A Revision of Parhyalella Kunkel (Crustacea: Amphipoda: Gammaridea) A io] This paper is dedicated to Willard D. Hartman, who retired in 1992 after serving nearly 40 years as a professor in the Department of Biology, Yale University, and as curator of invertebrate zoology at the Yale Peabody Museum of Natural History, where he was director from 1987 to 1990, and where he is presently curator emeritus. Both his tireless dedication to the Museum’s collections and the quality of his research have been an inspiration to us and many other scientists. Bulletin’ OF THE | PEABODY MUSEUM OF NATURAL HISTORY YALE UNIVERSITY 24 SEPTEMBER 2001 NEw HAVEN, CONNECTICUT A Revision of Parhyalella Kunkel (Crustacea: Amphipoda: Gammaridea) Eric A. Lazo-Wasem Division of Invertebrate Zoology Peabody Museum of Natural History Yale University New Haven, Connecticut, U.S.A. 06520 Michael F. Gable Department of Biology Eastern Connecticut State University Willimantic, Connecticut, U.S.A. 06226 Curatorial Affