known in the Actiniaria: e.g. Actinia spp. (Chia & Ros- tron 1960; Rossi 1971; Black & Johnson 1979; Ayre 1983; Manuel 1988; Russo et al. 1994; Yanagi et al. 1996, 1999), Aulac- tinia sp. (Dunn et al. 1980), Cereus pedun- culatus (Rossi 1971), Cnidopus japonicus (T. Uchida 1934, T. Uchida & Iwata 1954), Epiactis spp. (Dunn 1975, Fautin & Chia 1986, Edmands 1995), and Bunodactis her- maphroditica (McMurrich 1904). Aside from A. atodai, three species of Anthopleu- ra are reported to brood internally: A. handi Dunn, 1978, from the Philippines, Hong Kong, and Malaysia (Dunn 1978, England 1987); A. aureoradiata (Stuckey, 1909a) from New Zealand (Stuckey 1909a, 1909b; Carlgren 1949, 1954; Parry 1951); and A. hermaphroditica (Carlgren, 1899) from Chile (Carlgren 1899, 1927, 1949, 1959). Anthopleura atodai most closely resem- bles A. hermaphroditica. Because the anat- omy and cnidom of A. hermaphroditica is incompletely known, and its taxonomic sta- tus is unclear, we redescribe it to clearly VOLUME 117, NUMBER 3 Asamushi Hachinohe Katsuura Tateyama Pacific Ocean Fig. 1. species. Stars indicate records of Anthopleura sp. giv- en by Atoda (1958); circles indicate sites visited in this study. Distribution of Anthopleura atodai, new distinguish A. hermaphroditica from A. ato- dai and to evaluate the proposed synonymy between A. hermaphroditica and A. handi. We find that A. hermaphroditica and A. ato- dai can be distinguished based on color, number of tentacles, cnidom, and geograph- ic range, and that A. hermaphroditica 1s dis- tinct from A. handi. Materials and Methods Specimens of Anthopleura atodai were collected from high intertidal rocky shore around Asamushi (40°54’N, 140°51’E), Ot- suchi (39°22’N, 141°58’E), Katsuura (35°07'’N, 140°16’ E), and Tateyama (34°58'N, 139°46’E) (Fig. 1). Anatomical observations were made on 17 specimens of A. atodai; histological sections were made from 11 specimens. Anatomical ob- servations were made on 10 preserved specimens of A. hermaphroditica; histolog- ical sections were made from 5 animals. For specimens of both A. atodai and A. her- maphroditica, histological sections 6—8 4m thick were stained with hematoxylin and eosin or with Haidenhain’s Azan (Presnell and Schreibman, 1997). 409 Cnidae data were gathered following the method of England (1987) and Williams (1996). Cnidae were measured from both live and preserved specimens of A. atodai, and from preserved specimens of A. her- maphroditica. Cnidae were measured in smash preparations at 1000 X using differ- ential interference light microscopy. The terminology for cnidae follows Weill (1934), Mariscal (1974), and England (1991). The material examined was deposited in the Costal Branch of Natural History Mu- seum and Institute, Chiba (CMNH), Nation- al Science Museum, Tokyo (NSMT), Swed- ish Museum Natural History, Stockholm (SMNH), State Zoological Museum, Mu- nich (ZSM), and The University of Kansas Natural History Museum and Biodiversity Research Center (KUMNH). Systematic Account Family Actiniidae Rafinesque, 1815 Genus Anthopleura Duchassaing and Michelotti, 1860 Anthopleura atodai, new species Figs. 2—5 Anthopleura sp.—Atoda, 1954: 274, figs. 1-29, pls. 6—7.—Isomura et al., 2003: M3), 103, Ik. Holotype.—Kenashi-jima, Otsuchi, Iwate Pref., Honshu, Japan (39°21'30’N, 141°57’S50"E), 14 July 1997, collected by KY, | specimen, with histological sections and cnidae preparations (CMNH-ZG 64). Paratypes.—All from Honshu, Japan and collected by KY: Kenashi-jima, Otsuchi, Iwate Pref., 14 Jul 1997, 1 specimen with cnidae preparations (CMNH-ZG 65), 1 specimen with histological sections (NSMT-Co 1373), 1 specimen (NSMT-Co 1374), 1 specimen (KUMNH 1808), 1 spec- imen entirely sectioned longitudinally (CMNH-ZG 3692), 1 specimen entirely sectioned transversely (CMNH-ZG 3693); Banda, Tateyama, Chiba Pref., 28 Oct 1996, 1 specimen with cnidae preparations PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Y Fig. 2. A-C, Photographs of Anthopleura atodai, new species (A, collected at Banda, 24 Feb 1997; B, C, collected at type locality, Kenashi-jima, 14 Jul 1997): A, 2 specimens expanded and 1 specimen contracted; B, semi-expanded specimen; C, fully contracted specimen. D, E, Photographs of A. hermaphroditica (collected from Chiloe Island, Chile): D, Two specimens expanded; E, Typical oral disc patterning. Photographs in D, E courtesy of V. Haussermann. Scale Bars: A, D = 10 mm; B, C = 5 mm, E = 20 mm. VOLUME 117, NUMBER 3 (CMNH-ZG 115), 1 specimen (CMNH-ZG 200), 11 Dec 1996, 24 Feb 1997, 1 speci- men with cnidae preparations (CMNH-ZG 44), 1 specimen (NSMT-Co 1372), 5 Dec 1997, 1 specimen entirely sectioned longi- tudinally (CMNH-ZG 3695), 1 specimen entirely sectioned transversely (CMNH-ZG 3696); Hadaka-jima, Asamushi, Aomori Pref., 22 Jun 1998, 1 specimen with histo- logical sections and cnidae preparations (CMNH-ZG 209), 1 specimen with cnidae preparations (CMNH-ZG 210), 1 specimen (KUMNH 1809), 1 specimen entirely sec- tioned transversely (CMNH-ZG 3697). Non-type material examined.—All spec- imens collected from Honshu, Japan by KY: Kedo-ura, Katsuura, Chiba Pref., 1 May 1999, 4 specimens (CMNH-ZG 253); Banda, Tateyama, Chiba Pref., 11 Dec 1996, 4 specimens (CMNH-ZG 201), 24 Feb 1997, 6 specimens (CMNH-ZG 3694), 15 May 2000, 2 specimens (CMNH-ZG 906); Nojima, Otsuchi, Iwate Pref., 8 Aug 2001, 4 specimens (CMNH-ZG1060), 4 specimens (CMNH-ZG 1061); Hadaka- jima, Asamushi, Aomori Pref., 22 Jun 1998, 60 specimens (CMNH-ZG 3698), 5 speci- mens (KUMNH 1809-1810), 3 specimens (NSMT-Co 1375-1378). Description.—Column and pedal disc: Freshly collected specimens brown or blu- ish-green, proximal verrucae whitish (Fig. 2A-C). In living, expanded animals, col- umn width 6-12 mm, almost equal to height (Fig. 2A—C); oral and pedal disc of almost equal width. Column of contracted animals dome-like (Fig. 2A, C). Adhesive endocoelic verrucae in regular vertical rows from margin to limbus; in some individuals, becoming dense and irregular distally (Fig. 2B); number of rows 24—39 (37 in holo- type) distally, 24 proximally. Diameter of verrucae increases proximally: 0.4 mm at margin, 0.6—1.2 mm at limbus. In life, ver- rucae hold bits of gravel and broken shells. Marginal endocoels bear 9-32 pale, opaque, spherical acrorhagi that curve into fosse (Table 1). Pedal disc weakly adherent, »’ indicates that an attribute was not measured Table 1.—Morphological variability of 11 specimens of Anthopleura atodai, n. sp. collected from three localities. ** or counted for that specimen. Number of siphonoglyphs Number of pairs of directive mesenteries Number of pairs of mesenteries Number of (in mm) acrorhabi Distal column Proximal column Height of column Diameter of pedal disc (in mm) Specimen 28 32 23 10.7 12.0 ype (CMNH-ZG 64) ype (CMNH-ZG 65) 28 34 34 28 9.7 12.4 nN AN 34 34 14.4 10.0 11.9 11.7 39 24 20 28 12.1 9.3 N 7.9 9.8 6.1 28 15 9.5 38 34 35 38 34 35 9.7 11.8 7.8 N 11.9 N ype (NSMT-Co 1373) ype (KUMNH 1808) CMNH-ZG 3693) CMNH-ZG 209) KUMNH 1808) CMNH-ZG210) CMNH-ZG44) CMNH-ZG115) ype (CMNH-ZG 3696) ere ere ve eevee ie holo para para’ para para para para para para para para 411 412 circular in outline, paler in color than col- umn. Oral disc and tentacles: Diameter of oral disc of slightly contracted, fixed anemone approximately equal to that of pedal disc and column. Center of oral disc somewhat elevated into oral cone that bears mouth; mouth elongate along directive axis. Each siphonoglyph marked with a bright cobalt- blue spot in life (Fig. 2A, B); color fades in preservation. Tentacles marginal, slender, shorter than oral disc diameter, number 40 to 62 (59 in holotype). Each tentacle trans- lucent whitish to gray, with parallel longi- tudinal grayish streaks and/or white flecks on oral surface (Fig. 2A, B). Circular mus- cles of tentacles endodermal, longitudinal muscles of tentacles ectodermal (Fig. 3B). Numerous zooxanthellae in endoderm. Marginal sphincter muscle: Endodermal, circumscribed-pinnate to circumscribed-dif- fuse, with highly branched mesogleal pro- cesses (Fig. 4B, C). Mesenteries and internal anatomy: Actin- opharynx whitish, half to two-thirds length of column, with two siphonoglyphs each at- tached to a pair of directive mesenteries. Distinct marginal stomata; oral stomata not seen. Mesenteries in 24—39 pairs, arranged hexamerously in three to four cycles, same number proximally and distally (Table 1). Mesentery arrangement irregular in speci- mens that have regeneration scars. All older mesenteries, including directives, fertile; all specimens hermaphroditic, with gametes of both sexes on same mesenteries or not (Fig. 3C). Zooxanthellae more numerous in en- doderm of column than in endoderm of mesenteries. Each specimen may contain as many as 22 brooded young, early embryos through young adults with two cycles of mesenteries and tentacles (Fig. 3D—F); brooded young posses zooxanthellae. Mesenterial retractor muscles strong, dif- fuse to restricted (Fig. 4A). Parietobasilar muscles well developed, extend half to en- tire distance between column wall and re- tractor muscle, with small free pennon dis- tally (Fig. 4A). Basilar muscles distinct PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Fig. 3A). Cnidom: Spirocysts, basitrichs, holotrichs, heterotrichs, microbasic p-mas- tigophores, microbasic p-amastigophores (Fig. 5). Sizes and distribution of cnidae given in Table 2. Distribution and habitat.—Known from the middle to northern Pacific coasts of Honshu, Japan (Fig. 1). Found in high in- tertidal, attached to Mytilus or in crevices of rock. Typically forms dense populations. Etymology.—The species is named after Dr. K. Atoda, who first identified this as a new species. Anthopleura hermaphroditica (Carlgren, 1899) lags, Z, SD, © Bunodes hermaphroditicus Carlgren, 1899: 23). Anthopleura hermaphroditica Carlgren, 1899.—Carlgren 1927: 32.—England 1987: 245. Anthopleura hermafroditica Carlgr. Carl- gren 1949: 54.—1959: 22. non Cribrina hermaphroditica Carlgren, 1899.—MceMurrich, 1904: 287.—Daw- son, 1992: 38. Material examined.—SMNH 1177 (syn- type), SMNH 40829, 40830; ZSM (un- numbered) Description.—Column and pedal disc: Freshly collected specimens olive green to rosy pink, proximal verrucae paler (Fig. 2D). In living, expanded specimens, col- umn width 15—20 mm, height 17—25 mm. In contraction, column dome-like, width 4— 10 mm, height 3—12.5 mm. Adhesive, en- docoelic verrucae (Fig. 6A) in regular ver- tical rows from margin to limbus; number of rows 23—42. Verrucae larger and more prominent distally than proximally; maxi- mum diameter of distal verrucae 0.5 mm in preserved specimens. In life, verrucae hold small stones and pieces of shells. Margin denticulate, with endocoelic conical projec- tions that bear 1—3 verrucae on the outer surface; projection may bear a swollen ac- rorhagus on the inner surface. Acrorhagi VOLUME 117, NUMBER 3 413 Fig. 3. Anthopleura atodai, new species (A, B, holotype CMNH-ZG 64; C, paratype CMNH-ZG NSMT-Co 1373; D, paratype CMNH-ZG 3692; E, paratype CMNH-ZG 3695): A, cross section of proximal column showing directive mesenteries flanked by those of the second (II), third (IID) and fourth (IV) cycles; B—E, cross sections thorough circumscribed marginal sphincter. Scale Bars: A = 1 mm; B—E = 200 wm. Abbreviations.—4d, directive mesentery; p, parietobasilar muscle; 1, retractor muscle. Arrow indicating brooded young. 414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SRE: 2 iN GY _— Fig. 4. Anthopleura atodai, new species (A, E, paratype CMNH-ZG 3692; B-C, holotype CMNH-ZG 64; D, paratype CMNH-ZG 3969; G, paratype CMNH-ZG 3696; H, paratype CMNH-ZG 209): A, longitudinal section through pedal disc showing basilar muscles; B, longitudinal section through a tentacle; C—D, cross sections through a mesentery showing both spermatocysts and oocytes; E-H, internally brooded young in the enteron (E, EK H), and tentacles (G). Scale Bars: A, C, D = 200 pm; B, G, H = 500 pm; E, F = 100 pm (E- FE Abbreviations.—o, oocyte; s, spermatocysts. VOLUME 117, NUMBER 3 Fig. 5. Cnidae of Anthopleura atodai, new species (paratype CMNH-ZG65); see Table 2 for size ranges. The cnidae of A. hermaphroditica are identical in mor- phology and in distribution in the body, but differ in size; see Table 3 for sizes. A-C from tentacles, D-G from acrorhagus, H-J from column, K-M from actin- opharynx, N-R from filaments. A, spirocyst; B, basi- trich-1; C, basitrich-2; D, spirocyst; E, basitrichs; FE Holotrich; G, Holotrich; H, basitrich-1; I, S-basitrich; J, heterotrich; K, basitrich-1; L, basitrich-2; M, micro- basic p-mastigophore; N, basitrich-1; O, basitrich-2; P, S-basitrich; Q, microbasic p-mastigophore; R, micro- basic p-mastigophore. Scale Bar = 20 um. endocoelic, opaque, tan to white, approxi- mately 0.5 mm tall. Fosse deep. Pedal disc adherent, roughly circular in outline, paler in color than distal column. Oral disc and tentacles: Oral disc diam- eter of expanded individuals slightly greater than pedal disc diameter. center of dic ele- vated into an oral cone that bears mouth; mouth elongate along directive axis, pale gray to rosy pink in life. Oral disc with Opaque marks; marks grouped into six wedge-shaped zones or forming a stellate pattern of concentric, lighter and darker stripes (Fig. 2D, E); pattern fades in pres- ervation. Tentacles slender, marginal, coni- cal, shorter than oral disc diameter: approx- imately 4 mm long in an expanded pre- served individual; innermost tentacles 415 slightly longer than outermost tentacles. Tentacles number 34—80, in three to five cycles. In life, tentacles translucent, typi- cally with opaque white base and cross-bars on oral surface (Fig. 2D, E). Circular mus- cles of tentacles endodermal, longitudinal muscles ectodermal. Zooxanthellae in en- doderm. Marginal sphincter muscle: Endodermal, circumscribed-pinnate, pedunculate, asym- metrical, with closely spaced, highly branched mesogleal processes (Fig. 6C). Mesenteries and internal anatomy: Actin- opharynx one-half to two-thirds length of column, with two aborally prolonged si- phonoglyphs each attached to a pair of di- rective mesenteries. Marginal stoma slight- ly larger than oral stoma. Mesenteries in 24—48 pairs, arranged hexamerously into three to five cycles, same number proxi- mally and distally. Mesenteries of first three cycles typically perfect, those of fourth cy- cle imperfect. All perfect mesenteries, in- cluding directives, fertile, each typically bears both male and female gametes (Fig. 6D). Mesenteries of specimens that contain many brooded young typically lack gametic tissue. Zooxanthellae more numerous in en- doderm of column than in that of mesen- teries. A specimen may contain as many as nine brooded young; brooded young up to 2 mm long, with an oral disc diameter of 1 mm, and as many as 20 tentacles. Largest brooded young zooxanthellate, with small endocoelic verrucae and marginal projec- tions. Mesenterial retractor muscles diffuse-re- stricted; retractor typically abuts parietal muscle pennon (Fig. 6E). Parietobasilar muscles strong, each with a broad pennon and many short, thick, lateral processes. Pa- rietal muscle may span as much as half the distance between the column and the free edge of the mesentery. Basilar muscles strong (Fig. 6B). Cnidom: Spirocysts, basitrichs, hetero- trichs, holotrichs, microbasic p-amastigop- hores, microbasic p-mastigophores. Sizes PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 416 Uu u u u u u u (S1'0/6'7) TE-BT X) OO T/S 91) S 8I-S EI ‘(OE O/V' Py) O'S-6'E X ‘(L0'0/0'1) T 1-60 Xx (LT 0/9'€) O'P-TE X ‘(60'0/0'7) 1°81 X ‘(Or 0/9'b) TS—-O'V X “(LTO/ET) 8 T-O'T X OL = ¥ (80/07) 17-81 X Ol = Uu ‘(EE O/S E) BEBE X €=u4 Ol ‘(OL 0/07) S78 ‘(90'0/0'7) TT-0'T “(OS '0/6'€) 0'S-0'E “IT O0/C TC) €T-0'T ‘(Cr 0/V'T) O'E€-0'7T ‘(PL O/LO) SC-61 “(S0'0/0'7) 170° x x x x x x x x (TE O/0'E) LH-ST X (66'0/S 17) 0'€Z—0'07 (SET/S LE) S SE—0'8T (6 11/CVO) S STS 17 (OV O/EST) OLI-O'FI (€O' 1/861) € 1T-O'81 (870/67) T 8-0 CC (COI/ESI) € 17-081 (80'1/S 91) O'8I-S' FI (€S'E/S' ET) 0'9T—O'IT (SO T/E VL) €9I- Ol (L8'0/9' V1) L91-0'€EL (C1'€/6.9€) O'Cr-0'0E (L5'0/9'ST) O'91-C ST (87'€/6'ST) € €€—0'07 (€T'1/6'ST7) O'€T-C 81 (LIV 1) TEIl-001 (€9' 1/7 ST) S 87-07 sII Ol! Ol! cOl a (sues) euey (¢9DZ-HNWDO) 2dGoj0H Neen eee eee 6t-¢T X 8 CC-OTI CS-6E X OSC-ELI L1-60 X L9E-C VTE 97-6T X V8C-O'IC 9C-8 1 X 1 O0C-O'TI 9S-TE XK SOT-TSI CEO 1 XK T8C-0'CC CEPI &* Suge Iv-87 X 8 6I-S FI €C60 X 6 09-0'17C 9C-El X 88I-06 OC-LI X SLI-IT CI 8S-IT X T8t-E 0C €CT91 X 90C-8 OI 8 EOC X 6 EE-B TI 8C-6 1 X OEC-CSI €CSI X CEI-S 6 Tv-8' 1 X 8 6c-0'SI (wml) 9Z1S u t/v (qy) s10ydosysew-d o1seqo1syy v/v (O) e10ydosnseur-d otseqo1y| v/v (S) youIseg-S v/v (N) Z-YoLniseg v/v (N) [-YoLniseg JUS b/p (WN) er0ydos8nseure-d oiseqo1sljyy v/v (q) T-younseg vt (D [-youniseg xuAreyg v/V (f) YoINno1N10H v/v (1) yorniseg-s v/v (H) [-YoLnise g UUIN[OD [BUILXOIg E/E (H) Yorniseg UUINJOS [eIsIq V/V (D ‘4) YOINo[oH v/v (q) youtseg v/v (q) 3sAd011dg ISVYIOINY v/v (DO) c-YoLnIseg vv (dq) [-YoLniseg V/V (Vv) isAo0n1ds gjorua y, N epruo yo adAy, uoners07T “uUNTOS ayeIedas UI UDAIS OIe (PODZ-HNIND) edAlofoy 410}; veg -adkjoJoy WO BVP SuIpNjout ‘poinsvour sojnsdeo yo roquinu oy) st ,.U,, ‘oeprud Jo adAj yey) SuTUTeJUOS pouTUeXe suoUUToeds Jo JoquINU 9Y) SI ..N,, “sosoyjuored ul o1e sginsdeo [jews 10 o81e] AjeuoNdeoxe Jo s}uoWIaINsvoUT SYIPIA puL YVSus] Jo sosuvs sv USAIS av SOZIG “¢ “SIY 0} JoJor Jone] “poly vAnajdoysuy Jo sepluy—T PGP.L VOLUME 117, NUMBER 3 417 pe 4 “iy Uy Aa, », See ay Fig. 6. Internal anatomy and histology of Anthopleura hermaphroditica. A, longitudinal section through a verruca; B, longitudinal section through pedal disc showing basilar muscles; C, cross section through sphincter muscle; D, cross section through a mesentery showing both spermatocysts and oocytes; E, cross section through proximal to actinopharynx showing mesenteries of first (1), second (II), third (III), and fourth (IV) cycles. Scale Bars: A = 150 um; B, C = 100 um; D = 200 pm; E = 600m. Abbreviations.—p, parietobasilar muscle; r, retractor muscle; s, spermatocyst. Arrow indicating oocyte. 418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and distribution of cnidae given in Table 3; cnidae illustrated in Fig. 5. = ne ee : @ +t Distribution and habitat.—Known only bs a ¥ from high intertidal zone of Chile. il ii S Ri Seow 9 oO < as i és Ean onse) Rim sk CK WEA Discussion = Sent ae }+ < ere OE Seg ee ae nan ‘ . . xx WOKE silieff, 1908; A. pacifica Uchida, 1938; A. J 2 Al was © iridi. : iati a|\| ATA = TT SPO AT eer Juscoviridis Carlgren, 1949; A. asiatica E| ax Gn ex ma cx qa Uchida & Muramatsu, 1958; A. kurogane ~l ono dg Pet todamdtes . aa S| aanit CNW dn a Cada Uchida & Muramatsu, 1958; A. uchidai BN Fe ay eRe Cay England, 1992, and from most other nom- QRom~ Boqgnrannms : é : ae jawortoaetrrnHsoarka inal species of Anthopleura. It is distin- WAT YAAVTIAITP IDG ished fi A di by the ch aon Tenaee ttn ae guished from A. aureoradiata by the char- BOR HOOGR eC aaace Sn 0 ON OOO) al mea OS I Th a Th | acter of verrucae at the lower column and the coloration of the column: in A. aureo- radiata, the verrucae diminish in size prox- “ “1)5RSSSSeAnRSAoHA imally (Parry 1951), and “near the bottom oD Se ne of the column these become mere mark- ings” (Stuckey 1909a: 369); in A. atodai, the verrucae increase in size proximally. In A. aureoradiata, the coloration of the col- umn differs distally and proximally (Stuck- ey 1909a, Parry 1951), whereas in A. ato- dai, the coloration of the column is uni- form. Anthopleura atodai is distinguished from A. handi in its hermaphroditism, pos- Table 3.—Cnidae of Anthopleura hermaphroditica. Letters refer to Fig. 5. Sizes are given as a length by width range, in 1m; measurements of exceptionally large or small capsules in parentheses. ““N’’ is the number of specimens examined containing that type of cnidae, ‘‘n’’ is the number of capsules measured. The size range of cnidae for A. handi, combined from Dunn (1974) and England (1987) is given for comparison. eee g 62 £48 session of zooxanthellae, and circumscribed z & = & marginal sphincter muscle. Al Big ee ee. : 3 Anthopleura atodai and A. hermaphrod- a eleguzc¥d KEQODAR itica are both hermaphroditic, and brood = ase a 3 ao 2 BOS 2 2 young internally. However, they are distin- oe a s SEOEE ro Ele Ss 6 guished by number of tentacles, coloration, Baaee ae SEZI2a2So8 size of cnidae, and geographical distribu- wes Fe I tion. The maximum number of the tentacles observed in members of A. atodai is 62, al whereas Carlgren (1899) reported a maxi- aS mum of 90 in specimens of A. hermaphrod- 5 om 3 8 itica. The column of living specimens of A. Q x 3 cs ES 5 atodai is bluish-green or brown; in speci- gs ER: 4 5 mens of A. hermaphroditica, the column is eee At, ra VOLUME 117, NUMBER 3 419 Table 4.—Summary of differences in size and distribution of cnidae between Anthopleura atodai and A. hermaphroditica. Cnidae type and location Tentacle basitrichs Acrorhagus holotrichs Proximal column heterotrichs Proximal column basitrichs Filament basitrich-2 Shorter in A. atodai Shorter in A. atodai gray or pink. The nematocysts of the ten- tacles, acrorhagi, column, and filaments fur- ther distinguish the two (Table 4). Taxonomy of Anthopleura hermaphrodi- tica—The taxonomy of Anthopleura her- maphroditica has been confused because of a series of misidentifications and because of a proposed synonymy between A. herma- phroditica and A. handi. In the original de- scription of the species, as Bunodes her- maphroditicus, Carlgren (1899) mentioned two notable features: hermaphroditism and acrorhagi. McMurrich (1904) found speci- mens of a hermaphroditic actiniid from Chile that had pseudoacrorhagi, rather than true acrorhagi and identified these as Cri- brina hermaphroditica, changing the gener- ic assignment of Carlgren’s species and contesting Carlgren’s (1899) assertion that the species had acrorhagi. Carlgren (1927) transferred the species to Anthopleura, a ge- nus characterized as having acrorhagi, but maintained that the species he had origi- nally called Bunodes hermaphroditica and the specimens described by McMurrich (1904) as C. hermaphroditica were the same species. However, after examining ad- ditional material from Chile, Carlgren (1959) reversed this opinion, and erected a new species, B. hermaphroditica, which he attributed to McMurrich. Carlgren’s (1959) description constitutes a new combination for C. hermaphroditica McMurrich 1904, rather than an original description. According to the International Code of Zoological Nomenclature (ICZN 1999: Art. 11.6), the name C. hermaphrod- itica was made available by its subsequent use as valid (e.g., Clubb, 1908), and its au- Difference One size class in A. hermaphroditica; two distinct classes in A. atodai Narrower in A. atodai One size class in A. hermaphroditica; two distinct classes in A. atodai thorship dates from its publication by McMurrich (1904) as a synonym of Buno- des hermaphroditica (International Code of Zoological Nomenclature: Art. 50.7; ICZN 1999). Therefore, the specimens Mc- Murrich examined constitute the type series for C. hermaphroditica McMurrich, 1904; the type specimens of Bunodes herma- phroditicus Carlgren, 1899 (SMNH 1177) belong to Anthopleura as they have true ac- rorhagi with holotrichous nematocysts. The surviving material from the Lund University Chile Expedition includes two recognizable species: A. hermaphroditica (Carlgren, 1899) and Bunodactis herma- Dhroditica (McMurrich, 1904). There are many more specimens belonging to Buno- dactis hermaphroditica than to A. herma- phroditica; the difference in number of specimens collected reflects their abun- dance in the field (V. Haussermann, pers. comm.). Specimens belonging to Bunodac- tis hermaphroditica \ack holotrichous nem- atocysts in the distal column and in the proximal column; both of these features are diagnostic at the level of genus (e.g., Eng- land, 1987). Specimens of Bunodactis her- maphroditica have more prominent verru- cae than specimens of A. hermaphroditica, especially proximally. England (1987) suggested that A. her- maphroditica might be synonymous with A. handi. We disagree with this proposition of synonymy because A. hermaphroditica and A. handi differ in several important re- spects. Most importantly, members these two species differ in key life history fea- tures: members of A. hermaphroditica are hermaphroditic and zooxanthellate, mem- 420 bers of A. handi are gonochoric and azoox- anthellate. Furthermore, the basitrichs in both the distal and proximal column are larger in members of A. handi than in mem- bers of A. hermaphroditica. Finally, there is a considerable disparity in the geographic range and habitat of the two species: A. handi is found in the tropical Indo-Pacific around Malaysia, Singapore, and New Guinea (Dunn 1978, 1982b; England 1987; Fautin 1988); A. hermaphroditica is re- stricted to cold waters of the western Pacific (Carlgren 1899, 1959). Biology of Anthopleura atodai.—Antho- pleura atodai clearly corresponds to Ato- da’s Anthopleura sp.: the two have identical distributions, life history, and coloration. All specimens examined, regardless of size, were simultaneously hermaphroditic. In ac- tiniarians, hermaphroditism is unexpectedly rare (Shick 1991) in view of the “low den- sity model” of Ghiselin (1969). Among hermaphrodites, simultaneous hermaphrod- itsm is the most common mode; known ex- ceptions include the protandrous hermaph- rodite Sicyopis (= Kadosactis) commensal- is (Gravier, 1918) and the gynodioecious species Epiactis prolifera (Verrill, 1869) and Cereus pedunculata (Pennant, 1777) (see Bronsdon et al. 1993, Dunn 1975, Ros- si 1971). The reproductive biology of A. atodai re- mains ambiguous. Isomura et al (2003) were unable to find gametogenic tissue in any specimens that they identified as An- thopleura sp. sensu Atoda, although they regularly found brooded young. The mes- enteries of some specimens bore spherical protuberances proximally that were inter- preted to be early stages of the brooded young; from this they inferred that the brooded young were asexually produced (Isomura et al. 2003). None of our results refute an asexual origin for the brooded young. However, our finding of fertile spec- imens from the study site of Isomura et al. (2003), including those that contained both gametes and brooded young (e.g., Fig. 4A), indicates that the species is not exclusively PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON asexual, and lends support to the contention by Isomura et al. (2003) that the Mutsu Bay population is remarkable in lacking fertile individuals. In general, the gametes and the gametogenic region are small in A. atodai, making it possible that Isomura et al. (2003) overlooked them in the specimens they examined. The presence of gametes does not rule out an asexual origin for the brooded young; some species of Actinia have both gametes and asexually produced young in their enteron (Yanagi et al., 1999). Therefore, further investigation is necessary to definitively demonstrate the asexual ori- gin of the brooded young and to clarify re- productive ecology of A. atodai. Acknowledgments MD supported by NSF- DEB 9978106 (to D.G. Fautin). A part of this work also supported by Fujiwara Natural History Foundation (to the first author, KY). We thank D.G. Fautin (Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence KS, U.S.A.) and E. A. Robson (School of Animal and Microbial Sciences, The University of Reading, Read- ing, U.K.) for comments that improved this manuscript. We especially thank V. Haus- sermann (ZSM) for permission to use her photographs, for her generous loan of ma- terial, and for her insight into the ecology and biology of A. hermaphroditica. S.D. Cairns (USNM) and K. Sindemark (SMNH) also provided specimens. We thank the staff of Otsuchi Marine Research Center, Ocean Research Institute, Univer- sity of Tokyo; Asamushi Marine Biological Station, Biological Institute of the Faculty of Science of Tohoku University; Banda Marine Laboratory of Tokyo University of Fisheries for help in sampling. We are deep- ly grateful to the late E. 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Actiniaria (Cnidaria: Anthozoa) from Hong Kong with additional data on similar spe- cies from Aden, Bahrain and Singapore. Pp. 699-705 in B. Morton, ed., The marine flora and fauna of Hong Kong and southern China If. Proceedings of the Fourth International Ma- rine Biological Workshop; The Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong, 11—20 April 1989. Hong Kong University Press, Hong Kong. Fautin, D.G., & FE -S. Chia. 1986. Revision of the sea anemone genus Epiactis (Coelenterata: Actini- aria) on the Pacific coast of North America, with descriptions of two new brooding spe- cies.—Canadian Journal of Zoology 64:1665— 1674. . 1988. Sea anemones (Actiniaria and Coralli- morpharia) of Madang Province.—Science in New Guinea 14:22—29. . 2003. Hexacorallians of the world http://her- cules.kgs.ku.edu/hexacoral/anemone2/in- dex.cfm (Version 15 July 2003). Ghiselin, M. 1969. The evolution of hermaphroditism among animals.—Quarterly Review of Biology 44:189-208. Gravier, C. 1918. Note préliminaire sur les Hexacti- niaires recueillis au cours des croisiéres de la Princesse-Alice et de l’Hirondelle de 1888 a 422 1913 inclusivement.—Bulletin de 1’Institut Océanographique (Monaco) 346:1—24. International Commission on Zoological Nomencla- ture (ICZN). 1999. International code of zoo- logical nomenclature. International Trust for Zoological Nomenclature, London, 306 pp. Isomura, N, K. Hamada & M. Nishihira. 2003. Internal brooding of clonal propagules by a sea anem- one, Anthopleura sp.—Invertebrate Biology 122:293—298. MeMurrich, J. P. 1904. The Actiniae of the Plate col- lection (Fauna Chilensis 3).—Zoologische Jahr- bucher Jena. (Supplement 6):215—305. Manuel, R. L. 1988. British Anthozoa, Synopses of the British Fauna (New Ser.) No. 18 (Revised). The Linnaean Society of London, London, 241 pp. Mariscal, R. N. 1974. Nematocysts. Pp. 129-178 in L. Muscatine and H. M. Lenhoff, eds., Coelenter- ate biology: reviews and new perspectives. Ac- ademic Press, New York, 501 pp. Parry, G. 1951. The Actiniaria of New Zealand: a check-list of recorded and new species, a review of the literature and a key to the commoner forms, part 1—Records of the Canterbury Mu- seum 6:83-119. Pennant, T. 1777. A British Zoology. Benjamin White, London, 136 pp. Presnell, J. K., & M. P. Schreibman. 1997. Humason’s Animal tissue techniques, 5th edition. Johns Hopkins University Press, Baltimore, 572 pp. Rafinesque, C. S. 1815. Analyse de la Nature ou Tab- leau de l’Univers et des Corps Organisés. C. S. Rafineque, Palerme, 224 pp. Rossi, L. 1971. Thelytochous parthenogenesis in Ce- reus pedunculatus (Actiniaria).—Experientia 27:349-351. Russo, C. A. M., A. M. Solé-Cava, & J. P. Thorpe. 1994. Population structure and genetic variation in two tropical sea anemones (Cnidaria, Acti- niaria) with different reproductive strategies.— Marine Biology 119:267-276. Shick, J. M. 1991. A Functional biology of sea anem- ones. Chapman & Hall, London, 395 pp. Stuckey, W. 1909a. Notes on a New Zealand actinian, Bunodes aureoradiata.—Transactions of the New Zealand Institute 41:367—369, pl 17. . 1909b. A review of the New Zealand Acti- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON niaria known to science, together with a de- scription of twelve new species.—Transactions of the New Zealand Institute 41:374—398, pls 21-28. Uchida, T. 1934. A brood-caring actinian subject to a wide range of colour variation.—Journal of the Faculty of Science, Hokkaido University, Series 6 (Zoology) 3:17-31. . 1938. Report of the biological survey of Mut- su Bay. 33 Actiniaria of Mutsu Bay.—Science Reports of the Tohoku Imperial University, 4th Series (Biology) 13:281—317. , & EF Iwata. 1954. On the development of a brood-caring actinian.—Journal of the Faculty of Science, Hokkaido University, Series 6 (Zo- ology) 12:220—224. , & S. Muramatsu. 1958. Notes on some Jap- anese sea-anemones.—Journal of the Faculty of Science, Hokkaido University, Series 6 (Zool- ogy) 14:111—-119. Verrill, A. E. 1869. Review of the polyps of the west coast of America.—Transactions of the Con- necticut Academy of Arts and Sciences 1:377— 567. Wassilieff, A. 1908. Japanische Actinien. Pp. 1—52 in FE Doflein, ed., Beitrage zur Naturgeschichte Ostasiens.—Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematische- aturwissenschaftliche Abteilung (Munchen), Supplement B 1 (2). Weill, R. 1934. Contribution a I’ etude des cnidaires et de leurs nematocystes. II. Valeur taxono- mique du cnidome.—Travaux de la Staion Zoologique de Wimereux 11:340—701. Williams, R. B. 1996. Measurements of cnidae from sea anemones (Cnidaria: Actiniaria): statistical parameters and taxonomic relevance.—Scientia Marina 60:339-351. Yanagi, K., S. Segawa, & T. Okutani. 1996. Seasonal cycle of male gonad development of the inter- tidal sea anemone Actinia equina (Cnidaria: An- thozoa) in Sagami Bay, Japan.—Benthos Re- search 51:67—74. , & K. Tsuchiya. 1999. Early devel- opment of young brooded in the enteron of the beadlet sea anemone Actinia equina (Anthozoa: Actiniaria) from Japan.—lInvertebrate Repro- duction and Development 35:18. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):423—446. 2004. New species and new combinations in Rhysolepis (Heliantheae: Asteraceae) Harold Robinson and Abigail J. Moore (HR) Department of Botany, National Museum of Natural History, MRC-166, Smithsonian Institution, PO. Box 166, Washington, DC 20013-7012; (AJM) Department of Biology, University of Utah, Salt Lake City, UT 84112 Abstract.—A narrow circumscription of the genus Viguiera Kunth results in transfer of 58 species of Helianthinae with glabrous stamen filaments, exap- pendiculate style appendages, and a persistent pappus into Rhysolepis S.EBlake. Rhysolepis dillonorum from Peru, R. emaciata from Bolivia, and R. goyasensis, R. hatschbachii, R. laxicymosa, R. santacatarinensis, and R. sub- truncata from Brazil are new species. Viguiera pazensis and V. procumbens are placed in synonymy under Rhysolepis helianthoides, and V. misionensis 1s combined with R. pilosa. The present study began with a project by the junior author to clarify the limits of two Andean species of Viguiera Kunth in H.B.K. and join in the description of a spe- cies from Brazil known to be undescribed. The project was undertaken with the knowl- edge that none of the species involved were truly congeneric with the type species of Viguiera, V. helianthoides Kunth in H.B.K. = V. dentata (Cav.) Spreng. The arrival of additional material from Gert Hatschbach of the Museo Botanico Municipal de Curi- tiba, led to review of other species prob- lems and discovery of additional species needing description. In view of the number of species involved and because of the ge- neric redelimitations of Schilling and Pa- nero (2002), the decision has been made to abandon the long misapplied name Vigui- era and use a more phyletically appropriate generic concept for the species in this study. Viguiera traditionally has contained spe- cies related to Helianthus L., but differing by a more persistent pappus with squamel- lae. The most recent treatment of Viguiera in the broad sense was that of Blake (1918). Blake’s treatment excluded some genera such as Tithonia Desf. with broadened, fis- tulose peduncles (La Duke 1982); Syncre- tocarpus S. F Blake (1916) with a glabrous strip just inside the lateral margins of its achenes that was misinterpreted as a wing; and Rhysolepis S. E Blake (1917) with transverse corrugations on its paleae. The broad Blake concept of Viguiera included some elements now placed in Hymenoste- phium Benth. in Benth. & Hook.f., but ex- cluded others (Schilling and Panero 2002). Some single species once placed in Vigui- era have been moved to other genera, in example a Peruvian species named by Blake in 1918, Viguiera acutifolia, has been transferred to Pappobolus (Panero 1992) and a Mexican species included in Viguiera by Blake (1924) was subsequently trans- ferred to Stuessya (Turner & Davies 1980). In a brief review of members of the Hy- menostephium group, Robinson (1977) re- tained the broad concept of Viguiera in spite of the realization that the type species of Viguiera was individually distinctive with pubescent anther filaments and a small apical appendage on the branches of the style. Hymenostephium was retained in Vi- guiera because it had an apical appendage on the style branches and was technically closer to the type of Viguiera than most other species placed in the latter genus. The 424 needed generic revisions of the concept of Viguiera were fully initiated by Schilling and Panero (2002); but the South American species and their relatives in Mexico with exappendiculate styles have not yet been treated. The South American species are in need of transfer to some genus other than Vigui- era. The problem has been that none of the synonyms given by Blake (1918) seems to be applicable. Leighia Cass. belongs to the group, but that name is a later homonym of Leighia Scop. As noted by Blake, the type of Harpalium Cass., H. rigidum (Desf.) Cass. ( = Helianthus rigidus Desf.), is not a Viguiera. Other Blake synonyms, Heliom- eris Nutt. and Bahiopsis Kellogg, are con- sidered separate genera (Schilling & Panero 2002). The type of Gymnolomia Kunth in H.B.K., after some confusion, proved to be- long to Eleutheranthera Poit. ex Bosc. (Robinson 1992). Thus, none of the syno- nyms from Blake (1918) can be used. A name is found, however, outside the syn- onymy of Viguiera as circumscribed by Blake in 1918. His genus Rhysolepis, in spite of its sometimes weakly transversely corrugated paleae, is not distinct from the group treated here, and so the name can be applied. Rhysolepis S. F Blake, Contr. Gray Herb. 52: 36 (1917).—Type: Viguiera pal- merit A. Gray Leighia Cass. in E Cuvier, Dict. Sci. Nat. ed. 2. 25: 435 (1822).—Type: He- lianthus linearis Cav. Not Leighia Scop. (1777). = Ethulia L.f. Annual to perennial herbs or shrubs; of- ten with tubers or with fusiform nodes on roots. Stems and leaves usually strigose, pi- lose, or hispid. Leaves alternate or opposite, sessile or petiolate, filiform to ovate, lan- ceolate, oblong, or broadly rounded; blade often trinervate with secondary veins near and subparallel to lower margin; margins entire to serrate. Inflorescences usually with 1—6 heads, sometimes heads over 50; pe- duncles usually elongate, 3-30 cm long, of- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ten stout, not enlarged and fistulose distally; involucre broadly campanulate; bracts in 2— 5 series, gradate to subequal, oblong to ovate or lanceolate, at base usually with in- durated ribs, at tips herbaceous, appressed or reflexed, rounded to acute; receptacle convex to conical; pales persistent, partially enclosing achenes, mostly ribbed and in- durate, sometimes transversely corrugated, usually with blunt apex. Ray florets usually 8—24, sterile, sometimes lacking; corollas yellow, with yellow or orange resin ducts along veins. Disk florets usually 40—200, tightly packed, bisexual; corollas yellow or greenish-yellow, 5-lobed, with basal tube usually 0.5—1.0 mm long and narrow, usu- ally scabrid on abruptly broadened base of throat and on backs of lobes, with yellow or orange resin ducts along 5 veins of throat; anther filaments without hairs or pa- pillae; thecae blackish, shortly hastate at base; endothecial cells with nodes on trans- verse walls; apical appendage usually yel- low, blackish in some annual species, ovate, concave abaxially often with cluster of glands in concavity; style with resin ducts outside of veins not restricted to branches; style branches spreading radially, with tuft of hairs or papillae at tip, without apical appendage, with stigmatic papillae covering whole inner surface. Achenes compressed, with or without setulae, without differenti- ated intramarginal bare strip; walls with phytomelanin interrupted by striations of pale cells; pappus mostly persistent, with pair of awns usually longer than squamellae on margins between awns, but awns some- times not longer than squamellae. Chro- mosome numbers n = 17, 34. Rhysolepis was described from Mexico and has previously been credited with only three Mexican species as recognized by Robinson (1972): Rhysolepis kingii H. Rob., Phytologia 24: 210 (1972). Rhysolepis morelensis (Greenm.) S. E Blake, Contr ‘Gray silesba 523550 (1917). VOLUME 117, NUMBER 3 Viguiera morelensis Greenm., Proc. Amer. Acad. 40: 40 (1904). Rhysolepis palmeri (A. Gray) S. FE Blake, Contr. Gray Herb. n.s. 52: 37 (1917). Viguiera palmeri A. Gray in S.Watson, Proc. Amer. Acad. 22: 427 (1887). The broadened concept of Rhysolepis recognized here includes the rather overlap- ping Blake (1918) sections and series, Ten- uifoliae consisting of perennial herbs with linear leaves, solitary heads and involucral bracts 2-seriate and subequal; Revolutae with perennial herbs or subshrubs of the Chilean and Argentine Andes with large solitary heads and involucral bracts 2—5-se- riate, gradate and lanceolate; Grandiflorae with perennial herbs having one or few large, long-pedunculate heads and having few leaves with the lowest opposite and scale-like; Aureae, primarily Andean, in- cluding annuals to shrubby perennials with broad leaves and involucral bracts mostly 3—5-seriate, usually gradate, lanceolate, and with herbaceous tips not strongly differen- tiated; Bracteatae, mostly of Brazil and Paraguay, including herbaceous perennials similar to the Aureae but with involucral bract tips shortly and abruptly herbaceous and blunt; Leighia, mostly Mexican, but similar to the Aureae and Bracteatae with involucral bracts strongly gradate, oblong and usually with an abrupt herbaceous tip; Trichophylla consisting of slender virgate perennials with linear to filiform leaves, revolute leaf margins and involucral bracts lanceolate to linear-lanceolate; and subge- nus Verbalesia containing perennial herbs with pappus awns equalled in length by and partially fused to the squamellae. The fol- lowing new combinations agree, to a con- siderable extent, with species concepts of Blake (1918), although that work left ques- tions about the real distinctions of many species. As a result, more recent synony- mies are taken into account, and other syn- onymies are to be expected. Many poorly known species are omitted. 425 Rhysolepis anchusifolia (DC.). H. Rob. & A. J. Moore, comb. nov. Leighia anchusaefolia DC., Prodr. 5: 580 (1836). L. dissitifolia DC., Prodr. 5: 581 (1836). L. immarginata DC., Prodr. 5: 581 (1836). L. lomatoneura DC., Prodr. 5: 581 (1836). L. stenophylla Hook. & Arn., J. Bot. 3: 313 (1841). L. baldwiniana Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:365 (1841). Viguiera stenophylla (Hook. & Arn.) Gri- seb., Goett. Abh. 24: 193 (1879). V. anchusaefolia (DC.) Baker in Mart., FI. bras. 6(3): 222 (1884). Argentina, Brazil, Uruguay. Rhysolepis arenaria (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera arenaria Baker in Mart., Fl. bras. 6(3): 228 (1884). Brazil, north central Sao Paulo. Rhysolepis aspilioides (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera aspilioides Baker in Mart., FI. bras. 6(3): 228 (1884). Brazil, Matto Grosso. Rhysolepis atacamensis (Phil.) H. Rob. & A. J. Moore, comb. nov. Viguiera atacamensis Phil., Anales Mus. Nac. Chile, Segunda Secc., Bot. 1891: 48 (1891). Chile. Rhysolepis australis (S.EBlake) H. Rob. & A. J. Moore, comb. nov. Viguiera australis S. F Blake, Contr. Gray Herb. n.s. 54: 148 (1918). Chile. Rhysolepis bakeriana (S. FE Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera bakeriana S. F Blake, Contr. Gray Herb. n.s. 54: 130 (1918). Brazil, Minas Gerais. 426 Rhysolepis bishopii (H. Rob.) H.Rob. & A. J. Moore, comb. nov. Viguiera bishopii H. Rob., Phytologia 45: 458 (1980). Bolivia. Rhysolepis bracteata (Gardn.) H. Rob. & A. J. Moore, comb. nov. Viguiera bracteata Gardn., London J. Bot. 7: 404 (1848). Brazil. Distrito Federal, Goias, Minas Gerais. Rhysolepis breviflosculosa (S. FE Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera breviflosculosa S. F Blake, Contr. Gray Herb. n.s. 54: 158 (1918). Uruguay. Rhysolepis brittonii (Hochr.) H. Rob. & A. J. Moore, comb. nov. Viguiera brittonii Hochr., Bull. New York Bot. Gard. 6: 294 (1910). Peru. Rhysolepis discolor (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera discolor Baker in Mart., Fl. bras. 6(3): 228 (1884). Brazil, Minas Gerais. Rhysolepis ellenbergii (Cuatrec.) H. Rob. & A. J. Moore, comb. nov. Viguiera ellenbergii Cuatrec., Proc. Biol. Soc. Wash. 77: 146 (1964). Peru. A second specimen from the type locality is as follows: Peru. Cuzco: Prov. Urubamba, ruinas de Machu Picchu, high above Rio Urubamba, 80 km WNW of Cuzco, rock walls, rock piles, terraces & cliffs, Intyhuatana (Solar Observatory); 2500-2600 m, 27 May 1963, Ugent 5376 (US). Rhysolepis fabrisii (Saenz) H. Rob. & A. J. Moore, comb. nov. Viguiera fabrisii Saenz, Darwiniana 22: 50 (1979). Argentina. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rhysolepis fusiformis (S. E Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera fusiformis S. F Blake, Contr. Gray Herb. n.s. 54: 145 (1918). Bolivia. Rhysolepis gardneri (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera gardneri Baker in Martt., Fl. bras. 6(3): 224 (1884). Originally described from Brazil, Goias. Two more recent collections matching the type photograph are: Brazil. Goidas, Piren- Opolis (Morro da Caixa Dagua); cerrado seco, arborizado, com pedras no solo, su- jeito ao fogo periddico; planta com 80 cm, ramificada inflorescéncia terminais, flores roxas, 23 Apr 1976, Heringer 15560 (UB, US); Municipio de Niquelandia, entrada no km 8 da Rodovia Niquelandia/Uruagu; Fa- zenda Trairas. Morro. Relévo ondulado; 14°29'19"S, 48°33'19"W. Cerrado com mui- tas pedras de cor branca; arbusto, ca. 70 cm de altura; flores com corola amarela e an- teras alaranjadas. Nome comum: margarida; 13 Apr 1996; Mendonca, Marquete, Fon- seca & Oliveira 2453 (UB, US). Rhysolepis gilliesii (Hook. & Arn.) H. Rob. & A. J. Moore, comb. nov. Leighia gilliesii Hook. & Arn., J. Bot. (Hooker) 3: 313 (1841). Helianthus heteropappus Gill. ex Hook. & Arn., J. Bot. (Hooker) 3: 314 (1841), nom. nud. Viguiera gilliesii (Hook. & Arn.) Hieron., Actas Acad. Nac. Ci. Cordoba 4: 39 (1882). Flourensia hispida Phil., Anales Univ. Chile 36: 186 (1870). Argentina, Chile. Rhysolepis grandiflora (Gardn.) H. Rob. & A. J. Moore, comb. nov. Leighia grandiflora Gardn. in Field & Gardn., Sert. Pl. t. 54-55 (1844). Viguiera grandiflora (Gardn.) Gardn., Lon- don J. Bot. 7: 404 (1848). VOLUME 117, NUMBER 3 Rhysolepis guaranitica (Chod.) H. Rob. & A. J. Moore, comb. nov. Viguiera guaranitica Chod., Bull. Herb. Boiss. ser. 2, 3: 724 (1903). Argentina, Brazil, Paraguay. Rhysolepis helianthoides (L. Rich.) A. J. Moore & H. Rob., comb. nov. Fig. | Sanvitalia helianthoides L. Rich. in Willd., Sp. Pl. 3: 2190 (1803). Helianthus procumbens Pers., Syn. Pl. (Per- soon) 2: 475 (1807). Viguiera pazensis Rusby, Mem. Torrey Bot. Club 3(3): 59 (1893). Viguiera pflanzii Perkins, Bot. Jahrb. Syst. 49: 226 (1913).. Viguiera punensis S. FE Blake, Bot. Jahrb. Syst. 54, Beibl. 119: 48 (1916). Argen- tina, Bolivia, Peru. The present complex was maintained as two separate species by Blake (1918) based on longer, relatively narrower leaf shape and more prominent leaf venation in V. pa- zensis. Separation was maintained by Saenz as recently as 1979 based on larger, ovate- lanceolate rather than ovate to oblong leaves, multiple rather than single heads per stem, and smaller involucres in V. pazensis. We could not separate the species using these characters, nor pubescence type or shape of the involucral bracts. Tips of the involucral bracts were sometimes reflexed and thus looked different from bracts with- out reflexed tips, but the lengths and shapes were the same. The broadened concept of Rhysolepis he- lianthoides is characterized by leaves tu- berculate-pilose adaxially, pilose abaxially with hairs denser on veins; stems ribbed and villous; and involucral bracts oblanceo- late, subequal, often recurved, and with an indurate base and herbaceous apex. In ad- dition, the achenes tend to have rather read- ily deciduous awns and squamellae, a char- acter reportedly shared with R. lanceolata (Blake 1918). 427 The concept of Viguiera pazensis in this study includes two isotypes, Bang 44 (US). Some more southern material might prove distinct, and the name Helianthus ataca- mensis Phil. (not Viguiera atacamensis Phil.) is for the present omitted from the synonymy. For an additional specimen that was determined as V. pazensis, but is not this species, see Rhysolepis dillonorum A. J. Moore & H. Rob. below. Rhysolepis hilairei (S. EF Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera hilairei S. FE Blake, Contr. Gray Herb. 54: 153 (1918). Brazil, Minas Ger- ais. Rhysolepis hypoleuca (S. F Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera hypoleuca S. F Blake, Contr. Gray Herb. n.s. 54: 165 (1918). Brazil, Matto Grosso. Rhysolepis incana (Pers.) H. Rob. & A. J. Moore, comb. nov. Helianthus incanus Pers., Syn, Pl. 2: 475 (1807). Helianthus aureus Kunth in H.B.K., Nov. Gen. Sp., ed fol. 4: 176 (1818). Harpalium aureum (Kunth) Cass., Dict. Sci. Nat. 25: 438 (1822). Viguiera chimboensis Hieron., Bot. Jahrb. Syst. 29: 38 (1900). Viguiera lehmannii Hieron., Bot. Jahrb. Syst, 29: 38 (1900). Viguiera aurea (Kunth.) Hieron., Bot. Jahrb. Syst. 28: 608 (1901). Viguiera incana (Pers.) S. E Blake, Contr. U.S. Nat. Herb. 26: 252 (1930). Ecuador. Rhysolepis kunthiana (Gardn.) H. Rob. & A. J. Moore, comb. nov. Viguiera kunthiana Gardn., London J. Bot. 7: 399 (1848). Brazil, Goias. 428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON RE

SS QQ \\\ 5M n 4 i AR THGERIN( 2004 Fig. 1. Rhysolepis helianthoides (L. Rich.) A. J. Moore & H. Rob.., A. Habit. B. Head with ray florets removed and involucral bracts not recurved. C. Head showing ray florets. D. Receptacular pale. E. Ray corolla showing lack of style. F Disk floret showing striated achene with pappus of awns and squamellae. G. Disk corolla in section, showing filaments and anthers with small glands on outer surfaces of anther appendages. H. Disk style showing branches with continuous stigmatic area on inner surfaces and apex with hairs but no appendage. Drawn mostly from Bang 44 (US, isotype of Viguiera pazensis Rusby); C. from Buchtien 8579 (US). VOLUME 117, NUMBER 3 Rhysolepis lanceolata (Britton) H. Rob. & A. J. Moore, comb. nov. Viguiera lanceolata Britton, Bull. Torrey Bot. Club 19: 149 (1892). V. mandonii Sch.Bip. ex Rusby, Mem. Tor- rey Bot. Club 3(3): 60 (1893). Helianthus szyszylowiczii Hieron., Bot. Jahrb. Syst. 36: 491 (1905). Bolivia, Peru. Rhysolepis linearifolia (Chod.) H. Rob. & A. J. Moore, comb. nov. Viguiera linearifolia Chod., Bull. Herb. Boiss. ser. 2, 2: 392 (1902). Viguiera trichophylla Dusén, Ark. Bot. 9(15): 30, f. 12 & t. 7. f. 4 (1910). Brazil, Goids, Matto Grosso, Parana; Paraguay. Rhysolepis linearis (Cav.) H. Rob. & A. J. Moore, comb. nov. Helianthus linearia Cav., Icon., 3: 9, t. 218 (1794)[1795]. Helianthus squarrosus Kunth in H.B.K.., Nov. Gen. & Sp., ed. fol. 4: 174, t. 377 (1818). Leighia elegans Cass., Dict. Sci. Nat. 25: A435 (1822). Leighia linearis (Cav.) DC., Prodr. 5: 581 (1836). Viguiera linearis (Cav.) Sch.Bip. ex Hemsl., Biol. Centr-Amer., Bot. 2:178 (1881). Mexico. Rhysolepis macbridei (S. F Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera macbridei S. EF Blake, J. Wash. Acad. Sci. 16: 218 (1926). Peru. Rhysolepis macrocalyx (S. E Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera macrocalyx S. E Blake, Contr. Gray Herb. 54: 171 (1918). Brazil, Minas Gerais. 429 Rhysolepis macropoda (S. F Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera macropoda S. F Blake, Contr. Gray Herb. 54: 128 (1918). Brazil, Minas Gerais. Rhysolepis macrorhiza (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera macrorhiza Baker in Mart., FI. bras. 6(3): 225 (1884). Paraguay. Rhysolepis media (S. E Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera media S. FE Blake, Contr. Gray Herb. 54: 138 (1918). Ecuador. Rhysolepis mollis (Griseb.) H. Rob. & A. J. Moore, comb. nov. Viguiera mollis Griseb., Abh. K6nigl. Ges. Wiss. Gottingen 19: 183 (1874). Helianthus argentinus Saenz, Darwiniana 22: 64 (1979) Argentina. We do not know why Saenz (1979) ex- cluded the species from Viguiera in his treatment, creating the new name Helian- thus argentinus. Panero (1992) was correct in returning the species to Viguiera as then delimited. Rhysolepis nervosa (Gardn.) H. Rob. & A. J. Moore, comb. nov. Viguiera nervosa Gardn., London J. Bot. 7: 403 (1848). Brazil, Goias. Rhysolepis nudibasilaris (S. E Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera nudibasilaris S. FE Blake, Contr. Gray Herb. 54: 149 (1918). Brazil, Minas Gerais. Rhysolepis nudicaulis (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera nudicaulis Baker in Mart., FI. bras. 6(3): 228 (1884). Uruguay. 430 Rhysolepis oblongifolia (Gardn.) H. Rob. & A. J. Moore, comb. nov. Viguiera oblongifolia Gardn., London J. Bot. 7: 402 (1848). Rhysolepis oblongifolia was described from Brazil, Goias. Some more recent col- lections include: Brazil. Matto Grosso: Ser- ra do Roncador, Mun. de Barra do Gargas, 230 km along new road NNE of village of Xavantina, 6.0 km S of Coérrego dos Porcos, 30 km due S of 12°51’S, 51°45’W. ca. 450 m, 26 Nov 1969, Eiten & Eiten 9547 (SP, US); 209 km NNE of Xavantina; 9 Dec 1969; Eiten & Eiten 9818 (SP, US); Minas Gerais. 56 km along road NE of Barrocao, towards Porteirinha, 2400 ft.; 21 Jan 1981, King & Bishop 8585 (MO, US); Brasilandia de Minas, 1 Jun 2001, Soares 32] (BHCB, US); Maranhao, Balsas, approx. 25 km along road west from Balsas to fazenda of Sr. Damiao; 7°40’S, 46°10’W; 4 Dec 1981; Jangoux et al. 1783 (US). Rhysolepis obtusifolia (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera obtusifolia Baker in Mart., FI. bras. 6(3): 226 (1884). Brazil, Goias? Rhysolepis ovatifolia (DC.) H. Rob. & A. J. Moore, comb. nov. Leighia ovatifolia DC., Prodr. 5: 583 (1836). Viguiera ovatifolia (DC.) Baker in Mart., Fl. bras. 6(3): 226 (1884). The type is from Brazil, Sao Paulo. Ad- ditional specimens seen from Parana match the type photograph: Jaguariahyva, ad mar- ginem silvulae, 19 Apr 1910; Dusén 9723 (US)(det. Dusén as Viguiera robusta). Ja- guariahyva opp., in campo, 740 m.s.m, 5 May 1914, G. Jonsson 262a (US)(det. Mal- me as V. robusta). Rhysolepis peruviana (A. Gray) H. Rob. & A. J. Moore, comb. nov. Viguiera peruviana A. Gray, Proc. Amer. Acad. Arts 5: 124 (1861-62). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Viguiera weberbaueri S. FE Blake, Bot. Jahrb. Syst. 54, Beibl. 119: 49 (1916). Peru. Rhysolepis pilicaulis (S. FE Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera pilicaulis S. F Blake, Contr. Gray Herb. 54: 164 (1918). Rhysolepis pilicaulis was described from Paraguay. A recent collection has been seen from Brazil: Matto Grosso do Sul. Rod. BR-267, proximo do km 447, descida da chapada (Mun. Guia Lopes de Laguna, 9 Mar 2003, G. & H. Hatschbach & Barbosa 74393 (MBM, US). The Brasilian specimen is most like the Field Museum type photo- graph of the now destroyed, broad-leaved Berlin specimen. The inflorescence is char- acteristically rather profusely branched with short peduncles, and there are only 8 or 9 short, slender rays while Blake cited 10 to 11. The species has antrorse prorulosity in- side the disk corolla throat. Rhysolepis pilosa (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera pilosa Baker in Mart., Fl. bras. 6(3): 223 (1884). Viguiera malmei S. F Blake, Contr. Gray Herb. 54: 151 (1918). Viguiera misionensis Saenz, Darwiniana 2 O2 (D7). Viguiera misionensis of northern Argen- tina shows no obvious differences from R. pilosa from southern Brazil in Parana, Rio Grande do Sul, Santa Catarina. Rhysolepis pusilla (A. Gray) H. Rob. & A. J. Moore, comb. nov. Tithonia pusilla A. Gray, Proc. Amer. Acad. Arts 5: 124 (1861-62). Viguiera pusilla (A. Gray) S. E Blake, Contr. Gray Herb. 54: 160 (1918). Peru. VOLUME 117, NUMBER 3 Rhysolepis radula (Baker) H. Rob. & A. J. Moore, comb. nov. Viguiera radula Baker in Mart., Fl. bras. 6(3): 223 (1884). Brazil, Minas Gerais. Rhysolepis retroflexa (S. E Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera retroflexa S. F Blake, Contr. Gray Herb. 54: 146 (1918). Bolivia. Rhysolepis revoluta (Meyen) H. Rob. & A. J. Moore, comb. nov. Helianthus revolutus Meyen, Reise Erde 1: 311 (1834). Helianthus lanceolatus Meyen, Reise Erde 1: 311 (1834), not V. lanceolata Britton Flourensia corymbosa DC., Prodr. 5: 592 (1836). Viguiera poeppigii A. Gray, Proc. Amer. Acad. Arts 19: 6 (1883). Viguiera corymbosa (DC.) S. E Blake, Proc. Amer. Acad. Arts 49: 349 (1913). Viguiera revoluta (Meyen) S. E Blake, Contr. Gray Herb. 54: 121 (1918). Ar- gentina, Chile. Rhysolepis robusta (Gardn.) H. Rob. & A. J. Moore, comb. nov. Viguiera robusta Gardn., London J. Bot. 7: 403 (1848). Brazil, Goias. Rhysolepis rojasii (S. E Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera rojasii S. KF Blake, Contr. Gray Herb. 54: 179 (1918). Paraguay. Rhysolepis salicifolia (Hassl.) H. Rob. & A. J. Moore, comb. nov. Viguiera salicifolia Hassl., Repert. Spec. Nov. Regni Veg. 14: 274 (1916). Viguiera villaricensis S. FE Blake, Contr. Gray Herb. 54: 152 (1918). Argentina, Paraguay. 431 Rhysolepis simsioides (S. EF Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera simsioides S. F Blake, Bot. Jahrb. Syst. 54. Beibl. 119: 48 (1916). Peru. Rhysolepis sodiroi (Hieron.) H. Rob. & A. J. Moore. comb. nov. Helianthus sodiroi Hieron., Bot. Jahrb. Syst. 29: 41 (1900). Viguiera sodiroi (Hieron.) S. FE Blake, Contr. Gray Herb. 54: 139 (1918). Ec- uador. Rhysolepis speciosa (Hassl.) H. Rob. & A. J. Moore, comb. nov. Viguiera speciosa Hassl., Repert. Spec. Nov. Regni Veg. 14: 272 (1916). Viguiera simulans S. F Blake, Contr. Gray Herb. 54: 127 (1918). Rhysolepis speciosa has been known from Paraguay; Brazil, Matto Grosso. It also occurs in the Distrito Federal with specimens previously identified as Viguiera squalida as follows: Peunsula Norto, 1000 m, s. d., Valério de Carvalho dos grupos II (UB, US); Reserva Ecolégia do IBGE, 7 Nov 1977, Heringer et al. 249 (IBGE, US); Area do Cristo Redentor: 15°57'07"S, 47°53'37'"W, 19 Oct 1988, Azevedo 180 (IBGE, US); Reserva Ecologica do IBGE, Campo Limpo; 21 Aug 1990, Silva et al. 1009 (IBGE, US); Cristo Redentor, 10 Oct 1990, Brochado 70 (IBGE, US); Tampao das parcelas de campo sujo do Projeto Fogo—IBGE, 9 Dec 1991, Landim de Sou- za 83 (IBGE, US); Ecolégica do IBGE, 15°56’41"S, 47°53'07’W, 7 Nov 1994, Apa- recida da Silva 2457 (IBGE, US). Rhysolepis squalida (S. Moore) H. Rob. & A. J. Moore, comb. nov. Viguiera squalida S. Moore, J. Bot. 42: 37 (1904). Brazil, Goids, Matto Grosso, Matto Grosso do Sul. 432 Rhysolepis subdentata (S. FE Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera subdentata S. FE Blake, Contr. Gray Herb. 54: 131 (1918). Brazil, Minas Gerais. Rhysolepis tenuifolia (Gardn.) H. Rob. & A. J. Moore, comb. nov. Viguiera tenuifolia Gardn., London J. Bot. 7: 400 (1848). Brazil, Minas Gerais. Rhysolepis tuberculata (S. FE Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera tuberculata S. F Blake, Contr. Gray Herb. 54: 151 (1918). Brazil, Minas Gerais. Rhysolepis tuberosa (Griseb.) H. Rob. & A. J. Moore, comb. nov. Viguiera tuberosa Griseb., Abh. KO6nigl. Ges. Wiss. Gottingen 24: 192 (1879). Ar- gentina; Brazil, Rio Grande do Sul, Uru- guay. Rhysolepis tucumanensis (Hook. & Arn.) H. Rob. & A. J. Moore, comb. nov. Leighia tucumanensis Hook. & Arn., J. Bot. (Hooker) 3: 314 (1841) Viguiera stenophylla (Hook. & Arn.) Gri- seb. var. discoidea Griseb., Abh. Konig]. Ges. Wiss. Gottingen 24: 193 (1879). Viguiera discoidea (Griseb.) S. E Blake, Contr. Gray Herb. 54: 157 (1918). Viguiera oligodonta S. FE Blake, Contr. Gray Herb. 54: 146 (1918). Argentina. Rhysolepis weddellii (Sch.Bip. ex S. FE Blake) H. Rob. & A. J. Moore, comb. nov. Viguiera weddellii Sch.Bip. ex S. FE Blake, Contr. Gray Herb. 54: 126 (1918). Boliv- ia; Brazil, Goias-Matto Grosso. In addition to the species listed above we include the following seven previously un- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON described species. Of the new species, the ones from Bolivia and Santa Catarina, Bra- zil, seem to fit Blake’s series Aureae; whereas the others fit his series Bracteatae. One character seen in the new species seems to partially reenforce the distinction. All of the new members of the Bracteatae except R. subtruncata have bands of pro- rulose cells on the inner surface of the disk corolla throats, midway between the veins and often also along the veins. Prorulosity is the condition where elongate cells have the upper ends projecting as papillae. The two new species in the Aureae lack such prorulose bands. The specimens were de- scribed partially from dissections of florets mounted in Hoyer’s solution (Anderson 1954). Rhysolepis dillonorum A. J. Moore & H. Rob., sp. nov. Fig. 2 Type: Peru. Arequipa: Prov. Caraveli, Lomas of Atiquipa, ca. 10.5 km N of turn- off to Atiquipa, 584 km S of Lima; ca. 150—200 m, 1 Nov 1983, M. O. Dillon & D. Dillon 3775 (holotype US; isotype F). E speciebus omniibus in habitis fruticosis et in indumento appresse strigulosis et in bracteis involucri plerumque obtusis dis- tincta. Shrub to 1 m high, moderately and alter- nately branched at 30—45° angles; roots not seen; stem tan to dark brown, closely ap- pressed-strigulose, glabrescent with age. Leaves usually opposite in middle of branches, alternate at base of branches and distally, not decrescent except near heads; petioles none to 0.2 cm long, bases some- times continuous across node; blades ovate to oblong-ovate, 1.5—4.2 cm long, 0.7—2.2 cm wide, base broadly acute to rounded, margins entire, apex acute, both surfaces densely appressed-strigulose, abaxially with scattered glandular dots, triplinervate from near base, secondary veins reaching distal ¥Y%. Heads borne singly on long branches or with 2 or 3 heads on short branches; brac- VOLUME 117, NUMBER 3 UNITED STATES 3026289 NATIONAL HERBARIUM Fig. 2. Rhysolepis dillonorum A. J. Rhyselepis Aillenorum AdMoorer jit Holotype £ QAO 0073069; PLANTS OF PERU Field Museum of Natural History DEPTO: AREQUIPA PROV: CARAVELI ASTERACEAE Viguiera pazensis [Rusby Lomas of Atiquipa, ca. 10.5 km N of turn-off to Atiquipa. KM 584 S of Lima. ca. 150-200 m. % Erect shrub to 1 m.; ray & disc florets yellow. 1 Nov 1983 M. 0. Dillon & D, Dillon 3779 Planis collected under the sponsorship of the National Geographic Society (Grant No. 2706-83] Distributed by Field Museum of Natural History Moore & H. Rob., holotype, Dillon & Dillon 3775 (US). 434 teoles decrescent, oblong, 1.1—0.4 cm long; peduncles 3.5—17 cm long, 0.5—5.0 cm from last bracteole, appressed-strigulose. Invo- lucre 0.4—0.6 cm high, 1.2—1.5 cm diam.; bracts 2—3-seriate, obovate, gradate, 4—8 mm long, 2—4 mm wide, 3—5-nerved, tips obtuse to short-acuminate, indurate in prox- imal % to %, distally herbaceous, abaxially and adaxially appressed-strigulose especial- ly on tips; paleae obovate, indurate, ca. 7.5 mm long, ca. 1.0—1.5 mm wide, scabridu- lous on tip, apex short-acute. Ray florets 13-14; corollas yellow, tube ca. 1.2 mm long, sparsely scabridulous; limb oblong-el- liptical, 1.5 cm long, 0.5—0.8 cm wide, sparsely scabridulous abaxially, apex 3- lobed. Disk florets at least 50; corolla yel- low, ca. 5 mm long, tube | mm long, sca- bridulous; throat 3 mm long, slightly cam- panulate at base, scabridulous proximally, glabrous distally, with vertical bands of an- trorsely prorulose cells inside, lobes 1 mm long, glabrous outside, papillose inside es- pecially near margins; anther thecae 2 mm long; appendages yellow, 0.6—0.75 mm long, ca. 0.45 mm wide. Ray achenes ca. 3.5 mm long, ca. 0.7 mm wide, sericeous on margins, with pappus crown ca. 0.1 mm high. Disk achenes 3.5 mm long, 0.9 mm wide, sericeous with setulae over whole surface; pappus awns 2.0—2.2 mm long, fimbriate-margined, squamellae ca. 4, 1.0— 1.2 mm long, ca. 1 mm wide, margins fim- briate. Pollen 30—33 ym in diam. in Hoy- er’s solution. Rhysolepis dillonorum is_ presently known only from the type collection. The specimen was earlier identified as Viguiera Pazensis; it was seen as distinct in a recent review of the latter by the senior author of the new species. The low elevation at 150— 200 m near the coast, the shrubby habit, the appressed minute hairs on the stems, leaves, and involucral bracts, and the blunt tips of the involucral bracts are all distinctive. The supposed Andean relatives are found at 2000 m or above, are more herbaceous, have longer, mostly spreading hairs, and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON have more lanceolate, subequal involucral bracts. The blunt involucral bracts and the ver- tical bands of prorulose cells inside the disk corolla throat seem to relate the new species to members of Blake’s section Bracteatae that are most common in Brazil and distin- guish the species from the section Aureae to which V. pazensis ( = R. helianthoides) belongs. Opposite leaves are common on the specimen, but the branching is alternate, and the basal nodes of the branches have alternate leaves. Rhysolepis emaciata H. Rob. & A. J. Moore, sp. nov. Fig. 3 Type: Bolivia. Cochabamba: 10 NE; 2465 m; Campero, pajonal de Elyonurus tripsacoides, 2 May 1999, Antezana 1276 (holotype US, isotype MO). E speciebus aliis boliviensis in seriebus Aureis in ramis nullis in foliis dense spir- aliter insertis et in bracteis involucri ca. tris- eriatis gradatis differt. Slender subshrub or shrub 0.4—0.6 m tall, apparently unbranched above base; roots not seen; stems reddish-brown, densely his- pid with long hairs. Leaves rather densely spirally inserted, sessile; laminae herba- ceous, lanceolate, 1.5—2.8 cm long, 0.5—0.8 cm wide, base rounded, margins often with single blunt tooth near basal 4, subentire to remotely undulate distally, apex acute, mu- cronulate, adaxial surface densely scabrous with slender hairs, abaxially densely villous with white hairs and densely gland-dotted; triplinerved from near base, reaching to % leaf length. Inflorescence example seen with single terminal head, with leaves on 7 cm below head becoming smaller, upper- most bractlike; peduncle 2.5 cm long from last foliiform bract, densely villous. Head with involucre 1 cm high, 2 cm wide; bracts ca. 3-seriate, oblong-lanceolate, gradate, 6— 10 mm long, 1.5—2.0 mm wide, appearing herbaceous throughout, villous with white hairs abaxially, without distinct cilia on VOLUME 117, NUMBER 3 Fig. 3. UNITED STATES 3376064 NATIONAL HERBARIUM I@-"7 ays aoe ya HOM 00730693 BRS Rhysalepis emaciata HRbsAd. Hooxe- Heletype ; BOLIVIA ASTERACEAE Viguiera Cochabamba: Campero Pajonal de Elyonurus tripsacoides, 10 NE. Arbusto lenoso, inflorescencia amarilla. 2465 m 02-05-1999 Antezana, C. 1276 MISSOURI BOTANICAL GARDEN HERBARIUM (MO) Rhysolepis emaciata H. Rob. & A. J. Moore, holotype, Antezana 1276 (US). 435 436 margins distally, tips acute to slightly mu- cronulate, erect on inner bracts, shortly re- curved on other bracts, scabridulous on both surfaces; paleae yellowish-tan, indu- rate, oblong, 7.5 mm long, ca. 2.5 mm wide, tip minutely hispidulous, acute to, sometimes, trifid. Ray florets 17—18; corol- las yellow, tube 1.5 mm long, hispidulous, limb oblong, ca. 1.0—1.1 cm long, 0.4 cm wide, abaxial surface strigose and gland- dotted, apex 2- or 3-lobed. Disk florets 50 or more; corollas darker yellow, ca. 6 mm long, tube ca. 1 mm long, scabridulous, throat ca. 4 mm long, campanulate and sca- bridulous at base, smooth inside, lobes 1.0— 1.2 mm long, strigulose outside, papillose inside; anther thecae 2.5—3.0 mm long, with slender basal hastation much longer than collar, essentially short-tailed; appendage 0.58—0.70 mm long, 0.35—0.41 mm wide. Achenes ca. 2.5—3.0 mm long, 0.8 mm wide, sparsely strigulose with stiff setulae; pappus color a very light tan, awns 3.0—3.5 mm long, with fimbriate margins, squamel- lae ca. 5, 1.0-1.5 mm long, 0.2—0.5 mm wide, deeply fimbriate. Pollen grains 25—28 jum in diam. in Hoyer’s solution. Rhysolepis emaciata is known only from the type collection. Relation might be ex- pected to R. australis, R. fusiformis, R. he- lianthoides, and R. lanceolata of the Boli- vian Andes, but the latter all have thicker stems, more obvious branching, and only about 2 series of subequal involucral bracts. The spirally inserted leaves of R. emaciata characteristically seem to contract slightly in width near the basal fourth. The bases of the anthers seem unusually long and tailed for a member of the Heliantheae. Rhysolepis goyasensis H. Rob. & A. J. Moore, sp. nov. Fig. 4 Type: Brazil. Goids: Serra Geral do Pa- rana, ca. 3 km S of Sao Joao da Alianga, near Riacho, ca. 850 m, gallery forest and adjacent cerrado, 15 Mar 1971, /rwin, Har- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ley & G. L. Smith 31821 (holotype US; iso- types NY, UB, US). A R. breviflosculosam in pubescentibus caulis et involucri similis sed in foliis su- perioribus decrescentibus et in laminis base non cordatis differt. Subshrub to 1 m high; usually un- branched between base and inflorescence; part of rhizome seen, roots moderately stout, spreading, without evident fusiform enlargements; stem reddish-brown to tan, pilose to lanulose, denser above, hairs spreading to retrorse. Leaves alternate, re- duced to bracteoles distally, often gradually decrescent; petioles ca. 2 mm long; blades oblong-elliptical, 0.6—6.5 cm long, 0.3—2.1 cm wide, at base broadly acute, margins en- tire, apex short-acute to short-acuminate, adaxially villous with tubercle-based hairs, abaxially villous, triplinervate from near base, secondary veins reaching distal 4 of blade. Inflorescences unbranched or with 1— 3 branches on each side, conic to cylindri- cal when multibranched, elongate branches shorter than main axis, spreading at ca. 45° angles; bracteoles narrowly oblong-ellipti- cal 0.7—3.0 mm long; peduncles 3.5—5.0 cm long, 1-2 mm long from last bracteoles, lanulose as in stems. Heads usually 1—5; in- volucre 0.9—1.1 cm high, 1.2—1.9 cm wide; bracts ca. 3-seriate, oblong to oblong-lan- ceolate, gradate, 10-15 mm long, 3—4 mm wide, 3-nerved, tips abruptly acute to slightly acuminate, slightly indurate at base or herbaceous throughout; paleae rather ob- long, coriaceous, ca. 8 mm long, ca. 2 mm wide, apex short-acute to mucronulate. Ray florets 14-15; corollas yellow; tube 3.5 mm long, pilosulous; limb 1.5 cm long, 0.4 cm wide, pilosulous abaxially on veins, apex 2 or 3 lobed. Disk florets ca. 50; corolla yel- low, ca. 5 mm long, tube 0.7—1.0 mm long, nearly glabrous; throat 2.5—3.0 mm long, slightly campanulate at base, glabrous, in- side with vertical bands of antrorsely pro- rulose cells, lobes 0.7—1.0 mm long, acute, sparsely scabrid outside, papillose inside; anther thecae 2.5 mm long, appendages yel- low, 0.55—0.60 mm long, 0.3—0.4 mm wide. VOLUME 117, NUMBER 3 437 Rhy sole "$ goyasensis He p.zAs,Teore. Neletype: THES NEW YORK BOTANICAL GARDEN Plants of the Planalto do Brasil Extado do Golds No. 37821 Serra Geral do Parana Viguiera quinqueremis Blake (ex descript. of phyllaries) 6 det. John Pruski (NY), 1992 UNITED STATES Erect simple or few-branched subshrub Ga. Im tall. Ligules yellow; disc yellow- brown. Gallery forest and adjacent 3314050 cerrado, ca. 3km S. of Sao Joao da Aliansa, near riacho, ca. 850m. elev. NATIONAL HERBARIUM TT H. S, Irwin, R, M. Harley, G, L, Smith 15 March 4971 Field work condocted with the collaboration of tho Univers(dade do Brasilia and 06 tho Instituto de Pesquisas © Experimentaclo Agricola do Norte, supported In part by funds from the Natlonal Sclence Foundation. Fig. 4. Rhysolepis goyasensis H. Rob. & A. J. Moore, holotype, Irwin, Harley & Smith 31821 (US). 438 Achenes 3 mm long, 0.8—1.0 mm wide, gla- brous except few, small, marginal setulae; pappus awns 2.0—3.1 mm long, minutely scabrid on margins and keel, squamellae ca. 8, 1.1-1.7 mm long, 0.3-0.5 mm wide, margins fimbriate. Pollen 27-30 wm in diam. in Hoyer’s solution. Paratypes: Brazil. Goids: Sao Joao de Al- ianga, estrada para Vaozinho, campo cer- rado, solo rochoso, 9 Feb 1994, G. & M. Hatschbach 60230 & Silva (MBM, US); Corrente (Mun. Sao Joao da Alianga), cam- po cerrado, solo rochoso, 20 Feb 2000, G. & M. Hatschbach & O. S. Ribas 70471 (MBM, US). Rhysolepis goyasensis has pubescence of the stems and involucres reminiscent of that in R. breviflosculosa far to the south in Uru- guay. The new species differs by the leaf blades lacking cordate subamplexicaul ba- ses and by the decrescent size of the distal leaves of the stem. The new species seems related to the R. robusta species group, but has few or single heads or a narrowly conic to cylindrical inflorescence borne well be- yond the larger stem leaves. Rhysolepis hatschbachii H. Rob. & A. J. Moore, sp. nov. Bigs Type: Brazil. Matto Grosso do Sul: Ro- dovia Bonito, Campo dos Indios, proximo de Trés Morros (Mun. Bonito); encosta do morro; solo calcario, 10 Mar 2003, G. & M. Hatschbach & E. Barbosa 74469 (ho- lotype MBM, isotype US). A R. gardneri in formibus capituli similis sed in foliis abaxialiter dense pilosulis et in pedunculis ebracteatis longioribus et in lim- bis radii abaxialiter glabris distincta. Perennial herb or subshrub to 1 m high, with lateral branches ascending at 35—40° angles; roots not seen; stems tan to dark brown, densely hispid to strigose. Leaves of main stems alternate, 4.5—8.5(11) cm long, 1.5—2.8(4.2) cm wide, on branches often opposite, 2.0-5.5 cm long, 0.5—1.5 cm wide; petioles 1-2 mm long; laminae her- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON baceous, oblong-elliptical, base obtuse, margins entire to remotely serrulate, apex short-acute and apiculate, adaxial surface densely pilosulous with bases of hairs often enlarged, abaxial surface densely scabridu- lous on veins, less densely pilosulous be- tween veins, gland-dotted; triplinervate with strongly ascending secondary veins reaching middle or distal % of blade. Inflo- rescence of few heads terminal on stems and branches; peduncles 9-30 cm long without leaves or bracts, strigulose to hispid with white hairs, hairs denser below heads; involucre 10-13 mm high, 13-20 mm wide; bracts broadly 3—4-seriate, slightly unequal, oblong with obtuse to acute tips, 7-11 mm long, ca. 4 mm wide, bases of inner bracts indurate and strongly ribbed, abruptly shortly herbaceous and sometimes recurved at tips, outer bracts canescent with white, densely strigulose pubescence, mar- gins densely fimbriate with short cilia, inner surface usually glabrous, rarely strigulose near tip; paleae oblanceolate, ca. 9 mm long, 1.5 mm wide, acute, essentially gla- brous. Ray florets 9-14; corollas yellow, tube ca. 1 mm long, sparsely pilosulous; limb narrowly elliptical, ca. 1.7—2.4 cm long, 0.5—0.6 cm wide, abaxially glabrous, apex minutely bilobed. Disk florets 35—45 or more; corollas yellow, ca. 5 mm long, tube ca. 1 mm long, scabridulous, throat ca. 3 mm long, sparsely scabridulous on nar- rowly campanulate base, with vertical bands of antrorsely prorulose cells inside, lobes ca. | mm long, nearly glabous abax- ially, papillose inside; anther thecae ca. 2.3 mm long; appendage yellow, 0.6—0.7 mm long, ca. 0.4 mm wide. Sterile ray ovaries with pair of squamellae 0.5—0.9 mm long; disk achenes ca. 4 mm long, ca. 1.2 mm wide; awns 3.0—3.5 mm long, squamellae narrow, 0.5—0.8 mm long, fimbriate. Pollen grains ca. 32 pm in diam. in Hoyer’s so- lution. Paratype: Brazil. Matto Grosso do Sul: Serra de Bodoquena, Fazenda Bodoquena, Reserva da Tercola (Mun. Miranda); Mata, 5-8 m, Sopé de morro, solo argiloso raso, VOLUME 117, NUMBER 3 439 Your ad, ieee EG rys- A ‘a PREFEITURA MUNICIPAL DE CURITIBA | ERBARIO N: MUSEU BOTANICO MUNICIPAL Asteraceae Rodovia Bonito—Campo dos Indios, proximo de Trés Morros (Mun. Bonito) Mato Grosso do Sul UNITED STATES Rhysolepis htchbochi He 6 Haw. G. Hatschbach, M. Hatschbach & E. Barbosa 74469, 10.111.2003 Isofypes Ereta, Im, capitulo amarelo. Encosta do morro; solo calcario. 3441382 era keen WAC 00730695 PMC -DPP 004 Fig. 5. Rhysolepis hatschbachii H. Rob. & A. J. Moore, isotype, G. & M. Hatschbach & Barbosa 74469 (US). 440 encharcado, 17 Mar 1995, A. Pott & al. 7026 (US, MBM). Rhysolepis hatschbachii is known from only the two cited collections. The paratype was previously determined as near R. gard- neri of Goias, and the latter is possibly the closest relative. Differences include the long peduncles of the latter having many foliiform bracts, its involucral bracts being distinctly narrower and pale at the base, and its rays being shorter and puberulous abax- ially. The general habit of the new species is closer to R. ovatifolia of Sao Paulo and Parana, but that species has distinctive nar- rower involucral bracts that are essentially glabrous except for the densely ciliate mar- gins. Rhysolepis laxicymosa H. Rob. & A. J. Moore, sp. nov. Fig. 6 Type: Brazil. Minas Gerais: Serra do Ca- bral, estradad para Francisco Dumont (Mun. Joaquim Felicio); campo rupestre, 950 m, 16 May 2001, G. & M. Hatschbach & Barbosa 72088 (holotype MBM, isotype US). A speciebus novis R. goyasensem similis sed in caulibus non lanulatis et in inflores- centiis laxe cymiformibus et in limbis radii brevibus distinctis. Erect perennial herb or subshrub 50—90 cm high, apparently unbranched between base and inflorescence; roots not seen; stem reddish-tan, pilose to strigose or thinly vil- lous. Leaves alternate; petioles ca. 1 mm long; laminae coriaceous, oblong-elliptical, 3.7—1.5 cm long, 1.6—0.6 cm wide, decres- cent toward inflorescence, base rounded to broadly acute, margins entire or remotely 1—3-subserrulate, apex short-acute and slightly mucronulate, adaxial surface sparsely strigose and densely scabridulous, abaxial surface with prominent veins and prominulous veinlets, densely strigose on veins, strigulous to subsericeous between veins, gland-dotted; triplinerved from near base, secondary veins reaching distal % or PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON more of blade. Inflorescences are sparingly branched, cymiform, branches long, as- cending at ca. 30° angles; with bracteoles mostly at branch bases 1.5—0.7 cm long, 0.6—0.3 cm wide; peduncles 6—23 cm long, strigose, more densely villous near heads, with bracteoles 7-3 mm long, 3-1 mm wide. Heads ca. 5; involucre 0.5—0.6 cm high, 1.0—1.2 cm wide; bracts ca. 3-seriate, oblong, somewhat gradate, 3.0—6.5 mm long, 0.8-1.2 mm wide, tips obtuse to short-acute, outer bracts indurate in basal %, herbaceous in distal 4%, inner bracts almost completely indurate with broad sclerified bands between veins, exposed surfaces densely pilosulous; paleae pale tan, papery, lanceolate to oblong, ca. 7 mm long, ca. 1.5 mm wide, scaberulous at base and tip, gland-dotted at tip. Ray florets ca. 18; co- rollas yellow, tube ca. 1.2 mm long, sca- bridulous; limb broadly oblong, 6.5 mm long, 1.8—3.0 mm wide, puberulous abaxi- ally on veins, apex trilobed. Disk florets 30-35 or more; corollas yellow, 4 mm long, basal tube | mm long, scabridulous, throat ca. 2.5 mm long, base slightly campanulate and scabridulous, with vertical bands of an- trorsely prorulose cells inside, especially midway between veins, lobes ca. 0.7 mm long, pilosulous distally outside, papillose inside; anther thecae ca. 1.8 mm long; ap- pendages yellow, 0.4—0.5 mm long, 0.33— 0.38 mm wide. Achenes ca. 3.5 mm long, ca. 1.1 mm wide, sericeous with slender se- tulae; pappus whitish, awns mostly 2.0—2.5 mm long, fimbriate on margins and midrib; squamellae 5 or 6, ca. 1 mm long, 0.2—0.5 mm wide, margins fimbriate. Pollen grains 22-28 wm in diam. in Hoyer’s solution. Rhysolepis laxicymosa seems mostly closely related to R. goyasensis, but it 1s smaller in all parts. The pubescence of the stem is shorter, the inflorescence is more slender with fewer bracts, the involucre is smaller with narrowly oblong bracts, and the rays are scarcely twice as long as the involucre. In the length of its rays, R. lax- icymosa is closer to R. subtruncata, also of Goias, which has distinctive subtruncate VOLUME 117, NUMBER 3 44] nm oO, = a rex) fsz0 Rhysolepis laxicyreesa Hoh. - AS, Moore ie man Oo PREFEITURA MUNICIPAL DE CURITIBA _ | HERBARIO NS MUSEU BOTANICO MUNICIPAL aX Asteraceae Vequiers oblong ele Crate, re Serra a Cabral, estrada para Francisco Dumont (Mun, Joaquim Felicio) Q\e ayy Minas Gerais UNITED STATES G. Hatschbach, M. Hatschbach & E. Barbosa 72088, 16.V.2001 . Ereta, 50cm, capitulo amarelo. Campo mupestre. Alt.: 950m. 3407290 NATIONAL HERBARIUM PMC - OPP 004 Fig. 6. Rhysolepis laxicymosa H. Rob. & A. J. Moore, isotype, G. & M. Hatschbach & Barbosa 72088 (US). 442 leaves that are not decrescent below the in- florescence. Rhysolepis santacatarinensis H. Rob. & A. J. Moore, sp. nov. Fig. 7 Type: Brazil. Santa Catarina: Serra do Faxinal (Mun. Praia Grande), paredOes ro- chosos, 1200 m, 15 Apr 1993, G. & M. Hatschbach 59135 & J. M. Silva (holotype MBM, 2 isotypes US). A R. pilosam in foliis lanceolatis et brac- teis involucris lanceolatis similis sed in fo- liis distincte petiolatis et in nervis pinnatis et in caulibus densius lanulatis differt. Subshrub or shrub to 1 m high, moder- ately branched; roots not seen; stems tan to reddish-brown, villous, hairs denser near heads. Leaves alternate; petioles 0.2—1.7 cm long, sometimes slightly winged, villous; laminae herbaceous, lanceolate, 4-17 cm long, 0.4—3.5 cm wide, base and apex at- tenuate to acuminate, margins remotely cre- nate-serrulate, adaxially tuberculate-sca- brous, abaxially densely canescent, pilose to subvillous, denser on veins, with glan- dular dots; venation pinnate or essentially pinnate. Inflorescence with | or 2 heads per branch, often overtopped by leaves; pedun- cles 0.2—2.0 cm long. Heads 4—8; involucre 0.75—1.25 cm high, 2—3 cm wide, 3.5 cm wide in fruit; bracts 2—3-seriate, narrowly lanceolate to narrowly oblanceolate, 12—22 mm long, 2-3 mm wide, apices acuminate to mucronulate, tips strongly recurved, bas- al % to % indurate, 5-ribbed, distally her- baceous, abaxially villosulous, adaxially at tip pilosulous to subglabrous, sparsely gland-dotted, margins finely ciliate; paleae oblong, ca. 9-11 mm long, ca. 2 mm wide, indurate, to 7-ribbed, apex acute and mu- cronulate, sometimes with teeth, glabrous with scabridulous midvein. Ray florets ca. 23; corollas yellow, tube ca. 1 mm long, sparsely puberulous; limbs narrowly ellip- tical, 1.5-3.5 cm long, 0.3-0.4 cm wide, apex 1- or 2 -(3-) lobed, abaxially puberu- lous, gland-dotted. Disk florets to 120 or PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON more; corollas yellow, 5—6 mm long, tube 1.5 mm long, glabrous, throat 3.5 mm long, base moderately campanulate, scabridulous on base and veins, smooth inside, lobes 0.5—1.0 mm long, acute, sometimes sparse- ly scabridulous outside, papillose on distal Y% inside; anther thecae 2.5—3.0 mm long; appendage yellow, 0.7—0.8 mm long, 0.3— 0.4 mm wide. Achenes 4 mm long, ca. | mm wide, glabrous except for marginal se- tulae near pappus; awns 2-3 mm long, squamellae separated into broad segments, ca. 0.5 mm long, fimbriate. Pollen 25—28 ym in diam. in Hoyer’s solution. Paratypes: Brazil. Santa Catarina: Mun. Lauro Miller; 20 km west of Lauro Miller, lower and middle slopes of serra by Rio do Rastro, 700-1000 m, 3 Apr 1957, L. B. Smith & R. Klein 12339 (FLOR, US); Rod. SC-438, Serra do Rio do Rastro (Mun. Lau- ro Miiller); Pareddes rochosos; 1000 m, 7 Apr 1991, G. & M. Hatschbach & E. Bar- bosa 55311 (MBM, US). Rhysolepis santacatarinensis would be- long to the series Aureae of Blake (1918) on the basis of its lanceolate involucral bracts, and it would key to various species in the Blake key depending on the emphasis given to the dense canescent pubescence of the abaxial faces of its leaves. Its distribu- tion in southern Brazil and shape of its leaves suggest closest relation to R. pilosa, which has much sparser pilose pubescence, usually no petiole, and much smaller heads. The large heads with 120 or more disk flo- rets distinguish the new species from most other members of the genus in Brazil and elsewhere. The venation of the leaves is also distinctive, lacking strongly ascending lateral veins at the base. The basal second- ary veins are either strictly pinnate or only slightly more ascending. Rhysolepis subtruncata H. Rob. & A. J. Moore, sp. nov. Fig. 8 Type: Brazil. Goias: Chapada dos Vead- eiros, ca. 42 km N of Alto do Paraiso, ca. VOLUME 117, NUMBER 3 443 PREFEITURA MUNICIPAL DE CURITIBA MUSEU BOTANICO MUNICIPAL M. Asteraceae ISOTYPE OF: NC. Viguiers ~ inensis Rb. a AN. Moore Rhysolepis Sanlacaferinensis HRA» 4 \0C: Serra do Faxinal (mune Praia Grande) Santa Catarina L6G. G. & M- Hatschbach 59135 & JeMe Silva, 15-IV-1993 3422582 “Alte: 1200 me . UNITED STATES NATIONAL HERBARIUM IANO PMC - OPP 004 Fig. 7. Rhysolepis santacatarinensis H. Rob. & A. J. Moore, isotype, G. & M. Hatschbach 59135 & Silva (US). 444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON THB NEW YORK BOTANICAL GARDEN Plants of the Planalto do Brasil Estado do Golds Rhysolepi's subtrun cia HReb, 2 No. 33151A Chapada dos Veadeiros Li A,d.Aoore Viguiera sp. Hololype ‘ UNITED STATES Subshrub to ca. 2.5m tall. Ligules yellow; disc yellow-brown. Riacho margin in cerrado. Cerrado on rocky Slopes and 3313597 adjacent campo, ca. k2km N. of Alto do Paraiso, ca. 1250m. elev. NATIONAL HERBARIUM (ho Toatiluto de Derquisas 0 Experimentacio Awrfeola do Norte, supported In H, S, Irwin, R, M. Harley, G, L, Smith 25 March 1971 7 , HON NILALA Field work condveted with tho collaboration of tho Universidade do Drasilia and 4 davon part by funds from tho Natlonal Scteoce Foundation. Fig. 8. Rhysolepis subtruncata H. Rob. & A. J. Moore, holotype, Jrwin, Harley & Smith 33151A (US). VOLUME 117, NUMBER 3 125 m elev., riacho margin in cerrado, on rocky slopes and adjacent campo, 25 Mar 1971, Irwin, Harley & G. L. Smith 33151A (holotype US, isotypes NY, UB). E speciebus aliis in foliis coriaceis saepe subtruncatis et in ramis inflorescentibus longis valde ascendentibus et in floribus ra- diis brevibus differt. Subshrub to 2.5 m high, with few or no branches between base and inflorescence; roots not seen; stems tan to reddish-brown, strigose to stiffly pilose. Leaves alternate, petioles O-1 mm long, 1—2 mm broad, densely villosulous abaxially; laminae co- riaceous, obovate to cuneate, 1.8—4.5 cm long, 0.8—2.4 cm wide, scarcely smaller but more remote up to inflorescence, base cu- neate, margins slightly crenulate-serrulate above, apex subtruncate to scarcely retuse, adaxial surface nearly smooth, hairs stri- gose with enlarged bases, abaxial surface with prominulous veinlets, pilose to thinly sericeous, triplinervate from near base, lat- eral veins reaching distal 4. Inflorescence loosely corymbiform, with 2 or 3 long branches on each side, ascending at ca. 30° angles, pilose; bracts foliiform, mostly % to Y% as large as leaves, mostly at bases of branches, with few bracteoles on distal branches; peduncles 0.4—2.0 cm long be- yond bracteoles. Heads ca. 9; involucre 0.8 cm high, ca. 1.5 cm wide; bracts ca. 2-se- riate, lanceolate, 5-8 mm long, 1-2 mm wide, acute to slightly acuminate, basal 4% to % indurate, apices herbaceous, appressed to slightly spreading, abaxially puberulous, adaxially at tip pilosulous; paleae rather ob- long, obtuse, ca. 5.5 mm long, ca. 1.5 mm wide, indurate, glabrous or with midvein strigulose. Ray florets ca. 20; corollas yel- low, tube ca. 1.2 mm long, pilosulous; limb broadly oblong, 5—6 mm long, 3.5—4.0 mm wide, apex unlobed or 2-lobed, abaxially pilosulous mostly on veins, Disk florets ca. 50?; corollas yellow-brown, 4 mm long; tube 0.8 mm long, sparsely scabrid, throat 2.5 mm long, base scabrid, narrowly cam- panulate, glabrous distally, smooth inside, lobes deltate, ca. 1 mm long, scabrid out- 445 side; anther thecae 1.8—2.0 mm long; ap- pendage yellow, 0.35—0.40 mm long, 0.45— 0.55 mm wide. Achenes (immature) 2.5 mm long, 0.8—1.0 mm wide, setulae over whole surface, sericeous; pappus awns Ca. 1.5 mm long, squamellae ca. 0.5 mm long, deeply fimbriate. Pollen grains 22—26 wm in diam. in Hoyer’s solution. Rhysolepis subtruncata has distinctive cuneate, coriaceous leaves and ascending branches of the inflorescence reaching the level of the terminal central head. The rays are very short compared to many other spe- cies of the genus. The leaves below the in- florescence are not or are scarcely decres- cent. The throats of the disk corollas lack the vertical bands of prorulose cells found in other species of sect. Bracteatae. Any additional collections should be readily identifiable by the leaf shape and by the overall habit of the leafy plants and inflo- rescence. Acknowledgments A. J. Moore was supported in this study by the National Science Foundation Re- search Experience for Undergraduates pro- gram Award Number DBI-0243512. The extensive technical help of Marjorie Know- les is also acknowledged. The drawing of Rhysolepis helianthoides was prepared by Alice Tangerini, staff illustrator, Depart- ment of Botany. Literature Cited Anderson, L. E. 1954. Hoyer’s solution as a rapid mounting medium for bryophytes.—Bryologist 57:242-247. Blake, S. F 1916. Compositae novae imprimis andinae Weberbauerianae.—Bot. Jahrb. Syst. 54:47—51. . 1917. New and noteworthy Compositae, chief- ly Mexican.—Contr. Gray Herb. 52:16—59. . 1918. A revision of the genus Viguiera.— Contr. Gray Herb. n.s. 54:1—205, pl. 1-3. 1924. New American Asteraceae.—Contr. U.S. Nat. Herb. 22(8):—viii, 587-661, pl. 54— 63, 1X=x1. La Duke, J. C. 1982. Revision of Tithonia.—Rhodora 84:453—522. Panero, J. L. 1992. Systematics of Pappobolus (Aster- 446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aceae-Heliantheae).—Syst. Bot. Monogr. 36:1— 195. Robinson, H. 1972. Studies in the Heliantheae I. A new species of Rhysolepis.—Phytologia 24: 209-210. . 1977. Studies in the Heliantheae (Asteraceae). VIII. Notes on genus and species limits in the genus Viguiera.—Phytologia 36:201—215. 1992. New combinations in Elaphandra Strother (Ecliptinae-Heliantheae-Asteraceae).— Phytologia 72:144—151. Saenz, A. A. 1979. El género Viguiera (Compositae) en la Argentina.—Darwiniana 22:45—66. Schilling, E. E., & J. L. Panero. 2002. A revised clas- sification of subtribe Helianthinae (Asteraceae: Heliantheae). I. Basal lineages.—J. Linn. Soc., Bot. 140:65—76. Turner, B. L., & EF Davies. 1980. Stuessya (Asteraceae: Heliantheae), a new genus from southcentral Mexico.—Brittonia 32:209—212. Se 6 Vie i iiign 0 ow ie Pegh=. /aanap foe fatr i le ws yes NP ian @ijde Aelkg fe a een shee aes é Menem, et Mey Weak, # o f a ie” «: meyer ae i ral, 3 Bie tote inns — “ iy el Doane ts oF wi a Hewaiirsee ail ‘ — a ies & (aw an we, = a oe « - 6 wel hee “ we 7 eine -) ibid aa,/%i «a a 28 es hw Gy a x i * oa nm © _—?) ay P artis #6 my bh Oe te fide ee eee ay bad Wecterwast, ‘Uae Tew ee i “ 1 ee = ~ Hoa WRI ¢ tenn ‘ ~ denen ote & —_ fen taser pasty Dw | ” (3 avai uy ‘ e eo 7 @ in a Licongge Tae! Sete | ee ee a | * ig sh afin: Ping eed. F4 wlio Khan, wm re age Vo tsmumabiccoey 1 Wg » oe Gn goin a he hades i aes aera a Le 10 tie Tqyde 2. De Perla” aes : Lip i\ontmmes |, anger “ 7 in toem mijn ey 2-298 208 i 7 : ' INFORMATION FOR CONTRIBUTORS See the Society’s web page— www.biolsocwash.org Content.—The Proceedings of the Biological Society of Washington publishes original research bear- ing on systematics in botany, zoology, and paleontology, and notices of business transacted at Society meetings. Except at the direction of the Council, only manuscripts by Society members will be consid- ered. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in another language when appropriate. Submission of manuscripts.—Manuscripts may be submitted in one of three ways. You may mail three paper copies of the manuscript complete with tables, figure captions, and figures (do not submit original figures unless/until the manuscript is accepted for publication) to the Editor, Dr. Richard C. Banks, MRC-116, National Museum of Natural History, P. O. 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Front cover—from this issue, p. 304. CONTENTS A review of the North American subspecies of the Great Blue Heron (Ardea herodias) Robert W. Dickerman A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Forét des Mikea of southwestern Madagascar Steven M. Goodman and Voahangy Soarimalala Designation of the type species of Musaraneus Pomel, 1848 (Mammalia: Soricomorpha: Soricidae) Neal Woodman The mammals of Palawan Island, Philippines Jacob A. Esselstyn, Peter Widmann, and Lawrence R. Heaney A new species of Tropidonophis (Serpentes: Colubridae: Natricinae) from the D’ Entrecasteaux Islands, Papua New Guinea Fred Kraus and Allen Allison A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras James R. McCranie and Franklin E. Castafieda A new species of Kolpotocheirodon (Teleostei: Characidae: Cheirodontinae: Compsurini) from Bahia, northeastern Brazil, with a new diagnosis of the genus Luiz R. Malabarba, Flavio C. T. Lima, and Stanley H. Weitzman Astyanax biotae, a new species of stream fish from the Rio Paranapanema basin, upper Rio Parana sys- tem, southeastern Brazil (Ostariophysi: Characiformes: Characidae) Ricardo M. C. Castro and Richard P. Vari Tetragonopterus lemniscatus (Characiformes: Characidae), a new species from the Corantijn River basin in Suriname Ricardo C. Benine, Gabriela Zanon Pelicao, and Richard P. Vari Longipalpa saltatrix, anew genus and species of the meiofaunal family Nerillidae (Annelida: Polychaeta) from an anchihaline cave in Bermuda Katrine Worsaae, Wolfgang Sterrer, and Thomas M. Iliffe Neostrengeria lemaitrei, a new species of freshwater crab from Colombia (Crustacea: Decapoda: Pseudothelphusidae), and the vertical distribution of the genus Martha R. Campos A new species of Agostocaris (Caridea: Agostocarididae) from Acklins Island, Bahamas Fernando Alvarez, José Luis Villalobos, and Thomas M. Iliffe A new species of caridean shrimp of the family Stylodactylidae from the eastern Pacific Ocean Mary K. Wicksten and Joel W. Martin A new pedunculate barnacle (Cirripedia: Heteralepadidae) from the Northwest Atlantic L. Buhl-Mortensen and W. A. Newman Two new species of seven-spined Bathyconchoecia from the North Atlantic and Indian oceans (Crustacea: Ostracoda: Halocypridae) Louis S. Kornicker and J. A. Rudjakov The hermaphroditic sea anemone Anthopleura atodai n. sp. (Anthozoa: Actiniaria: Actiniidae) from Japan, with a redescription of A. hermaphroditica Kensuke Yanagi and Marymegan Daly New species and new combinations in Rhysolepis (Heliantheae: Asteraceae) Harold Robinson and Abigail J. Moore SMITHSONIAN INSTITUTION LIB t) 01118 3 OUI 242 251 266 271 303 311 37) 330 339 346 363 368 377 385 398 408 423 G2 H- ISSN 0006-324X BH X NH PROCEEDINGS oF THE BIOLOGICAL SOCIETY or WASHINGTON JAN 1 4 20d SHARES 20 DECEMBER 2004 VOLUME 117 NUMBER 4 THE BIOLOGICAL SOCIETY OF WASHINGTON 2003-2004 Officers President: Roy W. McDiarmid Secretary: Carole C. Baldwin President-elect: W. Ronald Heyer Treasurer: T. Chad Walter Elected Council Michael D. Carleton G. David Johnson Clyde Roper Michael Vecchione Marilyn Schotte Don Wilson Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: Richard C. Banks Associate Editors Plants: Carol Hotton Invertebrates: Stephen L. Gardiner Insects: Wayne N. Mathis Christopher B. Boyko Vertebrates: Gary R. Graves Janet W. Reid Ed Murdy Invertebrate Paleontology: Gale A. Bishop Membership in the Society is open to anyone who wishes to join. There are no prerequisites. 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With this in mind, the Council encourages members to consider making a U.S. tax- exempt contribution to the Society’s Endowment Fund (we are a 501(c)3 tax-exempt organization) and be recognized for doing so in the journal. The categories of contribution to the Endowment Fund are: Contributor ($100—499) Sponsor ($500—999) Benefactor ($1000 and over) Checks should be written to the Biological Society of Washington and sent to: Treasurer, Biological Society Washington National Museum of Natural History, MRC 163 P. O. Box 37012 Washington, D.C. 20013-7012 f : Shy ¢ VF Mil ery" it. e iz Ad ey ™ = 6 oer, yr o nO te Somme tia te oy - ud Voludes Ve od cam bY A ald . (ie lpkreruy raunn an8 i Gop Wilts a r } (repr bide eftay! mg Aine ps fVRe oa ut ave sn F 7 tit ' ; ie \ . 2 mages Stilt . Nae ft raga Oe i bing ih 7 = Cl thee © ruMheviekeee - na ecautathes, » o Sueel af ' Ve a er Kia als itt Hepes Pat Sal eae = Pere Sab palatine er maa: —— edn vinenl Alipied dy rl , rl ie nd ris? > (od vale i 4 a a WOE Opell _T, b eaehio). Sal Givin " ) rhe PY Lee thd j we oon fh | ot wie ut vuoigre eae, © PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):447—487. 2004. Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 5: The genera Plumarella Gray, 1870; Acanthoprimnoa, n. gen.; and Candidella Bayer, 1954 Stephen D. Cairns and Frederick M. Bayer Department of Zoology, National Museum of Natural History, Smithsonian Institution, PO. Box 37012, Washington, D.C. 20013-7012, U.S.A., e-mail: cairns.stephen@nmnh.si.edu Abstract.—The nine western Atlantic species belonging to three genera, Plu- marella, Acanthoprimnoa, and Candidella, are described and illustrated. Four new species of Plumarella are described, as well as one new species of Acan- thoprimnoa; the genus Acanthoprimnoa is also described as new, differentiated from Plumarella by lacking tubercles on the undersurfaces of its sclerites. Two western Pacific species are transferred to Acanthoprimnoa: A. serta and A. cristata. Three varieties are recognized of the common Plumarella pourtalesii, one previously described as a variety (P. p. robusta) and another proposed herein (P. p. var. obtusa). A dichotomous key and table of comparisons is provided for the species and forms of Plumarella, as are a table of comparisons for the two Atlantic species of Acanthoprimnoa, and an indented key to the eleven genera of western Atlantic Primnoidae. Specimens of these genera were found to be extremely common at lower shelf and upper slope depths primarily in the temperate western Atlantic; over 1500 specimens were examined in this study, including types of all included species. This is the fifth in a series of revisions (Cairns 2001; Cairns & Bayer 2002, 2003, 2004) of the western Atlantic deep-water octocorals, and the fourth dealing with the Primnoidae, a family consisting of about 205 species and 32 genera worldwide, of which approximately 33 species and 11 genera occur in the western Atlantic. Bay- er’s revision of western Atlantic Calyptro- Phora (2001) should also be considered as the first unnumbered part of this series, which also deals with primnoids. In order to facilitate identification at the generic lev- el within this family a key is provided be- low for those 11 genera that occur in the western Atlantic. Two genera occur twice in the key since they have both dichoto- mous and pinnate branching. In this part we review the genera Plumarella and Candi- della, as well as describe a new genus, Acanthoprimnoa, separated from Plumar- ella on the basis of its lacking tubercles on the undersurfaces of its sclerites. Specimens of Plumarella and Acanthoprimnoa are ex- tremely common at shelf and upper slope depths (137—1160 m) in the western Atlan- tic, occurring there opportunistically as weeds do on dry land. Ironically, species of these two genera were previously known from only 11 stations and as many speci- mens from the western Atlantic; this report lists approximately 1425 specimens from 145 localities. The third genus, Candidella, is known from deeper water (514—2139 m), and occurs farther north (New England Sea- mounts) as well as in the eastern Atlantic. Indented Key to the 11 Western Atlantic Genera of Primnoidae (fraction indicates number of western Atlantic species/total number of species; genera in bold face treated in this part) I. Colonies unbranched or extremely sparsely branched: Primnoella (4/14) 448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON II. Branching in the form of a bottle- brush: Thouarella (1+/25) Ill. Colonies pinnately branched A. Polyps arranged in whorls: Callo- gorgia (3/28) B. Polyps arranged biserially and al- ternately 1. Tubercles not present on un- dersurfaces of sclerites: Acan- thoprimnoa (2/4) 2. Tubercles present on undersur- faces of sclerites a. Undersurface of opercular scales with a prominent keel: Amphilaphis (1/6) _ b. Undersurface of opercular scales without a longitu- dinal keel: Plumarella (6/ 20) IV. Colonies dichotomously branched A. Polyps arranged in whorls 1. Polyps face outward from branch; 4 marginal scales: Candidella (1/4) 2. Polyps face up or down; 2 marginal scales a. Polyps encased by only two pairs of large abaxial body wall scales i. Members of two pairs of body wall scales inseparably fused to form a complete ring surrounding polyp; polyps face up or down: Calyptrophora (4/13) ii. Body wall scales not fused; polyps face downward: Paraca- lyptrophora (3/6) b. Polyps encased by 3 or 4 pairs of large abaxial body wall scales: Narella (7/25) B. Polyps arranged biserially and al- ternately 1. Tubercles not present on un- dersurfaces of sclerites: Acan- thoprimnoa (2/4) 2. Tubercles present on under- surfaces of sclerites: Plumar- ella (6/20) C. Polyps irregularly arranged: Prim- noa (1/3) Material and Methods This study was based on the examination of approximately 1505 specimens, collected at 161 deep-water stations by 18 research vessels (Appendix: Station data). Except for those reported from the Bibb and Atlantis, which were borrowed from the MCZ, the specimens are deposited at the National Museum of Natural History (USNM). Syn- onymies for all species are purported to be complete for all previously published re- cords. Unprefaced SEM stub numbers per- tain to the series made by Bayer; those pref- aced with C, to the series made by Cairns. The following abbreviations are used: Alb—U.S.EC.S. Albatross; Atl—R/V_ At- lantis; BM—British Museum (now The Natural History Museum, London); C/—R/ V Colombus Iselin; G—R/V Gerda; Gos— R/V Gosnold; H:W—height to maximum width of an opercular or marginal scale; JS—Johnson-Smithsonian Deep-Sea Expe- dition (Caroline); MCZ—Museum of Com- parative Zoology, Harvard, Cambridge; O—M/V and R/V Oregon and Oregon IT; P—R/V Pillsbury; SB—R/V_ Silver Bay; USNM—United States National Museum (now the National Museum of Natural His- tory, Smithsonian, Washington, D.C.). Subclass Octocorallia Order Gorgonacea Suborder Calcaxonia Family Primnoidae Gray, 1858 Genus Plumarella Gray, 1870 Cricogorgia Milne Edwards, 1857:6, pl. B2, fig. 6 (nom. nud.).—Gray, 1870:36— Bik Plumarella Gray, 1870:36.—Studer, 1887: 51.—Wright & Studer, 1889:73-74, 281.—Versluys, 1906:13—14.—Kinoshi- ta, 1908:6—8.—Kiikenthal, 1915:144; 1919:340—342; 1924:255.—Deichmann, 1936:155-156.—Bayer, 1956:F220.— Fabricius & Alderslade, 2001:244—245. VOLUME 117, NUMBER 4 Type species.—Gorgonia penna La- marck, 1815, by subsequent designation (Kiikenthal 1915:144). Diagnosis.—Primnoidae with a well-de- fined operculum; polyps usually inclined apically, each polyp completely surrounded by 8 rows of body wall scales; polyps ar- ranged biserially or irregularly, but never in whorls; 8 marginal scales, often pointed or spinose; undersurfaces of all sclerites tu- berculate, opercular scales not keeled; col- onies uniplanar, usually pinnately (plu- mose) branched but sometimes dichoto- mous. Distribution.—Western Pacific; Patagonia; western Atlantic; 10—1914 m. Remarks.—The only revision of the genus Plumarella was that of Kiikenthal (1919), reiterated in 1924: (Kiikenthal, 1924), which included the description and synonymy of all 17 species as well as a key to their iden- tification. He used the following characters to distinguish species, as emphasized in his key: shape of distal edge of marginal scales, presence of a longitudinal keel on the body wall scales, number of scales in the ab- and adaxial body wall rows, polyp size, and tex- ture of surface of body wall scales. These characters have also been used in this re- view (Table 1), along with the additional characters such as branching mode, termi- nal branchlet length and flexibility, number of polyps/em branch length, shape of the operculars, presence of tubercles on the un- dersides of the sclerites, ornamentation on the edges of the opercular scales, and coarseness of coenenchymal granulation, the last three characters being used to dis- tinguish a closely related new genus once confused with Plumarella. Key to the Species and Forms of the Six Species of Plumarella known from the Western Atlantic 1. Distal edges of marginal scales straight, gently rounded or only slightly angular 1’. Distal edges of marginal scales promi- nently spined 449 2. Branching alternate pinnate; colonies often large (up to 33 cm) ............ 3) 2’. Branching dichotomous; colonies fairly small (less than 11 cm) 3. Body wall scales smooth 3’. Body wall scales granular se An ems Mee a Bn err 5 (P. pourtalesii) 4. Closely-pinnate branching; opercular scales elongate and granular; 10—12 polyps/cm P. pellucida 4’. Loosely-pinnate branching; opercular scales shorter and smooth; 14—21 pol- yps/cm 5. Distal edges of marginal scales of some polyps in a colony slightly angled (but not spinose) ....P. pourtalesii var. obtusa 5’. Distal edges of marginal scales straight to slightly rounded 6. 11-13 polyps/cm; distance between pol- yps on one side of branch 1.0—1.2 mm aati tidtgce atengs. coeceeneea P. pourtalesti var. typical 6’. 14-16 polyps/cm; distance between pol- yps 0.5—0.8 mm Ne iv avec @ P. pourtalesii var. robusta 7. Hach marginal scale with a prominent spine 7’. Only 4—7 marginal scales with an elon- gate needle-shaped spine, those scales corresponding to operculars with at least one uncovered edge ..... P. aculeata ... P. dichotoma RE CEE te ke re es P. laxiramosa PE th Ue oho a ent a ene aed P. aurea Plumarella pourtalesii (Verrill, 1883) Figs. 1[A-B, 2A—C, 3A—C Primnoa Pourtalesii Verrill, 1883:28—29, pl. 2, figs. 2, 2a—e (S. Carolina).—Not Hargitt & Rogers, 1901:281, fig. D (prob- ably A. goesi). Plumarella pourtalesi.—Wright & Studer, 1889:73, 74, 280 (new comb.).—Ver- sluys, 1906:15—16 (comments only). Plumarella pourtalesii.—Versluys, 1906: 342 (key), 345-346 (German translation of Verrill); Ktikenthal, 1919:345—346; 1924:257-258 (same as 1919).—Deich- mann, 1936:156, pl. 25, figs. 17-18, pl. 26, figs. 10, 10a (two new records: Bibb 22, MCZ 4822 and Bibb 135, MCZ 4821).— Bayer, 1954a:281 (listed); 1956: F220, fig. 159-7; 1957:388 (two records, forma obtusa).—Bayer, Grasshoff & Ver- seveldt, 1983: fig. 53.—Bayer, 1973: fig. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 450 —<— ee OI-8 g[suv JYSI[S 10 WYsIeNnS pourfour Anystys “€I-11 “Sr —r'0 610-80 “epusts plooey ‘(wu ¢¢ 0}) SUuOT wo 7 {sSnowlojoysIp pue (ayeIA] 10) aJeuUId ssooT W CLS-8re {VUI[OIVD “S pue “NI yO SOX JOOS !y}OOUIS ‘sosplI Ou ‘yjOoUIS 6 1-81 S/L-9 popunol ‘yoours premdn pouryo “Ul “T7411 -S9'0 “TI-TI dun] ‘ww 0g OF, wd pz ‘oyeuurd -9S00] ‘aJeUIOI[e “Iv[NSoy S-V/9-S sourds Juoq “Iapueys ‘]][e1 / premdn Apysis pours “ul -CI-€I -0S'0 ‘C'1-8'0 IIGIxoy “WW Op OL (Wid QE) aIeuurd 0} (WD +) SNOWOJOYSIq W OOTI—6rs -seuey -eg ‘eployy Jo syie.ns Ysnosy} eurporeD “N Jjo SOX (ugon{sued}) Jejnuvis ‘yjoours ‘SOSPLI OU ‘Ie;NULIH (P9AInd) 7 7—-L'] c/9 popuno. 0} 1YSreNs “yJooUIS premMdn pourlfour -CT—-OT -S9'0 T'1 JJYS -WU 9 OL wid LJ ‘oyeuurd -9SO0[9 ‘o}BUIOIe “Ie[NSay V-e/v-€ sourds pedvys-atpoou “Suo[ (Lh) 9-S ATjensy Je[norpuadied ‘CI-Ol -8'0-L'0 ‘8 I-4'T IFAS sw OG OL Wd Z| ‘o1eIAT pue sjeuurd ‘snowojoys1q mw eyl E81 -Pplopy Jo syieng YSnoIy} eIBI0IDH Jo SOK “rea yeordAy ur sev “IWA yeoidAy ur sev “IwA [eordAy ul se (9[ Sue asnj}qQO) IJv[NSsueity, DISNGOA “I@A Ul Se DISNGOA “XeA Ul SB Wd 6] ‘oyeuuTd -9SO]9 ‘dIVUIO}V “Ie[NSay ISLA) sourds yeursieur ‘uourmoid g Ajjensp Je[norpusdied uayjo ‘CC-¥1 -6'0-8'0 ‘T1-8'0 GIXey Sw OT OL Wd €] ‘snowojoYsIG ul (C8 E81 -PpHojy Jo sens ysnoiy) euljoIeD “N Jyo SOX “IwA [eordAy ur se “eA [eoidAy ut sv “eA [eordAy ul sv “IBA [eordAy ur sev piemdn pouryo “ul -O[—-vI -$9°0 “TI-I'I Jys suru Og OL Wid QE ‘ayeuurd -9SO]9 ‘a}BUIOIe “Ie[Nsoy 9-S/9-S yJOOUIS soyouriq asi] UO Ie;noIpusd pode Oss O10) ele | duny 0) Aitm Suu gp OF wo [I ‘(e1e14]) snowojoysiq Ww C88—-961 seqnyD 0} eUI[OIeD ‘N JJo Sox (soded puvs ouy) Jepnuesis ‘1] -nuvis ‘sospii jeoide MoT 61-CT SV/9-S (pedojjvos) popunor 0} YSIeNs ‘yJooWS premdn pourjour “EI-I1 -9'0-S'0 “7 I-60 IIQIxoy ‘wu ¢¢ OF, wid ¢¢ ‘oyeuuld -9S0]9 ‘a]VUIO}[e “Ie[Nsay [eIxepe/jerxege ‘MOL [[@M Apog ad sapeag S[PUISIEU JO O3po [rIsIq uole} -udtIo ‘uis/sdAjod ‘(urur) Jojowieip ‘jysioy déjog AY -[IqIxo]} pue yjsue] yourig iysioy Auo -[09 WinWIIxRyyy ‘suryourig uOnNGLystG SOHADOS JO Ipissop -un uo juaseid sajoraqn J, yeu -Ayouous0s ‘mq ‘saejnoe -Iedo :uoneyuoureuIO a[eog sie[noiado Jo AA: H [eIxepesjerxege “MOI [JVM Apoqg sad sajeos S[PUISIVUI JO o3poa [eIsIq uonr} -udliIo ‘uio/sdAyjod ‘(urur) Ia}ouIvIp ‘Wystay dAjog Ay -[IQIxoy pue yysusy, yourig jysi1ey Auo -[09 winulxeyy ‘surpourig —_—_———————————————————————————— ‘ds -u ‘vsoup.xv] “qd ‘ds -u ‘ppionjjad ‘gq DSNIGO “eA usajpjinod ‘d (9€6| ‘uURWIYSIOG) DISNGO4 “A Usajpjinod “gq (881 T eordA] msajpiinod “gq SS ————————————____..____...u.._ ‘soneevA pur satseds vouulidoyjupoy pur vjJaiDUN]g MUL|W UIO\SAaM JO sUOSLIedUIOD JO aIGeI— | 21qR.1, VOLUME 117, NUMBER 4 Table 1.—Continued. A. goesi (Aurivillius, 1931) P. aurea (Deichmann, 1936) A. pectinata, n. sp. P. aculeata, n. sp: P. dichotoma, n. sp. 1.6—2.2 Ridged; granular; granu- lar 1.5—2.0 1.6—2.2 1.8-3.9 1.7-2.3 H: W of operculars Ridged; spinose; spinose Smooth; smooth; smooth All low granular (body Granular; smooth; granu- lar Ornamentation of opercu- lars; bw; coenenchymal scales wall sclerites may also be smooth) Yes No No Yes Yes Tubercles present on un- dersides of sclerites Distribution Off northeastern Yucatan Bahamas (northern Straits Straits of Florida to Yu- off S. Carolina to Florida; off S. Carolina to Cuba; 494-1065 m Peninsula (164476 m); Lesser Antilles, Straits of Florida (614-686 m) catan, Bahamas, Puerto Rico, Virgin Islands; 137-595 m of Florida); 400-900 m 310-878 m 451 16.—Bayer & Cairns (Verrill), 2004: pl. 26, fig. 1. Plumarella pourtalesii var. robusta Deich- mann, 1936:156—157, pl. 25, figs. 14—16, pl. 26, fig. 9. Material examined.—Typical form: Alb- 2416, 10 branches, USNM 10531; Alb- 2662, 4 branches, USNM 14609; Alb-2663, 10 branches, USNM 14479; Alb-2667, 4 branches, USNM 49425; Alb-2668, 11 col- onies, USNM 1019275; Alb-2669, over 50 colonies, USNM 14475; Atl-266-2, 6 colo- nies, USNM 1019276; Atl-266-7, 3 colo- nies, USNM 1021650; Ati-266-41, 7 colo- nies, USNM 1019277; Bibb 22, 2 branches, MCZ 4822 (reported by Deichmann 1936); Bibb 135, 2 branches, MCZ 4821 (reported by Deichmann 1936); Cape Hatteras SA6- 5, 13 colonies, USNM 79782 (topotypic); CI-123, 9 colonies, USNM 1019278; CI- 246, 26 branches, USNM 59491; Clelia 78, 1 colony, USNM 93910; Clelia 79A, 1 col- ony, USNM 93909; Combat 174, 1 colony, USNM 50799; Eastward 26017, 14 branch- es, USNM 59490; Eastward 26022, 2 branches, USNM 1019279; Eastward 26023, 10 branches (dry), USNM 1019280; Eastward 26028, 2 branches, USNM 1019281; Eastward 26052, 2 branches, USNM 1021652; G-170, 1 colony, USNM 1019305; G-177, 12 colonies, USNM 1019282; G-235, 1 branch, USNM 1019283; G-386, 3 colonies and SEM stub 250, USNM 53010; G-598, 1 colony, USNM 1019284; G-672, 1 colony, USNM 59498; G-785, 3 colonies, USNM 52994; Gos-2344, 10 branches, USNM 58448; Gos-2385, 10 colonies, USNM 56895; Gos- 2387, 1 colony, USNM 57308; Gos-2413, 8 colonies, USNM 1019285; Gos-2414, 17 colonies, USNM 1019286; Gos-2461, 2 branches, USNM 1019287; O-1343, 3 branches, USNM 50183; O-1349, 3 branch- es, USNM 50443 (Bayer 1958); O-11703, 2 colonies, USNM 59501; O-11717, 1 branch, USNM 59507; O-11725, 5 colo- nies, USNM 1019288; P-105, 2 branches, USNM 52999; SB-453, 2 branches, USNM PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aN 2416: C, P. aurea, 2662; B, P. pourtalesii var. robusta, Alb Alb- Alb 1 cm. 2354. All scale bars Acanthoprimnoa pectinata, D, > 452 aaa ae a (ANS . SZ : CS RiNsayg sNtraeS, ONG ot SA) Z A, Plumarella pourtalesii typical, Fig. 1. station unknown; ‘WU $77 = sieq gjeos [py ‘ddéjod v jo mat anbifgo oelajs ‘pl E[-9 ‘vsnigo eA nsajpjinod -g ‘q ‘S[eulsiew Sulpunosins pue winjnoiedo Jo MoIA Oara\s ‘gOE IWqWoD ‘DISngo.‘ ‘1eA nsajpjinod ‘q “DQ ‘déjod & Jo MatA onbi[qo oe19}s ‘gg¢E-5 “gq ‘dAjod e Jo MatA onbi[qo Oa19}s “Gps ZOIW ‘odAquas ‘vy :[eordAy msajpjinod nyjaspun)d *q-—V a ea ea 2 5 Z S a = 2 —] ie) > PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vee as Fig. 3. A-—C, Plumarella pourtalesii typical (A-B, G-386; C, syntype, MCZ 5749): A, upper and undersur- faces of 5 opercular scales; B, upper surfaces of 3 body wall scales; C, upper and undersurfaces of 4 coenen- chymal scales. D—E Plumarella pellucida (first opercular scale is from G-859, all other scales from the holotype): D, upper and undersurfaces of 5 opercular scales; E, upper and undersurfaces of 4 body wall scales; K upper and undersurfaces of 4 coenenchymal scales. All scale bars = 0.10 mm. VOLUME 117, NUMBER 4 51260; syntypes (MCZ, USNM, see be- low). Forma robusta: Alb-2416, over 25 colo- nies and | unnumbered SEM stub, USNM 49430, 49433: Alb-2666, 2 colonies, USNM 49422; Alb-2668, 10 branches, USNM 14474; Alb-2669, 8 branches, USNM 1019289; Anton Dohrn 6392, 3 col- onies, USNM 1019290; Atl-266-4, 1 colo- ny, 1 branch, USNM 1019291; Azi-266-40, 15 branches, USNM 1019293; Atl-266-41, 1 branch, USNM 1019292; Cape Hatteras SA6-1, 5 colonies, USNM 79783; CI-140, 1 branch, USNM 59497; Combat 368, 14 colonies and SEM stub 253, USNM 50800; Eastward 26023, 1 dry branch, 4 alcohol branches, USNM 1019294; G-170, 1 branch, USNM 1019295; G-177, over 20 colonies and 1 unnumbered SEM stub, USNM 52995; G-247, 1 colony, USNM 53012; G-261, 3 colonies, USNM 1019296; G-598, 7 colonies, USNM 52997; G-835, 3 branches, USNM 52998; Gos-2413, 5 branches, USNM 1019297; Gos-2461, 8 colonies, USNM 1019298; O-11726, 1 branch, USNM 73756; P-197, 1 colony, USNM 53013; holotype (see below). Forma obtusa: Alb-2416, 1 branch, USNM 1019299; Alb-2668, 8 branches, USNM 1019300; Alvin 77-761, 2 colonies, USNM 1019301; Alvin 1335, 2 colonies, USNM 73741; Atl-3780, 10 dry branches, MCZ 54321; Cape Hatteras SA6, 8 branch- es, USNM 79781; CI-140, 3 branches, USNM 59495; CI-246, 9 branches, USNM 1019302; Eastward 26022, 1 branch, USNM 76986; Eastward 26023, 1 branch, USNM 1019303; G-56, 1 colony and 1 un- numbered SEM stub, USNM 53005; G- 169, 7 colonies, USNM 53008; G-235, 8 colonies and SEM stub 260, USNM 53007; G-241, 9 colonies, USNM 53004; G-246, 2 branches, USNM 53003; G-261, 10 colo- nies, USNM 53009; G-386, 3 branches, USNM 1019304; G-391, 6 colonies and SEM 252, USNM 1019305; G-598, 1 col- ony, USNM 1019306; G-664, 3 branches, USNM 53001; G-679, 4 colonies, USNM 53002; G-1012, 1 colony, USNM 53011; 455 G-1314, 9 colonies and SEM stub 251, USNM = 53006; — Gilliss, 2a OUNE 79°24'36"W, 603 m, 25 May 1973, 1 branch, USNM 79515; Gos-2387, 5 colo- nies, USNM 57306; O-1328, 1 colony, USNM 50528; P-209, 4 branches, USNM 53000. Types and type localities.—TYwo colonies and several fragments of the typical form were mentioned by Verrill (1883) collected from Blake 318, all of which must be con- sidered as syntypes. Deichmann (1936) at- tributed catalog number MCZ 4821 to the syntype series, but this number was preoc- cupied by Deichmann by specimens col- lected from “off Florida’’, thus the syntype series was later re-cataloged as MCZ 42887, and consists of three small branches. Four fragments and SEM stubs 249 and C1084 of one of these syntypes are also de- posited at the USNM (5749). All syntypes are preserved in alcohol. Type Locality: Blake 318: 31°48'50"N, 77°51'50"W (Blake Plateau off South Carolina), 616 m. The holotype of forma robusta, two small branches in alcohol, the largest 9 cm high, is deposited at the MCZ (4823). It also bears Verrill’s personal number of 8032. It is also represented by an unnum- bered SEM stub at the USNM. Type Lo- cality: The type locality was stated to be “18 fms. off Alligator Reef, Florida”’ (Deichmann, 1936:157), but Deichmann correctly queried the extremely shallow depth of the collection. A newer label with this type indicates that it was collected at Bibb station 192 (24°48'05"N, 80°34'45’W: off Alligator Reef), at 216 m, this depth be- ing more consistent with the known bathy- metric range. Diagnosis.—Distal edge of marginal scales rounded (scalloped) or straight; branching close pinnate, branchlets of mod- erate length and stiff; body wall scales granular, surface of operculars ridged; 11— 13 polyps/cm (14—16 for forma robusta and obtusa). Description.—Colonies are flabellate and pinnately branched (plumose), consisting of 456 a main branch, which gives rise to 1—4 pri- mary branches (depending on the size of the colony), each main and primary branch giving rise to numerous stiff branchlets up to 55 mm in length, in an alternating se- quence (alternate pinnate). Although flabel- late, colonies may be uniplanar or slightly convex, such that the polyps are directed toward the convex side. A branchlet occurs on the main branch within 5 mm of the base of the colony; additional branchlets occur at regular intervals of 1.8—3.2 mm on the main and primary branches, producing a characteristic zig-zag pattern for the larger branches, the branchlets being straight and parallel to one another. The main stem is usually anchored by a dense, white, calcar- eous holdfast, which is often attached to the dead corallum of a deep-sea scleractinian. The axis is yellow-brown or gold and lon- gitudinally striate; overall the colony is white in alcohol. The largest colony (Com- bat 174) is 33 cm in height, 13 cm in width, and 2.7 mm in basal diameter, but most col- onies examined were considerably smaller. Polyps are arranged on all branches (1.e., main, primary, and branchlets) biserially in the plane of the flabellum in an alternating fashion, but angled upward such as to pro- duce a 30°—-45° angle with the branch, as well as being slightly curved toward the an- terior (convex) side of the flabellum. Two to six polyps occur on the internodes of the main and primary branches. Polyps are rarely more than | mm in height, slightly wider at the apex (0.5—0.6 mm) than the base, and fairly well spaced: polyps on the same side of a branchlet are separated by 1.0—1.2 mm. In general, 11—13 polyps oc- cur/em of branchlet length, this number sometimes lower in small colonies. Each polyp is encased by 8 opercular and 8 rows of body wall scales. The abaxial rows of body wall scales consist of 5 or 6 scales, the adaxial rows usually one less (4— 5 scales), the latter rows always shorter due to having less scales and being located on the shorter, concave, side of the upturned polyp. The marginal scales are up to 0.37 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mm in width, bearing sparse, low granules (10 wm in diameter) on their exterior sur- face and complex tubercles on undersurface (up to 13 pm in diameter). Body wall scales proximal to the marginals are progressively smaller. Body wall scales overlap one an- other within each row as well as overlap- ping the edges of the scales in both adjacent rows. Body wall scales, including margin- als, are somewhat crescent-shaped, wider than tall (H:W = 0.6-0.7), and slightly curved to accommodate the curvature of the polyp, the distalmost scales in each row (the marginals) having a straight or slightly rounded, finely serrate distal margin, pro- jecting only about 10—20 pm beyond the articulation with the base of the operculars, and, if rounded, producing a slightly scal- loped calyx margin. The 8 opercular scales are similarly-shaped (H:W = 1.5—1.9), symmetrical, isosceles triangles, having an apical angle of 41°-51° but with rounded tips. The abaxial operculars are up to 0.42 mm in height, the adaxials about 0.30 mm in height. The upper surface of the each opercular is covered with small spines on the lower half and 3 or 4 longitudinal ridges apically. The lower two-thirds of the un- dersurface is covered with complex tuber- cles of the same size as those of the body wall scales; the distal region is smooth, not keeled. The lateral edges of each opercular is finely serrate like the distal edges of the body wall scales. The 8 opercular scales fold together forming a moderately tall, conical operculum. Coenenchymal scales occur in one layer, are flat, and elliptical to elongate in shape, the largest scales being 0.40 mm in length. Like the other scales, they are sparsely granular above, tuberculate below, and bear finely serrate edges. Tentacular sclerites were not noted. Comparisons.—See Table 1. Distribution.—Typical form: Blake Pla- teau from North Carolina (34°45'N) through Straits of Florida (insular side) and north central part of Cuba; 196-882 m. For- ma robusta: Blake Plateau from off N. Car- VOLUME 117, NUMBER 4 olina (34°15'N) through Straits of Florida to Florida Keys; 183-877 m. Forma obtusa: Blake Plateau from off Georgia (31°26’N) through Straits of Florida and Northwest Providence Channel; 183—743 m. Remarks.—Deichmann (1936) described a variety of P. pourtalesii called robusta, which differs from the typical form in hay- ing a stouter corallum, thicker sclerites, a flatter operculum, and longitudinal ridges on the opercular scales. This form is well represented in our collection (listed sepa- rately above), and differs from the typical form (Table 1) in having a stouter, stiffer colony, and slightly larger and thicker pol- yps (1.0—1.2 mm tall, 0.65 mm in diameter) that are more closely spaced on the branch (distance between adjacent polyps on one side of a branch 0.5—0.8 mm), such that even though the polyps are slightly larger, there are more/cm, i.e., 14—16, the latter number achieved when polyps also bud from the anterior face of the branch. As Deichmann stated, in general, the opercu- lum is flatter, even concave in some highly contracted polyps, but both forms have lon- gitudinal ridges on the opercular scales. These several differences are fairty consis- tent but do not include any characters rou- tinely used to differentiate species of Plu- marella (see Ktikenthal 1919). Further- more, the distribution and bathymetric range of both taxa are virtually the same, 7 of the 22 records of robusta being from common stations. We thus concur with Deichmann in considering this form as just an environmental variation of the typical form. Another common form of P. pourtalesii, represented in our collection by 27 lots, is otherwise similar to forma robusta but dif- fers in that the marginal scales of at least some polyps of every colony have an an- gled or evei pointed distal edge (Fig. 2D). The angle o the distal edge is often a sharp right angle extending about 0.1 mm, or less commonly may be a discrete spine up to 0.25 mm in length. There is great variation in the expression of this character. In some 457 colonies all marginal scales of all polyps will have a short angled distal edge, where- as in other colonies only those polyps to- ward the distal branch tips will be so mod- ified, the marginal scales of the remaining polyps having a typically straight or round- ed edge. Furthermore, individual polyps may have one or all eight marginal scales with an angled edge. The long-spined mar- ginals are infrequent and when present only occur one to a polyp. All three forms some- times occur at the same station, and their bathymetric and geographic ranges are quite similar. Because of the great variation of this single character that separates this taxon from forma robusta and the typical form, it is considered to be an environmen- tal or genetic variation without taxonomic validity, but, in order to easily refer to this variation, the form name obtusa is applied to it, an allusion to the often obtuse angle formed by the distal edges of its marginal spines. Plumarella pellucida, n. sp. Figs. 3D—-E 4A-—B, 10A Material examined/types and type locali- ty.—Holot pe: G-647, 1 colony and SEM stubs C1079—1080, 1085-1086, USNM 52992. Paratypes: Alb-III 9-19, 1 colony, USNM 50573; Cape Hatteras SA6-5, 2 col- onies, USNM 1019404; CI-266, 3 colonies, USNM 59506; G-647, 2 colonies, 3 branch- es, USNM 1019403; G-808, | colony, USNM 59499; G-859, 3 branches and 1 un- numbered SEM stub, USNM 52993; O- 11705, 5 branches, USNM 1019405. Type Locality: 26°16'N, 79°43’W (Straits of Flor- ida off Fort Lauderdale), 520—549 m. Diagnosis.—Distal edge of marginal scales rounded or straight; branching close- ly pinnate, branchlets of moderate length and stiff; opercular scales granular but not ridged, other scales faintly granular, ap- pearing almost smooth and translucent; ab- axial and outer-lateral opercular scales quite long and curved; 10—12 polyps/cm. Description.—Colonies are flabellate and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 458 B JO MOIA [RIA}R] redAjoyoy DSOWDAIXD] Dj/JAMDUN] J ‘ a “WU QC'O Oo -uu 670 = A d‘d “ V J0J sieq a[R9S “MOIA Avjno1ado Os10}s pur ‘dAjod ev puv youriq —p ‘dAjod v Jo smata svjpnorsdo o910}s pue [e1oje] Oo19}s :adAjojoy ‘vpionjjad pyjasmunjg ‘gq-y ‘B14 VOLUME 117, NUMBER 4 closely pinnately branched, as in P. pour- talesii, the distance between successive branchlets 2.5—5.0 mm, and the entire fla- bellum is usually convexly shaped, the pol- yps curving toward the convex (anterior) face. Branchlets are up to 60 mm long and are fairly stiff. The main stem is anchored by a dense calcareous holdfast. The axis is yellow and faintly longitudinally striate; in alcohol the colony is light brown. The larg- est colony (holotype) is 17 cm in height, 16 cm in width, and 2.2 mm in basal stem di- ameter. Polyps are quite regularly arranged in a biserial, alternating fashion on all branch edges, and are inclined distally. Polyps on the same side of a branchlet are well sep- arated by 1.0—1.2 mm, resulting in 10—12 polyps/cm. Polyps are 1.1—1.2 mm in height and slightly flared distally (0.65 mm in diameter). Each polyp is covered with 8 opercular and 8 rows of body wall scales, the abaxial body wall scales numbering about 6, the ad- axial usually consisting of 5. The body wall scales, including the marginals, are similar in size and shape to those of P. pourtalesii; however, their exterior granulation is much reduced and the scales appear to be thinner, producing a smooth, almost translucent as- pect. Like P. pourtalesii, the distal edges of the marginals are straight to slightly round- ed, never spinose or angled. The 8 oper- cular scales are isosceles triangles (H:W = 1.7—2.4), having an apical angle of 20°-35°, ranging from sharply pointed to slightly rounded. The abaxial and outer-lateral oper- culars measure up to 0.54 mm in height, the adaxial and inner-lateral operculars only 0.37 mm in height. The distal third of the abaxial and outer-lateral operculars are at- tenuate, with slightly serrate edges; these scales are curved downward to follow the curvature of the polyp and almost reach the opposite side of the polyp, considerably overlapping the shorter adaxial and inner- lateral opercular scales. The opercular scales bear low sparse granules, lack distal ridges, and are tuberculate on the undersur- 459 face with no trace of a keel. In general, the opercular scales fold together in a conical operculum in a manner similar to that shown in Fig. 9f, the abaxial opercular hav- ing two exposed edges, the adaxial having none. Coenenchymal scales occur in one im- bricating layer, are flat, and elliptical to ir- regular in shape, the largest being about 0.41 mm in width. Their granulation is re- duced similar to that on the opercular scales. Tentacular scales were not noted. All sclerite types, body wall, opercular and coe- nenchymal bear complex tubercles on their undersurfaces, the largest of which mea- sures about 13 pm in diameter. Etymology.—The species name pellucida (Latin: pellucidus, transparent, translucent, clear) refers to the translucent nature of the body wall and coenenchymal scales when viewed in liquid, probably due to their thin- ness and sparse granulation, which allows a view of the outline of the branch axis and of 8 faint white longitudinal lines in the polyps corresponding to the mesenteries. Comparisons.—Plumarella pellucida be- longs to a closely related species complex characterized by having large, pinnately branched colonies and smooth to straight- edged (not spinose or pointed) marginal scales, this complex consisting of: P. pour- talesii, P. laxiramosa, and P. pellucida. It is probably most closely related to P. pour- talesii, but is distinguished by having non- ridged opercular scales, the adaxial and out- er-laterals of which are quite long and curved; and smooth, almost translucent body wall and coenenchymal scales. It dif- fers from P. laxiramosa in having closer pinnate branching; longer, more attenuate opercular scales that are granular; and fewer polyps/cm. (Table 1). All three species oc- cur in roughly the same geographic and bathymetric range and often occur at the same stations. Distribution.—North Carolina through Straits of Florida, Bahamas; 549-1160 m. 460 Plumarella laxiramosa, 0. sp. Figs. 4C—E, 5A—E, 10B Material examined/types and type local- ity.—Holotype: Cape Hatteras SA6-1, 1 colony and SEM stubs C1081-1083, USNM 1019406. Paratypes: Alb-2416, 7 colonies, over 50 branches and 1 unnum- bered SEM stub, USNM 50594 and 79458; Atl-266-41, 1 branch, USNM_ 1019407; Cape Hatteras SA6-1, 19 branches, USNM 79778; Cape Hatteras SA6-5, 8 colonies, USNM 79779: Combat 368, 1 branch, USNM 1019408; Gos-2387, 1 colony, USNM 1019409. Type Locality: 31°17'18"N, 79°00'39"W (Charleston Bump region, South Carolina), 572—575 m. Diagnosis.—Distal edge of marginal scales rounded or straight; branching loose- ly pinnate, branchlets long and flabby; body wall, opercular, and coenenchymal scales smooth; 14—21 polyps/cm, polyps often oc- curring on anterior face. Description.—Colonies are flabellate and pinnately branched, as in P. pourtalesii, but differ from that species in having more widely-spaced (loose pinnate) and thus few- er branchlets, each branchlet separated form their adjacent by 6-11 mm. Also, unlike P. pourtalesii, this species has flat, not curved, colonies, and its branchlets are longer (up to 80 mm) and less stiff, altogether produc- ing a limp or languid colony tension. The main stem is anchored by a dense, white calcareous holdfast, which often attaches to the corallum of a dead scleractinian or sty- lasterid coral with an encrustation up to 1 cm in diameter. The axis is golden-yellow and faintly longitudinally striate; overall the colony is light brown in alcohol. The ho- lotype is 18 cm in height, 17 cm in width, and has a main stem diameter of 2.7 mm, although the stem is broken from the sub- strate. The largest colony, which has an in- tact base (A/b-2416), is 24 cm in height. Polyps are closely arranged on all branches biserially in the plane of the fla- bellum and in an alternating fashion; how- ever, most larger colonies often have a third PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON row of polyps on the anterior side, which produces a very crowded arrangement of polyps that may number up to 21/cm. Up to 20 polyps may occur on the rather lengthy internodes of the main and primary branches. Polyps are 1.1—1.2 mm in height and are slightly flared distally (0.65 mm). Each polyp is covered with 8 opercular and 8 orderly rows of body wall scales, the abaxial body wall scales numbering 6 or 7/ row, the adaxial consisting of usually only 5. Body wall scales are roughly rectangular and slightly curved to fit around a segment of the polyp; the distalmost body wall scales (the marginals) are 0.30—0.34 mm in width and 0.20—0.25 mm in height, the scales becoming progressively smaller to- ward the branch. The distal edges of the marginals are finely serrate and straight to slightly rounded, never pointed or spinose. The undersurfaces of all scales are tuber- culate (tubercles 12—26 wm in diameter), whereas the upper surfaces of the body wall scales, as well as those of the opercular and coenenchymal scales, are virtually smooth. The 8 opercular scales are similarly-shaped (H:W = 1.75—2.20), symmetrical, isosceles triangles, having a blunt, rounded apical an- gle of about 30°. The abaxial operculars are up to 0.45 mm in height, the adaxials only slightly less tall (e.g., 0.37 mm). As men- tioned above, the surface of the operculars is smooth and without ridges, whereas their undersurface is tuberculate. When contract- ed, the operculars form a closely fitted, overlapping, low operculum. Coenenchymal scales occur in one im- bricating layer, are flat, and elliptical to ir regular in shape, the largest scales being about 0.25 mm in length. Tentacular scales were not noted. Etymology.—The species name laxira- mosa (Latin: laxus, loose, slack, and ra- mosus, branching) refers to the loose branching mode of the colonies as well as the limp tension of the branchlets. Comparisons.—Among those western Atlantic species of Plumarella having smooth-edged marginal scales (Table 1), P. VOLUME 117, NUMBER 4 Fig. 5. A—E, Plumarella laxiramosa, holotype: A, upper and undersurfaces of 4 opercular scales; B, tip of undersurface of an opercular showing fine serration of edge; C, upper and undersurfaces of 3 body wall scales; D, tubercles on underside of a body wall scale; E, upper and undersurfaces of 4 coenenchymal scales. F-I, Plumarella dichotoma, holotype: FE upper and undersurfaces of 4 opercular scales; G, tubercles on underside of an opercular scale; H, upper and undersurfaces of 5 body wall scales; I, upper and undersurfaces of 5 coenen- chymal scales. Scale bars for A, C, E-K H-I = 0.10 mm; B = 25 pm; D, G = 10 pm. 462 laxiramosa is most easily distinguished by its growth form (loose pinnate) and by hav- ing a third, anterior row of branchlet pol- yps, which leads to a high number of pol- yps/cm. It is also distinguished by having no surface granulation on any scales. Distribution.—Off North Carolina (to 34°15'N) and South Carolina; 348—625 m. Plumarella dichotoma, n. sp. Figs. 5F—I, 6A—-C, 10C Material examined/types and type local- ity.—Holotype, Gos-2387, 1 colony and SEM stubs C1087—1089, USNM 57307. Paratypes: Alb-2666, 10 colonies, USNM 49423; Alb-2667, 9 colonies, USNM 49431; Alvin 77-762, 1 colony, USNM 1019427; Alvin 1335, 1 dry colony, USNM 73739; Anton Dohrn 65-32, 3 dry branches, USNM 1019429; Cape Hatteras SA6-5, 5 colonies, USNM 79780; Eastward 26004, 2 colonies, USNM 1019430: Eastward 26023, 1 colony, USNM 1019431; G-169, 2 colonies, USNM 52990; G-170, 1 colony, USNM 52996: Gos-2344, 3 colonies, USNM 58447; Gos-2387, 10 colonies, USNM_ 1019428; Gos-2413, 7 colonies, USNM 1019432; Gos-2469, 1 colony, USNM 59021. Type Locality: 31°14'48”"N, 78°59'W (off South Carolina), 530 m. Diagnosis.—Distal edge of marginal scales straight or rounded; branching di- chotomous, sometimes lyrate, branchlets relatively short and wiry; opercular and body wall scales smooth; 8-10 polyps/cm (polyps widely spaced), standing perpendic- ular to branches on large-diameter branch- lets. Description.—Colonies are flabellate and usually slightly curved, as in P. pourtalesii, but consistently dichotomously branched. The main stem is attached to a substrate by a thin calcareous encrustation and rises only 12-16 mm before it bifurcates, producing an axil angle of 65°—85°. Subsequent equal, dichotomous branching occurs at intervals of 5—12 mm, although some end branches are up to 40 mm in length and are the result PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of 7-11 previous branching nodes. Termi- nal branchlets are wiry to limp in tension. Higher order axil angles are slightly small- er, 1.e., 40°—-45°, undamaged colonies usu- ally being slightly broader than tall. In some large colonies the two outermost branches remain slightly larger than their inner branchlets (unequal dichotomous branching), as in the holotype, producing a lyrate form. The largest colony (the holo- type) is 11 cm in height, 13 cm in width, and has a basal main stem diameter of 1.4 mm, although most colonies examined were considerably smaller. The axis is golden yellow and the colony appears white in al- cohol. As in all species of Plumarella, the pol- yps are biserially arranged in alternating fashion on the edges of all branches, angled slightly toward the anterior side of the fla- bellum, and standing perpendicular to large-diameter branches, but inclined dis- tally in smaller-diameter distal branches. Polyps are widely spaced, adjacent polyps on the same side of a branch separated by as much as 2.0 mm. Polyps are fairly tall and slender, up to 1.3 mm in height and 0.65 mm in apical diameter. The polyps are protected by 8 opercular and 8 rather disorganized rows of body wall scales, both ab- and adaxial rows containing 5 or 6 scales. Body wall scales are smooth, quickly decreasing in size from the margin- al to the more proximal ones. The distal edges of the marginals are straight to slight- ly rounded and the scales themselves are square to slightly rectangular. The opercular scales are modified isosceles triangles (al- most pentagonal), the two long sides of the triangle being parallel for much of the length, only the distal third having an apical angle of 45°—50°, culminating in a blunt tip. Abaxial opercular scales are up to 0.50 mm in height, adaxial, 0.35 mm; the H:W rang- es from 1.7—2.3. Operculars have a granular upper surface and a tuberculate lower sur- face, devoid of a keel. They infold to form an operculum as illustrated in Fig 9f. Coenenchymal scales are mildly granular ‘WW $77) = A-g ‘ww QS) = V JOJ sieq o[eog ‘syeursieur pourds Surpun S puv uInjnoiedo Jo MaIA Oa19}s puv dAjOd vB JO MOIA [vIO}L] OOIO}S (0b-997T-[1V ‘vainv vDjjaspunyg “q—q ‘dAjod Jo MOIA [eJO}e] Od19}S puv ‘MOIA Ie[NOIEdO ‘youRIg B JO MAIA [eIO}e] :odA\O[OY ‘VulojoYyoIp YJJasDUN]d “D-V ‘9 “BIA ~ za ea 2 2) Z is w 2 =) = 2) 2 464 above, tuberculate below, and irregularly elliptical in shape, rarely over 0.25 mm in greater diameter. Etymology.—The species name dichoto- ma (Greek: to be divided into two parts), is an allusion to the dichotomous branching of the colonies. Comparisons.—Among the western At- lantic species of Plumarella, P. dichotoma is unique in having both dichotomous branching and smooth-edged marginal scales (Table 1). It is also distinctive in hav- ing such widely spaced polyps that are of- ten oriented perpendicular to the branches. Distribution.—Off southeast coast of United States from South Carolina to off Dry Tortugas, Florida; 494—1065 m. Plumarella aurea (Deichmann, 1936) Figs. 1C, 6D-E, 7A—D Thouarella aurea Deichmann, 1936:165— 166, pl. 25, figs. 12-13, pl. 26, fig. 11.— Bayer, 1954a:281 (listed). Plumarella pourtalesii.—Deichmann, 1936:156 (Gin part: 2 of 4 specimens from Bibb 22, Bahia Honda). Plumarella aurea.—Bayer, 1981:934, fig. 70 (new combination).—Bayer & Cairns (Verrill), 2004: pl. 25, 6a, pl. 83, la. Material examined.—Alb-2666, 4 colo- nies, USNM 52984; Alb-2667, 5 colonies and | unnumbered SEM stub, USNM 52985; Alb-2668, 4 branches, USNM 1019312; Alvin 77-762, 3 colonies, USNM 1019313; Azl-3780, 15 dry colonies, MCZ; Atl-3782, 1 dry colony, MCZ 54327; Atl- 266-40, 5 colonies and SEM stub 390, USNM 58443; Atl-266-41, 5 colonies, USNM 1019314; Cape Hatteras SA 6-5, 2 colonies, USNM 1019315: Discoverer X, 1 colony and SEM stub 391, USNM 58446; Eastward 26004, 1 colony, USNM 1019316; Eastward 26022, 2 branches, USNM 1019317; Eastward 26023, 4 dry branches and | in alcohol, USNM 1019318; G-672, 1 colony, USNM 52974; G-679, 1 colony, USNM 1019319; G-936, 1 branch, USNM 1019320; Gos-2385, 4 colonies and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF V ASHINGTON SEM stub C1092, USNM 56892; Gos- 2414, 6 colonies, USNM 1019321; O- 11716, 1 colony, USNM 59500; P-105, 3 colonies, USNM 52976; SB-453, 1 colony, USNM 51265; syntypes (see below); spec- imens misidentified as P. pourtalesii by Deichmann (1936) from the type locality, Bibb 22 (MCZ 59442). Types and type locality.—Five small branches (syntypes) preserved in alcohol are deposited at the MCZ (4801), which also bear Verrill’s number 8042. An un- numbered SEM stub of one of these branches is also deposited at the USNM. Type Locality: 24°14’20"N, 80°59’40"W (off Bahia Honda, Straits of Florida off Florida Keys), 310 fathoms (=567 m). AI- though not stated in the original description, a label with the type specimens indicates they were collected at Bibb 22 (dredge 12), made on 4 May 1868. Diagnosis.—Distal edge of most margin- al scales prominently spinose; branching di- chotomous; opercular, body wall, and coe- nenchymal scales smooth; polyps crowded, sometimes on anterior face, 14—22 polyps/ cm. Description.—Colonies are flabellate and dichotomously branched. The main stem is attached to the substrate by a thin calcare- ous expansion and rises only 5—8 mm be- fore it bifurcates, producing an axil angle of about 55°; subsequent axial angles are 40°—45°. Branching is usually equal and di- chotomous, occurring at intervals of 5—10 mm, but some terminal branches are up to 10 cm in length. Branches and colonies are quite flexible in tension, almost limp. The largest colony examined (Gos-2385) is 13 cm in height, 12 cm in width, and has a main stem diameter of 1.5 mm. The axis is golden-yellow and the colony appears white in alcohol. Polyps are crowded, occurring biserially in alternating or opposite fashion on the branchlets and often with occasional polyps on the anterior side, resulting in 14—22 pol- yps/cm. Polyps are oriented perpendicular to the branches or tilted only slightly ante- VOLUME 117, NUMBER 4 465 Fig. 7. A—D, Plumarella aurea, Gos-2385: A, upper and undersurfaces of 3 opercular scales; B, upper and undersurfaces of 2 marginal scales; C, upper and undersurfaces of 4 body wall scales; D, upper and undersurfaces of 4 coenenchymal scales. E-I, Plumarella aculeata, paratype from G-252: E, upper and undersurfaces of 2 opercular scales; EK upper and undersurfiaces of 2 marginal scales; G, spination on marginal spine; H, upper and undersurfaces of 5 body wall scales; L. upper and undersurfaces of 5 coenenchymal scales. Scale bars for A—F H-I = 0:10 mm; G = 25 pm. 466 riorly. They are usually squat, cylindrical, and robust, 0.8—1.2 mm in height (depend- ing on contraction) but always 0.75—0.80 mm in apical diameter. Each polyp is protected by 8 opercular and 8 well-defined rows of body wall scales, the abaxial rows having 6 or 7 scales, the adaxial, 5 or 6. The distal edges of the marginal body wall scales are usually strongly spinose, the 8 tooth-like spines forming a small crown encircling the oper- culum and often rising above it. Occasion- ally 1 or 2 of the marginals of a polyp lack spines or have reduced spines, but most polyps have 8 prominent, equal-sized spines. The marginal spines are sharp (api- cal angle 20°—25°), often constituting half the height of the marginal scale, a large spine being up to 0.25 mm in length and 0.08 mm in basal diameter, contributing to a H:W for this kind of scale of up to 1.3— 1.5, the low value due to the wide base of the marginal scales. The marginal spines are circular in cross section and have finely serrate edges where they join the lower rectangular section of the scale (Fig. 7B). Opercular scales are fairly flat (not curved) and isosceles triangular in shape, the distal point being somewhat rounded, forming an angle of 33°—45°. Abaxial opercular scales are up to 0.55 mm in height, adaxial only 0.30 mm; the H:W ranges from 1.5—2.0. The upper surfaces of the body wall and opercular scales are smooth, the undersur- faces covered with complex tubercles that are up to 15—16 pm in diameter. Coenenchymal scales are also smooth above, tuberculate below, and irregularly elliptical, elongate, or circular in shape; and up to 0.40 mm in greater diameter. Comparisons.—Among the western At- lantic Plumarella having spinose marginal scales, P. aurea is most similar to P. acu- leata (see that description and Table 1). Distribution.—Blake Plateau from off South Carolina (32°10'N) through Straits of Florida to off Bahia Honda; Northwest Providence Channel, Bahamas; 310—878 m. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Plumarella aculeata, n. sp. Figs. 7E-I, 8A—B, 9a—g, 10E Material examined/types and type local- ity.—Holotype: G-707, USNM 52980, 1 di- chotomous colony and SEM 248. Para- types: Cape Florida X, 11 pinnate colonies, USNM 1019533; Eastward 26535, 15 pin- nate branches, USNM 1019534; Eastward 26547, 2 dichotomous colonies, USNM 1019535; G-241, 1 pinnate branch, USNM 1019536; G-252, 2 pinnate branches and SEM stubs 255 and C1090, USNM 52979; G-633, 2 pinnate colonies, USNM 52983; G-692, 8 pinnate colonies, USNM 52986; G-695, 1 dichotomous colony, USNM 52978; G-707, 5 dichotomous and 1 lyrate colonies, USNM 52981-52982; G-1125, 3 pinnate colonies, USNM 52977; G-1312, 1 lyrate colony, USNM 52975; SB-440, 4 dry pinnate branches, USNM 51292. Type Lo- cality: 26°27'N, 78°40'W (Northwest Prov- idence Channel, Bahamas), 514—586 m. Diagnosis.—Distal edges of 4—7 margin- al scales prominently spinose, the spines corresponding to those opercular scales that have one or both of their edges overlapped by flanking opercular scales; branching var- iable, including dichotomous, close-pin- nate, and lyrate; all scales covered with a low, often inconspicuous, granulation; 10— 12 perpendicularly oriented polyps/cm. Description.—Colonies are flabellate, slightly convex, and occur in three branch- ing forms: dichotomous, lyrate, and close- pinnate. The most commonly collected form is close-pinnate, colonies up to 12 cm in height and 11 cm in width, with a basal branch diameter of 1.7 mm, and consisting of 4 or 5 distinct plumes. Internodes are only 3—5 mm apart, producing a series of closely spaced, parallel, wiry branches that rarely exceed 4 cm in length. Dichotomous colonies are usually smaller, the holotype only 6 cm tall, 7.5 cm in width, and having a basal stem diameter of 0.8 mm. The first bifurcation occurs 7—11 mm above the sub- strate; subsequent branching occurs every 3-10 mm, distal branchlet rarely more than “WIL QS = Sivq g[eos [[V ‘sourds [eulsreul SuIpuNoOINs pue uNtno1edo JO MAIA Oala}s ‘EEQ-H ‘q ‘soutds [JVM Apog pedojoasp-jjam Surmoys dAjod & JO MOIA OdIO}S [eIO}IP] “6L9-D ‘OD 18208 vouunsdoyjuvoy “q—D ‘souids se[no1edo jo (46 ‘BIJ 99S) JuOWIASUBIIe ONSTIO}OvIeYS SUIMOYS WIn;NdIedoO uv JO MIA OdI9}s puv dAjod vB JO MAIA [RID}e[ ODIO}s :odAjO[OY ‘VIvajnov D]jJa/DUIN] g ‘G-—Y 8 BIA + ~% jaa [aa = 2) Z 5 jaa) = 2) =) oe) > 468 Fig. 9. 3 cm. The lyrate form is believed to be a variation of the dichotomous form. The axis is yellow-gold; polyps (in alcohol) are white. Polyps are arranged biserially in the plane of the flabellum in an alternating fashion and are well spaced (1 mm apart), resulting in 10—12 polyps/cm. Polyps are oriented away from the convex side of the flabellum (toward the anterior side) and usually perpendicular to the branchlet. Pol- yps are distally flared, and including the elongate marginal spines, measure up to 1.8 mm in height and 0.7—0.8 mm in distal di- ameter. Each polyp is protected by 8 operculars and 8 rows of body wall scales, both ab- and adaxial rows having the same number of scales (3 or 4) as the polyp is not curved toward the branch. Four to seven (usually 5 or 6) of the marginal scales bear extreme- ly long, slender, sharp (apical angle 8°—10°) spines, that are cylindrical in cross section. They are slightly curved over the polyp PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Diagrammatic representation of seven arrangements (a—g) of marginal and opercular scales in Plu- marella aculeata, as viewed from above the operculum. The eight large triangles of each figure are the 8 operculars; the smaller triangles are the corresponding marginal scale spines. The numbers on the operculars indicate how many of its two edges overlap an adjacent opercular, the arrows also indicating which edge overlaps an adjacent opercular. Spines occur only on those marginal scales corresponding to operculars having one or both of its edges overlapped by an adjacent opercular; marginal corresponding to operculars that are overlapped by both adjacent operculars do not have a spine. face and often lack granulation, thus ap- pearing translucent, or may be covered with aligned spinules (Fig. 7G). The spine por tion of the marginal scales constitutes 60— 65% of the length of the scale, resulting in a H:W of 1.8—2.8. The basal portion of the spined marginals is massive: rounded or shield-shaped. The number and position of marginal spines appears to be directly cor- related to the corresponding opercular scales that have one or both of their edges overlapped by flanking opercular scales. Opercular scales that overlap both adjacent operculars do not have a corresponding spi- nose marginal, their distal margins being only slightly rounded. Because every polyp has 8 operculars and thus 16 opercular edg- es, and every edge must either overlap or be overlapped by an adjacent opercular, it is mathematically possible for 4 to 8 oper- culars to have one or two edges overlapped, resulting in a polyp with 4—8 marginal spines (Fig. 9). Polyps having only 4—7 marginal spines have been observed; the VOLUME 117, NUMBER 4 nent Preteen ! 4 Fig. 10. A, Plumarella pellucida, holotype; B, P. laxiramosa, holotype; C, P. dichotoma, holotype; D, Candidella imbricata, Gos-2384; E, Plumarella aculeata, holotype; EK Acanthoprimnoa goesi, Atl-3465, MCZ 3741. Scale bars for A, B, F = 5 cm; C—-E = 2.5 cm. 470 hypothetical 8-spined polyp has not been seen. Body wall scales of the second and third tier are large, thick, rectangular, and have rounded upper edges; they are smooth or bear only low granules. The fourth tier of scales consists of small scales indistin- guishable from the coenenchymal scales. The opercular scales are isosceles triangular in shape with a broad base and attenuate rounded tips that form an apical angle of 20°—25°; their edges are finely serrate and their upper surface smooth to inconspicu- ously granular. Operculars are up to 0.62 mm in height and have a narrow range of H:W of 1.6—2.2. Coenenchymal scales are flat, irregular in shape, 0.20—0.40 mm in width or diameter, and have a conspicuously granular upper surface. The undersurface of all scales and the upper proximal sides of most where the scale is overlapped by an adjacent scale, are covered with complex tubercles up to 18 zm in diameter. Etymology.—The species name aculeata (Latin: aculeatus, sharp-pointed) is an al- lusion to the extremely long, sharp-pointed spines of the marginal scales. Comparisons.—Six species of Plumar- ella are characterized by having spinose marginal spines (Kiikenthal 1919); four of those six endemic to Japan. Plumarella aculeatus differs from these in having ex- tremely elongate and sharp marginal spines that occur only on those marginal scales that correspond to opercular scales that have overlapped margins (Figs. 8B, 9). Distribution.—Insular northern Straits of Florida; Northwest Providence Channel, Bahamas; 400—900 m. Acanthoprimnoa, n. gen. Type species.—Plumarella goesi Aurivil- lius, 1931, here designated. Diagnosis.—Primnoidae with a well-de- fined operculum; polyps usually inclined apically, each completely surrounded by 8 rows of body wall scales; polyps arranged alternately and biserially; 8 marginal scales, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON each with a spinose or finely serrate (pec- tinate) distal margin; no sclerites bear tu- bercles on their under surfaces; opercular scales not keeled; colonies uniplanar, usu- ally pinnately branched (plumose), dichot- omous, or lyrate. Brooding polyps are com- mon. Distribution.—Straits of Florida, Baha- mas, Yucatan Peninsula, Lesser Antilles; Japan; 60-1125 m. Remarks.—In his unpublished manu- script on the western Atlantic deep-water octocorals (Bayer & Cairns 2004), Verrill referred to the type species of this genus as Acanthoprimnoa aspera, a species later de- scribed by Aurivillius as Plumarella goesi. Because only Verrill’s plates and not the text survived we do not know what criteria he used to distinguish his new genus. We separate this genus from the morphologi- cally similar Plumarella by three criteria: the lack of tubercles on the undersurfaces of the sclerites, the distinctive pectinate dis- tal edges of the body wall and opercular scales, and the coarsely granular coenen- chymal scales. Two other species, previous- ly placed in Plumarella, also share these characteristics and are transferred to Acan- thoprimnoa: A. serta (Kiikenthal & Gor- zawsky, 1908), n. comb. and A. cristata (Kiikenthal & Gorzawsky, 1908), n. comb. Etymology.—The genus name Acantho- primnoa (Greek: acantha, a thorn + prim- noa, a common suffix used in this family) is an allusion to the spiny nature of the pol- yps of the type species. Acanthoprimnoa goesi (Aurivillius, 1931), n. comb. Figs. 8C—D, 10K 11A-—I, 12A—B ?Primnoa Pourtalesii.—Hargitt & Rogers, 1901:281, fig. D. Plumarella goési Aurivillius, 1931:244— 248, pl. 5, figs. 6a—b, text fig. 47, 3-5. — Bayer, 1957:388 (Cay Sal Bank). Thouarella goési Deichmann, 1936:164— 165, pl. 25, figs. 2, 19-23, pl. 26, fig. 8.—Bayer, 1954a:281 (listed). VOLUME 117, NUMBER 4 471 Fig. 11. A-I, Acanthoprimnoa goesi, G-633: A, upper and undersurfaces of 3 opercular scales; B, pectinate edge of an opercular; C—D, under and upper surfaces of 2 marginal scales; E, upper and undersurfaces of 3 body wall scales; F—H, upper and undersurfaces of 3 coarsely granular coenenchymal scales, F having a central spine; I, granules on upper surface of a coenenchymal scale. J-M, Acanthoprimnoa pectinata, holotype: J, upper and undersurfaces of 5 opercular scales; K, upper and undersurfaces of 4 butterfly-shaped body wall scales; L, upper and undersurfaces of 3 coarsely granular coenenchymal scales; M, coenenchymal scales on a branch. Scale bars for A, C-H, J-K, M = 0.10 mm; B, I, L = 25 pm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 472 “WUT QC'0 = q ‘wu 670 = D-V JJ seq ajeog “déjod v Jo MaIA [eASIe] ODIE}S “[EQ9T PLVMISDY “VIvoLqui vyapipuvD ‘q ‘dAjod & Jo MatA [eIA}v] ODIAIS ‘769-9 ‘YINUNIAad pouusdoyjupoy “> ‘sourds [eulss1eu SUIpuNnoMs puv winjnosiedo Jo MaIA O919}8 pur dATOd v Jo MoIA [eIO\L] O9I9)S :168-D ‘78208 nouumsdoyjupoy ‘g—yW *Z] “Biq I UO LT RES VOLUME 117, NUMBER 4 “‘Acanthoprimnoa aspera” Bayer & Cairns (Verrill), 2004: pl. 10, fig. 8, pl. 13, fig. 8a, pl. 27, figs. 5a—b, pl. 141, fig. 6. Material examined.—Alb-2342, 10 dry pinnate colonies, USNM 10236; Alb-2343, 1 pinnate colony, USNM 10243; A/b-2346, 1 pinnate colony, USNM 10783; Alvin 846, 3 dry pinnate colonies, USNM 79517; Alvin 77-764, 4 pinnate colonies, USNM 96825; Atl-2999, 10 dry pinnate branches, MCZ 54324 and 3832; Azl-3402, 4 dry pinnate branches, MCZ 3703; Atl-3403, 7 dry pin- nate branches, MCZ 54335; Az/-3438, 3 dry pinnate branches, MCZ 3751; Atl-3463, 7 dry pinnate colonies, MCZ 3604; A7l-3465, 54 dry pinnate colonies, MCZ 3744a, 3741, and 3737; Atl-3466, 17 dry pinnate colo- nies, MCZ 3603; .At/-3478, 20 dry pinnate colonies, MCZ 3605; Atl-3479, 17 dry pin- nate colonies, MCZ-3608 and 3759; Atl- 3480, 11 dry pinnate colonies, MCZ 3668 and 3762; Aftl-3482, 32 dry pinnate colo- nies, MCZ 3654 and 3663; Cape Florida X, 8 pinnate colonies, USNM 73932; JS- 43, 6 pinnate colonies and 1 unnumbered SEM stub, USNM 43801; JS-102, 1 dry pinnate colony, USNM 1011364 (topotyp- ic); JS-103, 3 pinnate colonies, USNM 50951 (topotypic); Eastward 26537, 2 pin- nate (USNM 98161) and 13 dichotomous colonies (USNM 98850); Eastward 26538, 10 pinnate branches, USNM 1019537; Eastward 26549, | pinnate colony (USNM 75064) and 24 dichotomous colonies (USNM 75065, 76987, and 79485); East- ward 26550, 53 pinnate colonies, USNM 94500; Eastward 26559, 8 dichotomous (USNM 98851) and 1 pinnate colony (USNM 98852); Eastward 31281, 15 pin- nate colonies, USNM 94522; G-235, 3 pin- nate colonies, USNM 98853; G-241, 16 pinnate colonies, USNM 52966; G-242, 1 pinnate colony, USNM 52968; G-251, 2 pinnate colonies, USNM 52969; G-252, 1 pinnate branch, USNM 98854; G-254, 1 pinnate colony, USNM 52962; G-387, 12 pinnate colonies, USNM 52970; G-533, 1 pinnate colony, USNM 52971; G-633, 13 473 pinnate colonies and SEM stubs 258 and C1091, USNM 52973 and 52983; G-679, 8 pinnate colonies and SEM stub 256, USNM 52963; G-680, 4 dichotomous colonies, USNM 1019538; G-696, 1 pinnate colony, USNM 52972; G-704, 1 pinnate colony, USNM 52965; G-706, 25 pinnate colonies, USNM 52967; G-707, 1 pinnate colony, USNM 98855; G-879, 2 dichotomous col- onies, USNM 76988; G-897, 10 dichoto- mous colonies and SEM stub 257, USNM 52964; P-594, over 50 pinnate colonies, USNM 52961, and 4 dichotomous colonies, USNM 98856: P-596, 2 dichotomous col- onies, USNM 52960; P-598, 1 pinnate col- ony, USNM 52957 and | dichotomous col- ony, USNM 98858; specimens reported by Deichmann (1936) and Bayer (1957); a syntype (USNM 44192). Types and type locality.—Three speci- mens are mentioned in the original descrip- tion, only one of which was figured; all three are considered to be syntypes. They are deposited at the Stockholm Museum (#28); a fragment of one of the colonies is also deposited at the USNM (44192). Type Locality: ““Virgin Islands”’, 457-548 m. Diagnosis.—Distal edges of 7 marginal scales (not one of the adaxial marginals) prominently spinose; branching dichoto- mous in shallow-water form and close pin- nate in deeper-water form, branchlets flex- ible but not flaccid; opercular scales cov- ered with numerous tiny spines, abaxial and outer-lateral body wall scales bear single, robust spines on distal margin; coenenchy- mal scales highly granular and sometimes bear a single tall spine; 13—15 polyps/cm; branch axis bronze; polyp brood chambers common. Description.—Colonies are flabellate and occur in two branching forms. The deeper water form is larger (up to 30 cm in height and equally broad), with closely pinnate branching colonies consisting of 2 or 3 reg- ular plumes. Pinnate branchlets begins within 5 mm of the base and are subse- quently arranged in a regular parallel fash- ion, the internodes being only 2—3 mm in 474 length; unbranched terminal branchlets are flexible and rarely exceed 4 cm in length. These large colonies are attached by a cal- careous holdfast that may reinforce the main stem as much as 15 mm above the base and attain a diameter of 4 mm. As with most species of Plumarella, the colony fla- bellum is slight convex, with the polyps di- rected slightly upward and toward the con- vex face. The shallow-water form is much smaller (rarely exceeding 4 cm in height) and is dichotomously branched, often re- sulting in a colony broader than tall. Inter- nodes are 2—4 mm in length; terminal branchlets are rarely more than 15 mm and number only about 10—15; the basal axis is about 0.5 mm in diameter. In both forms the axis is a rich bronze color, which con- trasts with the white (in alcohol) color of the polyps. Polyps are arranged biserially in the plane of the flabellum in an alternating fashion and are well spaced approximately 0.6—0.8 mm apart, resulting in 13—15 pol- yps/cm. Polyps of the deep-water form are 0.8—1.2 mm in height Gncluding the mar- ginal spines) and about 0.5 mm in diameter; polyps of the shallow water form are small- er, usually less than 0.8 mm in height. As mentioned in the remarks section, some polyps of both forms have brood chambers that greatly swell the base of the polyp. Each polyp is protected by 8 opercular and 8 rows of body wall scales, the abaxial row having 5 or 6 scales, the adaxial, 4 or 5. Seven of the 8 marginal body wall scales bear a prominent distal spine, the 8th (ad- axial) marginal having a very reduced spine, allowing the abaxial opercular scales to overlap the polyp edge at that point of the circumference (Fig. 8D). The marginal scales have a flat, rectangular to ellipsoidal base up to 0.4 mm wide from which the elongate, sharp-tipped (apical angle 7°—8°), often crooked spine emerges. The entire marginal scale may be up to 0.85 mm in height, the spinose part constituting 75— 85% of its height and contributing to a rath- er high H:W of 1.7—3.2. The elongate spi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nose part of each marginal scale is spinose itself, bearing prominent rows of smaller spines (25-30 wm in length), which are ar- ranged in rows on both the upper and lower surface of the spine. The smaller spines also cover the edges and upper surfaces of the base of the marginals, but the undersurfaces of the marginal base, covered by tubercles in all species of Plumarella, is smooth. The body wall scales of the abaxial and inner- lateral rows that lie proximal to the margin- als also have apical spines, but these are quite variable in size, some quite large (up to half the height of the scale), others in- conspicuous. The body wall scales of the adaxial and outer-lateral rows that lie prox- imal to the marginals have greatly reduced or no apical spines. The opercular scales are similar to the marginal scales in many ways but are isosceles triangular in shape, not having a rectangular base, and often have a notch on either side near the base. The ab- axial operculars are quite elongate (up to 0.7 mm) and, when closed, often complete- ly traverse the polyp. Their tips are pointed (apical angle 15°—25°), with a H:W ranging from 1.8—3.9. As with the marginal spines they are covered with prominent spines on the upper and undersurfaces, except for the undersurface of the base, which is smooth. All three edges of the opercular scales are serrate, but in the region of the proximal notches the serrations are developed into elongate (up to 40 pm long and 9 pm in diameter), finely granular pillars that often bi- and trifurcate (Figs. 11A—B). Coenenchymal scales are rather large (up to 0.5 mm) and have coarse granular edges and surfaces, the granules rounded and up to 12 wm in diameter and often twice as tall. Some coenenchymal scales also bear a prominent, centrally located, perpendicular spine up to 0.3 mm in height and 0.1 mm in basal diameter. These spines are orna- mented with smaller spines similar to those on the opercular and marginal scale spines. The undersurface of the coenenchymal scales is smooth. Comparisons.—See A. pectinata. VOLUME 117, NUMBER 4 Distribution.—Throughout the Straits of Florida to Arrowsmith Bank, Yucatan Channel; Northwest Providence Channel; Old Bahama Channel; Puerto Rico (Hargitt & Rogers 1901); Virgin Islands. In general, the dichotomous form occurs from 137—350 m and the pinnate form deeper, 320-595 m. Remarks.—The only differences between the two forms, aside from their different range of capture depths, are that the deeper- water form has close pinnate branching, a larger colony, and larger polyps, whereas the shallow-water form has dichotomous branching, a smaller colony, and smaller polyp size. All other characters are quite similar, unique characters including the brooding polyps, spinose body wall and coenenchymal scales, and long, spiny oper- culars. Several stations contain both forms (see material examined), but in general the forms occur at different depth ranges. Some colonies are transitional in form, beginning as dichotomous but with a tendency toward pinnate branching at least in part of the up- per colony. Such was the syntype illustrated by Aurivillius (1931: pl. 5, fig. 6a), al- though he unequivocally classified that col- ony as dichotomous. About one-third of the colonies exam- ined contained polyps with bulbous brood chambers in their base, this feature occur- ring in both the dichotomous and pinnate forms. Among the colonies containing pol- yps with brood chambers, approximately one in 50 polyps would be so modified, but oftentimes there would be 2 or 3 contiguous brooding polyps. There appears to be no seasonality regarding the presence of the brooding polyps. Acanthoprimnoa pectinata, n. sp. Figs. 1D, 11J—M, 12C Material examined/types and type local- ity.—Holotype: G-899, 1 colony and SEM stubs C1093—1095, USNM 1019539. Para- types: Alb-2354, 20 colonies and 1 unnum- bered SEM stub, USNM 43026 and 75112; Alvin 77-760, 5 dichotomous colonies, 475 USNM 1019540; Azl-3303, 3 dry pinnate colonies, MCZ 3627; G-692, 2 branches and SEM stub 254, USNM 52954; G-889, 4 colonies, USNM 52952; G-898, 1 pinnate colony, USNM 52956; G-899, 19 colonies, USNM 52955; O-4940, 2 colonies, USNM 52953; P-592, 20 colonies, USNM 52958; P-954, 1 colony, USNM 52959; SB-5190, 6 pinnate colonies, USNM 1019541. Type Locality: 20°57'N, 86°34'W (off Arrows- mith Bank, Yucatan, Mexico), 40—164 m. Diagnosis.—Distal edge of marginal scales straight or only slightly spinose; branching loosely pinnate, branchlets long and flaccid; opercular scales ridged and covered with numerous tiny spines; lateral edges of opercular and distal and proximal edges of body wall scales bear a series of comb-like spines; coenenchymal scales coarsely granular, but without a central boss; 11—13 polyps/cm; branch axis bronze; polyp brood chambers common. Description.—Colonies are flabellate and loosely pinnate, each colony consisting of 2 or 3 plumes, although one colony (P-594) is lyrate in branching. The first branchlets occur very near the base of the colony, suc- ceeding branchlets at a periodicity of every 4—5 mm (internode length), the branchlets up to 55 mm in length and flaccid in ten- sion. The main stem is anchored by a dense, calcareous, white holdfast, although the holdfasts of only five of the colonies are intact. Like A. goesi, the axis is bronze in color, which contrasts with the white pol- yps. The holotype is 18 cm in height, 7 cm in width, but lacks a base; the largest spec- imen (A@/-3303) is 22 cm tall. The largest main stem of an attached colony (G-899) has a diameter of only 1.1 mm. Polyps are arranged biserially on the branchlets and main stem (6—7 polyps/in- ternode on main stem) in an alternating fashion 0.8—1.1 mm apart, resulting in 11— 13 polyps/cm. Polyps are relatively small, only 0.8—0.9 mm in height and 0.40—0.45 mm in diameter. Colonies from all stations recorded contain some polyps with brood 476 chambers, which greatly swell the base of those polyps. Each polyp is protected by 8 opercular and 8 rows of body wall scales, the abaxial row consisting of 8—10 scales, the adaxial, 7-9. All body wall scales, including the marginals, are slightly curved to accom- modate the curvature of the polyp, and con- siderably wider than tall, such that a rela- tively high number occurs in the wall of a relatively short polyp. The upper surface of the body wall scales bears many small spines, especially toward the center of the scale, and their distal and proximal margins bear a series of fine, comb-like (pectinate) projections measuring up to 32 wm in length. Only rarely will the marginal body wall scale have a larger, projecting spine, the largest up to 0.25 mm in length and constituting about half the height of the scale. The opercular scales are isosceles tri- angular in shape (H:W = 1.6—2.2), and strongly curved in order to cover the top of the rounded polyp. Operculars are up to 0.38 mm in height and have an apical angle of 35°—45°. They are sculptured as in A. goesi. Coenenchymal scales are relatively small (0.09—0.21 mm in width) and circular to ir- regular in shape. As in A. goesi, they are densely covered on their upper surface with prominent, blunt granules measuring up to 15 wm in height and 10—12 pm in diameter, but smooth on the undersurface. Etymology.—The species name pectinata (Latin: pectinatus, comblike) refers to the comb-like serration of the edges of the opercular and body wall scales. Comparisons.—Acanthoprimnoa pectin- ata resembles A. goesi in the morphology of its opercular spines, color of the branch axis, coarsely granular coenenchymal scales, and the common presence of brood polyps. Further, A. pectinata differs (Table 1) in lacking distally spinose body wall scales (instead having pectinate distal and proximal margins), lacking a central boss on the coenenchymal scales, having more scales/body wall row, having much shorter PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON operculars, and in having a looser pinnate branching mode. A. pectinata is most sim- ilar to the Japanese A. cristata, but lacks the longitudinal ridges on the body wall scales. Distribution.—Off northeastern Yucatan Peninsula and northwestern Cuba (164—476 m); Straits of Florida; Mona Passage and off Montserrat, Lesser Antilles (614—686 m). Remarks.—All but two colonies of A. pectinata occur in relatively shallow water (164—476 m) off the Yucatan Peninsula, but the colonies from P-954 (off Montserrat, Lesser Antilles) and Alvin 77-760 (Straits of Florida) occur in deeper water (614—686 m) and are the only colonies to have non- pinnately (dichotomous, lyrate) branching colonies. Genus Candidella Bayer, 1954 Primnoa.—Johnson, 1862:245 (in part). Stenella Gray, 1870:48 (Gunior primary homonym of Stenella Gray, 1866, a ce- tacean).—Wright & Studer, 1889:56 (Gn part).—Ktkenthal, 1919:443—445 Gn part); 1924:303 (Qn _ part).—Aurivillius, 1931:289—290 (in part). Narella.—Studer, 1878:643 (Gn part). Stenella (Primnoa).—Roule, 1896:304. Stenella (Stenella).—Versluys, 1906:38—39. Candidella Bayer, 1954b:296 (nom. nov.); 1981:937.—Tixier-Durivault, 1987:171. Candidella (Candidella).—Bayer, 1956: F222. Type species.—Primnoa imbricata John- son, 1862, by monotypy. Diagnosis.—Primnoidae with a well-de- fined operculum; polyps stand perpendicu- lar to branch (not bent); polyp body wall completely surrounded by 2—4 rows of sclerites; polyps arranged in whorls; only four marginal scales; undersurfaces of all sclerites tuberculate, opercular scales strongly keeled; colonies dichotomously branched in one plane. Distribution.—North Atlantic, Ascension, central and western Pacific; 183-2139 m. Remarks.—Four species are known in VOLUME 117, NUMBER 4 this genus: C. imbricata (Johnson, 1862); C. johnsoni (Wright & Studer, 1889), As- cension; C. gigantea (Wright & Studer, 1889), Fiji; and C. helminthophora (Nut- ting, 1908), Hawaiian Islands. After its original description, the monographers Kiu- kenthal, Studer, and Aurivillius took a broad view of the genus Stenella, including similar species but some differing in having 5 or 8 marginal scales, these species later being transferred to Parastenella, Pteros- tenella, and Dasystenella. Versluys (1906) was the first to relegate what is now known as Candidella to a monophyletic group, the nominate subgenus of Stenella. After re- naming the genus Candidella (Bayer, 1954b), because the name Stenella was a junior homonym, Bayer (1956) also recog- nized it as a monophyletic subgenus: Can- didella (Candidella), subsequently elevat- ing it to generic rank in 1981. Characters used to distinguish species include the ar- rangement of polyps, colony branching, and polyp size (Kiikenthal 1924). Candidella imbricata (Johnson, 1862) Figs. 1OD, 12D, 13A—G, 14A—D Primnoa imbricata Johnson, 1862:245, pl. 31, figs. 2, 2a (Madeira); 1863:299 (ver- batim). Stenella imbricata.—Gray, 1870:48—49, 2 figs. (listed, new comb.).—Wright & Stu- der, 1889:56, 281 (listed).—Kiikenthal, 1919:448—449 (Blake from Cuba, first re- cord for western Atlantic); 1924:305—306 (diagnosis, key).—Thomson, 1927:32— 33, fl, 2, m3, GS, lla 3, ins, D5 Toll, DS, ws 5—6 (Azores, Morocco).—Aurivillius, 1931:290 (mentioned).—Deichmann, 1936:167—168, pl. 26, fig. 5 (West In- dies).—Bayer, 1954a:281 (listed for Gulf of Mexico); 1964:532 (Straits of Florida). Narella imbricata.—Studer, 1878:643 (list- ed, new comb.). ?Stenella (Primnoa) johnsoni.—Roule, 1896:304 (Gulf of Gascogne). Stenella (Primnoa) imbricata.—Roule, 1896:304 (comparison to C. johnsoni). 477 Stenella (Stenella) imbricata.—Versluys, 1906:42—43, 44, fig. 46 (redescription of type, key to spp.) Candidella imbricata.—Bayer, 1954b:296 (new. comb.).—Tixier-Durivault & d’Hondt, 1974:1412—1413.—Grasshoff, 1981:222, map 1 (mid-Atlantic Ridge sw of Azores); 1982a:738, maps 4, 20; 1982b:948—949, figs. 20—21.—Grasshoff & Zibrowius, 1983:119—-120, 122, pl. 2, fig. 7 (mid-Atlantic Ridge), pl. 3, fig. 14 (Biscay Bay).—Carpine & Grasshoff, 1985:6 (frontispiece), 33 (Musée Océan- ographique de Monaco catalog num- bers).—Pasternak, 1985:29 (Rockaway Seamount).—Pettibone, 1991:705, 707 (polychaete commensal). Candidella (Candidella) imbricata.—Bay- er, 1956:F222, fig. 159—4b. Candidella johnsoni.—Bayer, 1981:934, fig. 74. Stenella “‘florida” Bayer & Cairns (Verrill), 2004: pl. 13, figs. 1, la, pl. 25, fig. 13a— b, pl. 82, figs. 2, 2a, pl. 83, fig. 6, 6a. Material examined.—Alb-2753, 3 branch fragments, USNM 44126; Alvin 762, 6 branches, USNM 80939, 80940, and 1017255; Alvin 1335, 2 fragments (one dry), USNM 73744 and 73745; Alvin 3885- 5, 1 complete colony, USNM 1019238; Al- vin 3903-101-—2, 1 branch, USNM 1019273; Atl-266-47, branch fragments, USNM 60337; Atl-280-9, 3 branches and SEM stub 273, USNM 57552; CI-63, 1 col- ony, USNM 60223; CI-140, 1 branch, USNM 60341; Eastward 26019, 6 colonies, USNM 60338; Eastward 26022, 2 colonies, USNM 60340; Eastward 26023, dry branch fragments, USNM 1011365; Eastward 26031, 3 colonies (some dry) and SEM stubs 274 and C1071—1076, 1078, USNM 57553 and 60339; G-169, 6 colonies, USNM 52778; G-170, 2 colonies and 1 un- numbered SEM stub, USNM 52779; G- 177, 2 colonies, USNM 52780; G-386, 14 colonies and numerous branches, USNM 52784; G-660, 1 branch, USNM 52781; G- 661, 1 branch, USNM 52782; G-936, 2 478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 13. tuberculate undersurface of an opercular; C, upper surface of 2 basal body wall scales; D, K upper and under- surfaces of 3 medial body wall scales; E, undersurface of 2 marginal scales; G, upper and undersurfaces of 3 coenenchymal scales. Scale bars for A, C-G = 0.25 mm; B = 25 pm. branches, USNM 52783; G-965, 1 colony, USNM 52787; Gos-2383, 1 colony, USNM 57309; Gos-2384, 1 colony, USNM 57310; Gyre CO4, | colony (dry), USNM 89124; P-197, 2 colonies, USNM 52785; P-881, 2 colonies, USNM 52786; P-892, 2 colonies, USNM 52911; P-1146, 1 branch, USNM 52912; off Bermuda, 1200 m, 1 colony, USNM 75104. Types and type locality.—The holotype A-G, Candidella imbricata, Gos-26031: A, upper and undersurfaces of 4 opercular scales; B, is deposited at the BM (1863.1.31.1). Type locality: Madeira, depth unknown. Description.—Colonies consist of a ro- bust vertical main stem up to 9 mm in basal diameter, which supports a uniplanar fan achieved by dichotomous branching. The main stem is anchored by a dense, white, encrusting, calcareous holdfast, which often encrusts other calcareous Coelenterata, such as the scleractinians Enallopsammia pro- ‘WU SZ’ = q ‘wur gy, = q ‘wu Qs’9 = D-V JO} seq a[eog ‘sayeos Iejno1odo ouros Jo dn asojo ‘gq ‘dAjod & Jo apis [elxepe oY) JO MOIA OdIOIS ‘OD :[EQ9T PuvMIsD” “q ‘D ‘dn youvig ‘q ‘se[evos [eUISIeUI INOJ dy) SuIMOYs Ose ‘MIA Ie[NoIedo O919}s “gq ‘dAjod B JO MIA [eIO}V] OOIO}S “WV :6-O8T-4V ‘d ‘A-V :BIpoluqui DjjapipuvD, “p| “314 Se VES a= kee Le ERE s a aa aa) = 5) Z, eS ea = =) = © S 480 funda, Lophelia prolifera, Javania cailetti, the stylasterid Stylaster erubescens, and various bryozoans. The calcareous deposits may reinforce the basal stem as much as 2 cm upwards from the base. The holdfast and basal reinforcement are composed of 100% aragonite, consistent with the find- ings of Bayer & Macintyre (2001) for the congeneric C. helminthophora. The axis is yellow-gold in color and longitudinally stri- ate; overall the colony is white. The largest known colony (the holotype) is reputed to be 21.6 cm in height and 27.9 cm in width. Branching is dichotomous at intervals of 3— 12 mm, but unequal, resulting in asymmet- rical branching; there is little to no branch anastomosis. Polyps are arranged in whorls of 3 or 4 polyps (rarely as pairs); if in a whorl of 3, 2 polyps are usually directed in the plane of the fan in opposite directions, the third polyp standing perpendicular to the plane of the fan and thus at 90° to the other 2, the polyp projecting perpendicular to the fan defining the anterior face of the fan; few polyps originate from the posterior face of the fan. When 4 polyps constitute a whorl, the angular separation between polyps is not 90°, but about 60°, polyps avoiding the posterior face. Polyp whorls are closely spaced, about every 1.2—2.0 mm, 5—6 oc- curring per cm, polyps present even on the calcified region of the basal stem and hold- fast. Most polyps are 2.1—2.5 mm in height and slightly clavate (1.3—-1.4 mm in distal diameter), encased by the distal margin of the flared marginal scales, but some geo- graphic outliers have larger polyps (see Re- marks). Polyps are fairly rigid, projecting perpendicularly from the branches; howev- er, those in the plane of the fan are some- times slightly curved toward the anterior face. Polyps are protected by 4 marginal scales, 2—4 medial scales, 4—8 basal scales, and 8 operculars. The marginal scales are dimorphic in size and shape, consisting of 2 adjacent larger (0.9 mm in height, 1.1 mm in width), highly curved scales that define PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the abaxial side of the polyp and 2 adjacent smaller (0.65 mm in height, 0.62 mm in width), slightly curved adaxial scales, which overlap with the edges of the larger marginals. Three opercular scales corre- spond to each of the larger marginals, whereas about 1.5 operculars correspond to the smaller marginals, the number of oper- culars adding to more than 8 because of the overlap of marginal scales. The marginals are flared outward distally, rising about 0.15 mm above the junction with the opercular scales, but not enclosing the operculum. Medial body wall scales are roughly rect- angular and flat, with sides measuring 0.45— 0.65 mm in length; their lateral edges over- lap one another. Sometimes it appears that only 2 medial scales are present, these oc- curring on the adaxial side. Basal scales are dimorphic in size, consisting of 2 large, square to rectangular scales up to 0.65—0.70 mm in side length, and outwardly concave, as though squeezing the base of the polyp into a narrow opening. When polyps be- come abraded from the branches, these large scales often remain to mark the orig- inal position of the polyp. Between the 2 large basal scales, on the adaxial side, are 1 or 2 pairs of much smaller basal scales that are overlapped and overshadowed by the larger basals. The 8 operculars are elon- gate triangular, having a H:W of 1.6—2.0, pointed distally, highly convex above, and prominently keeled below. They form a tight conical operculum over the polyp, ris- ing well above the marginal scales. One of the 8 operculars is slightly larger (e.g., 0.8 mm tall, 0.5 mm wide) than the others and is positioned opposite the smallest opercu- lar (e.g., 0.6 mm tall, 0.3 mm wide), these two operculars defining the sagittal axis of the polyp. The remaining 6 operculars are of similar size, constituting 3 pairs mirrored across the sagittal axis. The 2 sagittal oper- culars are symmetrical, in that their keels are in a medial position, whereas the other 6 operculars are asymmetrical, their keels being offset toward the abaxial side (the side toward the large sagittal opercular), VOLUME 117, NUMBER 4 producing a longer and slightly upturned edge of their adaxial side. Each upturned adaxial opercular edge overlaps the abaxial edges of the adjacent operculars, the edges of the small sagittal opercular being over- lapped by both adjacent operculars and the large sagittal opercular overlapping both adjacent operculars (compare to Fig. 9f). Coenenchymal scales are large (up to 1.0 mm in length), occur in one layer, are po- lygonal in shape, and are usually slightly concave above. As mentioned below, they sometime orient perpendicular to the branch in order to contribute to the formation of the worm tube. The upper surfaces of all sclerites are finely and uniformly granular, the granules 11-13 sm in diameter; their undersurfaces are covered with complex tu- bercles 15-17 wm in diameter. Tentacular sclerites were not noted. Comparisons.—There is only one other species of Candidella known from the At- lantic, C. johnsoni (Wright & Studer, 1889), described from Ascension. As summarized by Versluys (1906), that species differs in having a very low operculum, marginal scales that are equal in size, and polyps that occur in pairs and singly. Although these two species are probably distinct, the only subsequent report of C. johnsoni is by Rou- le (1896) from the Gulf of Gascogne, which is probably C. imbricata, as he implied that his C. johnsoni might be a deep-water va- riety of C. imbricata. Candidella imbricata is morphologically more similar to the central Pacific C. hel- minthophora (Nutting, 1908), both species having dimorphic marginal scales and a similarly shaped polyp. However, C. hel- minthophora differs in having two rings of medial body wall scales, a larger colony with longer internodes (up to 4 cm), larger polyps, and more flexible branches. Distribution.—Western Atlantic: New England Seamounts (San Pablo, Rockaway, Kelvin, Muir), Bermuda, eastern coast of Florida, Bahamas, Greater and Lesser An- tilles, northern Gulf of Mexico; 514—2063 m. A rather large distributional gap exists 481 between the New England Seamounts and the coast of Florida. Eastern Atlantic: com- monly collected in the Bay of Biscay, off Morocco, Canary Islands, Madeira, Azores, and the mid-Atlantic Ridge southwest of the Azores; 815—2139 m (see Grasshoff 1982b for map). Remarks.—Colonies of even small size will usually host the commensal polynoid polychaete Gorgoniapolynoe caeciliae (Fauvel, 1913), larger colonies often host- ing 5 or 6 worms. The polychaete has es- sentially the same known distribution as C. imbricata, despite the fact that it occurs in at least two other gorgonians (Pettibone 1991). The gorgonian appears to be induced to form a tube that is slightly elliptical in cross section, the greater diameter being ap- proximately 2.3—2.5 mm and the length up to 25 mm, the tube always occurring on the anterior side of the fan; the length of the polychaete is about 11 mm. The tubes are formed predominantly of greatly enlarged and outwardly curved basal scales from two adjacent polyps. These basal scales, nor- mally only 0.7 mm in height, increase in size up to 1.6 mm in height and up to 2.9 mm in width. The curvature is such that basal scales from two adjacent polyps in the same whorl meet and sometimes fuse along the dorsal midline of the tube, whereas the proximal and distal edges of these enlarged basal scales meet and sometimes fuse with those of adjacent whorls, altogether form- ing a somewhat porous tube that is open at both ends. Occasionally, small coenenchy- mal scales that project perpendicular to the branch will fill in the spaces between basal scales of adjacent whorls. Although an ob- vious advantage is gained for the worm in this association, no advantage can be con- jectured for the gorgonian. Several specimens, collected at the mar- gins of the known distribution, show some variation in morphology. The single speci- men known from the northern Gulf of Mex- ico (USNM 89124) has a very low oper- culum, like that of C. johnsoni, but other- wise is similar to C. imbricata. The colo- 482 nies from Bermuda (USNM 75104) and San Pablo Seamount (USNM 57552) have unusually large polyps, 4.0 and 3.2 mm, re- spectively, but are otherwise similar to C. imbricata. Acknowledgments We wish to thank Ardis Johnston for the loan of Plumarella specimens deposited at the MCZ, and Elly Beglinger (Zoological Museum, Amsterdam) for the loan of typi- cal specimens of Plumarella penna. We thank Ian Macintyre for the mineralogical determination of the axis of C. imbricata. Molly Ryan, staff illustrator, produced Fig- ure 9, and Tim Coffer helped produce the plates. Specimens of C. imbricata from AI- vin stations made in 2003 were collected by the “‘Mountains-in-the-Sea’’ Expedition, Les Watling, Chief Scientist, funded by the NOAA Ocean Exploration program. Literature Cited Aurivillius, M. 1931. The Gorgonarians from Dr. Six- ten Bock’s expedition to Japan and Bonin Is- lands 1914.—Kungliga Svenska Vetenskaps— Akademiens Handlingar (3)9(4):337 pp., 65 figs., 6 pls. Bayer, EF M. 1954a. Anthozoa: Alcyonaria.—Fishery Bulletin of the Fish and Wildlife Service 89: 279-284. . 1954b. New names for two genera of Octo- corallia—Journal of the Washington Academy of Sciences 44(9):296. . 1956, Octocorallia. Pp. Fl166—189, 192-231 in R. C. Moore, ed., Treatise on Invertebrate Paleontology, University of Kansas Press, Lawrence, 498 pp. . 1957. Additional records of Western Atlantic octocorals.—Journal of the Washington Acad- emy of Sciences 47(11):379—390, 4 figs. . 1964. A new species of the octocorallian ge- nus Paragorgia trawled in Florida waters by R.V. ““Gerda’’.—Zoologische Mededelingen 39: 526-532, 3 figs. . 1973. Colonial organization in octocorals. Pp. 69—93, figs. 1-23 in R. S. Boardman, A. H. Cheetham, & W. A. 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Polynoids commensal with gor- gonian and stylasterid corals, with a new genus, new combinations, and new species (Polychae- ta: Polynoidae: Polynoinae).—Proceedings of the Biological Society of Washington 104(4): 688-713, 16 figs. Roule, L. 1896. Résultats scientifiques de la campagne du “‘Caudan” dans le Golfe de Gascogne— Aout-Septembre 1895. Coelentérés.—Annales de l'Université de Lyon 26:299-323. Studer, T. 1878. Ubersicht der Steinkorallen aus der Familie der Madreporaria aporosa, Eupsammi- na, und Turbinaria, welche auf der Reise S. M. S. Gazelle um die Erde gesammelt wurden.— Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1877: 625-654, 4 pls. . 1887. Versuch eines Systemes der Alcyonar- ia.—Atrchiv fiir Naturgeschichte 53(1):74 pp., 1 pl. Thomson, J. A. 1927. Alcyonaires provenant des cam- pagnes scientifiques du Prince Albert Ier de Monaco.—Résultats des Campagnes Scienti- fiques accomplies sur son yacht par Albert Ier, Monaco 73:77 pp., 6 pls. Tixier-Durivault, A. 1987. Sous-classe des Octocoral- liaires. Pp. 3-185, figs. 1-147 in D. Doumenc, ed., Traité de Zoologie. Volume 3: Cnidaires, Anthozoaires. Masson, Paris, 859 pp. , & M.-J. d’ Hondt. 1974. Les Octocoralliaires des la campagne Biagores.—Bulletin du Musé- um National d’histoire Naturelle, Zoologie (3)174(252):1361-1433. Verrill, A. E. 1883. Report on the Anthozoa, and on some additional species dredged by the “Blake” in 1877-1879, and by the U.S. Fish Commission Steamer “Fish Hawk” in 1880— 82.—Bulletin of the Museum of Comparative Zoology, Harvard 11:72 pp., 8 pls. Versluys, J. 1906. Die Gorgoniden der Siboga-Expe- dition. Il. Die Primnoidae.—Siboga-Expeditie 13a:187 pp., 10 pls., 1 map. Wright, E. P, & T. Studer. 1889. Report on the Al- cyonaria collected by H.M.S. Challenger during the years 1873—76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-76, Zoology 31(64):314 pp., 43 pls. Associate Editor: Stephen L. Gardiner 484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON APPENDIX: Station Data Station Latitude (°N) Longitude ((W) Depth (m) Date U.S.F.C.S Albatross 2342 23°10'39" 82°20'21” 384 19 Jan 1885 2343 23°11'35” 82°19'25" 510 19 Jan 1885 2346 23°10'39” 82°20'21” 366 20 Jan 1885 2354 20°59’30" 86°23'45" 238 22 Jan 1885 2416 31°26' 79°07' 505 1 Apr 1885 2662 29°24'30" 79°43' 794 4 May 1886 2663 29°39’ 79°49' 770 4 May 1886 2666 29°47'30" 79°49' 494 5 May 1886 2667 30°53’ 79°42'30" 499 5 May 1886 2668 30°58'30" 79°38'30" 538 5 May 1886 2669 31°09’ 79°33'30" 644 5 May 1886 2753 13°34’ 61°03’ 514 4 Dec 1885 IIIl9-19 off Beaufort, N. Carolina ? 27 May 1949 Alvin (submersible) 77-760 27°04.9' 79°20.1' 613-654 Jun 1977 77-761 27°04’ 79°18.8' 600 Jun 1977 77-762 27°03.3’ 79°20.0’ 600 Jun 1977 771-164 27°55.8' 79°09' 410 Jun 1977 846 26°26' TES. 525 3 Nov 1978 1335 27°05' 79°40' 608 21 Feb 1984 3885-5 33°46.17’ 62°33.9' 1821 4 Jun 2003 3903-101-2 38°47.33’ 64°07.95' 2063 15 Jul 2003 Anton Dohrn 6392 30°49’ 79°49' 400 2 65-32 Tortugas 1064 30 Jul 1932 R/V Atlantis (Atl) 266-02 31°58’ 77°18.5' 813 25 Jun 1961 266-04 31°56’ 77°26' 768 26 Jun 1961 266-07 31°53’ YDS! 750 28 Jun 1961 266-40 30°53’ T8°47' 804 13 Jul 1961 266-41 30°59’ 78°14’ 877 15 Jul 1961 266-47 30°53’ T8°47' 819 19 Jul 1961 280-09 38°51'18” 60°29'00" 1902 17 Jun 1962 2999 23°10’ 81°29’ 265-512 17 Mar 1938 3303 23°05’ 82°33’ 476 23 Mar 1939 3402 22°36’ 78°21' 421 28 Apr 1939 3403 22°36’ 78°22 384 28 Apr 1939 3438 23°05’ 79°37' 485 2 May 1939 3463 23°09’ 81°26 421 9 May 1939 3465 23°09’ 81°27’ 320 9 May 1939 3466 23°09’ 81°27’ 366 9 May 1939 3478 23°09’ 81°27'30" 240 11 May 1939 3479 23°10’ 81°26’ 384 11 May 1939 3480 23°10’ 81°28’ 366 11 May 1939 3482 23°09’ 81°27’ 348 11 May 1939 3780 30°27' USS) 458-485 24 Feb 1940 3782 30°10! 78°44’ 795-804 24 Feb 1940 U.S.C. S. S. Bibb 19 23°03’ 83°10/30" 567 4 May 1868 Dp) 24°14'20" 80°59'40" 567 4 May 1868 135 24°20'30" 81°58'30" 229 17 Feb 1869 VOLUME 117, NUMBER 4 485 APPENDIX: Continued Station Latitude (°N) Longitude (°W) Depth (m) Date Johnson-Smithsonian Deep-Sea Expedition (JS) 43 18°04’ 67°48’ 439-549 11 Feb 1933 102 18°51’ 64°33’ 90-500 4 Mar 1933 103 18°51’ 64°33’ 274-732 4 Mar 1933 R/V Cape Florida x 27°31’ 79°15' 350-400 Jul 1984 R/V Cape Hatteras SA6 31°18'08”" 79°00'08” 545-549 17 Nov 1985 SA6-1 31°17'18" 79°00'39”" 572-575 17 Nov 1985 SA6-5 31°49'40" 78°19'16" 625 18 Nov 1985 R/V Colombus Iselin (CI) 63 28°06’ 77°08' 1023-1153 21 Sep 1980 123 24°12'06" T7T18' 1435 24 Sep 1973 140 26°24 79°36' 738 28 Sep 1973 246 26°23’ 79°37' 743-761 29 Oct 1974 266 24°18.5' WP iy" ? 3 Nov 1974 Clelia (submersible) 78 32°43'38" 78°05'38” 175-196 5 Jul 1993 T9A 32°43'38" 79°05'50” 210 7 Jul 1993 M/V Combat 174 34°45’ 75°28' 320 14 Nov 1956 368 34°15’ T5231 348 16 Jun 1957 Discoverer x 32°10’ 78°07'18" 446 5 Oct 1967 R/V Eastward 26004 28°08’ 79°33' 785-830 Nov 1974 26017 26°38'30" 79°32'30" TIS Nov 1974 26019 27°16'48”" 79°25' 655-685 Nov 1974 26022 27°28'30" 79°25'18" 655-685 Nov 1974 26023 DIPS 79°22' 690 Nov 1974 26028 27°09.5’ 79°25' 635-700 Nov 1974 26031 27°00' 79°24'18" 645-690 Nov 1974 26052 25°42.7' 79°47.5' 660-770 Nov 1974 26535 DINGO! 79°15.6' 480 29 Mar 1975 26537 2TimAY 79°15’ 520 29 Mar 1975 26538 DPD! 79°13.7' 420 29 Mar 1975 26547 27°18’ 79°17' 520 Mar 1975 26549 27°17'30" 79°12'30" 370 30 Mar 1975 26550 27°16'24” 79°14'18" 440 30 Mar 1975 26559 26°30.3’ 79°14.7' 2 31 Mar 1975 31281 26°53'54" 79°07'18" 320 1977 R/V Gerda (G) 56 DSB" 79°20' 458 28 Aug 1962 169 27°01’ 79°21.5' 229-275 29 Jun 1963 170 27°06’ 79°32’ 659-677 29 Jun 1963 7 2T°17' 79°34 686 30 Jun 1963 235 25°44’ PDs 531 30 Jun 1963 486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON APPENDIX: Continued Station Latitude (°N) Longitude ((W) Depth (m) Date 241 25°26’ 79°18" 494-502 30 Jan 1964 242 25°36’ 79°21' 485-530 30 Jan 1964 246 26°57’ 79°12.5' 512 5 Feb 1964 247 27°07’ 79°21" 567 5 Feb 1964 251 Dif Dy 78°41’ 293-311 5 Feb 1964 252 DDO! 78°37.5' 485-496 5 Feb 1964 254 igs Amo 78°49' 488-516 6 Feb 1964 261 27°20’ TO D24 494-511 7 Feb 1964 386 27°09" 78°18’ 604 19 Sep 1964 387 DP 79°15’ 412 19 Sep 1964 391 27°20’ USD! iy 19 Sep 1964 533 26°27' 78°43’ 383-403 4 Mar 1965 598 24°47’ 80°26’ 183 15 Apr 1965 633 25°59’ I9A9’ 479-458 30 Jun 1965 647 26°16’ 79°43’ 520-549 15 Jul 1965 660 26°59’ 79°21’ 631 17 Jul 1965 661 27°07’ 79°32" 695-718 17 Jul 1965 664 DBS 79°22’ 567 17 Jul 1965 672 DSB) 79°03" 796 18 Jul 1965 679 25°56’ 78°09’ 595-711 20 Jul 1965 680 25°56’ 78°05’ 571-657 20 Jul 1965 692 26°34’ 78°25’ 329-421 21 Jul 1965 695 26°28’ 78°37' 535-575 22 Jul 1965 696 26°28’ 78°43’ 458-467 22 Jul 1965 704 26°29’ 78°40’ 275-366 22 Jul 1965 706 26°27’ 78°43’ 489-522 22 Jul 1965 707 26°27’ 78°40’ 514-586 22 Jul 1965 785 24°39" 80°40’ 205-210 16 Aug 1966 808 26°38’ 79°33’ 751 13 Sep 1966 835 24°22' 81°11’ 187-198 11 Jul 1967 859 23°54’ 81°57’ 1160-1190 21 Aug 1967 879 21°00 86°25’ 210 9 Sep 1967 889 20°55’ 86°28’ 175-220 10 Sep 1967 897 20°59’ 86°24" 210-290 10 Sep 1967 898 21°04’ 86°19" 340-360 10 Sep 1967 899 20°57’ 86°34’ 40-164 10 Sep 1967 936 26°35’ 79°20’ 600 1 Oct 1967 965 23°45’ 81°49" 1394-1399 1 Feb 1968 1012 23°35’ 79°33’ 509-531 14 Jun 1968 1125 26°45’ 79°05’ 900-950 17 Jun 1968 1312 26°38’ 79°02’ 505-527 31 Mar 1971 1314 26°52’ 79°11’ 532 ? R/V Gosnold (Gos) 2344 30°29’ 77°29.5' 882 ? 2383 30°56'24” 78°34'18" 869 27 Aug 1965 2384 30°54'24" 78°44'00" 820 27 Aug 1965 2385 30°57'12" 78°54'36" 379 27 Aug 1965 2387 31°14'48” 78°59'00" 530 27 Aug 1965 2413 30°14.5' 79°44.7' 585-622 2 Sep 1965 2414 30°16' 79°55.1' 494 3 Sep 1965 2461 28°14.4' 79°30.5' 850 15 Sep 1965 2469 29°43'12" 79°51'48" 640 16 Sep 1965 VOLUME 117, NUMBER 4 APPENDIX: Continued Station Latitude (°N) Longitude ((W) Depth (m) M/V, R/V Oregon and Oregon IT (O) 1328 24°33’ 83°34’ 366 1343 22°59' T9°1T' 457 1349 24°03’ 80°30’ 274 4940 20°30' 86°14’ 310-330 11703 30°28’ 79°51’ 494 11705 30°26 79°44! 640 11716 30°52’ 79°39' 576 11717 30°52’ 79°34’ 658 11725 31°44" 79°02' 543 11726 31°42’ 78°53’ 512 R/V Gyre CO4 27°28'06" 89°43'36" 1358-1518 R/V Pillsbury (P) 105 30°28’ 79°42’ 388-403 197 27°59’ 79°20' 567-586 209 26°59' 79°16' 550 592 215007 86°23’ 180 594 21°00.5' 86°23.0’ 330 596 24°42" 80°32’ 137 598 21°07' 86°21’ 155-205 881 13°20.8’ 61°02.5' 576-823 892 14°17’ 60°45'12” 1236-1313 954 16°55’ 62°43’ 686-1125 1146 20°08’ WG 1110-1189 M/V, R/V Silver Bay (SB) 440 D9 oa 79°15! 439-503 453 29°38’ 78°26! 879 5190 18°24’ 68°05’ 366 487 Date Jul 1955 Jul 1955 18 Jul 1955 12 Jun 1964 19 Jan 1972 19 Jan 1972 21 Jan 1972 21 Jan 1972 22 Jan 1972 12 Jan 1972 13 Apr 1984 27 Jul 1964 11 Aug 1964 12 Aug 1964 15 Mar 1968 15 May 1968 15 May 1968 15 May 1968 6 Jul 1969 6 Jul 1969 16 Jul 1969 14 Jun 1970 8 Jun 1958 12 Jun 1958 17 Oct 1963 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):488-504. 2004. A new species of the sea anemone Megalactis (Cnidaria: Anthozoa: Actiniaria: Actinodendridae) from Taiwan and designation of a neotype for the type species of the genus Adorian Ardelean and Daphne Gail Fautin (AA) Universitatea de Vest din Timisoara, Facultatea de Chimie-Biologie-Geografie, str. Pestalozzi, nr 16, Timisoara, 1900, Romania, e-mail: adorian@mynature.net; (DGF) Department of Ecology and Evolutionary Biology, and Division of Invertebrate Zoology, University of Kansas Natural History Museum and Biodiversity Research Center, University of Kansas, Lawrence, Kansas 66045 U.S.A., e-mail:fautin@ku.edu Abstract.—Megalactis comatus, new species, from Taiwan is the third species in this genus of sea anemones with highly branched tentacles. The others are M. hemprichii Ehrenberg, 1834, from the Red Sea, and M. griffithsi Saville-Kent, 1893, from the Great Barrier Reef. Size of nematocysts from acrospheres and column clearly separate M. comatus from the other species of Megalactis. One of us (A.A.) observed asexual blastulae in VM. comatus. This is the first record of asexual reproduction in the genus. Because type specimens of M. hemprichii have not been found and the original description cannot be used to distinguish this species from other species of Megalactis, we designate a neotype for the type species of the genus, M. hemprichii Ehrenberg, 1834. All the specimens of actinodendrids examined lacked basilar muscles; this calls into question the placement of family Actinodendridae among thenarian sea anemones. The family Actinodendridae is a group of three genera of exclusively tropical Indo- Pacific sea anemones: Actinodendron Blainville, 1830, Megalactis Ehrenberg, 1834, and Actinostephanus Kwietniewski, 1897. An actinodendrid has the oral disc drawn out into a number of branched ten- tacles that make it resemble a tree (Blain- ville 1830, 1834; Quoy & Gaimard 1833; Haddon 1898; Carlgren 1949). The last branches of tentacles terminate in acro- spheres that appear as white swellings of tissue; they are packed with nematocysts and spirocysts. Because the actinodendrids have been documented to sting humans badly (Saville-Kent 1893, Halstead 1970), knowledge of these animals is significant not only for taxonomy and phylogeny, but also for medicine and toxicology. Actinodendridae was considered by Carl- gren (1900, 1949) to belong to the supra- familial group Thenaria. Basilar muscles, which are structures “running along both sides of the base of the mesentery, close to the pedal disc” (Carlgren 1949, p. 8), were used by Carlgren (1899, 1900, 1942, 1949) to define two major groups in sea anemo- nes, Athenaria ““Nyantheae without basilar muscles” (Carlgren 1949, p. 21) and Then- aria ‘“‘Nyantheae with basilar muscles” (Carlgren 1949, p. 41). We did not find bas- ilar muscles in specimens of actinodendrids studied, which makes placement of Actin- odendridae among Thenaria questionable. The morphology of the tentacles of these sea anemones varies with environment, be- havior, and conditions of preservation. Al- though the number of species described in Actinodendridae is small, the lack of ter- minology for describing branched struc- tures and the enormous variety that can be found makes identification of species diffi- cult. In this paper we describe one species and redescribe two others of Megalactis, VOLUME 117, NUMBER 4 and standardize terminology for the branched tentacles of Actinodendridae. Actinodendrids are found in shallow wa- ter in sheltered places with sandy or muddy bottoms. Members of the genus Megalactis reportedly attach the pedal disc to hard sub- strata in sand or mud into which the anem- ones burrow (Saville-Kent 1893, Fishelson 1970). The new species of Megalactis de- scribed here lives in thickets of the scler- actinian coral Acropora in Taiwan; this might be the same species as that reported by den Hartog (1997) as an unidentified ac- tinodendrid living attached to coral branch- es in Indonesia. The description of Megalactis hemprichii Ehrenberg, 1834, the type species of the ge- nus, was diagnostic in the early 19th cen- tury. Mentioning only that a sea anemone had bipinnately branched tentacles was suf- ficient to distinguish M. hemprichii from all sea anemones known at that time. With the current state of knowledge, the original de- scription of M. hemprichii does not distin- guish it from other species of Megalactis: bipinnate disposition of the branches is ge- neric rather than specific. Type specimens of M. hemprichii Ehrenberg, 1834 have not been found (Klunzinger 1877, Favtin 2004 Hexacorallians of the World: http://hercules. kgs.ku.edu/hexacoral/anemone2/index.cfm). We designate a neotype for M. hemprichii in accordance with Article 75.3 of the In- ternational Code of Zoological Nomencla- ture (International Commission of Zoolog- ical Nomenclature 1999); no new species can be described within Megalactis without having a basis of comparison with the type species of the genus. In the course of this research, one of us (A.A.) found unusual gametogenic struc- tures in male specimens: nodes filled with spermatic packets that have a three-dimen- sional struc ire more voluminous than the thickened b nd typical for gametogenic tis- sue in members of Actiniaria. Male and small individuals of M. comatus had blas- tulae inferred to be of asexual origin among the mesenteries. This is the first record of 489 asexual reproduction in a member of Acti- nodendridae. The only female found con- tained no gametogenic nodes or blastulae among its mesenteries. Materials and Methods Specimens of the new species of Mega- lactis were investigated alive by diving and as preserved material; museum specimens of other species of Megalactis and Actino- dendron were investigated for internal mor- phology and histology (Table 1); results from this study are based on examination of more than 400 museum lots and photo- graphic documents of actinodendrids. Animals were recorded in situ on Hi8 videotape using a CanonES6000A video camera in an Amphibico underwater hous- ing. Live material was collected underwater by hand using gloves for protection against stinging. Geographic coordinates were read with an Eagle 12-channel GPS receiver at the point of collection. The animals were kept in aquaria with running seawater for two days; no food was given. Photographs were made in the aquarium using a Nikon Coolpix 950 digital camera. Archived vid- eotapes an 1 photographs are in the collec- tion of the Division of Invertebrate Zoolo- gy, University of Kansas Natural History Museum (KUNHM). Specimens were re- laxed with magnesium sulfate in seawater, then preserved in 10% seawater formalin. After at least two months, they were trans- ferred to 10% freshwater formalin. Undischarged cnidae from preserved an- imals were examined at 1000 in squash preparations using a light microscope equipped with differential interference op- tics. Squash preparations were made from acrospheres, the oral face of the main branches of the tentacles, the proximal, middle, and distal column, the actinophar- ynx, and the mesenterial filaments. Sigma Scan Pro version 4.01.003 measurement software was used to measure the length and the width of undischarged capsules pro- jected onto a Summa Sketch digitizing tab- 490 let (Summagraphics). Sampling nemato- cysts was done following the recommen- dations of Williams (1996). For histology, tissue was embedded in Paraplast, sectioned at 9 «wm, and stained with Heidenhain’s Azan or hematoxylin and eosin (Presnell & Schreibman 1997). Serial sections for three-dimensional reconstruc- tion were obtained from mesenterial struc- tures, column, and two entire juvenile in- dividuals. Images were obtained using a Ni- kon Coolpix 995 digital camera connected to an Olympus microscope through an Op- tem eyepiece digital coupler. Serial images were aligned manually using layers in Ado- be Photoshop. Three-dimensional recon- struction was done using the software Vay- tek VoxBlast Version 3.0 Light (http:// www.vaytek.com/). In the following discussion, as is conven- tional in sea anemones, the proximal direc- tion is toward the pedal disc and distal is the opposite. Tentacles are arranged in four cycles. Branches of the tentacles are or- dered by how close they are to the oral disc: a branch arising from the oral disc is con- sidered to be of the first order; a branch that ramifies from a branch of the first order is of the second order, etc. (Fig. 1). Abbreviations: CAS, California Acade- my of Sciences, San Francisco, CA, USA; KUNHM, University of Kansas Natural History Museum, Lawrence, KS, USA; NNM, Nationaal Natuurhistorisch Museum, Leiden, The Netherlands; NMNS, National Museum of Natural Sciences, Taichung, Taiwan; TAUI, Zoological Museum, Tel- Aviv University, Tel-Aviv, Israel. Taxonomic Account Order Actiniaria Family Actinodendridae Haddon, 1898 Diagnosis (modified from Carlgren 1949; see remarks below).—Limbus not well de- fined. No marginal sphincter muscle. Fosse absent. Up to 48 branched tentacles cycli- cally arranged. Terminal branches of tenta- cles with acrospheres. Two or more well de- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ramifications in a tentacle of the first cy- cle; view looking down a tentacle, 1.e., proximally, to the oral disc. The arrow labeled “‘d”’ indicates the dis- tal direction, that labeled “‘p” indicates the proximal direction. Abbreviations: enI, endocoel of the first cy- cle; enlII, endocoel of the second cycle; ex, exocoel; od, oral disc; TI, first order branch; TIl-a, lateral sec- ondary order branch; TII-b, oral face secondary order branch; TIIJ, third order branch; TIV, fourth order branch. veloped siphonoglyphs. Twenty-four pairs of mesenteries, all or almost all perfect and, apart from the directives, fertile. Retractor muscles diffuse, broad, band-like. Cnidom: spirocysts, basitrichs. Remarks.—Carlgren (1949, p. 67) indi- cated a “‘well developed disc” and “pairs of mesenteries up to 48” for Actinodendri- dae. Pedal disc size varies greatly: that of some specimens is wide, but that of others is narrow with a limbus that is hard to rec- ognize. None of the specimens studied had more than 24 pairs of mesenteries. Carlgren (1949) asserted that parietobasilar and bas- ilar muscles are distinct in actinodendrids, but we found them to be absent in all gen- era of the family. Genera.—Actinodendron Blainville, VOLUME 117, NUMBER 4 1830, type genus; Megalactis Ehrenberg, 1834; Actinostephanus Kwietniewski, 1897. Genus Megalactis Ehrenberg, 1834 Diagnosis (modified from Carlgren, 1949; see remarks below).—Actinodendri- dae with ramified tentacles having second- order branches arranged bipinnately. Last order branches with capitate acrospheres. Remarks.—Carlgren (1949, p. 68) stated that Megalactis has “the oral face of the arms [branches of the first order] free from tentacles.” All specimens of Megalactis we studied had two to three second-order branches on the oral face of branches of the first order. Carlgren (1949, p. 68) stated in his diagnosis for Megalactis that “the ulti- mate branches of the tentacles are simple and pointed.”’ One of us (A.A.) found spec- imens of Megalactis that have capitate ter- minal tentacles. Species.—Megalactis hemprichii Ehren- berg, 1834, type species by monotypy, Ras Kafil, Red Sea; Megalactis griffithsi Sa- ville-Kent, 1893, Warrior Reef, Torres Strait, Great Barrier Reef, 9°30’S, 143°06’E. Coordinates from Gazetteer of Australia, 2001 (http://www.ga.gov.au/). Megalactis hemprichii Ehrenberg, 1834 Megalactis Hemprichii Ehrenberg, 1834: 263 (original description). Megalactis Hemprichii Ehrenberg: Milne Edwards & Haime, 1851:11. Actineria Hemprichii Ehrb.: Klunzinger, 1877:90-91. Megalactis Hemprichii Ehr.: Andres, 1883: 308-309. Megalactis Hemprichii E.: Carlgren, 1899: 14. Megalactis Hemprichii Klunzinger: Delage & Hérouard, 1901:539. Megalactis hemprichii Ehrenberg, Carlgren, 1949:68. Megalactis hemprichi Ehrenberg: Fishel- son, 1970:109. 1834: 491 non Megalactis hemprichii Ehrenberg, 1834: Cutress & Arneson, 1987:53—62. Description.—Dimensions: column di- ameter 14—26 mm distally and 14-15 mm in the middle; pedal disc diameter 5—9 mm; column length 23—41 mm; oral disc diam- eter 21—23 mm; tentacles of the first cycle 45—51 mm long; tentacles of the fourth cy- cle 10-11 mm long. Color: Of live specimens unknown. Pre- served specimens beige to pale yellow. Column: Pyramidal to elongate with nar- row pedal disc; limbus hardly recognizable (Fig. 2A). Column smooth and mesenterial insertions clearly visible through column in relaxed specimens. In contracted speci- mens, column with circumferential folds (Fig. 2A). Oral disc and tentacles: Oral disc narrow. In preserved specimens, mesenterial inser- tions on oral disc visible as dark lines; ra- dial bumps near mouth mainly on exocoelic intervals (Fig. 2D). Forty-eight tentacles ar- rayed in four cycles (6 + 6 + 12 + 24). Tentacles of first, second, and third cycles ramified in branches of up to three orders. Proximal secondary branches of first, sec- ond, and third tentacle cycles short (Fig. 2B). Branches regularly oriented. Secondary branches pinnately disposed in one row on each side of a branch of the first order (Fig. 2E). Up to two long and broad secondary branches on aboral side of primary branch- es of tentacles belonging to first, second, and third cycles (Fig. 2E). Up to 45 sec- ondary branches on tentacles of first and second cycle; up to 25 secondary branches on tentacles of third cycles; up to 11 sec- ondary branches on tentacles of fourth cy- cle. Branches of last order relatively long. Large, round acrospheres. Internal structure: Actinopharynx short with two deep siphonoglyphs. Twenty-four pairs of mesenteries in three cycles (6 + 6 + 12); first two cycles usually perfect. Oral stomata large; marginal stomata very small. Retractor muscles diffuse and strong. Fila- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Megalactis hemprichii, external morphology (TAUI 21560). A, Aboral view of entire animal. B, Crown of tentacles, oral view of entire animal. C, Regenerated tentacles (TAUI 7812). Arrows indicate tentacles with missing secondary branches. D, Detail of oral disc and mouth. E. First order branch. Abbreviations: b, radial bumps on exocoelic intervals; co, column; od, oral disc; pd, pedal disc; s, siphonoglyph; TI, branch of the first order; TI, short proximal secondary branch; TIl-a, lateral secondary order branch; TII-b, oral face secondary order branch. Scale bars: A, B = 15 mm; C = 10 mm; D, E = 5 mm. ments absent on mesenteries proximally. Parietobasilar and basilar muscles not seen. Gonochoric. The only specimen sectioned was female (Fig. 3). Cnidae: Basitrichs densest in acrospher- es. Cnidom: spirocysts and basitrichs (Fig. 4). Measurements in Table 2. Type specimen and _ locality.—Neotype TAUI 31623, Red Sea, Gulf of Aqaba, Ei- lat, 29°30'N, 34°55’E. Coordinates from GEOnet Names Server of National Imagery and Mapping Agency (http://www.nima. mil). Voucher specimens.—Table 1. Megalactis comatus, new species Figs. 4—10 Description.—Dimensions: Diameter of column 2—38 mm distally and 5—21 mm in the middle, of pedal disc 2—8 mm; column length 8-26 mm; tentacles of the first cycle 9-11 mm long; tentacles of the fourth cycle 2-3 mm long; oral disc diameter 13—25 mm; tentacle crown diameter 5O—100 mm. Color: In live specimens, oral disc and tentacle color ranges from dark brown to pale orange or pink. Tentacles translucent, without pattern (Fig. 5). Oral disc with ra- VOLUME 117, NUMBER 4 Fig. 3. Megalactis hemprichii, histology (KUNHM 001948). Abbreviations: f, filament; m, me- soglea; 0, ova; r, retractor. Scale bar = 1 mm. dial rows of white spots aligned along ex- ocoelic spaces; radial spots may spread lat- erally onto adjacent endocoelic spaces (Fig. 5F). Insertions of mesenteries on oral disc visible as lighter lines (Fig. 5F). Column beige to white; distal column translucent tinged with brown or pale orange. Female gametogenic tissue purple and male game- togenic tissue white (Oscar Chen, currently at Institute of Oceanography, National Tai- wan University, pers. comm.). Preserved 493 specimens beige, column paler than oral disc or crown. Column: Pyramidal to elongate with a narrow pedal disc; limbus hardly recogniz- able (Fig. 5C). Pedal disc and proximal col- umn adhesive with strong ripples of ecto- dermal tissue in preserved specimens. Cir- cumferential folds resulting from contrac- tion of the column between pedal region and distal-most third of column (Fig. 5C). Distal-most third of column thinner and smoother than proximal column. Mesenter- ial insertions clearly visible through col- umn. Oral disc and tentacles: Oral disc narrow. Mesenterial insertions on oral disc visible as light lines in live specimens. Radial bumps close to mouth mainly on exocoelic intervals. Appearance of tentacle crown shaggy be- cause of numerous branches not regularly oriented (Fig. 5A, E). Forty-eight tentacles arrayed in four cycles (6 + 6 + 12 + 24). Tentacles of first, second, and third cycles ramified in branches of up to four orders. Proximal secondary branches of first, sec- ond, and third tentacle cycles long. Secondary branches pinnately disposed in One row on each side of a primary branch (Fig. 5D). On contracted tentacles, pinnate arrangement unclear: secondary branches appear to be arranged in two or more lateral rows on each side of a primary branch. Some large secondary branches occur on aboral side of primary branches of tentacles belonging to first, second, and third cycles. Secondary branches variable in length. Up to 48 secondary branches on each tentacle of first and second cycle; up to 40 second- ary branches on each tentacle of third cycle; up to 12 on each tentacle of fourth cycle. Branches of last order relatively long, ter- minate in small round to pointed acro- spheres. Internal structure and histology: Actino- pharynx short, with two deep siphono- glyphs (two specimens had three: Fig. 6), each connected to a pair of directive mes- enteries. 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N u (wi!) N u adXy oepiup anssly, YIPIM osue1 x YSU] osury UIPIM osuel x YSU] osuryY sorsadg nyoiuduay Ww SnIDUOD SIvIDsap “Hb “SI Ul p YOUMIseg 5D “dp “Stq Ul € YOUseg ‘gq “Cp ‘Sty Ul Z Youse ‘Wh “Shy Ul POILNSNI]I SI [| YOLMISeg ‘poyeSNsoAUt s[enprIAIpur Jo Joquinu pue jsAooyeuIOU Jo adA} v BUTUTLJUOD s[eNPIAIPUT JO JIoqUINU UsEMJeq ORI = N ‘poiNsvoul s}sAoo}eUWIOU JO Joquinu = u ‘sosoyjuoied ul poyeorpul oie poinsvoul o10M s}SAoO}eUIOU Op UY) DIOU YOTYA ut o[dures v IOJ sJUDWOINSeOU JO OSVIOAL :S}SADOJLUIOU JO 9ZIS—Z 2[GeL 496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF V ASHINGTON - eeet “See hah hh) Lh ALR ee AA RAee tye RS Fig. 4. Cnidae. Basitrichs of acrospheres (A, B), and middle column (C, D, E). Spirocyst (F). Image of a squash preparation from an acrosphere showing numerous basitrichs (G). Long basitrich (H) from filaments of M. hemprichii (TAUI 21560). Scale bars = 10 pm: Fig. 5. Megalactis comatus, external morphology. A, Crown of tentacles, oral view of entire animal. B, Tentacles of the fourth cycle oriented toward substrate. C, Column in a preserved specimen. Arrow indicates deep ripples in the pedal disc region. D, Secondary branches in bipinnate arrangement. E, Long proximal secondary branches (arrow). EK Oral disc. Scale bars = 10 mm. VOLUME 117, NUMBER 4 Fig. 6. Megalactis comatus, internal anatomy of a specimen with three siphonoglyphs (KUNHM 1664): transverse view. Abbreviations: f, filaments; g, game- togenic tissue; s, siphonoglyphs. Scale bar = 5 mm. in three cycles (6 + 6 + 12); first two cy- cles usually perfect. Stomata not seen. Re- tractor muscles diffuse and strong (Fig. 7A— C). Filaments absent on mesenteries proxi- mally. Parietobasilar and basilar muscles not seen. Gonochoric: Mesenteries in male speci- mens have nodes filled with spermatic packets. Each spermatic node formed through plications of mesentery along oral- aboral axis; node digitiform, closed on one side of mesentery and open on the other (Fig. 8). The only female specimen found had ova in arrangement typical of Actini- aria. Cnidae: Largest and densest basitrichs in acrospheres (Fig. 4G). Cnidom: spirocysts and basitrichs (Fig. 4). Measurements in Ta- blew Type specimens and locality.—Holotype KUNHM 1663, Pacific Ocean, Taiwan, Henchun Peninsula, Nanwan, power plant water intake basin, DNS 2 IN| 120°45.22’E. See Table 1 for paratype and voucher specimens. Etymology.—tThe epithet comatus, which means “with long hair, shaggy” in Latin (Brown 1978), refers to the hairy and irreg- ular aspect of the tentacle crown in this spe- cies. 497 Natural history.—Animals live in sym- biosis with zooxanthellae. We found speci- mens of M. comatus in water a few centi- meters to 4 m deep. Each specimen of M. comatus attaches to a coral skeleton with its pedal disc and proximal part of the col- umn. The color, similar to that of brown and red algae, and shaggy aspect of the tentacle crown make specimens difficult to find even when abundant. The water intake basin of the nuclear power plant from which the type specimens were collected was 18 years old at the time. It was inhabited by a large number of spec- imens of M. comatus and other species of sea anemones tentatively identified as Bol- oceroides memurrichi (Kwietniewski, 1898), Thalassianthus sp., and a species of family Actiniidae. The initially large pop- ulation of M. comatus had decreased in the previous decade (Dr. Keryea Soong, Na- tional Sun Yat-sen University, Kaohsiung, Taiwan, and Oscar Chen, pers. comm.), and has been replaced by the actinid. One of us (A.A.) found in nature speci- mens of M. comatus that appeared to be undergoing transverse fission; several spec- imens had their columns strongly constrict- ed. One specimen, KUNHM 1667, lacks a pedal disc, having a circular opening into the gastrovascular cavity (Fig. 9A, B); specimens KUNHM 1670 and KUNHM 1251 are sacciform, lack tentacles, and have a small opening rather than an oral disc (Fig. 9C), or have small undeveloped ten- tacles (Fig. 9D). Specimens of M. comatus are easy to collect, so it is not likely that the pedal or oral disc of a specimen was torn off during collection as can happen in other sea anemones that attach or are deeply buried in the substrate. Further observations in aquaria should be made to confirm trans- verse fission. Some sectioned individuals of M. coma- tus, including males and infertile individu- als, had blastulae among their mesenteries (Fig. 10). These larvae contained syncitial blastoderm (solid blastula or stereoblastula in Fautin et al. 1992) and were similar to 498 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Megalactis comatus, histology. A, Retractor muscle. B, Mesenteries. C, Detail of retractor muscle. D, Detail of mesenterial filament. E, Male gonads with the beginning of spermatic node. Abbreviations: c, column; cg, cnido-glandular tract; ct, ciliated tract; e, endoderm; ec, ectoderm; f, filament; gl, glandular cells; m, mesoglea; sn, spermatic node; sp, spermatic packet; zo, zooxanthellae. Scale bars: A, B, D, E = 100 pm; C = 50 pm. those depicted in Yanagi et al. (1999). A.A. vals (Fig. 10D, E). Juvenile stages were not also found larvae in an individual lacking found in histological sections. tentacles presumably because of transverse fission. Some larvae showed incipient blas- Discussion topores, indicating an early gastrula stage (Fig. 10B, C). In some larvae, the outer lay- Systematics.—Type specimens of Mega- er contained nematocysts at regular inter- lactis hemprichii have not been found VOLUME 117, NUMBER 4 Fig. 8. Three-dimensional reconstruction of sper- matic nodes in mesenteries of M. comatus from 20 serial slices each 9 wm thick. Abbreviations: f, fila- ment; 1, retractor muscle; sp, spermatic packet; sn, spermatic node. Scale bar = 0.5 mm. (Klunzinger 1877, Fautin 2004 Hexacoral- lians of the World: http://hercules.kgs.ku. edu/hexacoral/anemone2/index.cfm). To typify the genus, we designate a neotype for M. hemprichii. Specimens of M. hemprichii from the type locality of Ras Kafil in the Red Sea bordering Sinai (now part of Egypt) were unavailable and collection in this region is not feasible. We designate as neotype specimen TAUI 31623 from the Gulf of Aqaba in the Red Sea, a locality “as near as practicable from the original type locality” (Art. 75.3.6, International Commission of Zoological Nomenclature 1999). Because of poor descriptions and com- plex morphology of the tentacles, species of Megalactis are difficult to distinguish from each other. Ehrenberg’s (1834) description of M. hemprichii includes a very brief Latin 499 Fig. 9. B, Column without pedal disc KUNHM 1667. C, Specimen without oral disc (KUNHM 1670). D, Spec- imen with short tentacles (KUNHM 1668). Scale bars = 5 mm. Megalactis comatus, transverse fission. A, description and no illustration. The only il- lustration for M. hemprichii in Klunzinger (1877) is based on drawings left by Ehren- berg. Subsequent references to M. hempri- chii are translations of the original descrip- tion (Milne-Edwards 1857, Andres 1883, Delage & Hérouard 1901) and a distribu- tion record (Fishelson 1970). The specimen identified as M. hemprichii depicted in fig- ure 2A of Cutress & Arneson (1987) has secondary branches not bipinnately dis- posed, and therefore is probably a specimen of Actinodendron. Differences and similarities between the species of Megalactis are presented in Table 3. Type specimens of all the species de- scribed by Saville-Kent (1893), if they ex- isted, have not been located (Fautin 2004 Hexacorallians of the World: http://hercules. kgs.ku.edu/hexacoral/anemone2/index. cfm). The photograph and description of the color pattern of the oral disc in M. griffithsi PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON = : . on 78 Ne Lats paths S35) is > Fig. 10. Megalactis comatus, asexual larvae. A. Larva (arrow) among mesenteries. B. Late blastula (arrow). C. Three-dimensional reconstruction of a larva from 17 serial slides each 9 pm thick. D, E. Larva with nema- tocysts. Abbreviations: b, blastopore; c, column wall; m, mesentery; n, nematocyst. Scale bars = 0.25 mm. Saville-Kent, 1893, can be used to identify specimens and distinguish this species from M. comatus. Haddon (1898) used the shape of acros- pheres to distinguish M. griffithsi from M. hemprichii: clubbed for M. hemprichii and pointed for M. griffithsi. The shape of ac- rospheres cannot be used as a diagnostic character in either living or preserved spec- imens of Megalactis because it is influ- enced by behavior and preservation. It is common to find a museum specimen that has acrospheres of both shapes. Nematocysts from the acrospheres and middle column differ in size between spec- imens of M. comatus and M. griffithsi. The ratio between length and width of nemato- cysts shows a clear difference between the two species (Fig. 11). Three specimens of M. hemprichii from the Red Sea have a similar gross morphology to specimens of M. griffithsi but the nematocysts of the ac- rospheres have size values close to those of M. comatus. The nematocysts in the middle column of M. comatus are larger than those in M. hemprichii. The number of tentacles for all species of Megalactis is given as 10+10 for M. hemprichii by Ehrenberg (1834), Milne-Ed- wards (1857), Andres (1883), Delage & Hérouard (1901), and Klunzinger (1877) and 6+6+12 for M. griffithsi by Saville- Kent (1893) and Haddon (1898). We agree with Haddon (1898) that the number of ten- tacles indicated by Ehrenberg (1834) for M. hemprichii might be an individual peculiar- ity. One of the three specimens of M. hem- prichii studied (TAUI 7812) had only 41 VOLUME 117, NUMBER 4 missing data. Table 3.—Diagnostic characters of species of Megalactis. ? M. hemprichii M. hemprichii original description neotype M. griffithsi M. comatus Species/character Regular Regular Regular Irregular, shaggy Tentacle crown aspect Secondary branches Relatively short, usually ?, may be constricted proxi- Elongated, usually constrict- Relatively short, constricted constricted proximally Up to 45 Short mally proximally Up to 35 Short ed proximally Up to 48 Number secondary branches Proximal secondary branches Distal secondary branches Long to very long Present Present Present No pattern Complex pattern of radiat- Rows of white spots Oral disc pattern of live specimens ing lines and alternating dark and white regions Oral disc and tentacles Light brown Oral disc brick red and Oral disc and tentacles pink Color of live specimens gray; tentacles pale pink; column white brown or green; column beige to brown; column white, beige 501 width Fig. 11. Length in pm of basitrichs from acros- pheres of M. comatus (gray dots) and M. griffithsi (black dots). In the region delimited by the rectangle are measurements of nematocysts from the region where acrosphere (a) meets peduncle (p); open circles represent basitrichs type 2 (see Fig. 4B). tentacles, all of which showed signs of re- generation—lacking secondary branches, or having branches not bipinnately arranged (Fig. 2C). It is possible that M. hemprichii has predators that feed on its tentacles. We infer that in both previously described spe- cies of Megalactis, the fourth cycle of ten- tacles was overlooked, being probably con- sidered secondary branches on the adjacent tentacles. In situ, members of Actinoden- dridae usually orient the tentacles of the fourth cycle towards the substrate. All spec- imens of actinodendrids studied, including those belonging to Megalactis, had a typical tentacle arrangement in multiples of six (6 ae ©) ar IID ap Dab. Because we did not find basilar muscles in specimens of Actinodendron plumosum Haddon, 1898, A. glomeratum Haddon, 1898, Megalactis griffithsii Saville-Kent, 1893, and M. comatus, the position of fam- ily Actinodendridae among Thenaria as de- fined by Carlgren (1899, 1900, 1942, 1949) is questionable. It is possible that basilar muscles are reduced in size or have been lost in the family Actinodendridae; basilar muscles are reduced or absent in many bur- rowing sea anemones (Carlgren 1949, Daly 502 et al. 2002). Basilar muscles are absent in the thenarian family Aliciidae. Another ex- planation may be that the basilar muscles were not present in the ancestral lineage of Actinodendridae and this family does not belong to Thenaria. Spermatic nodes.—We report for the first time spermatic nodes in Actiniaria. Hyman (1940, p. 583) stated that generally the ga- metogenic tissues in actiniarians “occur as thickened bands on the septa behind the septal filaments.” Atypical organization of gametogenic tissue is reported in the hex- acorallian groups Actiniaria (Excoffon & Zamponi 1999), Zoanthidea (Ryland 2000), and Scleractinia (Harrison & Wallace 1990). The most similar structure to sper- matic nodes in M. comatus are the ““gonadal nodes” reported by Ryland (2000) that are lens-shaped folds in the perfect mesenteries of females of the zoanthid Parazoanthus anguicomus and of a male of P. axinellae. Spermatophores were described by Excof- fon & Zamponi (1999) in the sea anemone Sagartia troglodytes. The spermatic nodes in M. comatus are not stalked like the sper- matophores in S. troglodytes but have a three-dimensional structure more developed than a simple fold of the mesentery like the ‘““gonadal nodes’’ reported by Ryland (2000). Excoffon & Zamponi (1999) re- ported that spermatozoa in S. troglodytes were released from spermatophores through the stalk, the region by which the sper- matophores are attached to the mesenteries, and the mesogleal wall of the spermato- phores is continuous with that of adjacent mesentery. Thus, like spermatic nodes, spermatophores must develop from folds of mesenteries through evagination. We agree with Ryland (2000) that one function of the “gonadal nodes”’ is to increase the number of “gonadal packets” with no increase in length of body. Asexual larvae.—The origin of larvae found in the coelenteron of some sea anem- ones is uncertain (Fautin 2002). Chia & Rostron (1970) assumed that the larvae in- side Actinia equina (Linnaeus, 1758) were PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sexually produced, but Carter & Thorp (1979) found this to be unlikely because the phenotypes were identical between a brood and the adult host. In fungiid corals, any tissue fragment in the coelenteron is able to transform into a larva (Kramarsky-Winter & Loya 1996). Because one of us (A. A.) found blastulae in immature and male in- dividuals of M. comatus, they are consid- ered to be of asexual origin. Acknowledgments We especially thank Dr. Keryea Soong, National Sun Yat-sen University, Kao- hsiung, Taiwan, for bringing the specimens to our attention. His graduate student Oscar Chen was A.A.’s buddy and showed the an- imals in situ. Dr. M. Daly, and H.-R. Cha critically read the manuscript and made suggestions. W. N. Eschmeyer (CAS) pro- vided advice on designating a neotype. Thanks also go to Dr. Y. Benayahu and A. Shlagman for providing specimens from the collection of Zoological Museum, TAUI. N. E. Chadwick and G. Ayalon (The Interuni- versity Institute of Eilat, Israel) and Fan Tung Yung and Tsai Wan Hsu (National Taiwan University, Taiwan) collected or provided specimens used in this study. Sug- gestions from an anonymous reviewer im- proved the manuscript. This research was supported by NSF grants DEB-9521819 and DEB-9978106 in the PEET program to D.G.E and OCE-0003970 to D.G.EF and R. W. Buddemeier. Literature Cited Andres, A. 1883. Le Attinie (Monografia). Coi Tipi der Salviucci, Roma, 460 pp. de Blainville, H. M. 1830. Dictionnaire des Sciences Naturelles, vol. 60. Levrault, Paris, 631 pp. . 1834. Manuel d’ Actinologie ou de Zoophytol- ogie. Levrault, Paris, 644 pp. Brown, R. W. 1978. Composition of Scientific Words. Smithsonian Institution Press, Washington D.C., 882 pp. Carlgren, O. 1899. Zoantharien—Hamburger Magal- haensische Sammelreise 4:1—48. . 1900. Ostafrikanische Actinien. Gesammelt von Herrn Dr. FE Stuhlmann 1888 und 1889.— VOLUME 117, NUMBER 4 Mittheilungen aus dem Naturhistorischen Mu- seum 17:21—144. . 1942. Actiniaria Il—Danish Ingolf-Expedion 5:1-92. . 1949. 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Reproduction in British zoanthids, and an unusual process in Parazoanthus angui- comus.—Journal of the Marine Biological As- sociation of the United Kingdom 80:943—-944. 504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Saville-Kent, W. 1893. The Great Barrier Reef of Aus- Yanagi, K., S. Segawa, & K. Tsuchia. 1999. Early de- tralia; Its Products and Potentialities. WH Allen velopment of young brooded in the enteron of & Co., London, 387 pp. the beadlet sea anemone Actinia equina (Antho- Williams, R. B. 1996. Measurements of cnidae from zoa: Actiniaria) from Japan.—Invertebrate Re- sea anemones (Cnidaria: Actiniaria): statistical production and Development 35:1—8. parameters and taxonomic relevance.—Scientia Marina 60:339-351. Associate Editor: Stephen L. Gardiner PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):505—513. 2004. A new genus and new species of crab of the family Xanthidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the southwestern Gulf of Mexico Ana Rosa Vazquez—Bader and Adolfo Gracia (ARVB), (AG) Instituto de Ciencias del Mar y Limnologia, UNAM, Apdo. Postal 70-305, México, D.F, 04510, Mexico, e-mail: ana_rosav@yahoo.com.mx; gracia@mar.icmyl.unam.mx Abstract.—A new genus and new species belonging to the Euxanthinae sub- family, Batodaeus adanad, are described from the southwestern part of the Gulf of Mexico. Whereas the new genus is similar to Monodaeus Guinot, 1967 in the carapace ornamentation and shape of pereopods, it differs in the structure of the abdomen and male telson, sternoabdominal cavity, and shape and or- namentation of the first gonopod. Resumen.—Se describe un nuevo género y una nueva especie perteneciente a la subfamilia Euxanthinae, para el suroeste del Golfo de México. Este género nuevo es similar a Monodaeus Guinot, 1967, en la ornamentacion del capar- azon y forma de los pereopodos; sin embargo, difiere de éste en la forma y tamano del abdomen, telson y cavidad esterno-abdominal, asi como en la es- tructura de los apéndices sexuales. In the course of deep-water biodiversity surveys in the Cayo Arcas and west side of Triangulos in the southwestern part of the Gulf of Mexico, specimens of an unusual species of xanthid crab were obtained. Al- though superficially similar to species of Monodaeus Guinot, 1967, they possess sev- eral atypical features that suggest other- wise. They are here described as a new ge- nus and new species. Material and Methods Specimens were collected in 1998 during surveys investigating the marine fauna in the deep southwestern Gulf of Mexico, cruise BATO (Biota de los Arrecifes, de la Plataforma y Talud continental en el no- roeste del Banco de Campeche), carried out on board the R/V Justo Sierra by the Insti- tuto de Ciencias del Mar y Limnologia, UNAM. The samples were caught using a semicommercial otter trawl. The material was deposited in the refer- ence collection of the Instituto de Biologia, UNAM (CNCR). Measurements listed are in millimeters (mm): total carapace length (CL) and carapace width (CW). Batodaeus, new genus Diagnosis.—Carapace subhexagonal, broader than long; dorsal surface convex and granulated. Regions in male well de- marcated especially in the anterior half, front strongly deflexed; inner orbital teeth conspicuous; thoracic sternum relatively narrow. Anterolateral margins armed with 4 teeth (excluding outer orbital tooth), sube- qual in size to posterolateral margins; pos- terior margin of epistome triangular, median part depressed, with distinct median fissure, and a pair of shallow but clearly visible lat- eral notches; preorbital and postorbital lobes conspicuous, granulated. Incomplete endostomial ridges present. Basal antennal segment long, subcylindrical, almost touch- ing front, not filling space between front and inner orbital teeth; a small gap between basal antennal segment and suborbital mar- 506 gin. Third maxillipeds not filling buccal cavity; merus with deep, rounded depres- sion near mesial margin. A longitudinal tu- berculated ridge just below suture separat- ing subhepatic and subterygostomian re- gions. Pereopods 2-5 long, slender, with conspicuous spines on upper margin of merus and carpus. Sternoabdominal cavity relatively narrow and deep; thoracic sternite 4 with median part slightly raised, with short longitudinal furrow. Abdomen long, completely covering sternoabdominal cavi- ty. Second abdominal segment in each sex not reaching coxae of fifth pereopod, leav- ing a small portion of sternite 8 visible. Sternite 7 covering part of penis groove on sternite 8. First gonopod slender, slightly in- curved; a row of spines on mesial margin. Second gonopod very short, sigmoid, ter- minal process curved with short setae on distal end. Type species.—Batodaeus adanad, new species, by present designation. Gender.—Masculine. Etymology.—The name is a combination of Bato, the name of the cruise during which it was collected, and “‘daeus”’ to in- dicate its similarity with the genus Mono- daeus. Remarks.—TYhe taxonomic position of Batodaeus is uncertain. Although the spec- imens studied here share some characteris- tics with the subfamily Actaeinae in having sternite 4 with a longitudinal furrow and the male abdomen with locking mechanism on sternite 5, they also share some features with the Xanthinae in having the carapace with four teeth and with thoracic sternite 4 being rather long (see Serene, 1984). In fact, Batodaeus more closely resembles the euxanthinid genus Monodaeus Guinot, 1967 in the following characters: regions of the carapace well demarcated; hepatic re- gion inflated; presence of four anterolateral teeth; surface of carapace with several strong granules; basal antennal segment just touching the front; anterior border of buccal cavity with a conspicuous crest; third max- illipeds not completely closing buccal cav- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ity and merus with prominent distolateral angle; and thoracic sternite 4 with a median longitudinal furrow. However, the carapace of Batodaeus is more convex and the re- gions are less well demarcated and with acute spines, particularly on the hepatic re- gion; the posterolateral margins are sube- qual in length to the anterolateral; the front is more advanced, with a deep sulcus; the preorbital lobe is conspicuous; the sternoab- dominal cavity is relatively deeper and nar- rower; thoracic sternite 4 has a shallow but distinct longitudinal median furrow; the ab- domen is relatively narrow and longer; and the first male gonopod is long, slender and slightly incurved with few spines, not se- tose or stout as in Monodaeus. The bathymetric and geographic distri- bution of these two genera is also different. Monodaeus (type species Xantho couchii Bell, 1851) at present contains eight spe- cies: M. arnaudi Guinot & Macpherson, 1988; M. couchii (Bell, 1851); M. cristu- latus Guinot & Macpherson, 1988; M. guin- otae Forest, 1972; M. pettersoni Garth, 1985; M. rectifrons (Crosnier, 1967); M. rouxi (Capart, 1851); and M. tuberculidens (Rathbun, 1911). They all occur in the Western Indian Ocean, the Eastern Atlantic, the Mediterranean Sea and the Eastern Pa- cific, from 20 to 500 meters. Batodaeus adanad, however, was collected in the Western Atlantic at depths from 160 to 250 m. In comparison with Medaeops Guinot, 1967 (only known from the Indo-West Pa- cific), Batodaeus has a carapace which has the regions less inflated, the merus of third maxilliped with a prominent distolateral an- gle; the pereopods are more slender and longer, the thoracic sternum is not flat; and the first male gonopod lacks setae. It differs from the superficially similar Indo-West Pa- cific genus Alainodaeus Davie, 1992 in having a carapace which is subhexagonal, not ovoid; a straight frontal margin; a rounded telson (triangular in Alainodaeus) and an almost straight first male gonopod. Batodaeus, for the moment, is tentatively VOLUME 117, NUMBER 4 placed in the Euxanthinae, as it seems to fit there considering the primary character of the subfamily stated by Seréne (1984): the form of the anterolateral margin which has the anterior part gradually sloping down- wards via subhepatic region to meet the in- fraorbital margin. The similarity to Mono- daeus confirms that it can be placed as Eux- anthinae, even though the subfamily is now poorly defined. Batodaeus adanad, new species Figs. 1-4 Material examined.—Holotype: 1 ¢ CNCR (21023), 16.6 mm X 23.2 mm; Sta. 9, 22°17.19'N, 91°43.08’W (Banco de Campeche, off Cayo Arenas), 251 m, 23 May 1999. Allotype: 1 2 CNCR (22509), 11.8 mm X 16.6 mm, Sta. 9, 22°17.19'N, 91°43.08’W (Banco de Campeche, off Cayo Arenas), 251 m, 23 May 1999. Paratype: 1 2 CNCR (21024), 12.1 mm X 17.6 mm, off Sta. 8, 22°13.72’N, 91°46.64’W (Banco de Campeche, off Cayo Arenas), 162 m, 23 May 1999. Description.—Carapace (Fig. 1A) sub- hexagonal, about 1.4 to 1.5 times broader than long; dorsal surface convex, granulat- ed, granules coarse, more abundant on an- terolateral and posterolateral margins, with sparse short setae on frontal, hepatic and protogastric regions. Regions in male well demarcated, especially in the anterior half; orbital region with small, acute spines; he- patic region inflated, with 3—4 rows of spines; a depression between cardiac and gastric region; meso and urogastric regions less granulated; front straight, strongly de- flexed; margin dentate, at most %4 as long as CW, separated from protogastric region by a long transverse furrow. Deep notch be- tween frontal and preorbital lobes, the latter strongly granulated. Anterolateral margins armed with 4 teeth (excluding the outer or- bital), first small, with margins granulated, second and third longest, subequal in size, spinose and directed anteriorly, fourth big- ger than first, with borders and base gran- 507 ulated. Posterolateral and posterior margins of carapace almost straight, granulated. Pterygostomian region (Figs. 2A, 4C) densely granulated, with a longitudinal, spi- nose crest just below suture that divides subhepatic and pterygostomian regions; pterygostomian ridges present, marked with small tubercles. Orbits % as wide as front, separated from it by a deep, long notch, borders conspic- uously dentate; 2 large sutures on supraor- bital region; preorbital and postorbital lobes dentate. An acute granulated tooth on infra- orbital angle. Eyestalks completely fitting in orbits when retracted, with 4 sharp spi- nules and stiff setae. Antennules (Figs. 2A, 4C) with basal segment considerably inflated, and with a longitudinal crest of small granules; penul- timate and ultimate segments slender. Basal antennal segment subcylindrical, with 4 sharp spinules; second to fourth seg- ments mobile, longer than broad; flagellum long. Ischium of third maxilliped (Fig. 2B) longer than broad; outer surface with me- dian longitudinal furrow. Merus subquad- rate, outer surface coarsely granulated, me- sial margin dentate, setose; distolateral an- gle ending in a semiacute lobe, directed an- teriorly. Palp marginally setose. Exopod reaching to tip of distolateral angle of mer- us. Left cheliped shorter than right and more spinose (Figs. 4A—B); merus granulose, up- per and lower margins delimited by a row of strong, acute, inward spines; outer surface of carpus spinose, inner margin armed prox- imally with strong acute spine curving in- wards, junction between carpus and chelae fringed with setae, inner surface densely granulated. Palm of long chela with 3 upper rows of acute spines directed inwards, di- minishing in number and size towards lower margin, inner surface slightly punctate. Dac- tylus about as long as palm; fingers leaving small gap when closed, each terminating in inwardly curved corneous claw; movable 508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Batodaeus adanad, new genus, new species. A. Holotype male 16.6 mm X 23.2 mm (CNCR 21023): B, Allotype female 11.8 mm X 16.6 mm (CNCR 22509), dorsal view. VOLUME 117, NUMBER 4 Fig. 2. regions; B, third left maxilliped, outer view; C, sternoabdominal cavity. D, allotype female, sternoabdominal cavity. Abbreviations: g2, gonopod 2; s1—8, sternites 1—8. part with upper margin armed with small acute spines, cutting edges with blunt teeth. Pereopods 2—5, slender, subequal in length; pereopod 4 slightly long, and pereo- pod 2 slightly short; all segments with lat- eral and mesial faces spinose, covered by dense, thin setae. Dactylus slightly longer than propodus, terminating in corneous claw. Propodus with long thin setae on up- per border, outer surface punctate. Upper border of carpus with 4—6 small spines. Merus with 12 spines on upper border, which diminish in size proximally, directed anteriorly. Thoracic sternum in male (Fig. 2C) rel- atively narrow, densely granulated; sternal sutures 1—2 indistinct, 2—3 complete, 3—4 incomplete and confined to lateral regions; 509 Batodaeus adanad. A—C, holotype male. A, antennal, pterygostomian, suborbital, and epistomal 4—5 and 5—6 interrupted medially; 6—7 and 7-8 complete. Sternite 4 with slightly raised median part, with short longitudinal furrow. Locking mechanism on sternite 5 just be- low suture 4—5. Sternoabdominal cavity deep and relatively narrow. Male abdomen (Fig. 3A) with short mar- ginal setae on segments 1—6 and telson. First segment long, slender. Second as long as first, broadest, not reaching coxa of fifth pereopod, with small portion of sternite 8 visible (Fig. 3B). Third to fifth segments fused, punctate, longer than broad. Sixth segment as long as broad. Posterior margin of telson rounded. Male sexual openings coxal. First gonopod (Fig. 3C, D) long, reach- ing beyond suture separating sternites 4 and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. 5 and sternite 8; C, gonopod 1; D, tip of gonopod 1; E, gonopod 2. E allotype female, abdominal segments. Abbreviations: al—2, abdominal segments | and 2; cx 5, coxa 5; ep7, episternite 7. 5, when in situ, slender and with distal part slightly incurved. Second gonopod (Fig. 3E) very short, curved, tip sharp, recurved. Females with regions of carapace less de- marcated (Fig. 1B); front straight, less de- flexed; right cheliped longer than left, palm with less conspicuous spines, pereopods more setose, without spines on upper border of merus, carpus with spines more acute. Thoracic sternum (Fig. 2D) with longitu- dinal furrow on sternite 4 less marked; ab- dominal cavity less deep and broad; in lon- gest specimen, locking mechanism on ster- nite 5 not visible. Abdomen (Fig. 3F), with first and second segments as in male, leav- ing visible a small portion of sternite 8; seg- ments 3—6 free and subequal in size; pos- terior part of telson rounded. Pleopods long, Batodaeus adanad. A—E, holotype male. A, abdominal segments; B, abdominal segments 1—3, coxa slender, extending past edge of telson; gon- opores small, ovate. Color in life.—Cream, with tip of chelae fingers dark. Etymology.—This species name is formed from an arbitrary combination of ' the two first letters of each of our sons’ names: Adolfo, Andrés, and Adrian, and is used as a noun in apposition. Distribution.—Western Atlantic; south- western Gulf of Mexico; Banco de Cam- peche. Remarks.—The shape and ornamentation of carapace and pereopods of B. adanad are superficially similar to the species of Mon- odaeus, notably M. rouxi. Also, the ptery- gostomial region is densely granulated and the incomplete endostomial ridges of B. VOLUME 117, NUMBER 4 511 A Fig. 4. Batodaeus adanad, holotype male. A, right chela; B, left chela; C, ventral view of anterior part of carapace. SID adanad are similar to those seen in M. tub- erculidens and M. couchii. The morphology of the merus of the third maxilliped of B. adanad also resembles those of M. guinotae and M. tuberculidens. However, Batodaeus adanad is easily separated from all Monodaeus species in that the former has the posterolateral mar- gin almost as long as the anterolateral, the chelipeds are less stout, spinose, and are covered by strong tubercles; the sternoab- dominal cavity is deeper and with a less marked longitudinal furrow on sternite 4. The abdomen of Batodaeus is long, com- pletely covering the sternoabdominal cavi- ty, whereas in Monodaeus species the ab- domen does not completely cover the ster- noabdominal cavity, leaving a longitudinal furrow on sternite 4 exposed; the pereopods 2—5 without a granulated crest on the su- perior border of merus; and the telson of male is rounded. In addition, the morphol- ogy of the first gonopod in B. adanad dif- fers from that in any known Monodaeus species. The new species differs from Medaeops edwardsi Guinot, 1967, M. neglectus (Balss, 1922), and M. granulous (Haswell, 1882) in that all these species have a less convex carapace, with the regions hardly projecting; their pereopods are shorter and broader; and the fingers of their chelipeds are granulated. The first gonopod, thoracic sternum, and sternoabdominal cavity, too, are also different in morphology. Batodaeus adanad can be easily separat- ed from Alainodaeus akiaki Davie, 1992 and A. rimatara Davie, 1992 in that those species have a carapace that is transversally ovoid; the front is less deflexed; the cheli- peds are more robust; the first male gono- pod is stout with slightly twisted tip; and the second male gonopod is moderately longer. Acknowledgments Dr. Rafael Lemaitre and Dr. Janice Clark are greatly appreciated for their help and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON loan of specimens during our visit to the Smithsonian Institution. Special thanks are given to Dr. Michel Hendrickx for his kind review and valuable suggestions on the manuscript. The comments of Dr. P. Ng greatly improved the quality of the manu- script. We thank the crew and scientific staff of R/V Justo Sierra for field work dur- ing cruise BATO. We also thank Ana Elena Viniegra for drawings and Ana Isabel Bie- ler for taking the photographs. Literature Cited Capart, A. 1951. Crustacés Décapodes Brachyoures. In Expédition Océanographique belge dans les eaux cOtiéres africaines de 1’Atlantique Sud (1948—1949)—Bruxelles 3(1):11—205. Couch, R. Q. 1851. Notice of a Crustacean, New to Cornwall.—Transactions of Penzance Natural History Society: 13-14. Crosnier, A. 1967. Remarques sur quelques Crustacés Décapodes benthiques ouest-africains. Descrip- tion de Heteropanope acanthocarpus et Me- daeus rectifrons sp. noy.—Bulletin du Muséum national d’Histoire naturelle, Paris, (2), 39(2): 320-344. Davie, P. J. EF 1992. Deepwater Xanthid crabs from French Polynesia (Crustacea, Decapoda, Xan- thoidea).—Bulletin du Muséum national d’Histoire naturelle, Paris, 4°. sér., 14, section A, n° 2:501—561. Drach, P., & J. Forest. 1953. Description et répartition des Xantho des mers d’ Europe.—Archives de Zoologie expérimentale et générale, 90(1):1—35. Forest, J. 1972. Une espéce nouvelle de Xanthidae des eaux bathyales de Méditerrane: Monodaeus guinotae sp. nov.—Thalassia Jugoslavica 8(1): 63-69. Garth, J. S. 1985. On a small collection of brachyuran Crustacea from Easter Island obtained by the Scripps Institution of Oceanography Downwind Expedition of 1958.—Occasional Papers of the Allan Hancock Foundation (3):1—12. Guinot, D. 1967. Recherches préliminaires sur les groupements naturels chez les Crustacés Brach- youres. II. Les anciens genres Micropanope Stimpson et Medaeus Dana.—Bulletin du Mu- séum national d’ Histoire naturelle, Paris, 39(2): 354-374. , & E. Macpherson. 1988. Remarques sur le genre Monodaeus Guinot, 1967, avec la de- scription de deux espéces nouvelles (Crustacea Decapoda Brachyura).—Bulletin du Muséum national d’ Histotoire naturelle, Paris 4°. sér., 10: 731-757. VOLUME 117, NUMBER 4 MacLeay, W. S. 1838. On the brachyurous decapod Crustacea brought from the Cape by Dr.Smith. Tn Wlustrations of the Annulosa of South Africa: being a portion of the objects of natural history chiefly collected during an expedition into the interior of South Africa, under the direction of Dr. Andrew Smith, in the years 1834, 1835, and 1836; fitted out by “The Cape of Good Hope Association for Exploring Central Africa’’. London: 53-71. 513 Seréne, R. 1984. Crustacés Décapodes Brachyoures de l’‘Océan Indien Occidental et de la Mer Rouge. Xanthoidea: Xanthidae et Trapeziidae. Avec un Addendum par Crosnier (A):Carpillidae et Menippidae.—Faune Tropicale. Office de la Re- cherche Scientifique et Technique Outre-Mer. Paris, 24:1—400. Associate Editor: Christopher Boyko PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):514—522. 2004. A new anchialine shrimp of the genus Procaris (Crustacea: Decapoda: Procarididae) from the Yucatan Peninsula Richard v. Sternberg and Marilyn Schotte (R.v.S) NCBI--GenBank, Building 45, Room 6An, 18D-30, National Institutes of Health, Bethesda, Maryland 20892-6510 U.S.A. e-mail: sternber@ncbi.nlm.nih.gov (MS) Department of Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C., 20013-7012 U.S.A. e-mail: Schottem@si.edu Abstract.—A fourth species of the anchialine shrimp genus Procaris is de- scribed from Cozumel Island, Quintana Roo, México. The combination of char- acter states observed for the abdomen, antennal scale/stylocerite, second an- tennular segment, carapace, eyes, rostrum, and telson is unique in the genus. The new species appears to be morphologically most closely related to P. ascensionis from Ascension Island. Cladistic analysis of differentiating char- acter states supports a sister group relationship between P. ascensionis and the Mexican species, in two out of three most parsimonious hypotheses. In addi- tion, the Bermudan P. chacei and Hawaiian P. hawaiiana are positioned as sister taxa in all minimal length trees. While the discovery of a new Procaris species adds to our biogeographical knowledge of the genus, it has pointed to the possibility that the Atlantic taxa may be a paraphyletic assemblage. Shrimps of the family Procarididae are restricted to anchialine habitats, and occupy an unclear position within the Decapoda relative to the Caridea (Christoffersen 1988, 1990; Felgenhauer & Abele 1983; Kensley & Williams 1986; Schram 1986). The Pro- carididae contains two genera, Procaris and Vetericaris Kensley & Williams, 1986. Pro- caris has perhaps the most interesting dis- tribution of any anchialine decapod: P. as- censionis Chace & Manning, 1972 is re- stricted to Ascension Island in the mid- south Atlantic, P. chacei Hart & Manning, 1986 is endemic to Bermuda, and P. ha- watiana Holthuis, 1973 is found on the Ha- waiian archipelago. [A photograph of an undescribed “*Procarid sp.’ from Christmas Island in the Indian Ocean has been pub- lished (Jones & Morgan 2002), although the habitus of the pictured specimen looks more atyid than procaridid.] What is even more remarkable is the con- servative morphology of Procaris species, considering the disjunct biogeography of the taxa, as the three species differ in only a few characters (Hart & Manning 1986). Vetericaris is monotypic with the Hawaiian V. chaceorum Kensley & Williams, 1986 separated from any Procaris species by a plethora of character states. Despite the dis- tinctiveness of Procaris and Vetericaris, the monophyly of the family has not been ques- tioned. A recently described family for a genus of abyssal shrimp, the Galatheacari- didae Vereshchaka, 1997, overlaps with the Procarididae in several key (albeit plesio- morphic) character states, indicating that the hypothesized connection between an- chialine and abyssal caridean taxa of Hart et al. (1985) may not be entirely without merit. Interrelationships aside, an open question is how many additional anchialine and submerged caverniculous carideans await discovery that could, potentially, complete the known biogeographical gaps. Here we describe a fourth species of Pro- caris, from the Yucatan Peninsula. The dis- covery of this new species adds consider- VOLUME 117, NUMBER 4 Sea SE IE as era qa SS 515 Sree SERIE EES yy a , Fig. 1. of telson. ably to our biogeographic knowledge of the genus. The new Procaris material was col- lected by Drs. Dennis Williams and Jeff Bozanic who during the years 1988, 1989, and 1995 collected them from the cenotes of Quintana Roo, México. CL numbers re- fer to carapace length; USNM numbers de- note catalog numbers in the National Mu- seum of Natural History, Smithsonian In- stitution. WA Siac oo eos S SSeS ee Procaris mexicana, n. sp. A, habitus, lateral view; B, anterior region; C, telson and uropods; D, apex Procarididae Chace & Manning, 1972 Procaris mexicana, new species Figs. 1-3, Table 1 Procaris sp. Kensley, 1988:688. Material.—Holotype (USNM_ 1068789): México, Cueva Quebrada, Chankanaab Park, Cozumel, Quintana Roo, 25 September 1987, coll. Dennis Williams, CL 8 mm. Paratypes: USNM 1068790, 1 specimen, CL 516 6.5 mm, same locality as holotype, coll. Dennis Williams, 23 Sep 1987; USNM 1068791, 3 specimens (1 damaged), CL 5.1 mm, 5.5 mm, and 5.9 mm, Cueva Quebrada, depth of 25-30 feet, coll. Jeff Bozanic, 5 April 1988; USNM 1068792, 4 specimens, all CL 6 mm, Cueva Quebrada, coll. Dennis Williams, Feb. 1993; USNM 1068793, 1 specimen, CL 8 mm, Lagoon Cave, Cozu- mel, Quintana Roo, México, coll. Jeff Boz- anic, 3 Apr. 1988. Description.—Integument fragile and thin. Rostrum acutely triangular and lacking teeth, only reaching medial concavity of eyes. Carapace devoid of spines; anterior margin distinctly convex and slightly emar- ginate below distinct cervical sulcus; prom- inent anteroventral sulcus positioned parallel to ventral margin, and meeting ventral end of cervical sulcus; posterodorsal margin markedly concave. Eyestalk produced into two lobes, the me- dial lobe sharply triangular and extending beyond the more bluntly triangular lateral lobe; eye lacking facets and with irregular mass of pigment. Antennular peduncle does not reach distal one-third of antennal scale, broad; stylocer- ite tapering distally to acute apex, almost reaching distal margin of second antennular article; segments subequal in length; anterior margin of basal article with distinct V- shaped dorsomedial cleft. Antennal scale lacking distolateral tooth, distal margin convex, length approximately 2.5 times the width; distal margin of scale reached by antennal peduncle. Mandible pronouncedly developed, with three-segmented palp, molar and incisor pro- cesses forming one piece; incisor process subtrapezoidal, lacking distinct marginal teeth except for the two angular regions, scooplike. Paragnath sinuous, surrounding incompletely mandibular bases, distal end pointed, broadest around midlength. Endites of first maxilla well-developed, broad; palp simple. Second maxilla with two endites, distal endite with deep incision, palp pro- nounced and broader proximally, tapering PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON slightly distally, scaphognathite small in comparison to the endites and palp. Maxil- liped 1 with near tongue-shaped endite, well-developed palp; long, simple epipod; caridean lobe prominent. Maxilliped 2 en- dopod with seven segments of roughly sim- ilar width throughout; exopod long, strap- like; epipod simple, reduced. Maxilliped 3 with seven-segmented endopod, distal half of merus broader than all other parts of the appendage; exopod long, subequal to endo- pod length; epipod simple, small. Pereiopods 1—5 similar in organization, flexor margins lined with simple setae; dac- tyli approximately 0.12—0.13 times length of propodi, with strong, curved spines. All five pereiopod pairs with straplike exopod; pe- reiopods 1—4 with distinct simple epipod, and pleurobranch and setobranch; pereiopod 5 lacking epipod, pleurobranch, and seto- branch. Third abdominal somite with dorsal cap not reaching middle of fourth somite; pos- teroventral margin of the six anterior somites broadly rounded. Abdominal sternites 1—5 with median tubercle between coxae of ple- opods; sternite 6 with bulbous tubercle pos- teriorly directed between uropod bases. Tel- son approximately 1.4 times length of so- mite 6, not including posterior spines, armed with two pairs of dorsal spines; posterior margin armed with four pairs of spines, lat- eral spines shortest, two mesial pairs roughly half the length of sublateral spines. All pleopods similar in organization; en- dopods short and weakly developed; appen- dices internae and masculinae absent from all pleopods. Distribution.—Known only from anchia- line habitats of Cozumel, Quintana Roo, Yu- catan Peninsula, México. Remarks.—All Procaris species are re- markably similar in morphology, differing slightly but specifically in a set of characters (Table 1; Hart & Manning 1986). This is significant given the immense distances sep- arating all four taxa, especially P. hawaiiana vis-a-vis the three Atlantic species. On the basis of biogeography, one might expect the VOLUME 117, NUMBER 4 517 Fig. 2. Procaris mexicana: A, pleopod 4; B and C, mandible; D, second maxilliped; E, paragnaths; E first maxilliped; G, pleopod 1; H, first maxilla; I, second maxilla; J, third maxilliped; K, pleopod 3; L, pleopod 2; M, pleopod 5. 518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Procaris mexicana: A, pleopod 1; B, same, dactyl; C, pereopod 2; D, same, dactyl; E, pereopod 3; F same, dactyl; G, pereopod 4; H, same, dactyl; I, pereopod 5; J, same, dactyl. 519 VOLUME 117, NUMBER 4 () v1~ (1) pepunos Ajpeoiq (1) ot -OS YANO] JO oIp -prlui Suryovel jou (Z) JOuNSIp (7) Jussqe (1) juowises [euUuSjue JO pus 0} Jsowye (0) IasUuo] dqoOy uLIpSUr ({) AjtAvouo05 [eIpeul sayovel (1) vI~ (CZ) SC I~ (1) pepunor A][peoiq (Z) Jejnsue (€) oWuWIOS YANO JO a[ppru puokaq (Z) aquwIOs YLINO} Jo sPppru oO} (1) yeom (1) yeom (Q) quasead (Q) quasead Z JUOUIsES (1) Z Juowses jeuusjue Jo pua 0} jeuugjuRe Jo pus Suryovol jou ‘({) SSO] IO Z JUSUISES [RUUDJUR JO PUd oO} JOSUO] IqQOT [eJO}e] ‘(7 ) BqGo] uRIp ({) jenbo saqgoy] = -ou_~URY) IasUOT] IO 0} [enba 9qoy [e1A}e]| Soko SOyoeILIOAO {(Z) AyAvouos [BIpour sayore13aAo oqo] [eIpoul soyorol £(Z) AWIABOUOD [RIPOUI soyoRaIIDAO (€) SL I~ (Z) sejnsue (Z) emuios YyInoy JO [Pprur 0} (Z) 1OUNSsIP (1) woesqe (0) ZT JUOWISES [eu -Ud}UL SUTYIVILIOAO (0) IOSUO] 9qGO] ULIpoUT (1) Aqtaevouoo [eIpour soyovor (0) S I~ (0) popunor A[MoeU (Q) Juesqe (Q) Juesqe (Q) Juosoid (0) Z JUoWIZesS [eu -UdJU BUTYOROIIOAO (0) IOBUO] OqO] ULIPSUI (Q) ake Jo AjIAvoUOD je -Ipou SuTYyoReI JOU UOSTO} 0} O}TUIOS OUT -Opqe YIXIs :onei y3ueT ° SUNOS YD JO UISIeUI [eMUDAOIO\SOg ° deo o}TWIOS [RUTWIOpgR pIIUL, SNO[NS [VOTAIOD Y}00} o[eOs [eUUDIUY 9111990] A1S sokq UINA}]SORY No} wo + a) a — DUDIIXAU “I DUDIIDMDY ‘qd 12IDYO “I SIUOISUIISD “q SIUDIIAAIAA “€ pue ‘z ‘| = sorydiowrode ‘Q = saje}s o1ydIOWIOISe|g “siMvI01g JO soloeds INOF oY] BUOUTe SODUDIOFJIP o1e}1s Io}ORIeEYDO—'| 2IGRL 520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON B<2) = w oO D = = ow x < iS rr = aS) ® = S = = S AN fab) Y) es Igy} a ca} = ro} < > Qo o oO o 2 S oO 5 S = = o < 2 S Ss ro) o >< 2 = = D 2 Ke © = w = ro) = > oO oO o oO B Fig. 4. Most parsimonious cladograms obtained with the data matrix in Table 2. A, single shortest length tree found with characters 1—7 ordered; B, one of two alternative minimal step topologies identified with all characters unordered (the other tree identical to A). VOLUME 117, NUMBER 4 three Atlantic species to form a clade, with the Indo-Pacific P. hawaiiana as the sister group of the lineage. However, a comparison of the character states presented in Table 1 affords no clear-cut separation between At- lantic and Pacific congeners. Each Procaris species instead appears to be a mosaic of character states found in the other taxa; spe- cies differences are due then to specific char- acter state combinations as opposed to the presence of apomorphies. To test the possi- bility that P. ascensionis, P. chacei, and P. mexicana may be more closely related to each other than to P. hawaiiana, a data ma- trix was prepared for parsimony analysis (Table 2), and Vetericaris was used as the outgroup for character state polarization (Ta- ble 1). The purpose of the cladistic test was twofold: to identify a parsimonious hierar- chy of Procaris taxa, and to compare this hierarchy with biogeography. When characters 1—7 were treated as or- dered transformation series, one tree was ob- tained by Exhaustive Search using PAUP 3.1 software (Swofford 1993), with a length of 18 steps, consistency index (CI) value of 0.833, and a retention index (RI) .umber of 0.571 (Fig. 4A). This first hypotl esis indi- cates that P. ascensionis and P. :nexicana are sister species, with P. chacei and P. ha- waiiana forming a species pair. Placing the cladogram into the context of time and space, the split between Atlantic and Pacific Procaris species would have occurred after the emergence of two Atlantic clades: P. as- censionis and P. mexicana on the one hand, and the proto-P. chacei/P. hawaiiana ances- tor. A second exhaustive search was_per- formed though this time all characters were parameterized as unordered series. Two trees most parsimonious were found with lengths of 17 steps, CI = 0.882, and RI = 0.667. The topolo; y of one of the cladograms is identical in structure to the one in Fig. 4A. The second hypothesis is also a resolved hi- erarchy, though with P. ascensionis branch- ing off first, followed by P. mexicana, and 521 Table 2.—Data matrix used in the parsimony anal- ysis. See Table 1 for explanation of character states. Character 12345678 Vetericaris 00000000 P. ascensionis 10012223 P. chacei 21101222 P. hawatiana 21101311 P. mexicana 10112111 with P. chacei and P. hawaiiana positioned as sister taxa (Fig. 4B). Cladistic analysis of Procaris interrela- tionships indicates three things. First, P. chacei and P. hawaiiana are more closely related to each other on morphological grounds than either is to any other Procaris species. Second, relationships between P. ascensionis and P. mexicana are ambiguous. Parsimony searches conducted with ordered and unordered characters support a sister group relationship between the two (Fig. 4A). Yet the hypothesis that P. ascensionis is basal to the remaining Procaris species (Fig. 4B) cannot be dismissed. Finally, the Atlantic species appear not to form a clade; i.e., they are a paraphyletic assemblage mi- nus the inclusion of P. hawaiiana. One seri pus caveat of the parsimony study is the paucity of characters (eight) relative to the number of taxa (five). This reflects the extremely conservative morphology of Pro- caris species. Another caveat is the coding of character states (Table 1). Character states were coded to maximize hierarchical reso- lution given a limited number of characters. For instance, the rostrum character was di- vided into three character states: not reach- ing medial concavity of eyes (plesiomorph- ic); reaching medial concavity (apomorph- ic); overreaching medial concavity (also apomorphic). The way this character was coded for Procaris species, P. chacei and P. hawaiiana have the same state. Yet the ros- trum only reaches the median lobe in P. chacei although it overreaches the eyes in P. hawaiiana. The same critique applies to character 2. Nevertheless, if characters are recoded to reflect all the differences seen, 522 (SRE DU EO P. ascensionis 3(1) P. mexicana P. hawatiana Fig. 5. ings. Numbers to the right of a box denotes derived character states supporting a particular set of taxa. Venn diagram of apomorphy-based group- the tree obtained is identical to that shown in Fig. 4A (unpublished results). Figure 5 shows a Venn diagram of apo- morphy-based relationships in Procaris, un- derscoring the polythetic nature of species differences. Hart & Manning (1986) suggested that the remarkable similarity of Procaris species may be explained by the reduction of vari- ability by natural selection. The “reduced variability’ hypothesis appears rather weak considering that anchialine caridean taxa oc- curring with Procaris often exhibit consid- erable variability, morphs, and species-spe- cific apomorphies (e.g., Kensley & Williams 1986, Smith & Williams 1981). It may be that the distribution of Procaris is much more extensive than currently known, with gene flow over great distances occurring via semi-continuous populations distributed among shallow submerged ‘“‘crevicular”’ habitats (Hart et al. 1985, Maciolek 1983). Acknowledgments We are most grateful to Drs. Dennis Wil- liams and Jeff Bozanic who collected the new Procaris material, and to Drs. Charles Fransen and Mark Siddall for their com- ments on an earlier draft of the manuscript. Literature Cited Chace, F A., Jr, and R. B. Manning. 1972. Two new caridean shrimps, one representing a new family, from marine pools on Ascension Island (Crusta- cea: Decapoda: Natantia).—Smithsonian Contri- butions to Zoology 131:1-18. Christoffersen, M. L. 1988. Phylogenetic systematics of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the Eucarida (Crustacea Malacostraca).—Revista Brasileira de Zoologia 5(2):325-351. . 1990. A new superfamily classification of the Caridea (Crustacea: Pleocyemata) based on phy- logenetic pattern.—Zeitschrift fiir Zoologische und Systematik Evolutionsforschung 28(2):94— 106. Felgenhauer, B. E., and L. G. Abele. 1983. Phylogenetic relationships among shrimp-like decapods. In: E R. Schram, ed., Crustacean issues 1. Crustacean phylogeny, pp. 291-311. A. A. Balkema, Rotter dam. Pp. 1-372. Hart, C. W., Jr, and R. B. Manning. 1986. Two new shrimps (Procarididae and Agostocarididae, new family) from marine caves of the western north Atlantic—Journal of Crustacean Biology 6(3): 408-416. 5 , and T: M. Iliffe. 1985. The fauna of Atlantic marine caves: evidence of dispersal by sea floor spreading while maintaining ties to deep waters.—Proceedings of the Biological Society of Washington 98:288-292. Holthuis, L. B. 1973. Caridean shrimps found in land- locked saltwater pools at four Indo-West Pacific localities (Sinai Peninsula, Funafuti Atoll, Maui, and Hawaiian Islands), with the description of one new genus and four new species.—Zoolo- gische Verhandelingen 128:1- 48. Jones, D. S., and G. J. Morgan. 2002. A Field Guide to Crustaceans of Australian Waters. Western Aus- tralian Museum. Reed New Holland, Sydney, Australia, 224 pp. Kensley, B. 1988. New species and records of cave shrimps from the Yucatan Peninsula (Decapoda: Agostocarididae and Hippolytidae)—Journal of Crustacean Biology 8(4):688—699. , and D. Williams. 1986. New shrimps (families Procarididae and Atyidae) from a submerged lava tube on Hawaii.—Journal of Crustacean Bi- ology 6(3):417—437. Maciolek, J. A. 1983. Distribution and biology of Indo- Pacific insular hypogeal shrimps.—Bulletin of Marine Science 33(3):606—618. Schram, FE 1986. Crustacea. Oxford University Press, Oxford. Smith, M. J., and W. D. Williams. 1981. The occurrence of Antecaridina lauensis (Edmondosn) (Crusta- cea, Decapoda, Atyidae) in the Solomon Islands: an intriguing biogeographical problem.—Hybro- biologia 85:49—58. Swofford, D. L. 1993. PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1. Smithsonian In- stitution, Washington, D.C. Vereshchaka, A. L. 1997. New family and superfamily for a deep-sea caridean shrimp from the Gala- thea collections.—Journal of Crustacean Biology 17(2):361-373. Associate Editor: Christopher Boyko PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):523-528. 2004. Macrobrachium patheinense, a new species of freshwater prawn (Crustacea: Decapoda: Palaemonidae) from Myanmar Hla Phone and Hiroshi Suzuki Laboratory of Aquatic Resource Science, Faculty of Fisheries, Kagoshima University, 4-50-20 Shimoarata, Kagoshima 890-0056, Japan, e-mail: suzuki @fish.kagoshima-u.ac.jp Abstract.—A new species of freshwater palaemonid prawn, Macrobrachium patheinense, is described from Mayan Creek near Pathein City, Ayeyawaddy Division, Myanmar. The new species is most closely related to M. mirabile (Kemp, 1917), M. palaemonoides Holthuis, 1950, M. superbum (Heller, 1862) and M. inflatum Liang & Yan, 1985, but can be differentiated by the rostrum shape and dentition, telson shape, and the second pereiopod chela proportions. Like other South East Asian countries, Myanmar has a wealth of freshwater streams, lakes, ponds and rivers, but unlike its adjacent countries, the freshwater crus- taceans have been poorly studied. Important relevant investigations on freshwater deca- pods have been undertaken in India, Thai- land, China, Malaysia, Philippines, and In- donesia (Cai & Dai 1999, Cai & Ng 2001, Chace & Bruce 1993, Holthuis 1978, Jali- hal et al. 1988, Liang & Yan 1985, Shokita & Takeda 1989, Tiwari 1947, Wowor & Choy 2001, Yeo et al. 1999, and others). It was not until 1918 that the first signif- icant carcinological study of Myanmar’s fauna was conducted, and since then only 12 species of the shrimp genus Macro- brachium Bate, 1868 have been recorded (see Jalihal et al. 1988, Jayachandran 2001, Kemp 1918, Tiwari 1952). A major study by Cai & Ng (2002) reviewed the taxono- my of the Myanmar palaemonid freshwater prawns, reporting one new species and five new records of Macrobrachium for the country. Myanmar’s unique geographic position means that it has close connections with In- dia, China, and the rest of the Indo-Malay- sian region to the east and south. Thus, there is a strong likelihood that further in- vestigation will lead to more new taxonom- ic and zoogeographic discoveries. In Myan- mar, freshwater shrimps and prawns are im- portant components of inland fisheries, and further taxonomic and ecological studies must be made an urgent priority in order to ensure sustainable management and conser- vation of stocks. Specimens were collected from Mayan Creek near Thayet Kone village, about five miles west of Pathein City, Ayeyawaddy Division, on 6 September 2001. All speci- mens were preserved in formalin for ship- ping to Japan and examined at the Labo- ratory of Aquatic Resource Science, Fac- ulty of Fisheries, Kagoshima University. Among the collected specimens, 38 individ- uals of a Macrobrachium species possessed similar distinctive characteristics that could not be attributed to any known species, and are thus here described as a new species. The holotype and 33 paratypes are de- posited in the Laboratory of Aquatic Re- source Science, Faculty of Fisheries, Ka- goshima University, Kagoshima, Japan (KUMB). Additional paratypes are also de- posited in the Kitakyushu Museum of Nat- ural History, Kitakyushu, Japan (KMNM), and the Zoological Reference Collection (ZRC), Raffles Museum, National Univer- sity of Singapore. Numbers in parentheses in “Materials examined”’ indicate the post-orbital cara- 524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. 5 mm. pace length in millimeters. Abbreviations used include: M, male; FE female. Family Palaemonidae Rafinesque, 1815 Genus Macrobrachium Bate, 1868 Macrobrachium patheinense, new species Figs. 1-2 Materials examined.—Mayan Creek, Thayet Kone village, Pathein City, Ayeya- waddy Division, 6 Sep 2001: holotype, M (8.96), KUMBcr 1101, paratype, 2M (9.16, 8.85), KMNH IvR 400.100, KMNH IvR 400.101, 2M (7.21, 8.75), ZRC 2003.0324, 33M OAS, Vij, B47, SID, 8.50; 8.65, 8.85, 8.61, 8.33, 8.68, 8.48, 9.08, QM, BAS, BS, S07, BOS, 9.67, SS, 8.64, 8.10, 8.83, 7.97, 8.61, 8.16, 8.24, 7.92, 8.63, 9.10, 9.07, 7.91, 8.55), KUMBcer 1102-1134. Diagnosis.—Carapace smooth, with an- tennal and hepatic spine. Rostrum slender, long; dental formula 2+10/5. Mandible with 3-segmented palp. Scaphocerite broad, with slightly concave outer margin. First pereiopod slender, reaching to end of sca- phocerite. Second pereiopod equal, ex- tremely slender; carpus 2 times as long as merus, finger 1.8 times as long as palm, without teeth on cutting edge. Telson with 2 pairs of dorsolateral spinules; posterior Macrobrachium patheinense. Lateral view of holotype, KUMBcr 1101; male (8.9 mm). Scale equals margin ending in median tooth; 2 spines and 2 plumose setae on each side, inner spines well developed, outer spine very short, plumose setae shorter than inner spines. Description.—Rostrum (Figs. 1, 2a) long, slender, reaching beyond end of an- tennular peduncle almost to end of scapho- cerite, tip curving slightly upwards, upper margin with 12 teeth (mode 13, range 11— 17), of which 2 teeth (mode 2, range 2—3) are placed behind orbit; first tooth smaller than second, placed further from second than third; upper margin of rostrum with single row of setae between teeth; lower margin with 5 ventral teeth (mode 4, range 3-8), first tooth level with seventh and eighth teeth; ventral portion with single row of setae. Carapace (Fig. 1) has strong an- tennal spine below lower orbital angle, pro- duced anteriorly to broadly rounded lobe; hepatic spine smaller than antennal spine, placed below and some distance behind an- tennal spine; branchiostegal groove present. Abdomen smooth, glabrous, with broadly rounded first to third pleurites; fourth and fifth pleurites produced posteriorly, sixth abdominal somite about 1.5 times as long as fifth. Telson (Figs. 2b, c) 1.4 times length of sixth abdominal somite, with 2 pairs of VOLUME 117, NUMBER 4 S25 3 * t 3 4 z e specter s <* Fig. 2. Macrobrachium patheinense. Holotype, KUMBcr 1101; male (8.9 mm). a, lateral view of rostrum; b, dorsal view of telson; c, tip of telson; d, antennule; e, antenna; f, mandible; g, second pereiopod; h, chela of second pereiopod; i, dactylus and propodus of third pereiopod; j, first pleopod; k, second pleopod; 1, uropodal diaeresis. Scales equal 1 mm. 526 dorsolateral spinules; posterior margin end- ing in a median tooth, flanked on each side by 2 spines and 2 plumose setae; inner spine well developed, 4 times as long as median tooth, outer spine very short, 2 plu- mose setae slightly shorter than inner spine. Eyes well-developed, with cornea as long as stalk. Basal segment of antennular peduncle (Fig. 2d) broad, stylocerite very short, dis- tinctly pointed, not reaching middle of basal segment; anterolateral spine of basal seg- ment reaching about middle of second seg- ment; second segment as long as third seg- ment; anterior margin of basal segment strongly curved. Scaphocerite (Fig. 2e) 3.2 times as long as broad, not reaching tip of rostrum; outer margin slightly concave, ending in a tooth, not reaching end of la- mella. Mandible (Fig. 2f) with outer, lateral, 3-segmented palp. Other mouth parts typi- cal for genus. First pereiopod slender, reaching end of scaphocerite (Fig. 1); fingers slightly longer than palm, with numerous setae; carpus about twice as long as chela, broadest dis- tally, narrowing proximally; merus shorter than carpus; ischium about half as long as merus. Second pereiopods (Figs. 2g, h) equal in size and shape, extremely slender, carpus reaching beyond scaphocerite by half its length; chela gradually narrowing proximally, finger very long, slender, about 1.8 times as long as palm (mean 1.7, range 1.4-1.9), same width throughout length, cutting edge entire, tip curves inwards; car- pus as long as chela, unarmed, with distal portion broadest; merus about half as long as carpus (mean 0.7, range 0.5—0.9) but equal with ischium. Pereiopods 3—5 slender, subequal in size; third pereiopods over- reaching scaphocerite by length of entire dactylus; dactylus (Fig. 21) slender, concave on ventral, with numerous setae on dorsal surface, measuring about % of propodus length; propodus with eight spinnules on ventral surface, about twice as long as car- pus; merus nearly as long as propodus; is- chium about same length as carpus; fourth PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pereiopods shorter than fifth but longer than third. Exopod of first pleopod (Fig. 2j) oval- shaped, with small endopod, inner margin concave. Second to sixth pleopods nearly equal with endopods and exopods; endopod with a slender appendix interna. Second pleopod (Fig. 2k) with appendix masculina, placed between appendix interna and en- dopod; appendix masculina longer, stronger than appendix interna, bearing several stiff setae. Uropods reaching beyond end of tel- son; exopods ovate, outer margin straight, inner margin convex, uropodal diaeresis (Fig. 21) with a spine slightly longer than outer angle; endopods broadly ovate, small- er than exopods. Color.—Grayish white when live. Etymology.—The specific name is adapt- ed from the type locality (Pathein) where the specimens were collected. Distribution.—Macrobrachium pathei- nense inhabits freshwater and slightly brackish water habitats, known so far only from the type locality. Remarks.—Macrobrachium patheinense is similar to the Palaemon-like Macro- brachium species, that have slender and delicate pereiopods, especially M. mirabile (Kemp, 1917), M. palaemonoides Holthuis, 1950, M. superbum (Heller, 1862) and M. inflatum Liang & Yan, 1985. Macrobrach- ium patheinense is, however, distinguish- able from M. mirabile by the shape of the rostrum and telson. The rostrum of M. path- einense is slender and longer than the sca- phocerite, while that of M. mirabile is shorter than the scaphocerite, and has a high dorsal crest (Kemp 1917). The telson of the former has two pairs of plumose se- tae slightly shorter than the inner spine, but that of the latter has only one pair of plu- mose setae longer than the inner spine. The new species is also distinguished from M. palaemonoides by shapes of the rostrum and telson, and the proportions of chelae of the second pereiopods. The rostrum of M. patheinense is armed with teeth along the entire upper margin, but that of M. palae- VOLUME 117, NUMBER 4 monoides has an unarmed area on its distal half (Holthuis 1950, Kamita 1974). The tel- son terminates in a short median tooth in M. patheinense, but this tooth is longer in M. palaemonoides (Kamita 1974). The movable finger of the second pereiopod is 1.4—1.9 (mean 1.7) times as long as the palm in M. patheinense, but 1.3—1.4 times as long as the palm in M. palaemonoides (Chace & Bruce 1993, Holthuis 1950). The new species can easily be distinguished from M. superbum by the shape of the ros- trum and the second pereiopod chela pro- portions. The rostrum doesn’t reach beyond the distal end of the scaphocerite in M. su- perbum (Cai & Dai 1999, Holthuis 1950), but distinctly further in M. patheinense. In addition, the upper margin of the rostrum is generally straight in M. superbum, but dis- tally upcurved in M. patheinense. The mov- able finger of second pereiopod is 1.2—1.5 times as long as the palm in M. superbum, but 1.4—1.9 (mean 1.7) times in M. path- einense. The rostral shape and formula of M. patheinense is most similar to those of M. inflatum, but the second pereiopods and telson of both species are different. The movable finger of the second pereiopod of M. inflatum is subequal to the length of the palm (0.9-1.0 from the figures of Cai & Dai (1999) and Liang & Yan (1985)), but that of M. patheinense is much longer than the palm (1.4—1.9, mean 1.7). The telson of M. inflatum bears three pairs of plumose se- tae, these setae being longer than the inner spine on the posterior margin, but M. path- einense has only two pairs of plumose setae that are slightly shorter than the inner spine. The unique chela of M. patheinense re- sembles that of Leandrites stenopus Hol- thuis, 1950, and Pseudopalaemon bouvieri Sollaud, 1911, however the presence of a mandibular palp in M. patheinense confirms its placement in Macrobrachium and distin- guishes it from all Leandrites and Pseudo- palaemon species (Holthuis, 1993). Thus, the new species appears to occupy an interesting phylogenetic position and should be included in future studies inves- 527 tigating generic relationships within the family Palaemonidae. Acknowledgments We are grateful to Peter J. EK Davie of Queensland Museum, Australia, Peter K. L. Ng and Yixiong Cai of National University of Singapore, L. B. Holthuis of National Museum of Natural History, Leiden, The Netherlands, and an anonymous reviewer for their critical readings of the manuscript. Literature Cited Cai, Y., & A. Y. Dai. 1999. Freshwater shrimps (Crus- tacea: Decapoda: Caridea) from the Xishuang- banna region of Yunnan Province, southern China.—Hydrobiologia 400:211—241. , & PK. L. Ng. 2001. The freshwater decapod crustaceans of Halmahera, Indonesia.—Journal of Crustacean Biology 21(3):665—695. , & . 2002. The freshwater palaemonid prawns (Crustacea: Decapoda: Caridea) of Myanmar.—Hydrobiologia 487:59—83. Chace, E A., Jr., & A. J. Bruce. 1993. The Caridean Shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition 1907-1910. Part 6: Su- perfamily Palaemonoidea——Smithsonian Con- tributions to Zoology 543:1—52, pls. 1-7. Heller, C. 1862. Neue Crustaceen, gesammelt wahrend der Weltumseglung der k. k. Fregatte Novara. Zweiter vorlauliger Bericht.—Verhandlungen der kaiserlich-koniglichen zoologisch-botan- ischen Gesellschaft in Wien 12:519-528. Holthuis, L. B. 1950. The Decapoda of the Siboga Ex- pedition. Part 10. The Palaemonidae collected by the Siboga and Snellius Expeditions with re- marks on other species. 1. Subfamily Palae- moninae.—Siboga Expeditie 39(a9):1—268, figs. 1-52. . 1978. A collection of decapod Crustacea from Sumba, Lesser Sunda Islands, Indonesia.— Zoologische Verhandelingen 162:1—55. . 1993. The Recent genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda), with an Appendix on the Order Amphionidacea. Nationaal Natuurhistorisch Museum, Leiden, 328 pp. Jalihal, D. R., S. Shenoy, & K. N. Sankolli. 1988. Freshwater prawns of the genus Macrobrach- ium Bate, 1868 (Crustacea, Decapoda, Palae- monidae) From Karnataka, India.—Records of the Zoological Survey of India, Miscellaneous Publication, Occasional Paper No. 112:1—74. Jayachandran, K. V. 2001. Palaemonid Prawns: Bio- 528 diversity, Taxonomy, Biology and Management. Science Publishers, Enfield, 624 pp. Kamita, T. 1974. Four species of the Nepalese prawns.—Researches on Crustacea 6:1—16, pls. 1-2. Kemp, S. 1917. Notes on Crustacea Decapoda in the Indian Museum. IX. Leander styliferus, Milne- Edwards, and related forms.—Records of the Indian Museum 13:203—231, pls. 8-10. . 1918. Crustacean Decapoda of the Inle Lake Basin.—Records of the Indian Museum. 14:81— 102, pls. 15-16. Liang, X.-Q., & S.-I. Yan. 1985. New species and new record of palaemonidae from China (Crustacea Decapoda) (in Chinese with English summa- ry).—Acta zootaxonomica Sinica, 10(3):253— 258. figs. 1—4. Shokita, S., & M. Takeda. 1989. A new freshwater prawn of the genus Macrobrachium (Decapoda, Caridea, Palaemonidae) from Thailand.—Bul- letin of the National Science Museum, Tokyo, Series A (Zoology), 15(3):147-154. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sollaud, E. 1911. Pseudopalaemon Bouvieri, nouveau genre, nouvelle espece, de la Famille des Palae- monidae.—Bulletin du Museum National d’ Histoire Naturelle, Paris 17:12—16. Tiwari, K. K. 1947. Preliminary descriptions of two new species of Palaemon from Bengal.—Re- cords of the Indian Museum 45(4):329—331. . 1952. Diagnosis of new species and subspe- cies of the genus Palaemon Fabricius (Crusta- cea: Decapoda).—Annals and Magazine of Nat- ural History 5:27-32. Wowor, D., & S. C. Choy. 2001. The freshwater prawns of the genus Macrobrachium Bate, 1868 (Crustacea: Decapoda: Palaemonidae) from Brunei Darussalam.—The Raffles Bulletin of Zoology 49(2):269—289. Yeo, D. C. J., Y. Cai, & P. K. L. Ng. 1999. The fresh- water and terrestrial decapod Crustacea of Pulau Tioman, Peninsular Malaysia.—The Raffles Bulletin of Zoology, Supplement No. 6:197— 244. Associate Editor: Christopher Boyko PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):529-540. 2004. A new species of Enhydrosoma Boeck, 1872 (Copepoda: Harpacticoida: Cletodidae) from the Eastern Tropical Pacific Samuel Gomez Instituto de Ciencias del Mar y Limnologia, Unidad Académica Mazatlan. Joel Montes Camarena s/n, Ap. Postal 811, Mazatlan 82040, Sinaloa, México, e-mail: samuelgomez @ola.icmyl.unam.mx Abstract.—Some enhydrosomids were found while sorting samples taken from the Urias system during a short-term study on the effects of organic enrichment on the abundance and diversity of benthic copepods. Upon careful examination, these specimens proved to belong to a new species, Enhydrosoma brevipodum, of the species-group defined by the lack of sexual dimorphism on the male P3 and can be separated by the reduced exopod of female P5. En- hydrosoma brevipodum, whose full description is herein provided, constitutes the fourth record of the genus from the Pacific Mexican coast. The genus Enhydrosoma Boeck, 1872 is a group of harpacticoid copepods common- ly found in shallow brackish and marine coastal systems worldwide. Some Enhydro- soma specimens were found in sediment samples from two shallow brackish systems in central (Ensenada del Pabellon lagoon) and southern (Urias system) Sinaloa during the course of two short-term studies about the effects of organic enrichment on the dis- tribution and abundance of meiofauna (see Gomez-Noguera & Hendrickx 1997) and on the diversity of benthic harpacticoids. Some of these specimens belong to three species recently described by Go6mez (2003), whereas some specimens constitute the Pacific counterpart of Enhydrosoma la- cunae Jakubisiak, 1933 (Gomez 2003), originally described from Cuba and rede- scribed by Fiers (1996) from the Yucatan Peninsula. While sorting samples taken from Urias system, some specimens of a different species of Enhydrosoma were found. These specimens proved to belong to a new species mainly characterized by the reduced exopod of female P5. A de- tailed description of this species is herein provided. Materials and Methods Quantitative sediment cores were taken for the analysis of the effects of organic en- richment on benthic copepods along a pol- luted estuary (Urias system) in southern Sinaloa (north-western Mexico) during 2001 and 2002. Sediment samples were taken with an Eckman box corer with a sampling area of 225 cm?, and subsamples were taken using plastic corers with a sam- pling surface of 7 cm?. Sediment cores were subdivided vertically into separate 1 cm slices to a depth of 5 cm. Each slice was fixed with 10% formalin, and sieved through 500 and 63 pm sieves to separate macro- and meiofauna. Meiofauna was pre- served in 70% ethanol and stained with Bengal Rose until further inspection. Mei- ofaunal major taxa were quantified and co- pepods (cyclopoids, poecilostomatoids and harpacticoids) were separated from the rest of meiofauna and stored in 70% ethanol for further investigation. Observation and drawings of the species described herein were made from whole and dissected spec- imens mounted in lactophenol, under 100 oil immersion objective using a Leica com- pound microscope equipped with drawing 530 tube and phase contrast. The type material was deposited in the collection of the Insti- tuto de Ciencias del Mar y Limnologia, Ma- zatlan Marine Station. The terminology proposed by Huys & Boxshall (1991) for the general description and armature for- mulae was adopted. Abbreviations used in the text and tables: P1l—P6, first to sixth swimming leg; EXP, exopod; ENP, endo- pod. Family Cletodidae T. Scott, 1904 sensu Por (1986) Genus Enhydrosoma Boeck, 1872 Enhydrosoma brevipodum, new species Type material—One female holotype preserved in 70% ethanol (EMUCOP- 090301-73), one dissected male allotype (EMUCOP-090301-62), and one dissected female paratype (EMUCOP-090301-61); collected from station 10; 9 Mar 2001; leg. S. Gomez. Type _ locality.—Urias system, Sinaloa, northwestern Mexico (23°09’—23°13'N, 106°20’—106°25'W). Etymology.—The specific name alludes to the reduced exopodal lobe of female P5. Female.—Body (Fig. 1A, 2A) tapering from posterior margin of cephalothorax, curved in lateral view; length of holotype, 420 wm from tip of rostrum to posterior margin of caudal rami. Cephalic shield about % total length, with strongly folded lateral and dorsal surface, posterior margin plain, with sensilla arising from distinct cones. Rostrum triangular, fused to cephalic shield, with rounded tip, with two sensilla. Dorsal surface of free thoracic somites (P2— P4) smooth, with sensilla arising from dis- tinct cones along plain posterior margin. First urosomite (P5-bearing somite) as pre- ceding somites except for fewer sensilla. Surface of genital double somite smooth, with dorsolateral division between first and second genital somite (second and third urosomites), posterior margin of both gen- ital somites plain, first somite with sensilla arising from distinct cones along posterior PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON margin, second somite as first one except for two additional tube pores (arrowed in Fig. 1A), both somites with additional sen- silla arising from paired bulbous structures laterally; genital somites completely fused ventrally, first somite bearing pair of P6 and genital pore, the former each bearing a short spinulose spine, and with an associ- ated tube-pore (arrowed in Fig. 2A), copu- latory pore covered by integumental fold, ventral surface of second segment smooth, except for spinules and fragile setules along posterior margin between pair of sensillum- bearing cones. Dorsal surface of fourth and fifth urosomite as in preceding somite, ex- cept for lack of central pair of sensilla on posterior margin of fourth somite, and lack of sensilla along posterior margin of fifth urosomite, both somites with pair of tube pores (arrowed in Fig. 1A); ventral surface of fourth and fifth urosomite smooth, fourth urosomite ornamented with spinules and setules as in second genital somite, fifth urosomite with only spinules along poste- rior margin. Anal segment smooth, rounded anal operculum without ornamentation and flanked by pair of sensilla. Caudal rami cy- lindrical and about 8.3 times as long as wide, with seven setae in all, setae I and II located in proximal fifth, the former arising ventrally and about % total length of seta II, the latter dorsal to seta I, seta III slightly longer than seta II and arising in the middle along outer margin of ramus, seta IV and V fused, seta VI located in distal inner cor- ner and as long as seta II, seta VII arising in proximal third at the level between seta II and III. Antennule (Fig. 2B). 5-segmented; sur- face of segments smooth except for spinular row on first, third and fourth segment. Ar- mature formula 1-(1), 2-(7), 3-(7+ae), 4- (1), 5-(11+ae). Antenna (Fig. 3A, B) with proximal and distal set of spinules on inner margin of al- lobasis; with very small distal abexopodal seta close to distal set of spinules, the latter difficult to see and can be easily mistaken for a setule (arrowed in Fig. 3A). Exopod a gS NET ea i aS Diy N Se ie) = ee awe AI Cs Gee ae Lo ry ; ¥ ! \ SS ia a s,s Se 3 1 3e We. ms i i an Ake, Pc a \ footy i “ & SS 532 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ | = = | BB i — Z | ec — . >= = LZ LEE Rg )\ Fig. 2. Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, urosome, ventral (P5 bearing- somite omitted; tube pores in genital field arrowed); B, antennule. Scale bar: A, 100 wm; B, 75 wm. VOLUME 117, NUMBER 4 Fig. 3. B, normal exopod of antenna; C, maxilliped; D, Pl. Scale bar, 50 pm. 1-segmented and armed with two bipinnate elements (Fig. 3B). Endopodal segment or- namented with two strong spines subdistal- ly along inner margin; distal margin five se- tae/spines (outer pectinate spine seemingly without fused small seta), and ornamented with two hyaline frills on outer margin. Mandible (Fig. 4A, B) with slender gna- thobase; biting edge with uni- and multi- cuspidate teeth, and one bare seta at distal 533 Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, antenna with aberrant exopod; inner corner. Palp well-developed, 1-seg- mented and armed with one endopodal and two basal setae (Fig. 4B). Maxillule (Fig. 4C). Arthrite with five distal and two lateral elements, and two sur- face setae; coxal endite fused to basis and represented by one seta, basis represented by two distal setae, endopod and exopod represented by one seta each. Maxilla (Fig. 4D) with short spinular row 534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Enhydrosoma brevipodum, female paratype EMUCOP-090301-61. A, mandible; B, mandibular palp; C, maxillule; D, maxilla. Scale bar: A, B, 50 wm; C, D, 25 pm. on distal inner corner of syncoxa; proximal syncoxal endite with two slender and bare setae and one bipinnate element; distal syn- coxal endite with two elements (inner strongly pinnate, outer anvil shaped, fused to endite and with only one pinnule). Al- lobasal endite with non-articulated spine and two setae. Endopod represented by two setae fused at base. Maxilliped (Fig. 3C) prehensile, with short and unarmed syncoxa; basis with spi- nules along inner margin; claw slender and curved distally, with accessory seta. P1 (Fig. 3D). Coxa and basis ornamented as depicted, the latter with inner and outer setae. Exopod three-, endopod 2-segment- ed, the latter reaching distal third of last ex- opodal segment. P2—P4 (Figs. 5A, B, 6A) with coxa and basis ornamented as shown, the latter with outer plumose seta. Exopod three-, endopod 2-segmented. Endopod of P2 slightly longer than, of P3 as long as, of P4 clearly shorter than first and second exopodal segments combined. Armature formulae of P1—P4 as follows: 235) VOLUME 117, NUMBER 4 g Sl, —— : he Se aN eS ‘i — ee EN 3 —< = ; => ri ae . J 2S SSS iS : : SSS A Ss 5 ig < a ee oa Sus se 5 Z Se a PZ : a SX te a S == AY SO \y \ 8, 3 = Ss NX ( 2 oS NR Sa \ Ny Ay : 3 Ce Gr IN 2 ee. \ . Os E _ a : & 5 y S < & Ss Ns : oe Z . 8 aeitig E/4, a > 2»? 5, 377 RF SO? ae > => ae ‘ = SSD OLY? > Le Re —_— > 3 4 “vt N ay EIEN 7TS 4 ot = 23 =>?? > pre ° 4% Ses = >> dd > f 2 ae — 34 a 4 >) 2 = iL = ‘S > rs’ QIN? ? Sy >) ee > 10mm Cranial spines of holotype of Sunagocia sainsburyi, 26230-007. WAM Fig. 2. VOLUME 117, NUMBER 4 547 5 mm Fig. 3. below suborbital ridge. the first records of the genus taken with trawling gear, the other species having typ- ically been taken with rotenone and SCU- BA. Methods Counts and measurements were taken ac- cording to Hubbs & Lagler (1949). Mea- surements were made with calipers and rounded to the nearest mm. Vertebrae were counted from radiographs. Terminology of head spines follows Knapp et al. (2000). In- 1mm Fig. 4. LL scale, circa 22nd scale from right side of holotype of Sunagocia sainsburyi. Head of holotype of Sunagocia sainsburyi, right side, showing lack of sensory tubules on cheek stitutional acronyms follow Leviton et al. (1985) except for South African Institute for Aquatic Biodiversity (SAIAB), formerly RUSI. Standard length and head length are abbreviated as SL and HL, and lateral-line aclele: Sunagocia sainsburyi, new species Sainsbury’s flathead Fig. 1 Holotype.-—WAM _ 26230-007, 86 mm SL, Western Australia, 125 km NE of Port Hedlund, 19°07’S, 119°25'’E, EV. Coura- geous, 28 May 1978, 73-74 m, K. Sains- bury et al. Paratype.—CSIRO H 5856-01, 97 mm, Northern Australia, near Darwin, 11°53’'S 131°15’E, EV. Soela, Cr. 5, Sta. 49, 6 July 1980, trawl, 20—22 m. Other material examined.—Sunagocia arenicola, USNM 362804 (12, 37-117 mm), Western Indian Ocean, Amirantes Is., D’Arros I., R/V Anton Bruun Cr. 9, 5°24'S, 53°13"E, 8 Dec. 1964, rotenone, 4—8 m, RS-40, R. D. Suttkus et al.; SAIAB 8219 (8, 46-131), Mozambique, Pinda Reef, Bay of Bocage, 14°10’S, Sept. 1956, M. M. Smith. S. carbunculus, USNM 99703 (8. 548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON CH TS CN ye 25 3mm Fig. 5. 70-116), Malaysia, Sabah, North Borneo, Sandakan Bay, 2 Mar. 1908, U.S. Fisheries Steamer Albatross, seine. S. otaitensis, USNM 366402 (6, 106-151), Northern Philippines, Babuyan Is., Fuga I., circa 18°51’N 121°22’E, coral and tide pools, 11 Mar. 1990, rotenone, A. Ross; USNM 366403 (5, 83-159), Central Philippines, Negros, Apo I., 9°4.5'N 123°16.4’E, 18 May 1979, LK 79-20, rotenone, 0—2.4 m, L. W. Knapp et al.; ROM 42303 (10, 37— 80) Indian Ocean, Chagos Arch., Peros Banhos, Isle du Coin, 5°26’21”S 71°46'52”E’, 6 Feb. 1979, WE 79-06, ro- tenone, O—7 m, R. Winterbottom et al. 3mm Fig. 6. Sketch of iris lappet from right eye of ho- lotype of Sunagocia sainsburyi. Area around LL scales 19—27, right side of holotype of Sunagocia sainsburyi. Diagnosis.—A species of Sunagocia with 4—5 preorbital spines; 5 total gill rak- ers on the first arch; a bony expansion of the suborbital ridge upper base on cheek bearing 1—2 rows of small spines; maxilla reaching to below middle of eye; no papil- lae on upper surface of eye; a series of spines on the ethmoid and several pairs of nasal spines (Fig. 2); and smaller, more nu- merous spines on the supraorbital and sub- orbital ridges. Sensory tubules are absent from the cheek area below the suborbital ridge (Fig. 3). Description.—Data for holotype given, followed by that of paratype in parentheses when differing. Dorsal-fin damaged in holo- type, last 1-2 spines missing, VII(IX), 11; anal-fin rays 12; pectoral-fin rays 2 un- branched + 14 branched + 3 unbranched (2+13+4) = 19; pelvic fin with 1 spine and 5 rays, innermost is unbranched; caudal-fin branched rays 8; vertebrae 27; total gill rakers on first arch 5; pored scales in LL 52, ante- rior 3 scales bearing a small spine; 6 rows of scales between 2nd dorsal fin origin and LL. Number of oblique scale rows above LL about equal to number of LL scales. LL VOLUME 117, NUMBER 4 9 8 @ td ® e @ O ® Snout Length / Interorbital Width 0 20 40 60 80 Standard Length (mm) Fig. 7. holotype, solid diamond). scale pores with two openings to the exte- rior (Fig. 4). Relationship of LL scales to adjacent scale rows is shown in Fig. 5. Iris lappet bears short branches with bifurcate tips (Fig. 6). Lip margins without papillae. Body depressed, upper body covered with ctenoid scales, breast scales largely cy- cloid. Interopercular flap lacking. HL 2.8 (2.9) in SL; orbit going 1.1 times in snout. Ratios of least interorbital width into snout length for the four species of Sunagocia ap- pear in Fig. 7. Villiform teeth in bands on jaws and palatines, in two separate patches on vomer. Top and sides of head armed with nu- merous spines (Fig. 2). Preopercular spines 3, uppermost longest, not bearing an acces- sory spine on base; a pair of stout nasal spines, with 2—3 smaller spines running an- teriorly to each; base of opercular spines 549 @ carbunculus Mi arenicola Ootaitensis © sainsburyi 100 120 140 160 180 Ratios of least interorbital width into snout length for the four species of Sunagocia (S. sainsburyi covered by scales, not bearing serrae. Sub- orbital ridge with about 17—20 serrae. Color observations were taken on the paratype after it thawed, prior to preserva- tion. Dorsum brownish, with about six darker bands crossing back, venter whitish. Two brown infraorbital bands and two brown suborbital bands present. Cheek be- low suborbital ridge with a series of brown blotches. A brown band angling back from anterior ethmoid to front of eye. Dorsal-fin spines and rays bearing small dark spots; pectoral fin with several vertical brownish bands above, clear below; pelvic fin with four reddish-brown bands; and caudal fin with about four vertical dark brown bands. Etymology.—The species is named in honor of Keith J. Sainsbury, collector of the holotype and other flatheads later during the EV. Soela cruises. 550 Acknowledgments We thank the following individuals for the loan of specimens or other assistance: P. R. Last and A. Graham (CSIRO); J. B. Hutchins (WAM); R. Winterbottom and M. Rouse (ROM); P C. Heemstra and V. Mthombeni (SAIAB) and S. Jewett (USNM). Thanks are also due J. Finan and S. Raredon, USNM, for considerable tech- nical assistance. The fine drawing of the paratype was made by Keiko Hiratsuka Moore. Literature Cited Cuvier, G., & E. Griffith. 1832. The Animal Kingdom, Arranged in Conformity with its Organization, Vol. 15, Insecta 2. Whittaker, London, 796 pp. , & A. Valenciennes. 1829. Histoire Naturelle des Poissons. EF G. Levrault, Paris, vol. 4: i— xxvi + 2 pp. + 1-518 pp. , & . 1833. Histoire Naturelle des Pois- sons. EF G. Levrault, Paris, vol. 9: i—xxix + 3 pp. + 1-512 pp. Hubbs, C. L., & K. FE Lagler. 1949. Fishes of the Great Lakes region.—Bulletin of the Cranbrook Insti- tute of Science 26: 186 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Imamura, H. 1996. Phylogeny of the family Platyce- phalidae and related taxa (Pisces: Scorpaenifor- mes).—Species Diversity 1(2):123—233. . 2003. Sunagocia, a replacement name for the platycephalid genus Eurycephalus (Actinopter- ygii: Percomorpha), with taxonomic comments on the species of the genus.—Species Diversity 8(3):301—306. Knapp, L. W., H. Imamura, & M. Sakashita. 2000. Onigocia bimaculata, a new species of flathead fish (Scorpaeniformes: Platycephalidae) from the Indo-Pacific.—J. L. B. Smith Institute of Ichthyology Special Publication No. 64:1—10. Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. Dawson. 1985. Standards in herpetology and ichthyology: part 1. Standard symbolic codes for institutional resource collections in herpe- tology and ichthyology.—Copeia 1985(3):802— 832. Ogilby, J. O. 1898. New genera and species of fish- es.—Proceedings of the Linnean Society of New South Wales 23:32—41. Schultz, L. P., E. S. Herald, E. A. Lachner, A. D. We- lander, & L. P. Woods. 1966. Fishes of the Mar- shall and Marianas Islands.—Bulletin of the United States National Museum 202, Vol. 3: 45-62. Associate Editor: Carole Baldwin PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):551—563. 2004. A new species of Nannocharax (Characiformes: Distichodontidae) from Cameroon, with the description of contact organs and breeding tubercles in the genus Richard P. Vari and Carl J. Ferraris, Jr. (RPV) Department of Zoology, Division of Fishes, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A., e-mail: vari.richard@nmnh.si.edu; (CJF) Research Associate, Department of Zoology, Division of Fishes, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A., e-mail: cferraris@msn.com Abstract.—Nannocharax reidi, new species, is described from several local- ities in the upper Cross River basin in Cameroon. The species possesses the synapomorphies of the clade comprising Nannocharax and Hemigrammochar- ax. It is assigned to Nannocharax on the basis of its possession of a completely- pored lateral line, a feature distinguishing that questionably monophyletic ge- nus within the clade composed of these two genera. Nannocharax reidi is distinguished from its congeners on the basis of a combination of meristic and morphometric features and details of pigmentation on the body. Comparative studies revealed the presence of hook-shaped contact organs on the pectoral fins of some species of Nannocharax and epidermal breeding tubercles on the head, body, and fins of at least one species of the genus. These observations represent the first reports of contact organs and breeding tubercles in African members of the order Characiformes. Some species of Nannocharax were found also to possess variably-developed fields of hook-shaped contact organs on the exposed surfaces of scales of the midlateral portion of the body posterior of the pectoral girdle. This latter feature has not been previously reported among fishes. Species of the African distichodontid ge- nus Nannocharax are relatively small-sized fishes inhabiting the Nile River and many sub-Saharan rivers, with the greatest spe- cies-level diversity of the genus occurring in West Africa and the Congo River basin. Species of Nannocharax share a number of distinctive modifications relative to phylo- genetically-proximate taxa including trans- versely-flattened ventral surfaces of the head and body, a down-turned mouth, and, in many species, expanded pelvic and pec- toral fins; these features apparently corre- late with their habits of resting on, and feeding off of, the substrate or vegetation (e.g., Nannocharax fasciatus, see Géry 1977:89). The most recent comprehensive treatment of Nannocharax was that of Bou- lenger (1909:279) who discussed seven nominal species that recent authors have as- signed to the genus. Subsequent decades saw the progressive descriptions of addi- tional species of Nannocharax, resulting in 24 species being recognized in the compen- dium of the genus by Daget & Gosse (1984: 200). Two treatments of the West African species of Nannocharax have been pub- lished (Daget 1961, Gosse & Coenen 1990), and recent decades have seen the de- scription of several new species of the ge- nus from that region (Vari & Géry 1981, Coenen & Teugels 1989, Van den Bergh et al. 1995). Numerous uncertainties, nonethe- less, remain concerning the species-level 552 diversity within Nannocharax. Perhaps the major question is whether the geographi- cally widespread N. fasciatus, whose distri- butional range in West Africa reportedly ex- tends from Guinea to Gabon (Daget & Gosse 1984:201), is a single widely-distrib- uted species or a complex of similar spe- cies. Reid (1989:24, 56), followed by Teu- gels et al. (1992:43), noted that population samples of an N. fasciatus-like form from the upper Cross River system in Cameroon differed from the more-typical N. fasciatus populations from that region, but those au- thors deferred from pursuing the question of the identity of these samples. Studies of the species of Nannocharax in the lower Guinea region encompassing Cameroon, Rio Muni, Gabon, and the coastal portions of the Republic of Congo, Brazzaville demonstrate that some popula- tions of an N. fasciatus-like form from the upper Cross River represent an undescribed species that we describe herein. We also de- scribe unusual modifications of the scales, fins, and epidermis in some species of Nan- nocharax that were discovered during our comparative studies. These noteworthy modifications are either elaborations of some body scales of a form unique to the species of Nannocharax within the Chara- ciformes (and perhaps fishes), or are elab- orations of the fin rays and epidermis that were previously thought to be restricted to New World members within that order. Materials and Methods Measurements are given as a percentage of standard length (SL) except for subunits of the head that are presented as percent- ages of head length. Lateral-line scale counts include all pored scales along that series, including scales located posterior to the hypural joint. In fin-ray counts, lower- case Roman numerals indicate unbranched rays, and Arabic numerals _ indicate branched rays. The two posteriormost anal- fin rays, which are joined at their bases, were counted as one element. Morphomet- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ric and meristic data were taken following the procedures outlined in Fink & Weitz- man (1974). Counts of gill-rakers, teeth, cteni, and branchiostegal rays were taken from two specimens that were cleared and counterstained following the method of Taylor & Van Dyke (1985). Vertebral counts were acquired via radiographs and include the four vertebrae of the Weberian apparatus and the terminal centrum. Insti- tution abbreviations are: AMNH, American Museum of Natural History, New York; CU, Cornell University, Ithaca, New York; MRAC, Musée Royal de |’ Afrique Centra- le, Tervuren, Belgium; and USNM, Nation- al Museum of Natural History, Smithsonian Institution, Washington, D.C. Nannocharax reidi, new species Fig. 1 Nannocharax sp. 1, Reid, 1989:24, 56 [Cameroon, upper Cross River system]. Nannocharax sp., Teugels et al., 1992:43 [upper Cross River system]. Holotype.—USNM 304046, 62.7 mm SL; Cameroon, Cross River system, col- lecting points on southern Munaya River draining northern Korup, on Basep River at junction with Munaya River (5°49'30’N, 9°03'30"E); collected by Gordon McG. Reid, 22 February 1988. Paratypes.—20 specimens, 34.3—59.0 mm SL. USNM 375193, 16 specimens (2 cleared and counterstained for cartilage and bone), 34.3-59.0 mm SL; AMNH 233622, 2 specimens, 37.3—41.6 mm SL; MRAC A3-47-P-1-2, 2 specimens, 37.1—43.1 mm SL; collected with holotype. Non-type specimens examined.—19 specimens, 33.3-44.7 mm SL. USNM 375195, 3 specimens, 34.5—38.8 mm SL; Cameroon, Manya, Cross River system, collecting points on main Cross River, downstream of Mamfé, Mam River, junc- tion with Cross (3°50'30"N, 9°14’50"B). USNM 375196, 1 specimen, 41.7 mm SL; Cameroon, Cross River system, collecting points on main Cross River below Mamfé VOLUME 117, NUMBER 4 Fig. 1. points on southern Munaya River draining northern Korup, on Basep River at junction with Munaya River (05°49'30"N, 09°03’30"E); lateral and ventral views. SSL 25 N, QIU SO). WSINIM S7SIOT, Z specimens, 36.4—44.5 mm SL; Cameroon, Cross River system, collecting points on southern Munaya River draining northern Korup, southern Munaya River, junction with Cross River (5°53'N, 9°00’E). USNM 375194, 6 specimens, 36.3—36.7 mm SL; Cameroon, Cross River system, collecting points on southern Munaya River draining northern Korup, on Basep River at junction with Munaya River (5°49'30'N, 9°03'30"E); collected with holotype. MRAC 88-053-P- 0163-0168, 6 specimens, 33.3—44.7 mm SL; Cameroon, mainstream of Cross River, 5—15 km downstream of Mamfé (approxi- mately 5°46’N, 9°17’E). MRAC 88-053-P- 0170, 1 specimen, 42.4 mm SL; Cameroon, mainstream of Cross River, approximately 5 km downstream of Mamfé (approximate- ly 5°46’N, 9°17’E). Diagnosis.—Nannocharax reidi is distin- guished from all congeners by the combi- nation of: the lack of a large, dark, rounded spot extending from the posterior portion of the caudal p: duncle to the basal portions of the middle c iudal-fin rays; the lack of a dis- tinct, dark, midlateral stripe extending from the snout at least to the rear of the caudal peduncle; the absence of a series of very narrow, vertical, dark bars positioned along 353) Nannocharax reidi, holotype, USNM 304046, 62.7 mm SL; Cameroon, Cross River system, collecting the lateral surface of the body; the location of the origin of the dorsal fin posterior to the vertical through the insertion of the pel- vic fin; the possession of 47 to 49 scales along the lateral line, 5, rarely 6, scales dor- sal of the lateral line to the origin of the dorsal fin, and 4 scales ventral of the lateral line to the origin of the anal fin; and the overall body form. Descript on.—Morphometric values for holotype and paratypes are presented in Ta- ble 1. Body elongate, relatively wide trans- versely in region from rear of head to ver- tical through posterior terminus of base of dorsal fin and increasingly transversely- compressed posterior to latter region. Trans- verse widening of anterior portion of body proportionally more pronounced in larger examined individuals. Ventral region of head and body anterior to insertion of pel- vic fin distinctly flattened; degree of flatten- ing more pronounced in larger examined specimens. Dorsal profile of head gently convex from tip of snout to vertical through posterior margin of orbit, straight or very slightly convex from that point to posterior limit of supraoccipital spine. Predorsal pro- file of body slightly convex in all examined specimens. Dorsal profile of body slightly posteroventrally-inclined along base of dor- 554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphometrics and meristics of holotype and paratypes of Nannocharax reidi, new species, n = 21. Standard length is expressed in mm; measurements | to 14 are percentages of standard length; 15 to 18 are percentages of head length; mean includes holotype. Holotype Paratypes Mean Morphometrics Standard Length 62.7 34.3-59.0 1. Snout to anal-fin origin 75.6 72.1—16.3 74.3 2. Snout to pelvic-fin insertion 40.7 37.2-41.4 39.3 3. Snout to pectoral-fin insertion 23.4 23.2—26.3 24.8 4. Snout to dorsal-fin origin 43.7 42.1-45.9 44.3 5. Dorsal-fin origin to hypural joint 54.7 53.7—-59.6 56.3 6. Dorsal-fin origin to anal-fin origin 35.4 32.3-36.7 34.3 7. Dorsal-fin origin to pelvic-fin insertion 20.2 17.9-20.0 18.8 8. Dorsal-fin origin to pectoral-fin insertion 26.2 23.9-27.1 25.7 9. Caudal-peduncle depth 10.2 9.8-10.4 10.1 10. Pectoral-fin length ADA) 23.9-27.2 25.6 11. Pelvic-fin length 25.4 22.4-25.6 23.8 12. Dorsal-fin length AN) 21.5—25.4 ABI 13. Anal-fin length 17.1 15.9-17.8 16.8 14. Head length 25.8 24.5—27.5 25.8 15. Postorbital head length 37.8 37.8-44.7 42.1 16. Snout length 34.6 31.5-35.3 33.3 17. Bony orbital diameter Die 27.1—31.9 29.9 18. Interorbital width 21.0 16.3—20.9 19.2 Meristics Lateral-line scales 47 47-49 47.8 Scale rows between dorsal-fin origin and lateral line S) 5-6 5.1 Scale rows between anal-fin origin and lateral line 4 4 4.0 Predorsal median scales 10 10-12 11.2 Branched dorsal-fin rays 9 8-10 9.4 Branched anal-fin rays 6 6-8 7.0 Branched pelvic-fin rays 7 6-8 WM Pectoral-fin rays 14 13-15 14.1 sal fin and slightly convex from posterior terminus of base of fin to caudal peduncle. Ventral profile of head straight and slightly posteroventrally-inclined. Ventral profile of body nearly straight along prepelvic region and slightly convex from insertion of pelvic fin to caudal peduncle. Mouth slightly subterminal. Lower jaw comparatively wide relative to condition in many congeneric species, with width of posterior portion of jaw equal to height of orbit in larger specimens. Jaw teeth elon- gate, bicuspid, and slightly expanded dis- tally (see Daget 1961, fig. 4 for shape of teeth in genus), with single series of func- tional teeth in each jaw. Dentary with 5 or 6 teeth. Dentary teeth gradually decreasing in size posteriorly with terminal tooth in se- ries approximately one-half as long as tooth proximate to dentary symphysis. Replace- ment teeth on dentary arranged in single se- ries within enlarged dentary replacement tooth trench. Dentary lacking segment of laterosensory canal system and movably at- tached to lateral surface of anterodorsal sur- face of angulo-articular. Contralateral den- taries immovably attached syndesmotically along medial surfaces. Premaxilla with 5 or 6 teeth of same morphology as dentary teeth. Premaxillary teeth gradually decreas- ing in size posteriorly with terminal tooth in series approximately one-half as long as tooth proximate to premaxillary symphysis. Premaxillary replacement teeth arranged in VOLUME 117, NUMBER 4 single row embedded in fleshy covering of inner surface of premaxillae. Contralateral premaxillae immovably attached syndes- motically along medial surfaces; but with premaxillary complex vertically mobile on mesethmoid. Maxilla edentulous, with pos- terior portion of bone flat, plate-like, and extending nearly entirely under first infra- orbital bone when mouth closed. Pupil ovoid, with pronounced emargination of an- terior portion of iris (see Vari & Géry 1981, fig. 2, for illustration of this condition in Nannocharax maculicauda). Snout and dor- sal portions of upper lip and head covered with small papillae-like processes in holo- type and to lesser extent in larger paratypes. Such papillae may represent intermediate developmental stages of well-developed breeding tubercles present in those regions and elsewhere on head, body, and fins in at least one congeneric species (see comments on breeding tubercles under “Contact or- gans and breeding tubercles in species of Nannocharax’”’ below). First gill arch with 13 or 14 gill rakers in 2 cleared and stained specimens. Bran- chiostegal rays 4. Scales ctenoid (sensu Johnson 1984, Roberts 1993), with cteni formed by series of independent ossifications positioned along posterior margin of scale. Scales of lateral surface of body with 23 to 28 cteni along scale margin. Lateral-line scale series completely pored, with last scale in series horizontally elongate. Body scales extend- ing onto base of middle rays of caudal fin in triangular pattern. Many smaller individ- uals with scaleless region on median por- tion of prepelvic region immediately pos- terior of ventral margin of pectoral girdle. Largest examined specimens with lateral surface of scales in region posterior and posterodorsal to insertion of pectoral fin bearing some scattered, elongate, contact organs (sensu Collette 1977). Contact or- gans of scales most concentrated in region proximate to posterior margin of pectoral girdle and best developed in holotype, the largest examined specimen. Contact organs 555 elongate, laterally-directed, and with ante- riorly-directed distal tips. Field of contact organs neither as dense as, nor as extensive as, pattern of hook-shaped processes pre- sent on that region of body in at least one congeneric species (see “Contact organs and breeding tubercles in Nannocharax” below). Dorsal-rays 11,8 to 10. Distal margin of dorsal fin nearly straight. Anal-fin rays 11,6 to 8 or rarely ii1,7. Distal margin of anal fin concave. Individual lepidotrichia of un- branched anal-fin rays anteroposteriorly ex- panded with overall form of distal portion of fin rays somewhat club-shaped; such rays proportionally more expanded in larger individuals. Anterior and lateral surfaces of first unbranched dorsal-fin ray and distal portions of second unbranched ray envel- oped by overlying, thick, fleshy layer in many, but not all, specimens; fleshy cov- ering more developed in holotype and larg- er paratypes. Caudal fin distinctly forked. Pectoral and pelvic fins proportionally longer than in many congeneric species. Pectoral-fin rays 11,13 to 15. Dorsal surface of basal portions of second unbranched, and first through fifth branched rays of pectoral fin with basally-directed, hook-shaped, bony contact organs on basal one-half to two-thirds of rays in holotype, the largest examined specimen. Larger male paratypes with such contact organs developed to less- er degree, but still obvious. Hook-shaped processes on pectoral-fin rays apparently limited to mature males. Unbranched pec- toral-fin rays and distal portions of first branched pectoral-fin ray with individual lepidotrichia widened, more so on distal portions of fin rays; expanded portion of fin rays consequently somewhat club-shaped. First unbranched pectoral-fin ray distinctly shorter than second ray, with second un- branched pectoral-fin ray somewhat shorter than first branched ray; latter ray longest of fin. Ventral and lateral surface of first un- branched pectoral-fin ray, and distal por- tions of second unbranched, and first branched, pectoral-fin rays with thick, 556 fleshy covering. Fleshy layer on fin rays thicker and extending farther basally along rays in larger individuals, and particularly well-developed in holotype. Tip of pectoral fin extending distinctly beyond vertical through insertion of pelvic fin in specimens of all sizes. Pelvic-fin rays 11,6 to 8. Pelvic fin with unbranched rays and distal portion of first branched ray with individual lepidotrichia thickened, more so on distal portions of fin rays that consequently have a somewhat club-shaped form. Ventral and lateral sur- face of first unbranched, and distal portions of second unbranched and first branched pelvic-fin rays with thick, fleshy covering; fleshy layer also extending over dorsal sur- face of distal portions of ray. Fleshy layer on pelvic-fin rays thicker and extending fur- ther basally along rays in larger examined individuals. First unbranched pelvic-fin ray distinctly shorter than second unbranched ray; second unbranched ray somewhat shorter than first branched ray with medial branch of latter ray longest in fin. Tip of longest pelvic-fin ray reaching vent in smaller individuals, but falling slightly short of opening in larger specimens. Vertebrae 36 to 38 [37 in holotype]. Coloration in alcohol.—Ground colora- tion light dusky-brown, with scattered dark chromatophores in holotype and paratypes; tan with fewer dark chromatophores in some more lightly-pigmented, non-type specimens. Lateral and dorsal surfaces of head with irregular field of small, dark chromatophores; chromatophore field more concentrated on upper lip, snout, and dorsal surface of head. Some larger individuals with concentration of dark chromatophores located posterior to orbit and on dorsal two- thirds of operculum. Ventral surface of head ranging from unpigmented to having scat- tered, small, dark chromatophores. Body with pattern of relatively wide, ir- regular bars on dorsal and sometimes ven- tral surfaces; bars often narrowing towards midlateral region with dorsal and ventral bars variably in contact in that region. Re- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gions of contact between bars sometimes appearing as irregular, darker, midlateral patches of dark pigmentation. Smaller, more lightly-pigmented, non-type speci- mens with deep-lying, diffuse region of dusky pigmentation positioned along mid- lateral region, particularly on posterior two- thirds of body. Ventral surface of body ranging from pale with few, scattered, small, dark chromatophores in some small- er non-type specimens to dusky in holotype and paratypes. Some darker specimens with variably-shaped, unpigmented, typically scaleless area on anteroventral portion of body immediately posterior of margin of pectoral girdle. Dorsal fin with transverse band of dark pigmentation located slightly dorsal of ba- ses of fin rays and second, more distally- positioned, wider band of dark pigmenta- tion extending across entire fin. Wider band of pigmentation distinctly separated from distal margin of fin anteriorly, but angling toward and reaching margin of fin along distal portions of first or second branched dorsal-fin rays. Anal fin with variably-de- veloped patch of dark pigmentation on bas- al portions of anterior rays and with dark band extending from more distal portions of anterior rays across fin to its posterior margin. Caudal fin with patch of dark pig- mentation situated basally and with vari- ably-shaped and -positioned patches of dark pigmentation located on both lobes of fin. Pectoral-fin rays overlain dorsally by small, dark chromatophores; dark pigmentation most intense on lateral most fin rays, more so distally. Pelvic fin with pattern of dark chromatophores on distal portion of un- branched rays and more central sections of branched rays. Individual patches of dark pigmentation forming broad, interrupted, dark band across pelvic fin. Adipose fin dark distally in smaller, more lightly-pig- mented individuals, dark throughout in larger specimens. Coloration in life-—Photos of specimens taken soon after capture show that the spe- cies has the same pattern of dark pigmen- VOLUME 117, NUMBER 4 tation as described above, but with the hy- aline regions of head, body, and fins of pre- served specimens having a rosy tint in life. Remarks.—Vari (1979:332) noted that the monophyly of Nannocharax had yet to be demonstrated. That author further com- mented that although Nannocharax and Hemigrammocharax share a series of hy- pothesized synapomorphies (Vari 1979: 331), the single feature that has been uti- lized to distinguish Nannocharax from Hemigrammocharax (the possession of a completely- versus incompletely-pored lat- eral line, respectively) may not serve to de- limit monophyletic assemblages in light of the independent reduction in the degree of development of the lateral line in various groups of characiforms. The possession of various derived characters in a subset of the species of Nannocharax and Hemigram- mocharax to the exclusion of other mem- bers of each genus (Vari & Géry 1981: 1082), furthermore, apparently delimits a monophyletic lineage. That hypothesis sug- gests that both Nannocharax and Hemi- grammocharax as now defined are non- monophyletic. More recently, Coenen & Teugels (1989: 317) documented that population samples of some nominal species within the Nan- nocharax-Hemigrammocharax clade dem- onstrated a continuum between distinctly- shortened and fully-developed lateral lines. Such continuity in the degree of develop- ment of the poring of the lateral-line scale series bridges the gap that purportedly dis- tinguished Nannocharax from Hemigram- mocharax, thereby casting further doubt on the utility of a complete versus incomplete lateral line as a generic delimiter for these taxa. Above and beyond the uncertainty about the naturalness of Nannocharax and Hemigrammocharax, we are also encum- bered by the limitation that the phylogenet- ic relationships within the clade formed by these two genera are yet to be critically ex- amined within the context of a comprehen- sive analysis. In the absence of such a phy- logenetic study, we follow current taxo- S57) nomic practice and assign the new species to Nannocharax on the basis of its com- pletely-pored lateral line, in conjunction with the possession of the synapomorphies for the clade formed by Nannocharax and Hemigrammocharax (see Vari 1979:331, synapomorphies 96 to 107). Distribution.—All examined population samples of Nannocharax reidi were col- lected in the upper portions of the Cross River basin in Cameroon. Etymology.—The specific name, reidi, is in honor of the collector of the specimens that served as the basis of the description of the species, Dr. Gordon McGregor Reid, of the North of England Zoological Society, who first reported that these samples might represent an undescribed form and who has contributed broadly to our knowledge and conservation of African freshwater fishes. Ecology.—Nannocharax reidi was typi- cally captured in the swiftly-flowing main- stream portions of rivers, usually in asso- ciation with submerged logs and branches. In all such localities the water is clear and brown-tinged. The new species was col- lected together with N. fasciatus at the type locality and at three other localities in the upper Cross River. Those two species of Nannocharax were sympatric with N. lati- fasciatus at two of those sites. Contact Organs and Breeding Tubercles in Species of Nannocharax Examination of other species of Nanno- charax revealed the presence of hook- shaped contact organs on the pectoral-fin rays and on the scales in the region of the body proximate to the insertion of the pec- toral fin, along with the occurrence of breeding tubercles on the head, body, and fins of at least some species. The hook- shaped contact organs on the fin rays and breeding tubercles of these species of Nan- nocharax had been previously reported within the Characiformes only in some Neotropical members of the order. The hook-shaped contact organs on some scales 558 of the body in the genus are unknown in any other member of the Characiformes. Presence of hooks on pectoral-fin rays.— In their recent analysis of the phylogenetic relationships of various groups of Neotrop- ical characids, Malabarba & Weitzman (2003:73) enumerated a series of generic and suprageneric taxa within the Characi- formes that bear hook-shaped processes on various combinations of the paired and un- paired fins, including the pectoral fin. These hook-shaped processes were termed contact organs by some authors (e.g., Wiley & Col- lette 1970, Collette 1977), who discussed the distribution and possible functions of these structures. In their commentary on contact organs on the fins of characiforms, Malabarba & Weitzman (2003) noted that such bony processes on individual segments of lepidotrichia were known to be present in diverse Neotropical components of the order, but were unknown in Old World characiforms, including the family Disti- chodontidae. Although Malabarba & Weitz- man (2003) were correct that contact organs on the fins had not been previously reported for Old World characiforms, we found that larger, apparently male, individuals of Nan- nocharax reidi have a series of hook- shaped, distally slightly anteriorly-bent bony processes arranged in a single series along the dorsal surface of the basal one- half to two-thirds of the medial rays of the pectoral fin. As is the case with many Neo- tropical characids, each lepidotrichium of the pectoral-fin rays with a hook-shaped contact organ bears a single such process. The extent of the field of hook-shaped contact organs on the pectoral-fin rays dif- fers both among specimens of N. reidi and between species of Nannocharax. In N. rei- di, the hook-shaped processes on the dorsal surface of the pectoral-fin rays are more highly-developed in larger individuals, but even at their maximum observed degree of development these structures are limited to the basal one-half to two-thirds of the sec- ond unbranched and first through fifth branched pectoral-fin rays. Mature males of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON N. rubrolabiatus (MRAC 95-022-P-001- 007), in contrast, have hook-shaped contact organs on a greater number of fin rays (sec- ond unbranched through eighth branched) and have these processes nearly to the distal tips of the fin rays. The presence of these hook-shaped contact organs on the pectoral fin in specimens of N. rubrolabiatus is cor- related with other apparently breeding-as- sociated modifications of the scales and fins (the presence of hook-shaped contact or- gans on the lateral surface of the scales in the region medial to the pectoral fin and the possession of epidermal breeding tubercles distributed over the head, body, and fins; see discussion of next two characters). This correlation of apparently sexually-dimor- phic features in conjunction with the rela- tively few examined specimens of the spe- cies of Nannocharax that demonstrate these modifications indicate that the hook-shaped contact organs on the pectoral-fin rays of N. reidi and N. rubrolabiatus may be restricted to mature males only during the height of the breeding season. Hook-shaped processes on scales.—TYhe holotype and larger male paratypes of Nan- nocharax reidi possess a form of contact organ (sensu Wiley & Collette 1970, Col- lette 1977) involving an elaboration of the scales in the region of the body medial to the adpressed pectoral fin that is apparently unique not only within the Characiformes, but perhaps throughout bony fishes. The typical form of the scales among members of the Distichodontidae (sensu Vari 1979) is a laterally-unelaborated, relatively flat os- sification with the posterior margin of the main body of each scale bearing a series of smaller, independent ossifications (see Vari 1979, figs. 38b and c), that form a distinct- ly-serrate posterior margin to the scale. These independent ossifications, which con- stitute true cteni (sensu Johnson 1984, Rob- erts 1993:70), vary both in number and form across the members of the Disticho- dontidae, but such elaborations, nonethe- less, are nearly invariably limited to the posterior margin of the scale. The single ex- VOLUME 117, NUMBER 4 559 * Fig. 2. Nannocharax rubrolabiatus, MRAC 95-022-P-001-007, 56.1 mm SL; showing breeding tubercles on the head, anterior two-thirds of the body, and dorsal, pectoral, and pelvic fins. Arrow indicates the midlateral region of the body lacking breeding tubercles, but with hook-shaped contact organs on the lateral surface of the scales. ception to that generality that we have dis- covered, involves the form of the scales on the portion of the body medial to the pec- toral fin in some species of Nannocharax. In larger specimens of N. reidi the scales of the portion of the body medial to the basal portion of the pectoral fin (see description above), particularly those scales in the re- gion of the body immediately dorsal to the posteriorly-directed process of the cleith- rum, have the cteni along their posterior margins complemented by hook-shaped processes arising from the lateral surface of the scales. These scale processes have the form of moderately elongate spines with slightly anteriorly-directed distal hooks. Al- though such processes are obvious in the larger examined specimens of WN. reidi, they nonetheless are somewhat scattered across, and fail to completely cover, the lateral sur- face of the involved scales. A dramatically greater degree of devel- opment of such contact organs in the region of the body posterior to the insertion of the pectoral fin characterizes mature males of Nannocharax rubrolabiatus (Fig. 2). Con- trary to the situation in all other examined distichodontids, males of N. rubrolabiatus lack distinct cteni along the posterior mar- gins of the scales on the anterior portion of the midlateral surface of the body. More strikingly, the specimens in this population sample have the lateral surface of the scale variously covered by fields of laterally-di- rected, elongate, hook-shaped contact or- gans with anteriorly-curved distal tips. Continuity between the fields of hook- shaped processes of adjoining scales varies across the portion of the body with such lateral elaborations of the scales. Those scales positioned closer to the posterior margin of the pectoral girdle have patches of contact organs that together with those of adjoining scales form a nearly uninter- rupted, brush-like expanse continuing ap- proximately five scales posteriorly from the 560 posterior margin of the pectoral girdle and extending dorsally to the horizontal running through the dorsal margin of the opercular opening. Farther posteriorly, the hook- shaped processes on the lateral surface of the scales are restricted to the posterior one- half of the exposed portion of the scale and, thus, form discrete patches of such contact organs, with these patches distinctly sepa- rated from each other. These posteriorly-po- sitioned scales with separate patches of con- tact organs on their lateral surfaces also dif- fer from the more anterior scales character- ized by the possession of such processes in retaining independent, variably posteriorly- directed cteni along at least a portion of the posterior margin of the scale. Such cteni are, however, often somewhat more later- ally-directed than are the homologous os- sifications in other members of the Disti- chodontidae. We are unaware of any laterally-posi- tioned, hook-shaped contact organs of a comparable morphology on the body scales, elsewhere either within the order Characi- formes or among other groups of fishes. The only other report of an African fresh- water fish with laterally-directed hook- shaped processes on the scales involves the gonorhynchiform Phractolaemus ansorgéi, an ostariophysan that is phylogenetically distant from the Characiformes. Phracto- laemus differs significantly from Nanno- charax in the distribution, morphology, and number of such hook-shaped processes per scale (see Thys van den Audenaerde 1961a, fig. 2; 1961b, fig. 2) and the elaborations of the scales in the two genera, thus, are ap- propriately considered to be non-homolo- gous. As a consequence of their apparent unique morphology, the presence of the dense patches of hook-shaped processes on the anterior portion of the midlateral scales of Nannocharax is a likely synapomorphy for at least a subunit of that genus, albeit one perhaps restricted to fully mature, sex- ually-active males during the height of the breeding season. Breeding tubercles.—The presence of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF W ASHINGTON epidermal breeding tubercles has been re- ported in a number of New World members of the Characiformes including the families Characidae, Parodontidae, and Lebiasinidae (Wiley & Collette 1970:164—167, Collette 1977:236—241), but not within any of the African families within that order, an ap- parent absence that included the Disticho- dontidae. One series of Nannocharax rub- rolabiatus (MRAC 95-022-P-001-007) ex- amined during this study has, however, very well-developed epidermal breeding tuber- cles distributed over the head, body and fins (Fig. 2). The degree of development of the tubercles correlates somewhat, albeit not absolutely, with the size of the specimens. The smallest specimen in the lot (45.5 mm SL) has both fewer tubercles than most of the larger conspecific individuals captured with it and, furthermore, those tubercles are proportionally less-developed than those in larger specimens. In larger, apparently male, individuals of N. rubrolabiatus, the breeding tubercles are broadly distributed in large numbers across the snout and the dor- sal and lateral surfaces of the head (Fig. 2). On the ventral surface of the head, the tu- bercles are arranged in discrete rows along the ventral surfaces of the branchiostegal rays. Scales on the surface of the body have one to four tubercles, other than those scales medial to the pectoral fin whose sur- faces are covered with the hook-shaped contact organs (described in the previous section and indicated by white arrow of Fig. 2). When present, the tubercles on the scales are positioned toward the posterior margin of the scale, and when three or four tubercles occur on an individual scale, these structures are arranged in an arch parallel- ing the posterior margin of the scale. The size and number of tubercles tend to be re- duced on the scales of the ventrolateral por- tion of the body. An extensive series of tu- bercles occurs, however, on scales of the prepelvic region of the body, with a less concentrated field of tubercles present in the region from the insertion of the pelvic fin to the origin of the anal fin. VOLUME 117, NUMBER 4 Breeding tubercles are present on all fins of Nannocharax rubrolabiatus with the ex- ception of the adipose fin. The tubercles on the caudal fin are less developed than those on the remaining fins, being apparent solely as small, raised areas along the basal and middle portions of the caudal-fin rays. Tu- bercles are present on all of the dorsal-fin rays with the exception of the first un- branched and last branched rays. At their maximum degree of development, such breeding tubercles extend along nearly the entire length of each dorsal-fin ray. Some larger examined specimens of N. rubrola- biatus have indications of poorly-developed breeding tubercles on the basal portions of the second unbranched anal-fin rays, with better-developed tubercles present on all but the terminal branched anal-fin ray. The pec- toral fin has tubercles on the dorsal surface of the unbranched rays, but tubercles are absent on the portions of the second un- branched through eighth branched rays with anteriorly-directed, hook-shaped contact or- gans. The ventral surface of the pectoral fin has at most a few tubercles distributed along the unbranched rays, but such struc- tures are completely absent in some indi- viduals. Variably-developed series of tuber- cles extend along the length of the ventral surfaces of each of the branched pectoral- fin rays. The pelvic fin has a series of tu- bercles arranged along the dorsal surface of the branched rays, and along the ventral surfaces of the last unbranched fin ray and all of the branched fin rays with the excep- tion of the medialmost branched ray. Our comparative studies failed to reveal any comparably well-developed breeding tubercles in the other examined species of Nannocharax. Larger examined specimens of N. reidi do, however, have a pattern of small, papillae-like processes on the upper lip, snout, and dorsal surfaces of the head that have an arrangement comparable to the pattern of the breeding tubercles that occur in those regions in most examined speci- mens of N. rubrolabiatus. It will be nec- essary to examine additional population 561 samples of N. reidi captured during the height of the breeding season in order to determine whether the papillae-like pro- cesses present in that species would devel- op into the distinctly larger breeding tuber- cles that typify the examined sample of UN. rubrolabiatus. Broader comparative studies would possibly also yield insight in the range of the distribution of breeding tuber- cles across the species of Nannocharax. Comparative material examined.—Nan- nocharax altus: MRAC 78-22-P-801-804, 4 specimens, Republic of the Congo, Mayala, Niola Creek. Nannocharax fasciatus: USNM 303754, 5 specimens; USNM 303756, 2 specimens; USNM 303811, 1 specimen; USNM 303847, 3 specimens; USNM 303867, 2 specimens; USNM 303908, 4 specimens; USNM 303995, 2 specimens; USNM 304081, 3 specimens; USNM 375192, 5 specimens; Cameroon, upper Cross River system. Nannocharax intermedius: CU 80570, 2 specimens, Gabon, Motoboi Village, Kine- né Creek; CU 90276, 3 specimens, Gabon, Okolville; MRAC 91-79-P-202-206, 4 specimens, Gabon, Riviere Loukénini; MRAC A2-006-P-0826-0828, 3 specimens, Gabon, Ivindo basin, Balé Creek. Nannocharax maculicauda: USNM 224524, 3 paratypes; Gabon, upper Ivindo River (1°20’N, 13°12’E); CU 80621, 1 specimen, Gabon, Woleu-Ntem, Ngomo River (1°42'N, 11°38’E). Nannocharax parvus: CU 80148, 19 specimens, Gabon (0°34’S, 10°12’E). CU 80163, 1 specimen; CU 80185, 2 speci- mens; CU 80184, 3 specimens; Gabon, Bi- roundou Creek (2°13’S, 11°28’E). CU 80191, 1 specimen, Gabon, Mimboumbou Creek, near Franceville (1°38’S, 13°31’E). CU 80279, 5 specimens, Gabon, Okolo- ville. CU 80607, 6 specimens, Gabon, stream at Okolville (1°29’S, 13°31’E). Nannocharax rubrolabiatus: MRAC 95- 22-P-001-007, 7 specimens, Cameroon, Sanaga River basin, Mi River (6°12'N, 14°23'E). 562 Acknowledgments Research associated with this project was supported by the Herbert R. and Evelyn Axelrod Chair in Systematic Ichthyology in the Division of Fishes of the National Mu- seum of Natural History, Smithsonian In- stitution. We thank Melanie L. J. Stiassny, Scott A. Schaefer, Barbara Brown, and Radford Arrindell (AMNH), John Friel (CU), and Emmanuel Vreven and the late Guy Teugels (MRAC) for the loan of spec- imens and other assistance. Assistance at USNM was provided by David Smith and in particular Sandra Raredon who also pre- pared Figs. 1 and 2. Gordon McG. Reid, North of England Zoological Society, pro- vided information on the collecting locali- ties of the type-series and coloration of re- cently captured specimens. The paper ben- efitted from the comments and suggestions of Thomas A. Munroe. Literature Cited Boulenger, G. A. 1909. Catalogue of the fresh-water fishes of Africa in the British Museum (Natural History). Volume 1. British Museum (Natural History), London, 373 pp. Coenen, E. J., & G. G. Teugels. 1989. A new species of Nannocharax (Pisces, Distichodontidae) from South-East Nigeria and West Cameroun, with comments on the taxonomic status of Hemigrammocharax polli Roman, 1966.—Cy- bium 13(4):311-318. Collette, B. B. 1977. Epidermal breeding tubercles and bony contact organs in fishes. Pp. 225-268 in R. I. C. Spearman, ed., Comparative Biology of the Skin.—Symposia of the Zoological Society, London, 39. Zoological Society of London, London. Daget, J. 1961. Note sur les Nannocharax (Poissons Characiformes) de l’Ouest africain.—Bulletin de I’Institute Frangais d’Afrique Noire, Dakar, series A 23(1):165-181. , & J. P Gosse. 1984. Distichodontidae. Pp. 184-211 in J. Daget, J.-P Gosse, & D. EF E. Thys van den Audenaerde, eds., Check-list of the freshwater fishes of Africa——Musée Royal de l’Afrique Centrale, Tervuren, Belgium and Office de la Recerche Scientifique et Technique Outre-Mer, Paris. Fink, W. L., & S. H. Weitzman. 1974. The so-called cheirodontin fishes of Central America with de- scriptions of two new species (Pisces: Characi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dae).—Smithsonian Contributions to Zoology 172:1—42. Géry, J. 1977. Characoids of the World.—T.FH. Pub- lications Inc., Neptune City, New Jersey, U.S.A., 672 p. Gosse, J.-P., & E. J. Coenen. 1990. Distichodontidae. Pp. 237-260 in C. Lévéque, D. Paugy, & G. C. Teugels, eds., Faune des poissons d’ eaux douces et saumatres d’ Afrique de l’Ouest. Volume 1.— Musée Royal de 1’Afrique Centrale, Tervuren, Belgium and Office de la Recerche Scientifique et Technique Outre-Mer, Paris. Johnson, G. D. 1984. Percoidei: development and re- lationships. Pp. 464—498 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahey, & S. L. Richardson, eds., Ontogeny and systematics of fishes.—American Society of Ichthyologists and Herpetologists, Special Publication No. 1. Malabarba, L. R., & S. H. Weitzman. 2003. Descrip- tion of a new genus with six new species from southern Brazil, Uruguay and Argentina, with a discussion of a putative characid clade (Teleos- tei: Characiformes: Characidae).—Comunica- ¢does do Museu de Ciéncias e Tecnologia da PUCRS, Porto Alegre, Série Zoologia 16(1): 67-151. Reid, G. M. 1989. The Korup project; the living waters of Korup Rainforest—W.W.F (U.K.) Report 3206/A8:1. 72 p. Roberts, C. D. 1993. Comparative morphology of spines scales and their phylogenetic significance in the Teleostei.—Bulletin of Marine Sciences 52(1):60-113. Taylor, W. R., & G. C. Van Dyke. 1985. Revised pro- cedures for staining and clearing small fishes and other vertebrates for bone and cartilage.— Cybium 9:107—119. Teugels, G. G., G. M. Reid, & R. P. King. 1992. Fishes of the Cross River basin (Cameroon—Nigeria). Taxonomy, zoogeography, ecology and conser- vation.—Annales Sciences Zoologiques, Musee Royal de 1’ Afrique Centrale 266:1—132. Thys van den Audenaerde, D. F E. 1961a. Lanatomie de Phractolaemus ansorgei Blgr. et la position systématique des Phractolaemidae.—Annales, Musee Royal de |’ Afrique Centrale, Sciences Zoologiques 103:99—170. . 1961b. Existence de deux races géograph- iques distinctes chez Phractolaemus ansorgei Blgr. 1901 (Pisces, Clupeiformes).—Bulletin des Sciences. Académie Royale des Sciences d’Outre-mer 7(2):222-251. Van den Bergh, E., G. G. Teugels, E. J. Coenen, & FE Ollevier. 1995. Nannocharax rubrolabiatus, a new species of distichodontid fish from the San- ga River basin in Cameroon, Africa (Teleostei: Distichodontidae).—Ichthyological Explora- tions of Freshwaters 6(4):349-—356. VOLUME 117, NUMBER 4 Vari, R. P. 1979. Anatomy, relationships and classifi- cation of the families Citharinidae and Disti- chodontidae (Pisces, Charcoidea).—Bulletin of the British Museum (Natural History), Zoology Series, 36(5):261—344. , & J. Géry. 1981. Nannocharax maculicauda, a new species of African characoid fish (Char- acoidea: Distichodontidae) with comments on the genus Hemmigrammocharax.—Proceedings 563 of the Biological Society of Washington 94(4): 1076-1084. Wiley, M. L., & B. B. Collette. 1970. Breeding tuber- cles and contact organs in fishes: their occur- rence, structure, and significance.—Bulletin of the American Museum of Natural History 143(3):143-216. Associate Editor: Edward O. Murdy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):564—5S74. 2004. Rhamdia guasarensis (Siluriformes: Heptapteridae), a new species of cave catfish from the Sierra de Perija, northwestern Venezuela Carlos DoNascimiento, Francisco Provenzano, and John G. Lundberg (CDN) Seccién de Ictiologia, Museo de Historia Natural La Salle, Fundacion La Salle de Ciencias Naturales, Apdo. 1930 Caracas 1010-A, Venezuela, carlos.donascimiento @ fundacionlasalle.org.ve; (FP) Laboratorio de Biosistematica de Peces, Instituto de Zoologia Tropical, Universidad Central de Venezuela, Apdo. 47058 Caracas 1041-A, Venezuela, fprovenz @strix.ciens.ucv.ve; (JGL) Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, Pennsylvania 19103, USA, lundberg @acnatsci.org Abstract.—Rhamdia guasarensis n. sp. is described from subterranean waters in the Rio Guasare drainage of northwestern Venezuela. The new species is distinguished from congeners by its concave head profile; medially sutured frontal bones; small, circular vestige of the anterior cranial fontanelle; and troglomorphic characters such as absence of eyes and pigmentation, wide ce- phalic laterosensory pores, and wide fossae of preoperculomandibular sensory canal in preopercle and dentary. Cave catfish diversity in the Sierra de Perija region of Venezuela is reviewed and compared to cave catfish diversity else- where in South America. Resumen.—Se describe Rhamdia guasarensis sp. n. proveniente de aguas subterraneas de la cuenca del Rio Guasare en el noroccidente de Venezuela. La nueva especie se diferencia de las restantes especies que conforman el género por su perfil dorsal de la cabeza concavo; huesos frontales suturados medialmente; fontanela craneal anterior reducida a un pequeno foramen cir- cular; y caracteres troglomorficos tales como ausencia de ojos y pigmentacion, poros cefalicos latero sensoriales anchos, fosas ensanchadas del canal sensorial preoperculomandibular en el preopérculo y dentario. La diversidad de bagres cavernicolas de la Sierra de Perija es revisada y comparada con la diversidad de bagres cavernicolas de otras regiones de Suramérica. The family Heptapteridae has invaded and adapted to hypogean waters multiple times. Among Neotropical catfish families, heptapterids have the greatest diversity of truly troglobitic taxa: Phreatobius cister- narum, Pimelodella kronei, Rhamdia lalu- chensis, Rhamdia laticauda typhla, Rham- dia macuspanensis, Rhamdia quelen urichi, Rhamdia redelli, and Rhamdia zongolicen- sis. Trajano & Bockmann (2000) described the ecology and behavior of Taunayia sp., a troglobitic catfish, inhabiting caves of northeastern Brazil, but the species has not been formally named. Pimelodella spelea Trajano, Reis & Bichuette, 2004 is a re- cently described troglophile without marked specializations for hypogean life. Taxonomic practice has shifted away from assigning supra-specific rank to cave-dwell- ing fishes solely on account of their trog- lobitic adaptations. Among Heptapteridae, the nominal monotypic genera Caecorham- dia, Caecorhamdella, and Typhlobagrus have long been treated as synonyms of Rhamdia and Pimelodella respectively. Fur- thermore, Silfvergrip (1996) synonymized all cave populations of Rhamdia described as separate species with R. quelen or R. VOLUME 117, NUMBER 4 laticauda, both wide-ranging epigean spe- cies. In this paper we describe a new troglob- itic heptapterid species in the genus Rham- dia. Our placement of the new species is more a matter of convenience than firm phylogenetic resolution. Rhamdia is taxo- nomically complex. In the latest revision of the genus, Silfvergrip (1996) consolidated its approximately 100 nominal species into eight and he described three new species. In 1998, Weber & Wilkens described the blind species R. macuspanensis, and in 2003, Weber et al. described Rhamdia lal- uchensis, another troglobitic species from Mexico. In the most thorough phylogenetic study of Heptapteridae to date, Bockmann (1998) concluded that Rhamdia is non- monophyletic but he did not attempt to re- solve the genus into phylogenetically di- agnosable units. As it stands, Rhamdia is a non-monophyletic assemblage of common fishes with an immense geographic distri- bution in South and Middle America from the lower Parana Basin in Argentina to cen- tral México. The new species, from a cave in the Si- erra de Perija region of northwestern Ve- nezuela, is distinct both in its typical trog- lobitic specializations and other apomorph- ic features, but overall it is most similar to other Rhamdia. Discovering the relation- ships of the new species and, more gener- ally, resolving the relationships of Rhamdia species are major problems quite beyond our present scope. Our immediate concern is to name and describe this previously un- seen species that has a highly restricted dis- tribution in a marginal and potentially frag- ile habitat. We comment also on the sub- terranean catfish fauna of the Perija region. Material and Methods Morphometric measurements follow the criteria set out by Lundberg & McDade (1986) and Bockmann (1994). Terminology of cephalic laterosensory canals and branches follows Arratia & Huaquin’s de- 565 scription of Diplomystes and Nematogenys (1995). However, our numbering of sensory pores in Rhamdia reflects anteroposterior or mesiolateral pore order, and does not imply individual homologies of pores among cat- fishes. All measurements were made on the left side of the specimens using a Mitutoyo digital, needlepoint caliper at a precision of 0.1 mm. For osteological observation one paratype (101.1 mm SL) was cleared and stained using the method of Taylor & Van Dyke (1985). A second paratype (93.4 mm SL) was radiographed. Only these two specimens were used for counts of verte- brae, branchiostegal rays, ribs, and ptery- giophores. The vertebral count includes the first five vertebrae incorporated into the Weberian apparatus whereas the compound caudal centrum is counted as one. Institu- tional abbreviations follow Leviton et al. (1985). Other abbreviations are: SL—stan- dard length, HL—head length, CS—cleared and stained skeletal preparation, alc— whole specimen preserved in alcohol. Rhamdia guasarensis, new species Figs. 1-4 Holotype.—MBUCV-V-29604: 106.8 mm SL; Surgencia del Tigre at 2.5 km W of Cerro Yolanda, Rio Guasare basin, Sierra de Perija, Estado Zulia, Venezuela (10°52’53"N, 72°30'03”W). Elevation 200 m asl; collected by J. Lagarde, 3 April 1999. Paratypes.—All collected with the ho- lotype: MBUCV-V-29622, two specimens, 87.2—101.1 mm SL, the second cleared and stained; ANSP 179878, one specimen, 93.4 mm SL. Diagnosis.—Rhamdia guasarensis 1s dis- tinguished from congeneric species by two characters: dorsal profile of head concave (Fig. 1, vs. convex or straight); and frontal bones broadly sutured to each other anterior to small, circular remnant foramen of an- terior cranial fontanelle that is anteriorly ad- jacent to epiphyseal bar (Fig. 2, vs. frontals separated by anterior fontanelle -widely 566 Fig. 1. of head; C, ventral view of head. open from mesethmoid to epiphyseal bar). Rhamdia guasarensis differs from all epi- gean Rhamdia by the following troglo- morphic characters: absence of eyes, com- plete depigmentation, widened cutaneous pores of the cephalic laterosensory system, preoperculomandibular sensory canal form- ing wide fossae in the dentary and preo- percle (Fig. 3, vs. narrow pores and canals). In addition to these characteristics, R. guasarensis can be distinguished from other species of the genus by the following com- bination of characters: pectoral fins with a spine and ten branched rays (vs. modally PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON i i j i i if i hy ie I | Rhamdia guasarensis. Holotype MBUCV-V-29604, 106.8 mm SL. A, lateral view; B, dorsal view eight or nine soft rays in other species, data from Silfvergrip 1996); both lobes of the caudal fin pointed (vs. at least one lobe rounded); caudal skeleton with three hy- pural plates, PH; 1 + 2; 3 + 4 + 5 Ws. modally four PH; 1 + 2; 3 + 4; 5 in the other species with the exception of R. lau- kidi and R. jequitinhonha, see Silfvergrip 1996). Description.—Morphometric data are presented in Table 1. Body elongate, strong- ly depressed anteriorly and gradually more compressed from origin of pectoral fins to caudal peduncle. Shape approximately tri- VOLUME 117, NUMBER 4 Fig. 2. Rhamdia guasarensis. Skull roof illustrat- ing reduction of the anterior cranial fontanelle and midline contact of frontal bones. MBUCV-V-29622, 101.1 mm SL. Abbreviations: ACE anterior cranial fontanelle; FR, frontal bone contact on midline; LET, lateral ethmoid; MES, mesethmoid; PCE posterior cra- nial fontanelle; PT, pterotic; SPH, sphenotic. angular in transverse section at dorsal-fin origin. Dorsal profile sinusoidal anterior to dorsal fin, then approximately straight to middle of adipose fin, then slightly concave along caudal peduncle. Ventral profile near- ly straight to anal-fin origin, then slightly concave posteriorly. Head depressed, its dorsal profile con- cave, its lateral and ventral profiles nearly straight. Mouth terminal, upper jaw slightly in advance of lower jaw. Rictal folds little developed. Upper and lower lips with weak sulci, slightly evident in holotype, forming single labial fold. Premaxillaries with single band of diminutive teeth, arranged in ten irregular tooth rows, the posterolateral cor- ners rounded, not produced. Dentition of 567 Fig. 3. culomandibular laterosensory canal and associated fo- ramina. MBUCV-V-29622, 101.1 mm SL. Abbrevia- tions: ANG, anguloarticular; D, dentary; HYO, hy- omandibula; MPT, metapterygoid; OP, opercle; POP, preopercle; Q, quadrate. Rhamdia guasarensis. Enlarged preoper- lower jaw similar to premaxillary teeth, in six irregular tooth rows. Palatine and vomer edentulous. Maxillary barbels long, extend- ing beyond base of pelvic fins. Mental bar- bels relatively short, inner mentals scarcely reaching posterior border of branchial membrane; outer mentals surpass pectoral fin bases. Inner mental barbel bases inserted slightly in advance of outer mental barbel bases. Anterior nares tubular, near border of snout. Posterior nares with elongated orific- es, bounded anterolaterally by membrane of fine skin. Internarial length less than width between posterior nares. Eyes completely absent. Branchial membranes overlapping medially; united to isthmus only anteriorly. Cephalic lateralis sensory system with paired supraorbital (SO), infraorbital (IO), preopercular (POP), mandibular (MA), otic (OT), and post-otic (POT) canals, without tubular commissure connecting supraorbital canals. Sensory pores simple, not branched and multiple. SO canal with six pores: SO1—SO3 associated with nasal bone, SO1 medially adjacent to anterior naris, wide and delimited by membrane of fine skin, SO2 between anterior and posterior nares, slightly closer to first, SO3 posteromedially near posterior naris. SO4 near dorsal mid- line at end of short medial tube and separate from its counterpart of opposite side. SOS lateral to its canal midway between SO4 and union of SO and IO canals. SO6 medial to its canal a little posterior to union of SO and IO canals. 568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1—Measurement data for the type series of Rhamdia guasarensis. Measurement | expressed in mm. Proportional measurements expressed as thousandths of standard length (2-19; 26—28) or head length (20—25). Holotype MBUCYV-V-29604 1. Standard length 106.8 2. Total length 1057 3. Body depth 162 4. Body width 171 5. Predorsal length 351 6. Preanal length 645 7. Prepelvic length 462 8. Preadipose length IS 9. Caudal peduncle length 211 10. Caudal peduncle depth 97 11. Dorsal fin spine length 85 12. Length of first branched dorsal fin ray 172 13. Dorsal fin base 115 14. Adipose fin length 387 15. Dorsal fin to adipose fin 99 16. Anal fin base 141 17. Pectoral fin spine length 121 18. Length of first branched pectoral fin ray 204 19. Pelvic fin length 169 20. Head length 259 21. Head width 661 22. Head depth 550 23. Internarial length 137 24. Anterior internarial width 173 25. Posterior internarial width 156 26. Maxillary barbel length 541 27. Outer mental barbel length 205 28. Inner mental barbel length 98 IO canal with four pores; [O1—3 wide like SO1. IO1 posterior to anterior nostril; [O02 emerges dorsal to groove for maxillary barbel, posterior to base of barbel; IO3 near point where IO canal curves dorsally; 104 at tip of short posterior tube near union with SO canal. Holotype and one paratype (87.2 mm SL) have different single supernumer- ary IO pores; extra pore of holotype from left canal between the IO2 and IO3; extra pore of paratype from right IO canal be- tween IO3 and [O04. POP canal with four pores; MA canal with seven pores; all except MAI and POP4 originate from much enlarged cavi- ties in dentary and preopercular bones. MAT in mental position near to midventral line at tip of its branch from lower jaw sym- physis. Paratype Paratype MBUCV-V-29622 Paratype MBUCV-V-29622 (CS) ANSP 179878 87.2 101.1 93.4 1093 1083 178 167 180 168 173 367 352 359 653 659 656 458 490 545 530 210 214 224 104 105 86 108 176 183 187 121 104 113 411 432 394 2 68 127 127 143 111 123 197 5 216 153 159 169 262 256 276 645 631 633 S22 538 528 144 147 169 187 152 165 610 557 578 262 258 262 121 108 121 POT canal with two pores, POT1 over pterotic dorsal to gill opening; POT2 dorsal to supracleithrum and above main lateralis canal at level of first pore. First pore of la- teralis canal at end of ventral branch dorsal to postcleithral process. Several following pores also at tips of short postero-ventral branches. Lateral line canal complete to base of middle upper-lobe caudal rays. Dorsal fin with a spinelet, spine and six branched rays; its margin rounded. Dorsal spine weakly developed, only its basal part rigid and unsegmented; dentations diminu- tive and scarcely visible, limited to basal part of anterior margin. The distal two- thirds of dorsal spine flexible and obliquely segmented. Adipose fin long and low, its origin near tip of depressed dorsal fin, and extending posteriorly to approximately 80% VOLUME 117, NUMBER 4 =S x au et 7 5 mm Fig. 4. Rhamdia guasarensis. Pectoral spine in dorsal view. Holotype MBUCV-V-29604, 106.8 mm SL. of caudal peduncle length; posterior end of adipose fin adnate to caudal peduncle with- out a free fleshly tab. Caudal fin deeply forked; both caudal lobes pointed; upper lobe slightly longer than lower; membrane uniting innermost caudal rays complete. Principal caudal rays 1,7-8,1, except 1,7-7,1 in one paratype. Anal fin with 12 rays, an- teriormost two or three rays simple, others branched; its margin rounded. Pectoral fins with a spine and ten branched rays. Pectoral spine (Fig. 4) with weak dentations proximally on anterior margin; distal half of spine flexible and obliquely segmented. First branched pec- toral-fin ray longest, posterior branched rays diminishing in length. Postcleithral process short, sharp. Pelvic fins with one simple ray and five branched rays, its origin posterior to end of dorsal fin. Skull roof (Fig. 2) with anterior cranial fontanelle extremely reduced to a small cir- cular foramen located in front of epiphyseal bar; mesethmoid posteriorly lacking con- cave notch of fontanelle, and frontals meet- ing medially along most of their length. Posterior cranial fontanelle reduced to oval foramen near center of supraoccipital. Su- praoccipital process short, its length about equal to length of supraoccipital body. Anus and urogenital papillae separated, anus located equidistant between medial edge of pelvic-fin base and urogenital pa- pilla, approximately at midlength along pel- vic fins; urogenital papilla conspicuous and elongated, located closer to base of anal fin than to base of pelvic fin. Total vertebrae 40—42; neural spines of vertebrae 6—10 bifid; hemal arch closed in vertebra 12 or 13, first hemal spine on ver- tebra 14, 15 or 16; eight pairs of ribs borne on vertebrae 6—13. Seven dorsal-fin ptery- 569 72W = Rio Guasare Lago de Maracaibo Fig. 5. The Lago de Maracaibo—Sierra de Perija region, Venezuela, showing type locality (star) of Rhamdia guasarensis. Map based on shaded relief im- age PIA03388, Shuttle Radar Topography Mission, National (NASA). Aeronautics and Space Administration giophores preceded by small supraneural; first dorsal-fin pterygiophore inserted be- tween rami of neural spine of fourth ver- tebra. Eleven anal-fin pterygiophores, first inserting posterior to hemal spine of verte- bra 21. Caudal skeleton with three hypural plates: rectangular parhypural; triangular hypurals 1 + 2; triangular hypurals 3 + 4 sD Color in alcohol.—Body and fins com- pletely depigmented; most of skin, rayed- and adipose-fin membranes hyaline and translucent; musculature appearing yellow- ish, particularly jaw adductors and dorsal trunk myomeres; parts of head and fin bases whitish. Distribution and habitat.—R. guasaren- sis 1s known only from the Surgencia del Tigre (Zu. 23), in the middle basin of the Rio Guasare, north of the Sierra de Perija in northwestern Venezuela (Fig. 5). The cave is near the margin of Rio Guasare and is the source of a spring during seasonal rains (Sociedad Venezolana de Espeleologia 570 1991). The cave’s lower conduit has a 280 m course, 2—3 m wide and 1—2 m high, nar- rowly communicating with the access gal- lery. The underground river is permanently fed by a spring about 60 m into the lower gallery. At the time the cave was surveyed, the average depth of this water course was 1.5 m with deeper pools along its course where the catfishes were observed (Socie- dad Venezolana de Espeleologia 1991). Etymology.—The name is based on Rio Guasare, parent stream of the subterranean waters in which this endemic catfish species lives. Discussion Rhamdia guasarensis is placed in Hep- tapteridae by its possession of four syna- pomorphies identified for the family (Lund- berg & McDade 1986, Ferraris 1988, Bock- mann & Guazelli 2003): posterior limb of fourth transverse process expanded and notched; posterodorsal corner of hyoman- dibula greatly expanded for attachment of levator operculi muscle; dorsal margin of quadrate free, not sutured to hyomandibula and metapterygoid; ventrolateral corner of mesethmoid anteriorly recurved. However, the new species lacks a fifth synapomorphy of heptapterids: a straight-edged vertical bony lamina on the Weberian complex cen- trum. Instead, the vertical lamina has a con- cave margin in R. guasarensis. Except for the obvious lack of a free or- bital rim, R. guasarensis possesses the character combination presented as diag- nostic of Rhamdia by Silfvergrip (1996:74). This includes: three pairs of barbels, double lip fold, vomer without teeth, transverse processes of fourth vertebra expanded branched distally, supraoccipital process not contacting anterior nuchal plate, adi- pose fin with free posterior margin, poste- rior fontanelle closed and postcleithral pro- cess well developed. However, none of these are unambiguous synapomorphies of the group of species comprising Rhamdia. Instead, some characters are heptapterid or PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON higher level synapomorphies, and others, some of uncertain polarity, have wider and variable distributions among heptapterids. Thus, placement of this new species in Rhamdia must be considered provisional because the genus has not been supported as monophyletic. Bockmann’s (1998) phy- logenetic analysis of Heptapteridae placed one representative species, R. laticauda, sister to Pimelodella but a second species, R. quelen, is deeper in his cladograms. In this context R. guasarensis has one derived, although non-unique, feature listed by Bockmann as diagnostic for R. quelen. This is the highly reduced posterior cranial fon- tanelle, long used as one of the diagnostic characters of Rhamdia. Indeed, we find the posterior fontanelle closed or reduced to a small foramen in the supraoccipital in most other Rhamdia examined: R. laukidi, R. ni- caraguensis, R. quelen (including speci- mens originally identified as R. guatema- lensis, R. hilarii, R.vilsoni, R. wagneri) and some R. laticauda. Silfvergrip (1996) re- ported the fontanelle to be variably open or reduced in R. laticauda, and our sample also shows such variability among speci- mens. We find that R. muelleri has an open posterior fontanelle. The fontanelle is also closed in the heptapterids Brachyglanis, Brachyrhamdia, Leptorhamdia, and Myog- lanis (Bockmann 1998, pers. obs.). Fur- thermore, R. guasarensis has an uncinate process on hypobranchial 1, unlike R. que- len that lacks the process (listed as a second non-unique derived feature of R. quelen by Bockmann 1998). Accordingly, we do not take the foregoing as evidence for a partic- ularly close phylogenetic relationship be- tween R. guasarensis and R. quelen. The midline union of frontal bones (Fig. 2) and concomitant extreme reduction of the an- terior fontanelle are a distinctive apomorph- ic character of R. guasarensis. Although not all species have been examined for this fea- ture, we have not observed it in any Rham- dia nor has it been previously reported, and in his description of the genus, Silvergrip (1996:74) reported the anterior fontanelle to VOLUME 117, NUMBER 4 be invariably open. This is at least a diag- nostic autapomorphy of the species, al- though these features are potentially infor- mative about relationships. Bockmann (1998) illustrated a variety of conditions of anterior fontanelle narrowing and closure in other heptapterids including Myoglanis, Taunayia, Imparfinis pristos, and an unde- scribed species, but all of these are struc- turally different from that in R. guasarensis. Another peculiar character of the new species is the concave dorsal profile of the head. In general, Rhamdia species, includ- ing most cave populations, have convexly rounded heads. The cave species R. macus- Panensis recently described from Mexico (Weber & Wilkens 1998) has a straight dor- sal head profile, somewhat more similar to that of R. guasarensis than to other con- geners. Rhamdia macuspanensis is readily distinguished from-R. guasarensis by its strong development of pectoral spine den- tations and rounded tips of the caudal lobes. Rhamdia guasarensis possesses typical reductive characteristics in common with other cave-dwelling species and popula- tions of the genus. Furthermore, the greater relative length of the head and elevated number of pectoral-fin rays are also shared by other troglobitic species of the genus (Weber 1996). It has been suggested that larger head size is related to an increase in the development of the cephalic laterosen- sory system (Langecker & Longley 1993, Weber 1996), and the greater number of pectoral-fin rays is possibly correlated with the increased mass of the anterior part of the body, compensating this increase with a greater fin area for hydrodynamic lift (We- ber 1996). If there is a functional relation- ship between head size and pectoral-fin area in cave dwelling Rhamdia, it does not ex- tend to the heptapterid genus Pimelodella, wherein the large-headed P. kronei has eight or nine pectoral-fin rays (Trajano 1997, Trajano & Britski 1992) and the small-headed P. spelaea has ten pectoral- fin rays (Trajano et al. 2004). No Rhamdia species are known from the 571 surface waters of Rio Guasare, thus R. guasarensis 1s not a cave-dwelling ecotype of a proximate epigean species. Two Rham- dia species have been reported from north- western Venezuela. From the Lago de Ma- racaibo Basin, Schultz (1944) published on specimens now identified as R. quelen (Sil- fvergrip 1996). Fernandez-Yépez & Martin (1953) reported R. wagneri based on spec- imens collected in the Rio Negro in the southern part of the Perija range. One of us (CDN) has reidentified these specimens at the Museo de Historia Natural La Salle as R. quelen. As noted above, there is no ev- idence for a uniquely close relationship be- tween R. guasarensis and R. quelen. The fauna of troglobitic catfishes of the Sierra de Perija region includes: Ancistrus galani Pérez & Viloria, 1994, Trichomyc- terus spelaeus DoNascimiento, Villarreal & Provenzano, 2001, and Rhamdia guasar- ensis. There is another hypogean population of Trichomycterus, possibly an undescribed species, living in a cave drained by the Rio Yasa (Rio Negro system) in the southern part of the Sierra de Perija (DoNascimiento, in prep.). The diversity of three cave catfishes of the Rio Guasare system is among the high- est of any Neotropical karst region, al- though the species are not found syntopi- cally in the same cave. By contrast Bichu- ette & Trajano (2003) list five troglobitic species in caves of the Sao Domingos karst area, Goias, Brazil. Ancistrus cryptophthal- mus and the trichomycterid /tuglanis pas- sensis coexist in the Sao Vicente cave, To- cantins Basin, Goiads, Brazil (Trajano & Souza 1994, Fernandez & Bichuette 2002). Also, the inundated caves of the Formosin- ho karst region of Bodoquena, Mato Grosso do Sul, southeastern Brazil, are co-inhabit- ed by Ancistrus formoso and an unde- scribed troglomorphic population of T7i- chomycterus (Sabino & Trajano 1997). Cave-dwelling and specialized troglobitic neotropical catfishes belong to the families Astroblepidae, Heptapteridae, Loricariidae, and Trichomycteridae. Within the last three 572 of these families the genera Rhamdia, An- cistrus, and Trichomycterus are most com- monly represented in cave faunas. Their prevalence suggests possession of morpho- logical, physiological, behavioral, and eco- logical features (preadaptations) that facili- tate existence in cave waters (Eigenmann 1919, Norman 1926, Hubbs 1936). Wilkens (1986) proposed a correlation between degree of morphological reduction and time of subterranean evolution based on a neutral mutation model for the regres- sive evolution of eyes and pigmentation in cave fishes and crustaceans. The subterra- nean catfishes of the Sierra de Perija, es- pecially Trichomycterus spelaeus and R. guasarensis, are highly advanced in their troglobitic features, suggesting that they are not new arrivals in their subterranean en- vironment. Ocular and pigmentation reduc- tion of R. guasarensis and T. spelaeus are complete. These species exhibit additional autapomorphies such as extremely elongate barbels in Trichomycterus and much en- larged head laterosensory organs in Rham- dia. These characters, too, may indicate a long period of hypogean evolution. Indirect evidence suggests the availability of an am- ple period of time for the evolution of the Perija cave fishes. Paleogeographic recon- structions of northwestern Venezuela sug- gest that uplift of the Sierra began in the early Cenozoic (Gonzalez de Juana et al. 1980). It is reasonable to assume that these fishes originated in situ after subterranean waters carved out their habitat within Cre- taceous limestones of the Sierra de Periya. On the fish side of the equation, the only fossil record of Rhamdia are fin spines of relatively young Pleistocene age (Cione 1982). However, based on much older Mio- cene fossils of phylogenetically related pi- melodid and pseudopimelodid catfishes, the heptapterids are expected to have originated and diversified long before the late Pleis- tocene (Lundberg 1998). Thus it is possible that subterranean aquatic habitats and cave fishes have been present in this region for tens of millions of years. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Comparative material.—Rhamdia lati- cauda: ANSP 104034, one specimen, X-ray and alc, 86 mm SL, Panama, Cocle: UMMZ 197078, two dry skeletons, 131— 146 mm SL, Honduras. R. laukidi: ANSP 139184, one of three specimens, X-ray and alc, 127 mm SL, Colombia, Meta. R. muel- leri: ANSP 162521, two of four specimens, X-ray and alc, 109-110 mm SL, Venezuela, Amazonas. R. nicaraguensis: ANSP 8444, one specimen, alc, 135 mm SL, Nicaragua, Lago Nicaragua. Rhamdia quelen: ANSP 141578, two of five specimens, X-ray and alc, 100-105 mm SL, Venezuela, Bolivar; ANSP 45365 (original identification R. guatemalensis), one specimen, X-ray and alc, 120 mm SL, Panama, Canal Zone; ANSP 172138 (original identification R. hi- larii), two of 37 specimens, X-ray and alc, 107-110 mm SL, Brazil, Minas Gerais; ANSP 16020 (original identification R. vil- soni), one specimen, alc, 200 mm SL, Trin- idad; ANSP 71621 (original identification R. wagneri), one specimen, X-ray and alc, 125 mm SL, Colombia, Magdalena; DU- F1021, one dry skeleton, 202 mm SL; MBUCV-CT-561, eight specimens, CS, 23— 57 mm SL, Venezuela, Zulia; MHNLS- 1645, two specimens, alc, 59-223 mm SL, MHNLS-1734, three specimens, alc, 107— 228 mm SL, Venezuela, Zulia. Acknowledgments We are indebted to the members of So- ciedad Venezolana de Espeleologia, espe- cially O. Villarreal who both brought us the specimens described here and assisted with illustration of the skull. K. Luckenbill ably prepared the final figures. H. H. Ng gener- ously provided us with character data on comparative skeletal specimens at UMMZ. We are indebted to two anonymous and careful reviewers for many useful com- ments on the manuscript. N. Milani de Ar- nal photographed the holotype, and M. Litt- mann radiographed the ANSP paratype. Partial support of publication costs was pro- vided by the All Catfish Species Inventory VOLUME 117, NUMBER 4 (NSF DEB-0315963) and an NSF research award to JGL (DEB-0089612). 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 systematics and ecological consider- ations.—Bonner Zoologische Monographien 36:1—110. Bichuette, M. E., & E. Trajano. 2003. Epigean and subterranean ichthyofauna from the S40 Dom- ingos karst area, Upper Tocantins River basin, central Brazil.—Journal of Fish Biology 63(5): 1100. Bockmann, EF A. 1994. Description of Mastiglanis aso- pos a new pimelodid catfish genus from north- ern Brazil, with comments on phylogenetic re- lationships inside the subfamily Rhamdiinae (Siluriformes: Pimelodidae)—Proceedings of the Biological Society of Washington 107(4): 760-777. . 1998. Andlise filogenética da familia Heptap- teridae (Teleostei, Ostariophysi, Siluriformes) e redefenig¢ao de seus géneros. Unpublished doc- toral dissertation, Universidade de Sao Paulo, 599 p. , & M. G. Guazzelli. 2003. Family Heptapter- idae. Pp. 406—431 in R. E. Reis, S. O. Kullan- der, & C. J. Ferraris, Jr, eds., Check List of the Freshwater Fishes of South and Central Amer- ica. EDIPUCRS, Porto Alegre, Brazil, 729 pp. Cione, A. 1982. Peces del Pleistoceno tardfo de la Provincia de Buenos Aires. Consideraciones biogeogrdficas.—Circular informativa de la Asociacion Paleontolo6gica Argentina 8:12. DoNascimiento, C., O. Villarreal, & E Provenzano. 2001. Descripcion de una nueva especie de ba- gre anoftalmo del género Trichomycterus (Sil- uriformes, Trichomycteridae), de una cueva de la Sierra de Perija, Venezuela.—Boletin de la Sociedad Venezolana de Espeleologia 35:20— 26. Eigenmann, C. H. 1919. Trogloglanis pattersoni a new blind fish from San Antonio, Texas.—Proceed- ings of the American Philosophical Society 58(6):397—400. Fernandez, L., & M. Bichuette. 2002. A new cave dwelling species of Jtuglanis from the Sao Domingos karst, central Brazil (Siluriformes: Trichomycteridae).—Ichthyological Exploration of Freshwaters 13(3):273-278. Ferndndez-Yépez, A., & E Martin Salazar. 1953. Apuntes sobre la ictiologia de Perija.—Memo- tia de la Sociedad de Ciencias Naturales La Salle 13(35):227-243. Ferraris, C. J., Jr. 1988. Relationships of the neotrop- 37/3} ical catfish genus Nemuroglanis, with a descrip- tion of a new species (Osteichthyes: Silurifor- mes: Pimelodidae).—Proceedings of the Bio- logical Society of Washington 101(3):509-516. Gonzalez de Juana, C., J. M. Iturralde de Arozena, & X. Picard. 1980.—Geologia de Venezuela y de sus Cuencas Petroliferas. Foninves, Caracas, 2 vols. 1021 pp. Hubbs, C. L. 1936. Fishes of the Yucatan Peninsula.— Carnegie Institution of Washington Publication 457:157—287, pls. 1=15. Langecker, T., & G. Longley. 1993. Morphological ad- aptations of the Texas blind catfishes Troglog- lanis pattersoni and Satan eurystomus (Siluri- formes: Ictaluridae) to their underground envi- ronment.—Copeia 1993(4):976—-986. Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. Dawson. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.—Copeia 1985(3):802—832. Lundberg, J. G. 1998. The Temporal Context for Di- versification of Neotropical Fishes. Chapter 2. Pp. 49—68 in L. R. Malabarba, R. E. Reis, R. P. Vari, C. A. S. Lucena, & Z. M. S. Lucena, eds., Phylogeny and Classification of Neotropical Fishes. Museu de Ciéncias e Tecnologia, PUCRS. Porto Alegre, Brazil, 603 pp. , & L. McDade. 1986. On the South American catfish Brachyrhamdia imitator Myers (Siluri- formes, Pimelodidae), with phylogenetic evi- dence for a large intrafamilial lineage.—Notula Naturae, Academy of Natural Sciences, Phila- delphia 463:1—24. Norman, J. R. 1926. A new blind catfish from Trini- dad, with a list of the blind cave-fishes—An- nals and Magazine of Natural History (Ser. 9) 18(106):324—331 Pérez, A., & A. Viloria. 1994. Ancistrus galani n. sp. (Siluriformes, Loricariidae), with comments on bioespeleological explorations in western Ve- nezuela.—Meémoires de Bioespéologie 21:103— 107. Sabino, J., & E. Trajano. 1997. A new species of blind armoured catfish genus Ancistrus, from caves of Bodoquena region, Mato Grosso do Sul, south- western Brazil (Siluriformes, Loricariidae, An- cistrinae).—Revue Frangaise d’ Aquariologie et Herpetologie 24:73—-78. Schultz, L. P. 1944. The catfishes of Venezuela, with descriptions of thirty-eight new forms.—Pro- ceedings of the United States National Museum 94(3172):173-338. Silfvergrip, A. 1996. A systematic revision of the neo- tropical catfish genus Rhamdia (Teleostei, Pi- melodidae). Department of Zoology, Stockholm University and Department of Vertebrate Zool- 574 ogy, Swedish Museum of Natural History, Stockholm, 156 pp. Sociedad Venezolana de Espeleologia. 1991. Catastro Espeleologico de Venezuela: Surgencia del Ti- gre (Zu. 23).—Boletin de la Sociedad Venezo- lana de Espeleologia 25:30—31. Taylor, W., & G. Van Dyke. 1985. Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study.—Cy- bium 9:107-119. Trajano, E. 1997. Threatened fishes of the World: Pi- melodella kronei (Ribeiro, 1907) (Pimelodi- dae).—Environmental Biology of Fishes 49: 332. , & E Bockmann. 2000. Ecology and behavior of a new cave catfish of the genus Taunayia from northeastern Brazil (Siluriformes: Pime- lodidae).—Ichthyological Exploration of Fresh- waters 11(3):207—216. , & H. A. Britski. 1992. Pimelodella kronei (Ribeiro,1907) e seu sindnimo Caecorhamdella brasiliensis Borodin, 1927: morfologia externa, taxonomia e evolugao (Teleostomi, Silurifor- mes).—Boletim de Zoologia, Sao Paulo 12:53— 89. , R. E. Reis, & M. E. Bichuette. 2004. Pime- lodella spelaea: a new cave catfish from central PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Brazil, with data on ecology and evolutionary considerations (Siluriformes: Heptapteridae).— Copeia 2004(2):315—325. , & A. Souza. 1994. Behaviour of Ancistrus cryptophtalmus, an armoured blind catfish from caves of central Brazil, with notes on syntopic Trichomycterus sp. (Siluriformes, Loricariidae, Trichomycteridae).—Mémoires de Bioespéolo- gie 21:237-243. Weber, A. 1996. Cave dwelling catfish populations of the genus Rhamdia (Pimelodidae, Siluroidei, Teleostei) in Mexico.—Mémoires de Bioespéo- logie 23:73-85. , G. Allegrucci, & V. Sbordoni. 2003. Rhamdia laluchensis, a new species of troglobitic catfish (Siluriformes: Pimelodidae) from Chiapas, Mexico.—Ichthyological Exploration of Fresh- waters 14(3):273—280. , & H. Wilkens. 1998. Rhamdia macuspanen- sis: a new species of troglobitic pimelodid cat- fish (Siluriformes: Pimelodidae) from a cave in Tabasco, Mexico.—Copeia 1998(4):998—1004. Wilkens, H. 1986. The tempo of regressive evolution: studies of the eye reduction in stygobiont fishes and decapod crustaceans of the Gulf Coast and West Atlantic Region.—Stygologia 2:130—143. Associate Editor: Edward O. Murdy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):575—-581. 2004. Taxonomic review of the fossil Procellariidae (Aves: Procellariiformes) described from Bermuda by R. W. Shufeldt Storrs L. Olson Division of Birds, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A., e-mail: olsons@si.edu Abstract.—The literature and specimens relevant to the three new species of petrels (Procellariidae) proposed by R. W. Shufeldt from Quaternary fossils from Bermuda were re-examined. A case is made for citing all three binomials as dating from Shufeldt’s earlier preliminary publication (1916) rather than his later monograph (1922). Aestrelata vociferans Shufeldt, 2 October 1916, was correctly synonymized with Aestrelata cahow Nichols & Mowbray, 31 March 1916, and a lectotype is designated here. Puffinis mcgalli Shufeldt, 1916, was correctly synonymized with Puffinus puffinus Briinnich, 1764, with the holo- type evidently representing a casual occurrence. A lectotype is designated for Puffinus parvus Shufeldt, 1916. This taxon is not synonymous with Puffinus lherminieri.:Lesson, 1839, being much smaller, and is provisionally retained until its status relative to other taxa in the Puffinus assimilis/lherminieri com- plex can be assessed. Because seabirds of the family Procellar- lidae are usually the most prevalent mem- bers of the fossil avifaunas recovered in Bermuda, it is desirable to resolve several taxonomic and nomenclatural problems that were introduced in two papers by R.W. Shufeldt (1916, 1922) in which he named three new species of petrels and shearwa- ters from fossil remains of uncertain age obtained in several caves in Bermuda. Al- though his names were all subsequently synonymized, these actions were taken without reference to Shufeldt’s original ma- terial, most of which is now to be found in the Carnegie Museum of Natural History, Pittsburgh (not the British Museum, as sur- mised by Brodkorb, 1963). The objectives of this review are: (1) to establish the orig- inal citation for each of Shufeldt’s names; (2) to attempt to identify at least parts of the type series upon which each species was based and designate lectotypes where ap- propriate; and (3) to determine autoptically the identity and validity of each of Shu- feldt’s taxa. Considering the deficiencies of the com- parative osteological material available to Shufeldt, his studies of Bermudan fossils are quite exemplary. Regardless of the ul- timate fate of Shufeldt’s names, his analysis of the specimens and his conclusions were for the most part meritorious—something that cannot be said for many of his other studies of fossil birds. Shufeldt’s first con- tribution to Bermudan paleontology (Shu- feldt 1916) was intended only as a prelim- inary introduction to a larger work. He had progressed at least as far as mounting the plates for this proposed monograph, as at this point he refers specifically to the plate and figure numbers of the unpublished larg- er manuscript. The figure numbers men- tioned at this time correspond exactly with those published later (Shufeldt 1922), al- though the plates were renumbered accord- ing to the sequence necessitated by the jour- nal in which they appeared. Publication of the definitive paper was originally to have been through the American Museum of Natural History, but this never took place; 576 the paper was delayed (7 years) and even- tually was issued in the Carnegie Museum series. That a delay was forthcoming must have been apparent to Shufeldt in 1916, as he included an addendum to his preliminary paper in which he named his new taxa, al- though the descriptions accompanying the names were very spare. Some of the names have been construed as nomina nuda at this point (e.g., Brodkorb 1963:246), but for reasons given below I consider all of Shu- feldt’s names to date from the 1916 publi- cation. There were several collections of Ber- mudan fossils upon which Shufeldt based his descriptions of Aestrelata vociferans, Puffinus mcgalli, and P. parvus. The orig- inal one, upon which he had been invited to work by EF A. Lucas “Director of the American Museum of Natural History” (Shufeldt 1916:623), had been obtained ‘by L. L. Mowbray. Material from this collec- tion was identified by Shufeldt (1922) as being from the American Museum (AMNH). Another collection was obtained by Edward McGall and was referred to in Shufeldt (1922) as the McGall Collection. Apparently the AMNH material was never returned and most of Shufeldt’s material that has been traced so far is in the collec- tions of the Carnegie Museum. Further- more, at least one specimen identified in Shufeldt (1922) as coming from the AMNH collection was exchanged from the Carne- gie Museum to the Smithsonian Institution in 1932 (USNM 320059, accession no. 117209). (All USNM and CM catalog num- bers refer to series in the ornithological rather than paleontological collections.) Identifying Shufeldt’s type material is made more difficult by the fact that none of the specimens involved had been cataloged or numbered. It should be noted that McGall and Anthony Tall evidently sent ad- ditional specimens to Harvard University, the British Museum, and perhaps elsewhere (Shufeldt 1922:384), but Shufeldt never ex- amined these specimens and they certainly have no claim as types. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pterodroma cahow (Nichols & Mowbray, 1916) Aestrelata cahow Nichols & Mowbray, 1916 (31 March):194. Aestrelata vociferans Shufeldt, 1916 (2 Oc- tober):633, Shufeldt, 1922:365. Oestrelata vociferans: Lambrecht, DY Ne Pterodroma cahow: Bent, 1922 (19 Octo- ber):112 (new combination with A. vo- ciferans in synonymy); Brodkorb, 1963: 246. IQ32 Lectotype (here designated).— Aestrelata vociferans Shufeldt 1916, skull (neurocra- nium with attached maxillary rostrum and right quadratojugal) included with USNM 320059. Measurements: total length 74.7 mm; cranium length 40.2, cranium width at postorbital processes 29.5, cranium depth 21.1; least width interorbital bridge 10.4, width at naso-frontal hinge 10.3; length of rostrum from naso-frontal hinge 36.2; length of nostril 11.4; length of premaxilla anterior to nostril 20.0. This specimen can be identified unequiy- ocally as the fossil of Aestrelata vociferans illustrated in Shufeldt (1922) as Figure 5 on Plate 16, by the shape of the small flange of bone projecting ventrally nearly across the ventral interorbital fenestra. This flange is extremely variable in Pterodroma cahow and may range from a small pointed pro- jection to a continuous bridge across the fe- nestra. The distinctive shape in USNM 320059 is exactly as shown in Shufeldt’s figure (Fig. la, b), and all other variations, such as positions of small foramina, corre- spond exactly as well. In Shufeldt (1916: 635) it is stated that ‘““The differences in the osseous mandibles of a Petrel (4strelata vociferans) and a Shearwater (Puffinus lherminieri) are easily appreciated upon comparing those parts in figs. 5 & 6 of pl. i.” This reference is to figures in the then unpublished manuscript. The plates were renumbered in Shufeldt 1922, so that plate 1 became plate 16g in which Fig. 5 is the specimen designated here as lectotype. In VOLUME 117, NUMBER 4 577 Fig. 1. A, lectotype of Aestrelata cahow Shufeldt (1916), USNM 320059; the quadratojugal and quadrate were separated from the rest of the skull subsequent to Shufeldt’s photograph and may not have been rejoined in exactly the same position; the quadrate is not necessarily from the same individual as the skull and is not to be considered as part of the lectotype. B, Shufeldt’s illustration (1922: fig. 5, plate 16) of the same specimen; arrow indicates the diagnostic flange of bone in the interorbital foramen that identifies the photograph with USNM 320059. C, left humerus of Puffinus Iherminieri USNM 428934 from Bermuda. D, left humerus, lectotype of Puffinus parvus Shufeldt (1916), CM 16539. E, Shufeldt’s illustration (1922: fig. 56, plate 25) of the same specimen; the markings on the shaft and bit of matrix in the olecranal fossa identify the photograph with CM 16539. the legend, this was identified as being part of the series that was supposed to be in AMNH (see above). USNM 320059 was received from the Carnegie Museum in exchange in 1932. The label with this specimen reads “‘Skel- eton of adult ‘Cahow’ | Aéstrelata vociferans sp. nov. Shuf. | Made as perfect as the bones in the | collection would allow R. W. S[hufeldt]. | 11 Dec. ‘15.” Paralectotypes.—Because of adhering matrix, discolorations, or individual osteo- logical variation, the following specimens can be identified with photographs in Shu- feldt (1922) and are therefore unequivocally part of his type series. Shufeldt’s figure number follows the current museum num- ber: skulls CM 16533 (fig. 1), 16534 (fig. 2), 16535 (fig. 3); sterna 16537 (fig. 26), 16538 (fig. 27). Skull CM 16536 may be the one illustrated in fig. 4, but if so, both quadratojugals are now lacking and I did not detect any peculiarity of the specimen that would allow it to be certainly identified with the figure. Remarks.—Of the new names for Ber- mudan petrels introduced by Shufeldt, the citation for Aestrelata vociferans presents the most difficulties, as no characters of the species itself are actually mentioned and no specimens were illustrated in Shufeldt (1916). Nevertheless, he did discuss osteo- logical characters of the fossils that defi- nitely refer them to Aestrelata (= Ptero- droma) as opposed to Puffinus. Only one species of Pterodroma has ever been found 578 in fossil deposits on Bermuda, and Shufeldt identified his new species with the “‘ca- how” of legend, which was later definitely established as being a species of Ptero- droma (Murphy & Mowbray 1951). Fur- thermore, Shufeldt specifically refers to bones of the new species illustrated in plates prepared for his monograph pub- lished later (Shufeldt 1922) and unequivo- cally identifies them by figure number and plate number. Therefore, it is now possible to identify particular specimens of Shu- feldt’s new species based on information given in the 1916 publication. Thus, it may be argued, as I believe, that Aestrelata vo- ciferans is valid as of Shufeldt 1916 rather than Shufeldt 1922. It is a moot point, how- ever, as A. vociferans Shufeldt 1916 is still a junior synonym by 6 months of A. cahow Nichols & Mowbray, 1916. If A. vociferans is dated from Shufeldt 1922, Bent (1922: 114), who had access to Shufeldt’s manu- script, effectively synonymized Shufeldt’s name 17 days later by saying that it was “apparently the same bird” as A. cahow of Nichols & Mowbray. The unravelling of the identity of the bird known to Bermuda’s early settlers as the “cahow” is well summarized by Murphy & Mowbray (1951). This bird was once in- credibly abundant and provided the early colonists with a ready supply of food. But it was so overexploited by man and intro- duced mammals that it had seemingly dis- appeared before its identity could be made known to naturalists. A living example of a Pterodroma was taken in Bermuda in 1906 by L. L. Mowbray, but was referred to a species that breeds in New Zealand (Brad- lee 1906). Not until a decade later was this specimen described as the type of a new species, Aestrelata cahow (Nichols & Mowbray 1916), almost simultaneously with Shufeldt’s (1916) preliminary note. Shufeldt deserves a fair amount of credit for developing our knowledge of the Ca- how, as his paleontological studies were as seminal as any in providing documentation that the Cahow was one of the gadfly pet- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rels now recognized in the genus Ptfero- droma. Puffinus puffinus puffinus (Briinnich, 1764) Puffinus puffinus bermudae Nichols & Mowbray, 1916 (31 March):195. Puffinus mcgalli Shufeldt 1916 (2 October): 630; Shufeldt, 1922:354. Puffinus puffinus puffinus: Dwight 1927: 243 (with P. p. bermudae in synonymy). Puffinus puffinus: Wetmore, 1931:407 (foot- note; suggested synonymy of P. mcgalli); Lambrecht, 1933:269; Wetmore, 1962: 16; Brodkorb, 1963:246. Holotype.—Puffinus mcgalli Shufeldt 1916, sternum CM 16531, with a split in the carina from which a piece of bone is missing, also lacking the tip of the carina and tips of some of the posterior processes. Referred specimen.—In an addendum, Shufeldt (1922:381, footnote) identified what he believed to be a pedal phalanx 2.8 cm in length that he thought “belonged to an adult specimen of Puffinus mcgalli, and possibly to the same individual” as the hol- otypical sternum. This specimen (CM 16532) is still in the same box with the ho- lotype and measures 28.7 mm. It is actually the left tibiotarsus of a juvenile passerine bird with the proximal end quite porous and incompletely ossified. It has no status what- soever as a type. Remarks.—Shufeldt (1916) based Puffi- nus mcgalli on a sternum that was stated to be larger than that of P. lherminieri and smaller than that of P. major (= P. gravis), in addition to which a measurement of the holotype was provided. This is quite suffi- cient to establish the name P. mcgalli at this point. Wetmore (1931:407), presumably on the basis of size and geographical proba- bility, suggested that P. mcgalli was prob- ably synonymous with P. puffinus and was followed by Lambrecht (1933). Later, Wet- more (1962:16) considered that Shufeldt’s figures of the sternum of P. mcgalli “‘agree exactly with a sternum of a female Puffinus puffinus puffinus.” Brodkorb (1963) fol- VOLUME 117, NUMBER 4 lowed Wetmore’s lead, but no one since Shufeldt had ever critically examined the specimen. The shape of the manubrial area, the an- gle of the sterno-coracoidal processes, and other features establish that the holotype is correctly referred to the genus Puffinus, as opposed to Pterodroma. In size, it is within the range of Puffinus puffinus puffinus: length along midline 58.0 mm, width across posteriormost costal facets 25.4 mm. In a series of 10 skeletons of Puffinus puffinus puffinus the length was 52.2—58.0 (avg. 55.1) and width 23.9—27.2 (avg. 25.7). This is larger than Puffinus Iherminieri but smaller than any of the other Atlantic spe- cies of Puffinus. Thus Puffinus mcgalli Shu- feldt, 1916, was correctly synonymized with Puffinus puffinus Briinnich, 1764. This occurrence of Puffinus puffinus as a fossil in Bermuda is unique, as no other fos- sils of the species have ever been encoun- tered among the thousands of bones of sea- birds collected so far. Although this species is a common offshore visitor to Bermuda, there are only three records of attempted breeding (Bradlee et al. 1931, Bourne 1957). The first was a specimen “‘captured while sitting on its solitary egg in a rocky hole on a small island in Castle Harbor, in April, 1864” (Reid 1884:274). The second record, more doubtful, was another bird sit- ting on an egg in an island in Castle Harbor in May 1877 tentatively recorded as Puffi- nus opisthomelas (Reid 1884:276). The fi- nal record was a specimen taken “March 10, 1905, sitting on a single white egg in a crevice in Gurnet Head Rock’’ (Nichols & Mowbray 1916). This was described as a new subspecies, Puffinus puffinus bermudae Nichols & Mowbray, 1916, that was later definitively synonymized with Puffinus puf- finus puffinus by Dwight (1927). In an instance perhaps similar to those on Bermuda, a single incubating Manx Shear- water was found in June 1973 on Penikese Island, Massachusetts, west of Martha’s Vineyard (Bierregaard et al. 1975), but breeding evidently did not continue there 579 (Lee & Haney 1996). The first North Amer- ican breeding colony of the species was es- tablished in 1977 on Middle Lawn Island, southern Newfoundland, and by 1981 the population had grown to an estimated 350 individuals (Storey & Lien 1985). There is no evidence that Puffinus puffinus was ever able to establish such a colony on Bermuda at any time in the last 400,000 years and all the records, including the fossil sternum de- scribed as Puffinus mcgalli, appear to have resulted from single individuals or pairs. Puffinus parvus Shufeldt, 1916 Puffinus parvus Shufeldt, 1916:632; Shu- feldt, 1922:356. Puffinus lherminieri: Wetmore, 1931:407 (footnote; suggested synonymy of P. par- vus); Lambrecht, 1933:270; Wetmore, 1962: Brodkorb, 1963:246. Lectotype (here designated).—Puffinus parvus Shufeldt, 1916, left humerus, CM 16539 (fig. 56 of Shufeldt 1922). Measure- ments: Total length 58.8 mm; proximal width 10.7, depth of head 3.3, width and depth of shaft at midpoint 3.8 X 2.6, distal width 7.9. Paralectotypes (figure numbers from Shufeldt 1922 in parentheses).—CM 16540 right humerus (fig. 55), 16541 right humer- us, 16542 left humerus, 16543 left humer- us, 16544 right ulna (fig. 43), 16545 right ulna, 16546 left ulna (fig. 44), 16547 left radius (fig. 45), 16548 right carpometacar- pus (fig. 67), 16549 right phalanx 1 of ma- jor alar digit (fig. 74), 16550 left coracoid (fig. 92), 16551 incomplete furcula (fig. 79), 16552 right tibiotarsus (fig. 119), 16553 left tibiotarsus (fig. 120), 16554 right tarsometatarsus (fig. 107), 16555 right fe- mur, 16556—58 left innominates. Remarks.—TYhe name Puffinus parvus dates from Shufeldt (1916), as there this taxon was specifically characterized as be- ing smaller than P. [herminieri and as be- longing to a group of small shearwaters having a short, rather than an elongate ster- num. The type material he listed (p. 632) 580 as 12 bones from what he called the AMNH series (of which only one certainly, and three probably, can now be accounted for) and the following from the McGall collec- tion: “five perfect humeri, three ulnae, a ra- dius, a carpo-metacarpus, a proximal joint of an index digit, a coracoid, an inferior mandible, an imperfect os furculum, a tar- so-metatarsus, an Os innominatum of the left side; subsequently there also came to light an imperfect cranium.” These lists were repeated nearly verbatim in Shufeldt (1922:356) save that the last imperfect cra- nium is omitted and that specimen is no longer present, so perhaps he subsequently re-identified it. In an addendum, Shufeldt (1922:385) listed and identified a further se- ries of 77 specimens of Puffinus parvus col- lected by McGall and Tall that also was de- posited in the Carnegie Museum, where all but the 5 sterna and 2 of the fragmentary furculae may still be found. It is very clear from Shufeldt’s statements (e.g., 1922:385), however, that the first two collections con- stituted the type series and that the addi- tional specimens were referred only subse- quent to his 1916 paper and thus have no status as types. In the CM collections was a container of bones labelled in Shufeldt’s hand ““McGall Collection | Puffinus parvus Shuf. sp. nov | Noy. 27 1915 | Fragile.” This series cor- responds exactly to Shufeldt’s list of this collection, less the cranium mentioned above, except that it has been augmented by a right and left tibiotarsus, a right femur, and an additional two innominate bones. Although no tibiotarsus was listed for the McGall collection in either of Shufeldt’s publications, the legend for Shufeldt’s (1922) fig. 119 of a right tibiotarsus iden- tifies it as being from the McGall collection, whereas the left tibiotarsus in fig. 120 is identified as being from the AMNH series, in which there was only a single tibiotarsus. The femur and the additional two innomi- nates are doubtless the femur and two of the four innominates listed for the AMNH series, which has otherwise disappeared. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON I think that there can be no question that all 21 of these bones may be safely regard- ed as syntypes of Puffinus parvus Shufeldt. Several can be identified with photographs in Shufeldt (1922) and from these I have selected as lectotype a humerus with dis- tinctive markings making it individually identifiable (Fig. Id, e). All of the remain- ing bones in this series may be considered paralectotypes and have been listed above with their current catalog numbers and ref- erence to the figure numbers in Shufeldt (1922) where appropriate. Without having seen the material, Wet- more (1931) suggested in a footnote that Puffinus parvus was probably the same as the living Audubon’s Shearwater Puffinus lherminieri Lesson, 1839, in which he was followed by Lambrecht (1933). Later, in ex- amining a few remains of small Puffinus found in 1958 on Cockroach Island, Har- rington Sound, Bermuda, Wetmore (1962) noted what seemed to be two size classes but considered that the smaller one consist- ed of juveniles. Although he stated (p. 16) that ““Shufeldt (1916 p. 632) noted two ap- parent size groups and named the smaller one Puffinus parvus,” 1 cannot interpret anything in Shufeldt’s publication as indi- cating that he thought there were two size classes. Wetmore also noted that Shufeldt’s (1922) photographs of the bones of P. par- vus were not to the scale indicated, as Shu- feldt himself had pointed out, however (p. 362 footnote). Wetmore concluded that P. parvus was not a valid taxon and synony- mized it with P. lherminieri, and he was followed by Brodkorb (1963). After having examined Shufeldt’s type- series and much more extensive fossil ma- terial from Bermuda dating from the middle Pleistocene onward, I have concluded that Puffinus parvus is indeed a much smaller species than P. /herminieri (Fig. 1c, d). The systematics of the Puffinus lherminieri/P. assimilis assemblage is very complex and imperfectly understood. Puffinus parvus needs comparison with the Atlantic taxa known as Puffinus affinis baroli, which oc- VOLUME 117, NUMBER 4 curs in the Azores, Madeira group, and Ca- nary Islands, and Puffinus lherminieri boydi of the Cape Verde Islands (Jouanin & Mougin 1979). Unfortunately, there is al- most no skeletal material of these taxa available for comparison. Apparently, P. parvus was exterminated after human arriv- al in Bermuda, after which P. lherminieri was able to colonize the island for a brief period before it became extinct itself as a breeding: bird in the late 20th century. [ron- ically, both species are present in the Cock- roach Island material. Further investigation of the small shearwaters of Bermuda is un- der way, but for now Puffinus parvus Shu- feldt, 1916, is retained as a taxon that is clearly distinct from P. lherminieri. Acknowledgments I thank Kenneth _C. Parkes and Robin Panza, Carnegie Museum of Natural His- tory, Pittsburgh (CM), for making Shu- feldt’s material available and for supplying catalog numbers. The figure is by Brian Schmidt, Division of Birds, National Mu- seum of Natural History, Smithsonian In- stitution (USNM). Literature Cited Bent, A. C. 1922. Life histories of North American petrels and pelicans and their allies —United States National Museum Bulletin 121:1—343. Bierregaard, R. O. Jr., A. B. David, Il, T. D. Baird, & R. E. Woodruff. 1975. First northwestern Atlan- tic breeding record of the Manx Shearwater.— Auk 92:145-147. Bourne, W. R. P. 1957. The breeding birds of Bermu- da.—Ibis 99:94—105. Bradlee, T. S. 1906. Audubon’s Shearwater and Peale’s Petrel breeding in Bermuda.—Auk 33:217. , L. L. Mowbray, & W. EF Eaton. 1931. A list 581 of birds recorded from the Bermudas.—Pro- ceedings of the Boston Society of Natural His- tory 30:279-382. Brodkorb, P. 1963. Catalogue of fossil birds. Part 1 (Archaeopterygiformes through Ardeifor- mes).—Bulletin of the Florida State Museum, Biological Sciences 7:179—293. Dwight, J. 1927. The “new” Bermuda shearwater proves to be Puffinus puffinus puffinus.—Auk 44:243. Jouanin, C., & J.-L Mougin. 1979. Order Procellari- iformes. Pp. 48-121 in E. Mayr & G. W. Cot- trell, eds., Check-list of Birds of the World. Vol- ume 1, 2nd ed. Cambridge, Massachusetts, Mu- seum of Comparative Zoology, 547 pp. Lambrecht, K. 1933. Handbuch der Palaeornithologie. Gebrueder Borntraeger, Berlin, 1022 pp. Lee, D. S., & J. C. Haney. 1996. Manx Shearwater Puffinus puffinus—Birds of North America 257:1—28. Murphy, R. C., & L. S. Mowbray. 1951. New light on the Cahow, Pterodroma cahow.—Auk 68:266— 280. Nichols, J. T., & L. L. Mowbray. 1916. Two new forms of petrels from the Bermudas.—Auk 33:194— 195. Reid, S. G. 1884. The birds of Bermuda.—U.S. Na- tional Museum Bulletin 25:163—279. Shufeldt, R. W. 1916. The bird-caves of the Bermudas and their former inhabitants.—Ibis series 10, 4: 623-635. . 1922. A comparative study of some subfossil remains of birds from Bermuda, including the “Cahow”’.—Annals of the Carnegie Museum 13:333-418. Storey, A. E., & J. Lien. 1985. Development of the first North American colony of Manx Shear- waters.—Auk 102:395—401. Wetmore, A. 1931. The fossil birds of North America. Pp. 401—472 in Check-list of North American Birds, 4th ed. American Ornithologists’ Union, Lancaster, Pennsylvania. . 1962. Bones of birds from Cockroach Island, Bermuda. Pp. 15-17 in A. Wetmore, Notes on fossil and subfossil birds. Smithsonian Miscel- laneous Collections 142(2):1—-17. Associate Editor: Gary R. Graves PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):582-593. 2004. Revision of the genus Squamigera (Insecta: Zygentoma: Nicoletiidae) with descriptions of two new species Luis Espinasa and Bethany Burnham Natural Sciences, Shenandoah University, 1460 University Drive, Winchester, Virginia 22601, lespinas@su.edu or espinas!@yahoo.com Abstract.—The genus Squamigera was described in 1999 from a single male. Understanding of the genus was therefore limited. After several unsuccessful expeditions, new material has finally been collected from the same cave. New material of related Squamigera species was also found while reviewing mu- seum collections. From these specimens two new species, S. cumcalcaris and S. jaureguii, are described, and a better description of the diagnostic characters of the genus is provided. In 1988, a single male thysanuran was collected by R. Espinasa-Closas in a Mex- ican cave (Cueva de las Pozas Azules). The specimen is unique in many ways. Measur- ing 22 mm, it is one of the largest speci- mens in the family Nicoletiidae, but more diagnostically, it has spines on the cerci and scales cover its body and head. All other members of the subfamily Cubacubaninae lack this combination. Despite many sub- sequent visits to the same locality, no other specimens were found. Eleven years after the original discovery, the specimen was described (Espinasa 1999a) and the new ni- coletiid genus Squamigera, was established. By necessity, the description of Squamigera lacked a description of the female mor- phology or of postembryonic development. Comparison of the genus with other mem- bers of the subfamily was difficult because it was unclear which characters were unique to the specimen (species variation) and which characters had phylogenetic/taxo- nomic value. Fortunately, the situation has changed. A revision of the nicoletiid collection of the American Museum of Natural History pro- vided a single female from a surface local- ity collected in 1976 by Reddell and Grubs. Also, the Sbordoni collection of cavernicole organisms from Chiapas provided two males and one female from two caves. And finally, an additional male has been col- lected from the type locality. This male is considerably larger than any other Ameri- can nicoletiid described. From these specimens, two new species are described and a revision of the taxo- nomic characters for the genus is provided. Materials and Methods The live specimen was found crawling on the cave wall and was preserved in 96% ethanol. Dissections were made with a ste- reo microscope and the body parts were mounted in fixed preparations with Hoyer’s solution. The female and juvenile male from Chiapas, and the new Pozas Azules specimen were not dissected. All illustra- tions were made with aid of a camera lucida attached to a compound microscope. The types were deposited in the Zygentoma col- lection of the American Museum of Natural History. Squamigera Espinasa, 1999 Diagnosis (amended).—A member of the subfamily Cubacubaninae with mucronate to emarginate scales with smooth to serrate borders. Cerci of males with modified VOLUME 117, NUMBER 4 spines. Parameres without a cleft on the apex. Description (amended).—Body propor- tions normal to robust. Head, thorax, ab- domen, and proximal articles of legs with scales and setae. Distal articles of legs, mouthparts and abdominal stylets only with setae. Scales numerous and multiradiate, their form mucronate to emarginated, with smooth to highly serrated borders. Pedicellus of adult males with unicellular glands and apparently with a spur on its base. Mouthparts not specialized. Mandi- bles strongly sclerotized apically with usual teeth. Galea apically with several sensory pegs. Lacinia heavily sclerotized distally. First process of lacinia pectinate. Labium without prominent lateral lobes. Tarsi with four articles. Praetarsi with three simple claws. Middle claw glabrous, slender and smaller than lateral claws. Urosterna II—VII subdivided into coxites and sternite. Urosterna VIII and IX of male entire. Middle portion of sternites with 1 + 1 sublateral macrochaetae at hind borders, as well as | + 1 near suture at about middle of segment. Coxites on segments II-LX with stylets. Eversible vesicles on segments [— VI, pseudovesicles on VII. Urosterna III of adult males sometimes with modified cox- ites. Urosterna IV apparently without artic- ulated submedian appendages. Urosterna VIII with a wide and not too deep posterior emargination. Posterior projections acute to slightly round, pointing slightly outward. Tergum X very protruding, almost straight on posterior border. Posterior angles with several subequal macrochaetae. Point of insertion of parameres relatively deep and with modified setae on internal face of coxal processes. Parameres with specialized setae on apex, but without a clef or other modifications. Stylets [IX apparent- ly without spines. Opening of penis longi- tudinal. Cerci of male with modified spines. Median filament with or without spines. Fe- males with a subgenital plate and gonapo- physes of adult females apparently with nu- merous articles. 583 Rica le (larger individual, dorsal view) and Squamigera sp., juvenile male (smaller individual, ventral view). Com- parison of body proportions to illustrate the large size of S. latebricola. Squamigera latebricola, male topotype Type species.—Squamigera latebricola (Fig. 1). Distribution.—All specimens to date come from south-central Mexico. It is cur- rently unknown but likely that members of the genus occur in South America and the Antillean islands. Their distribution is prob- ably restricted to the neotropics. Remarks.—Several amendments were made to the original description of the ge- nus: 1. Body proportions are not always ro- bust. 2. Scales are not only slightly serrated, but can be highly serrated. 3. Size of spur on male pedicellus can be variable. 4. Uros- terna II subdivided into coxites and sternite. In the fixed preparation of the holotype it was unclear if the urosterna was divided. 5. Urosterna III of adult males can have mod- ified protuberances similar to those found in some Cubacubana (Espinasa 1991) and Prosthecina (Espinasa 2000). 6. Number of macrochaetae in posterior angles of tergum X can be variable. 7. Point of insertion of parameres relatively deep and with modi- fied setae on internal face of coxal process- es. 8. Parameres without a cleft. In the fixed preparation of the holotype, the parameres were broken as an artifact of the prepara- tion, giving the impression of a cleft (The 584 cleft/break was not represented in the orig- inal figures, it was only mentioned in the text). 9. Central filament sometimes with spines. 10. Females with a subgenital plate and gonapophyses of adult females appar- ently with numerous articles. There were no female samples available when the original description was made. Squamigera belongs to a group of nico- letiid genera, the Cubacubaninae (Mendes 1988), characterized by subdivided uroster- na II-VII and fused coxites VIII and IX of males. Squamigera is distinguished from al- most all genera of this subfamily by having scales. It differs from Texoreddellia (Wy- godzinsky 1973), the only other genus with scales, by the morphology of scales (in Tex- oreddellia scales have three pointed borders instead of smooth to serrated borders), and by having scales in the head and modified spines in cerci, which are both absent in Texoreddellia. Squamigera cumcalcaris, new species Figs. 2A—G, 3A—E Type material.—“*Grotta I Finca S. Ani- ta’? cave, Finca S. Anita, Simojovel de Al- lende, Chiapas, México. 830 m above sea level. 10/LX/1973 V. Sbordoni col. Male ho- lotype, female paratype. Description.—Body length 15 mm. Max- imum conserved length of antennae 12 mm and of caudal appendages 11 mm. General color: light yellow to white. Morphology of the body as in generic description. Scales similar to Fig. 5B. Male antennae as in Fig. 2A—B. Pedicel- lus slightly more than one half as long as basal article. On ventral side with approxi- mately six clusters of unicellular glands ar- ranged in two long rows, surrounded by mi- crochaetas forming a “U” shape. Outside this microchaetae, another two clusters of unicellular glands and a downward pointing robust spine, opposite to an extension of the basal article (Fig. 2B). Base of female an- tennae simple and pedicellus half as long as basal segment. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Head with approximately 5 + 5 macro- chaetae on border of insertion of antennae (Fig. 2A). Mouthpart appendages relatively short. Labial palp as in Fig. 2D. Apical ar- ticle slightly wider than long and barely longer than penultimate article. Penultimate article with bulge containing macrochaetae. Labium and first article of labial palp with macrochaetae. Maxilla as shown in Fig. 3D. Last article slightly longer than penulti- mate. Apex of maxillary palp with two con- ules of similar width and a 3rd minute extra conule similar to Fig. 4G. Two teeth on la- cinia. Mandibles chaetotaxy as in Fig. 2C. Thoracic nota with scales and macro- chaetae on lateral borders apart from sev- eral setae of varied sizes (Fig. 3B), but no small sclerotized spines on posterior bor- ders. Legs relatively short and stout. Tibia on 2nd leg with five macrochaetae, some of them stout, and approximately 3.5 longer than wide and % shorter than tarsus (Fig. 2E). Tibia on 3rd leg with five macrochae- tae, and approximately 4.5 longer than wide and ¥; shorter than tarsus (Fig. 2F). Claws relatively short. Urosterna III and IV of male without modifications in the samples examined. It is currently unknown if more fully adult spec- imens will develop them. Urosterna VIII posterior projections acute to slightly rounded, subtriangular (Fig. 3A). Urotergite X posterior angles with several long macro- chaetae and setae of different sizes. On the borders some prominent scales (Fig. 2G). Urosterna IX of male similar to some Anelpistina; point of insertion of parameres deep and setae slightly more sclerotized on internal face of coxal processes and above insertion of parameres (Fig. 3A). Stylets [IX bigger than the others, with 4—5 macro- chaetae and an extra subapical pair. In both males and females without spines or other modifications. Other stylets have only three macrochaetae plus the subapical pair. Penis and parameres as shown in Fig. 3A. Parameres attaining % of stylets IX. Parameres globular and with a distinct group of microchaetae on the tip. Overall VOLUME 117, NUMBER 4 585 Ni { me Ww \ i) wea TARAS Fig. 2. Squv umigera cumcalcaris. Male holotype. Scales and microchaetae partially shown; A, Head; B, Pedicellus; C, IWandible; D, Labial palp and labium; E, Tibia of 2nd leg; K 3rd leg; G, Urotergum X. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON onl 586 me Tm SS = \n ~ S ~ & ok WSS 2 IFFY rae : B TOO 0 2, I aN a) aN 1 RNY = 2 ne ve COLE Q Ss S ASS Sa eS, 2 SS Pr) shee J, + ~ ANS) EIS: SSS RNS ew SS = gy Fig. 3. Squamigera cumcalcaris. Male holotype except C, female paratype. Scales and microchaetae partially shown; A, Genital area; B, Thoracic tergum; C, Ovipositor; D, Maxilla; E, Median filament (left) and cercus (right). VOLUME 117, NUMBER 4 appearance similar to some Prosthecina (Wygodzinsky 1946). Subgenital plate of female parabolic (Fig. 3C). Ovipositor sur- passing apex of stylets IX by thrice the length of stylets (Fig. 3C). Gonapophyses with approximately 38 articles. Male caudal appendages as in Fig. 3E. Inner side of cerci in males with spines of varied sizes. Some spines arranged even in a double row. The central filament also with spines of subequal size arranged on multi- ple rows facing both cerci. Female caudal appendages without modifications. Postembryonic development unknown because of the scarcity of samples. It is as- sumed that specimens examined are adult based on the development of sexual sec- ondary characters. Comparison to other species within the subfamily indicates that in younger instars we could expect that spines, modifications of antenna, and size of parameres to be reduced in younger males. In females a smaller ovipositor could be expected. Known range.—Known only from the type locality. Etymology.—The name is derived from the Latin “‘cum-+calcaris” for with+spur, alluding to the prominent curved spur in the pedicellus of the antennae in males. Remarks.—Squamigera cumcalcaris can be differentiated from all species of sub- family Cubacubaninae by the spines on the central filament. Such spines until now were described only in Nicoleiids in the subfamilies Coletiniinae and Subnicoleti- inae (Mendes 1988). Adult males can be further differentiated by the large curved spur oriented toward the base of antennae in the pedicellus, which in S. latebricola is reduced to a small spine. Adult females can be differentiated from S. jaureguii by a par- abolic instead of trapezoidal subgenital plate and by a considerably less subdivided gonapophyses. Squamigera jaureguii, new species Figs. 4A—-H, 5A-K 6A—D Type material.—Puente Actopan, 5 km SE Actopan, Veracruz, Mexico. 25 Dec 587 1976. J. Reddell and A. Grubbs cols. Fe- male holotype. Description.—Body length 9.5 mm. An- tennae and caudal appendages broken. Maximum conserved length of antennae 4 mm and of caudal appendages 5 mm. Body proportions as in Fig. 4A. General color: light yellow to white. Morphology of the body as in the generic and S. cumcalcaris descriptions, unless otherwise stated. Scales as in Figs. 4D and 5B. Antennae as shown in Fig. 4B. Basal ar- ticle without projections. Pedicellus slightly less than one half as long as the basal ar- ticle. Head with approximately 8 + 8 ma- crochaetae on border of insertion of anten- nae (Fig. 4C). Labial palp as in Fig. 4E. Maxilla as shown in Fig. 4F—G. Last article ¥. longer than penultimate. Apex of maxil- lary palp with two conules of similar width and a 3rd minute extra conule (Fig. 4G). Mandibles chaetotaxy as in Fig. 4H. Tho- racic nota as in fig. 5A. Legs relatively short and stout (Fig. SC—D). Tibia on 2nd leg with five macrochaetae, some of them stout, and approximately 3.2 longer than wide and % shorter than tarsus, and five ma- crochaetae. Hind leg broken in the speci- men. Claws relatively short (Fig. 5E). Urosterna I and II as in Fig. 5K Uroter- gite X posterior angles with 2—3 long ma- crochaetae and setae of different sizes and on borders some prominent scales (Fig. 6A). Stylets [X bigger than the others, with 5—6 macrochaetae and an extra subapical pair (Fig. 6B). Subgenital plate of female trapezoid, outer border almost straight (Fig. 6B). Ovipositor surpassing apex of stylets IX by thrice the length of stylets (Fig. 6B). Apex as in Fig. 6C. Gonapophyses with ap- proximately 53 articles. Cerci without mod- ifications (Fig. 6D). Males unknown. Postembryonic devel- opment unknown because only a single fe- male individual could be examined. It is as- sumed that this individual is an adult based on its large ovipositor. Comparison to other species within the subfamily indicates that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 588 kit LE | Pee i | Es / ——— pare rey £4) A mm { Sy -\\,. > sepa REEFS EH Dyyy, = Fig. 4. Squamigera jaureguii. Female holotype. Scales and microchaetae partially shown. A, Body; B, Basal portion of antennae; C, Head; D, Scales on head; E, Labial palp and labium; EK Maxilla; G, Apical portion of maxilla; H, Mandible. VOLUME 117, NUMBER 4 Re lial aa f SS SS I ES = Z Z gS Ae. ene RK iT PPPS ai ppp eh RO FD. ae oe bs a GRE == 4 ? : ye & ,? x Pn RB a OMe ONG ee Se ‘ 4 ae eT oN LAW 2x5 fi Ti V, x y fp 7. ef ae xX nee | ie Fig. 5. Squamigera jaureguii. Female holotype. Scales and microchaetae partially shown. A, Thoracic ter- gum; B, scales of urotergum I; C, 2nd leg; D, Apex of 2nd tibia; E, Claws of 2nd leg; K Urosternum I and II. 590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oy) } ee Fig. 6. Squamigera jaureguii. Female holotype. Scales and microchaetae partially shown. A, Urotergum X: B, Subgenital plate and ovipositor; C, Apex of ovipositor; D, Caudal appendages. VOLUME 117, NUMBER 4 in younger instars we could expect a small- er Ovipositor. Known range.—Known only from the type locality. Etymology.—this species is dedicated to Sergio Jauregui to recognize his enthusias- tic, long time participation in cave nicole- tiid collecting and field work. Remarks.—Squamigera jaureguii can be differentiated from other described Squa- migera by having more macrochaetae on the head at the border of the insertion of antennae, and a shorter body and append- ages. Adult females can be differentiated from S$. cumcalcaris by the trapezoidal in- stead of parabolic subgenital plate and by a considerably more subdivided gonapophys- es. No S. latebricola females are available for comparison. . Squamigera latebricola Espinasa Fig. 7A—G Topotype.—““Cueva de las Pozas Azu- les”’ cave (Espinasa-Perefia 1989), Taxco de Alarcon Municipality, Guerrero State, Méx- ico, 18°36'40"N, 99°33'25”’W. April 2001. L. Espinasa col. Male. Description.—Body length 29 mm. Max- imum conserved length of antennae 29 mm and of caudal appendages 35 mm. Body proportions as in Fig. 1. General color: light yellow to white. Morphology of body sim- ilar to S. cumcalcaris and S. jaureguii, un- less otherwise stated. Scales with slightly less serrated borders. Pedicellus with clusters of unicellular glands and a small spur (Espinasa 1999a; Fig. 1C) instead of long hooked spine of S. cumcalcaris. Mouthpart appendages rela- tively thin and long. Apical article of labial palp barely longer than wide and barely shorter than penultimate (Espinasa 1999a; Fig. 1D). Penultimate article’s bulge not too prominent. Maxilla as in Fig. 7D. Last ar- ticle shorter than penultimate. Apex of maxillary palp with two conules of similar width and a 3rd small extra conule (Fig. TB). 591 Thoracic nota with small sclerotized spines on lateral and posterior borders (Es- pinasa 1999a; Fig. 3A). Legs relatively long (Espinasa 1999a; Fig. 2A). Tibia on 2nd leg with seven thin macrochaetae, and approx- imately 4.5 longer than wide and ¥% short- er than tarsus. Tibia on 3rd leg with eight thin macrochaetae, and approximately slightly over 5X longer than wide and 4 shorter than tarsus. Trochanter on 3rd leg with a protuberant spine projection (Fig. 7B) which is not present in the smaller sized (22 mm) holotype. Claws of normal Size. Coxites in urosterna III (Fig. 7C) with protuberances similar to those found in some Cubacubana (Espinasa 1991) and Prosthecina (Espinasa 2000). Urosterna III in smaller holotype also with a slight pro- tuberance (not reported in original descrip- tion), similar to nascent protuberance found in some immature individuals of the afore- mentioned Cubacubana and Prosthecina. Urosterna IV without modifications (Fig. 7A). Urosternum IX as in Fig. 7E In this specimen the point of insertion of paramer- es is slightly deeper than in the holotype and closer in appearance to some Anelpis- tina (Espinasa 1999b). Stylets [X with five macrochaetae and an extra subapical pair but otherwise without any other modifica- tions. Penis and parameres as shown in Fig. 7F Parameres attaining less than % of sty- lets [IX and curved outward. Cerci as in Fig. 7G. Females unknown. Postembryonic development only partial- ly understood since only two fairly large male individuals are available, the holotype (22 mm) and this new topotype (29 mm). In the smaller specimen, projections of urosterna III are only starting to develop, spines in cerci are less prominent and tro- chanter of hind leg has no projection. Known range.—Known only from the type locality. Remarks.—Being 3 cm in length (10 cm if antennae and caudal appendages are in- cluded), S. latebricola can easily be differ- entiated from all species of the subfamily 592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Squamigera latebricola. Male topotype. Scales and microchaetae partially shown. A, Urosternum IV; B, 3rd leg. Notice projection in trochanter (scales not shown); C, Urosternum III; D, Maxilla; E, Apical portion of Maxilla; EK Genital area; G, Spines in cercus. VOLUME 117, NUMBER 4 Cubacubaninae by its large size. This is the longest species among the nicoletiids, which typically measure 1 cm or less. En- largement of body and appendages is com- mon among cave adapted organisms and it is certainly the case for this species. This species can further be differentiated from S. cumcalcaris and S. jaureguii by the small sclerotized spines on lateral and posterior borders on thoracic nota (Espinasa 1999a; Figs. 1G and 3A), and by the morphology of its sexual secondary characters. Squamigera sp. Fig. 1 Material examined.—**E] Chorreadero”’ cave, Chiapa de Corzo Municipality, Chia- pas, México. 650 m above sea level. 10/1 1- VIII- 73. V. Sbordoni col. Male. Description.—Body length 10 mm. An- tenna and caudal appendages broken. Mid- dle filament missing. Scales as in other members of the genus. No apparent spines in pedicellus, sterna III, or cerci. Parameres curved outward, similar to the holotype of S. latebricola (Espinasa 1999a, Fig. 2C), but attaining less than Y; of stylets IX. This single individual is probably not a mature adult. Chorreadero cave is visited relatively often by speleologists and hopefully more samples will be available one day for a for- mal description of this population. Acknowledgments We thank Dr. Randall T. Schuh, curator and chair of the Division of Invertebrate Zoology of the American Museum of Nat- ural History, for kindly giving access to the 593 museum collection and facilitating exami- nation of the specimens. We also thank Val- erio Sbordoni for facilitating acquisition of specimens from Chiapas. Work was done with support from CEAMISH-Universidad Aut6noma del Estado de Morelos, in facil- ities of the American Museum of Natural History and Shenandoah University. Literature Cited Espinasa, L. 1991. Descripci6n de una nueva especie del género Cubacubana (Zygentoma: Nicoleti- idae) y registro del género para América Con- tinental.—Folia Entomolo6gica Mexicana 82:5— 16. . 1999a. A new genus of the subfamily Cuba- cubaninae (Insecta: Zygentoma: Nicoletiidae) from a Mexican cave.—Proceedings of the Bi- ological Society of Washington 112(1):52—58. . 1999b. Two new species of the genus Anel- pistina (Insecta: Zygentoma: Nicoletiidae) from Mexican caves, with redescription of the ge- nus.—Proceedings of the Biological Society of Washington 112(1):59—-69. . 2000. A new species of the genus Prosthecina (Insecta, Zygentoma, Nicoletiidae).—Pedobiol- ogia 44:333-341. Espinasa-Perefia, R. 1989. El resumidero del Isote y la Cueva de las Pozas azules.—Tepeyollotli: Gac- eta de la Sociedad Mexicana de Exploraciones Subterraneas 4:24—27. Mendes, L. EF 1988. Sur deux nouvelles Nicoletiidae (Zygentoma) cavernicoles de Gréce et de Tur- quie et remarques sur la systématique de la fam- ille—Revue Suisse de Zoologie 95(3):751— V2: Wygodzinsky, P. 1946. Sobre Nicoletia (Anelpistina) Silvestri 1905 e Prosthecina Silvestri, 1933.— Ciencia 7:15—25. . 1973. Description of a new genus of cave thy- sanuran from Texas (Nicoletiidae, Thysanura, Insecta)—American Museum Novitates 2518: 1-8. Associate Editor: Wayne Mathis PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):594—596. 2004. BIOLOGICAL SOCIETY OF WASHINGTON 131st Annual Meeting, 15 June 2004 President Roy McDiarmid called the meeting to order at 10:30 a.m. in the Waldo Schmitt Room, National Museum of Natu- ral History (NMNH). Council members and editorial staff present: Marilyn Schotte (Elected Council), Ron Heyer (Acting Pres- ident Elect), Chad Walter (Treasurer), Car- ole Baldwin (Secretary), Richard Banks, Stephen Cairns, Bruce Collette, and Storrs Olson (Past Presidents), and Steve Gardi- ner, Carol Hotton, and Ed Murdy (Associate Editors). Minutes of the 130th Annual Meeting were summarized by Secretary Baldwin. Those minutes are scheduled to appear in Volume 117(1) of the Proceedings, which had not been published at the time of the annual meeting. Following approval of the minutes, McDiarmid summarized recent Society activities. McDiarmid announced that Proceedings Editor Richard Sternberg submitted his resignation as Editor on 9 Oc- tober 2003 but agreed to remain in the po- sition until a replacement could be found. Past President Richard Banks has agreed to serve as interim Editor beginning | July 2004. In view of declining manuscript sub- missions to the Proceedings, McDiarmid is appointing a committee to investigate elec- tronic publishing. To investigate declining Society membership, he is re-establishing a Membership Committee. McDiarmid also noted that he met recently with NMNH Di- rector, Cristian Samper, and new NMNH Associate Director for Research and Col- lections, Hans Sues, to inform them of the existence of the Society and its historical relationship with and support from the mu- seum. Those administrators are in favor of the museum’s continued support of the So- ciety and are interested, in principal, in hosting the Society’s website on the muse- um server, but they indicated that a final decision about the website should not be made until the museum’s new information- technology director is hired. Associate Ed- itor Steve Gardiner, who has produced the Society’s web pages on the server at Bryn Mawr College, announced that his institu- tion is agreeable to leaving the Society’s website on its server if necessary. Mc- Diarmid concluded his summary of recent Society activities by noting that the Society will publish a special Bulletin this year en- titled Study of the Dorsal Gill-Arch Mus- culature of Teleostome Fishes, with Special Reference to the Actinopterygii, by Victor G. Springer and G. David Johnson. This 800+ page Bulletin will comprise two vol- umes and be published in an 8%” X 11” format. President McDiarmid then called on Chad Walter for the Treasurer’s Report (Ta- ble 1). Income for the period 1 January 2003 to 31 December 2003 was $93,105.92, and expenses for the same pe- riod were $74,024.02. Total Society assets as of 15 April 2004 were $99,705.40. The value of the endowment fund increased by $12,307 in 2003. The Audit Committee, Don Wilson and Neal Woodman, indicated that they had reviewed the books and led- gers of the Treasurer and found all financial records to be accurate and in good order. The Treasurer’s report was approved. Proceedings Editor Richard Sternberg re- ported at the Society’s Council meeting on 17 May 2004 that four issues of Volume 116 were published comprising 75 papers and 1007 pages. As of 1 June 2004, there were 34 submissions, but neither 117(1) nor 117(2) had yet been published. Issue 117(1) was submitted in January 2004, but because of the low number of submissions (8), that VOLUME 117, NUMBER 4 Table 1.—Summary Financial Statement for 2003. General Fund Assets: January 1, 2003 18,365.23 Total Receipts for 2003 78,473.46 Total Disbursements for 2003 71,698.63 Assets: December 31, 2003 25,140.06 Net Changes in Funds 6,774.83 a: Annual gain in value of Endowment. b: Annual loss in value of Endowment. issue would have been unusually small. Publication was delayed until more manu- scripts were ready for publication. A deci- sion was then made to split 117(1) into 117(1) and 117(2), both to be published ap- proximately the same time and very soon. Sternberg acknowledged that the decrease in submissions: reported last year finally caught up with us. Furthermore, he noted that the delayed publication of the first is- sues of volume 117 also was attributable to slower-than-normal production of page proofs and page-proof mailing errors. Issue 117(3) is on track for timely production. The Editor’s report was approved by the Council. Custodian of Publications Storrs Olson reported little activity with back issues but noted that he had filled a few orders. The print run of the Proceedings was reduced previously from 1000 to 850 copies, but since membership is 730, the print run could be reduced again, perhaps to 800. Frank Ferrari noted that the Finance Committee (Stephen Cairns, Oliver Flint, Chad Walter, and Ferrari) had consulted an attorney about tax laws regarding member contributions to the Society. The Finance Committee recommends three categories of gifts: Contributor ($100—$499), Sponsor ($500—$999), and Benefactor ($1000 and higher). Donors will receive a letter from the Society that indicates the donor re- ceived nothing for his/her contribution, and names of donors will be listed in four con- secutive issues of the Proceedings. The Committee is currently working on an an- nouncement of the gift-fund categories. Endowment Fund Total Assets 66,277.07 84,642.30 14,631.94a 93,105.40 2,325.39b 74,024.02 78,583.62 103,723.68 12,306.55 19,081.38 Secretary Baldwin indicated that a vote was needed on a change to Article 8 of the Bylaws proposed last year by the Finance Committee. Regarding the Society’s En- dowment Fund, the first sentence of Article 8 currently states: “There shall be an En- dowment Fund which shall consist of con- tributions from members, miscellaneous gifts, and surplus funds from operations.” The proposed change would remove “and surplus funds from operations,” and the amended first sentence would read: ““There shall be an Endowment Fund which shall consist of gifts from members and miscel- laneous gifts.’ The proposed change was unanimously approved. Baldwin also noted that results of the 2004 election of officers could not be an- nounced as usual at the annual meeting be- cause the ballots, which are part of Pro- ceedings issue 117(1), have not been mailed. Results of the election will be add- ed as an addendum to these minutes. President McDiarmid then announced that in hopes of increasing attendance at the annual meetings, he had decided this year to add a program at the conclusion of the annual meeting. Uncharacteristically for Society meetings, the Waldo Schmitt room was nearly full as McDiarmid introduced the scheduled program. McDiarmid re- marked that the purpose of the program was to honor Past President Bruce Collette, whose activities in our Society are a reflec- tion of his attitude, activities, and devotion to promoting science. The program began with Division of Fishes ichthyologist David Smith presenting The Natural History of 596 Bruce B. Collette, a talk Dave had written as an introductory talk for a symposium honoring Collette held at the May 2004 meetings of the American Society of Ich- thyology and Herpetologists in Norman, Oklahoma. Following Dave’s thoughtful and entertaining presentation, Bruce was in- vited by McDiarmid to speak. Bruce’s com- ments reflected his love of his job and the thrill of being able to work on so many in- teresting fish groups, such as tunas (“‘warm- blooded fish!”’). At the core of his message was the fact that seeking answers to simple natural history questions, rather than hy- pothesis testing, had led him to so many years of study and a lot of publications. Bruce concluded with a plea for Society members to work to conserve diversity and habitats. The meeting was adjourned at 12: 15 p.m. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Respectfully submitted, Carole C. Baldwin Secretary Addendum to Minutes.—Results of the 2004 election of officers are as follows: President-Elect—W. Ronald Heyer; Secre- tary—Carole C. Baldwin; Treasurer—T. Chad Walter; Elected Council—Michael D. Carleton, W. Duane Hope, Marilyn Schotte, E Christian Thompson, Jeffrey T. Williams, and Neal Woodman. Additionally, a pro- posed amendment to Article 5 was passed. This amendment allows the Council to elect a replacement President-Elect to finish the term if the President-Elect is unable to car- ry out the duties of office, and then both the President and President-Elect positions are voted on at the next scheduled election. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(4):597—600. 2004. THE BIOLOGICAL SOCIETY OF WASHINGTON CONSTITUTION AND BYLAWS Adopted 3 December 1884 (As amended August 2004) Article 1. Name The name of this Society shall be the Bi- ological Society of Washington Article 2. Purpose The purpose of this Society shall be for the furtherance of taxonomic study of or- ganisms and for the increase and diffusion of biological knowledge among interested persons. Article 3. Membership Membership in this Society shall be open to persons and organizations interested in the promotion of systematic biology. The following classes of members shall be rec- ognized: Associate Members, Active Mem- bers, Life Members and Emeritus members. Changes of status of membership may be effected at any time by the payment of ap- propriate dues. Membership shall become effective upon payment of dues. Associate Members shall pay annual dues, shall receive notices of meetings and be eligible to vote at meetings of the So- ciety and in ballots by mail. Active Members shall pay annual dues, shall receive the publications of the Society, shall receive notices of meetings, and shall be eligible to vote at meetings of the So- ciety and in ballots by mail. Life Members shall be recognized as such by the payment of a fee established by the Council. This fee shall be paid either in one lump sum or in four equal, consecutive annual installments. During their lifetime, Life Members shall receive the publications of the Society, shall receive notices of meetings and shall be eligible to vote at meetings of the Society and in ballots by mail. Emeritus Members. Any member who has been an Active or Associate Member may, at the discretion of the Council, be accorded the privileges of Emeritus Mem- bership. These persons shall then be granted the same status as a Life Member. An organization which is a member may designate a representative who may cast a single vote in its behalf. Article 4. Dues Annual dues for Associate and Active Members shall be fixed by the Council and may be changed by the Council. Article 5. Officers and Elections The Officers shall be a President, a Pres- ident-Elect, a Secretary and a Treasurer. The President-Elect shall succeed the Pres- ident upon the expiration of the latter’s term of office. The President-Elect, Secretary, and Treasurer shall be elected for a term of two years by a majority of the members voting by means of a mail ballot. The of- ficers shall take office at the end of the an- nual business meeting. A slate of candidates shall be prepared by a Nominating Com- mittee appointed by the President. Ballots shall be mailed to all members at the time of billing for annual dues in an election year. If, for any reason, the President shall be unable to carry out the duties of the office, he/she shall be succeeded by the President- Elect until the Council judges him/her to be competent to resume the duties of the of- 598 fice. If, for any reason, the President-Elect shall be unable to carry out the duties of office, the Council will elect a replacement to fill the term; both the President and Pres- ident-Elect positions will be voted on at the next scheduled election. Vacancies in the other offices shall be filled temporarily or until the next election by a majority vote of the Council. Article 6. Council The Council shall consist of the Presi- dent, the President-Elect, the Secretary, the Treasurer, the Chairmen of Standing Com- mittees, the ex-Presidents, and six addition- al members who shall be nominated by the Nominating Committee and elected at the same time the Officers are elected and have two year terms of office. The Council shall be the governing body of the Society. It shall be responsible for matters of policy and procedure. It shall meet at the discretion of the President and shall always meet prior to the annual busi- ness meeting. It shall receive and act on re- ports from the Committees of the Society. It shall receive and act on the annual budget prepared by the Finance Committee. It shall fix the time and place of the annual busi- ness meeting. Actions of the Council may be emended at any annual meeting of the Society by a three-fourths vote of the members present. Actions of the Council may be approved or rejected at any annual meeting of the So- ciety by a majority vote of the members present. The President, with the approval of the Council, shall appoint ad hoc committees which shall report to the Council. Article 7. Meeting The Society shall hold at least one sched- uled meeting each year except in an emer- gency as decided by a three-fourths vote of the Council members present. Reports of Standing Committees, the Treasurer, the Auditor, and the Council shall PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON be presented to the members at the annual meeting. Should the Council declare an emergen- cy, these reports may be made to the mem- bers in printed form. The Council shall in- stall officers in the event there shall be no annual meeting. Article 8. Publications The publications of the Society shall be The Proceedings of the Biological Society of Washington and any other publication that the Council authorizes. The Proceed- ings shall be managed by an Editorial Com- mittee, consisting of an Editor, who shall serve as Chairman, and not less than three Associate Editors. Article 9. Bylaws The Society may enact bylaws which in- terpret and implement this Constitution. Such bylaws, when approved by the Coun- cil, may be adopted, amended, or repealed by a two-thirds majority of those voting at an annual meeting of the Society or in a mail ballot, provided that in either case, no- tice of the proposed action shall have been sent to each voting member of the Society at least thirty (30) days before the date of the vote. Article 10. Amendments This Constitution may be amended by a two-thirds majority of members voting, ei- ther at an annual meeting of the Society, or in a mail ballot, provided that in either case notice of the proposed action, when ap- proved by the Council, shall have been sent to each voting member of the Society at least thirty (30) days before the date of the vote. Article 11. Limitation The purposes of the Society are listed in Article 2 of the Constitution. Lobbying or activities specifically designed to influence legislation are not among the objectives of VOLUME 117, NUMBER 4 the Society and no official group within the Society shall engage in such activity. Article 12. General Prohibitions Notwithstanding any provision of the Constitution or Bylaws which might be sus- ceptible to a contrary construction: a. The Biological Society of Washington shall be organized exclusively for sci- entific and educational purposes; b. The Biological Society of Washington shall be operated exclusively for scien- tific and educational purposes; c. No part of the net earnings of the Bio- logical Society of Washington shall or may under any circumstances inure to the benefit of any private shareholder or individual; d. No substantial part of the activities of the Biological Society of Washington shall consist of carrying on propaganda, or otherwise attempting to influence legis- lation; e. The Biological Society of Washington shall not participate in, or intervene in (including the publishing or distribution of statements) political campaigns on be- half of any candidate for public office; f. The Biological Society of Washington shall not be organized or operated for profit; g. The Biological Society of Washington shall not: 1) lend any part of its income or corpus; without the receipt of ade- quate security and a reasonable rate of interest to; 2) pay any compensation, in excess of a reasonable allowance for sal- aries or other compensation for personal services actually rendered, to; 3) make any part of its services available on pref- erential basis, to; 4) make any purchase of securities or any other property, for more than adequate consideration in money or money’s worth from; or 6) en- gage in other transactions which result in substantial diversions of its income or corpus to; any officer, member of the Council, or substantial contributor to the 599 Biological Society of Washington. The prohibitions contained in this subsection (g) do not imply that the Biological So- ciety of Washington may make such loans, payments, sales or purchases to anyone else, unless such authority be given or implied by any other provisions of the Constitution or Bylaws. Article 13. Distribution or Dissolution Upon dissolution of the Biological So- ciety of Washington, the Council shall dis- tribute the assets and accrued income to one or more organizations as determined by the Council, but which organization or organi- zations shall meet the limitations prescribed in subsections (a)—(g) inclusive, of Article 12, immediately preceding. BYLAWS 1. Quorum. Five Council members shall constitute a quorum at a meeting of the Council. 2. The Secretary. The Secretary shall keep minutes of the meetings of the Council and of the Society and shall present a yearly summary to the Society and Council. He/she shall issue notices for the meetings of the Society and the Council, shall notify members of their election, and shall conduct the corre- spondence of the Society and Council. 3. The Treasurer. The Treasurer shall be in charge of the funds and keep the fi- nancial records of the Society. He/she shall be authorized by the Council to make necessary disbursements, within the limits set by the budget. The Trea- surer shall preserve a receipted bill, or bill and cancelled check for each pay- ment. He/she shall present a statement of financial accounts, audited by the Fi- nance Committee at the time of the an- nual business meeting. 4. The President. The President shall pre- side at meetings of the Council and of the Society, and perform such other functions that may adhere to the office. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5. The President-Elect. In cases of illness, incapacities, death or absence of the President, the President-Elect shall as- sume all duties incumbent on the Presi- dent until the Council judges the Presi- dent to be competent to resume the du- ties of the office. In the event of the death of the President, the President- Elect shall automatically become Presi- dent. . Budget. Prior to the annual meeting, a budget for the next year shall be pre- pared by the Finance Committee and submitted to the Council for action. No financial obligation against the Society may be contracted by any officer or member except as specified in the annual budget or as provided for by special ac- tion of the Council upon recommenda- tion of the Finance Committee. . Committees. The Society shall maintain the following committees. They shall be provided with such needed financial sup- port, to be designated in the budget, as the funds of the Society may warrant. The chairmen of the standing commit- tees shall be appointed by the President following the annual meeting. A. The Finance Committee shall con- sist of the Treasurer and two members to be appointed by the President. The Treasurer shall not serve as the Chair- man. It shall prepare the annual budget for submission to the Council and shall advise the Council in all matters affect- ing the finances of the Society, including the deposit and investment of funds, en- dowments, and long-term financial pol- icies. B. The Editorial Committee shall be composed of the Editor, who shall serve as Chairman, and not less than three As- sociate Editors. This Committee shall advise the Council on all matters affect- ing publication. The Editor shall be ap- pointed by the Council. The Associate Editors shall be appointed by the Editor. Terms for the members of the Editorial Committee shall be at the discretion of the Editor. C. The Membership Committee shall consist of a Chairman and not less than three members and shall be responsible for the Society’s effort to increase or maintain the membership. The Chairman shall be appointed by the President, the other members shall be appointed by the President upon recommendation of the Chairman. . Endowment Fund. There shall be an Endowment Fund which shall consist of contributions from members and miscel- laneous gifts. At the discretion of the Council, the principal of this fund may be used in publishing the Society’s jour- nal or for the general operations of the Society. At the discretion of the Council, the principal of this fund may also be used in the publication of symposia, monographic studies, or other special publications; however, such a decision must be reached only during a regularly scheduled meeting of the Council. PROCEEDINGS of the Biological Society of Washington VOLUME 117 2004 Vol. 117(1) published 1 June 2004 Vol. 117(3) published 7 December 2004 Vol. 117(2) published 4 August 2004 Vol. 117(4) published 20 December 2004 WASHINGTON PRINTED FOR THE SOCIETY EDITOR RICHARD V. STERNBERG RICHARD C. BANKS ASSOCIATE EDITORS Classical Languages Invertebrates FREDERICK M. BAYER STEPHEN L. GARDINER CHRISTOPHER B. BOYKO JANET W. REID Plants Vertebrates CarROoL HOTToNn Gary R. GRAVES CAROLE C. BALDWIN EDWARD O. Murpy Insects Invertebrate Paleontology WAYNE N. MaTHIS GALE A. BISHOP All correspondence should be addressed to the Biological Society of Washington, National Museum of Natural History Washington, D.C. 20013 Printed by ALLEN PREss INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 2004—2005 OFFICERS President ROY W. McDIARMID President-Elect W. RONALD HEYER Secretary CAROLE C. BALDWIN Treasurer T. CHAD WALTER COUNCIL Elected Members MICHAEL D. CARLETON F. CHRISTIAN THOMPSON W. DUANE HOPE JEFFREY T. WILLIAMS MARILYN SCHOTTE NEAL WOODMAN Parra ve r _ -— a f, The bey, tt an). Ae i AUG OM 4é ASR : nah hi ; a LOTOMINGAW IO CTO JAG A HES © 7 4 7 (te Ul iw ‘ ' BIL 0% AE ge wa a (an ange. 7 7 horny = inet y é ce nie ¢ Parris 4c SL a a se cary LG. SA ‘i te | f on, eet < » '¢ sie PRA nmibekeg.y , é “+ . - CHag RNS: be. WP ie wus 7 , Hiei ini) 4 34) uy bine ) ~ a = aiid y ci a \ 1 aa ¥ Ta yoeryTs is vig 1 AT ! ty INFORMATION FOR CONTRIBUTORS See the Society’s web page— www.biolsocwash.org Content.—The Proceedings of the Biological Society of Washington publishes original research bear- ing on systematics in botany, zoology, and paleontology, and notices of business transacted at Society meetings. Except at the direction of the Council, only manuscripts by Society members will be consid- ered. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in another language when appropriate. Submission of manuscripts—Manuscripts may be submitted in one of three ways. You may mail three paper copies of the manuscript complete with tables, figure captions, and figures (do not submit original figures unless/until the manuscript is accepted for publication) to the Editor, Dr. Richard C. Banks, MRC-116, National Museum of Natural History, P. O. 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Front cover—from this issue, p. 546. CONTENTS Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 5. The genera Plumarella Gray, 1870; Acanthoprimnoa, n. gen.; and Candidella Bayer, 1954 Stephen D. Cairns and Frederick M. Bayer A new species of the sea anemone Megalactis (Cnidaria: Anthozoa: Actiniaria: Actinodendridae) from Taiwan and designation of a neotype for the type species of the genus Adorian Ardelean and Daphne Gail Fautin A new genus and new species of crab of the family Xanthidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the southwestern Gulf of Mexico Ana Rosa Vazquez-Bader and Adolfo Gracia A new anchialine shrimp of the genus Procaris (Crustacea: Decapoda: Procarididae) from the Yucatan Peninsula Richard v. Sternberg and Marilyn Schotte Macrobrachium patheinense, a new species of freshwater prawn (Crustacea: Decapoda: Palaemonidae) from Myanmar Hla Phone and Hiroshi Suzuki A new species of Enhydrosoma Boeck, 1872 (Copepoda: Harpacticoida: Cletodidae) from the Eastern Tropical Pacific Samuel Gomez New record of Ophiosyzygus disacanthus Clark, 1911 (Echinodermata: Ophiuroidea: Ophiomyxidae) in the Caribbean Sea Giomar Helena Borrero-Pérez and Milena Benavides-Serrato Sunagocia sainsburyi, a new flathead fish (Scorpaeniformes: Platycephalidae) from northwestern Australia Leslie W. Knapp and Hisashi Imamura A new species of Nannocharax (Characiformes: Distichodontidae) from Cameroon, with the descrip- tion of contact organs and breeding tubercles in the genus Richard P. Vari and Carl J. Ferraris, Jr. Rhamdia guasarensis (Siluriformes: Heptapteridae), a new species of cave catfish from the Sierra de Perija, northwestern Venezuela Carlos DoNascimiento, Francisco Provenzano, and John G. Lundberg Taxonomic review of the fossil Procellariidae (Aves: Procellariiformes) described from Bermuda by R. W. Shufeldt Storrs L. Olson Revision of the genus Squamigera (Insecta: Zygentoma: Nicoletiidae) with descriptions of two new species Luis Espinasa and Bethany Burnham Minutes of the 2004 Annual Meeting Constitution and Bylaws SMITHSONIAN INSTITUTION LIBR IVAN 8 01118 6020 447 488 505 514 523 529 541 545 551 564 575 582 594 597 ‘an ore tae bra ee hi ie Ne de Sob ne ly salle dat yi Nuc comet fat age eee Wh Bhilai Me eal % ial be bal uy enpin ia aise ape yee ‘gg! |hiteatpy Ak: gee intl pl Pil, etiea ids garcabwe Se" Wig, Pelgame Deets ahi R Ee a ‘iy ‘ato 7 Abi anus Gick, ike: Tee. r ihm Weirg mh peer ira hs ; 2 i i =~ wi ne day hone ian ~ indy aby : i lag Ws lhe @ 3 ae os Pemiiad wi © Aye ’ gre PN ty Cee G ree ee : Ps sad: ra) ree Tybet bie wei SMITHSONIAN INSTITUTION LIBRARIES INULIN NI 3 9088 01481 725 wed ay egy ae reel wer LAL tyes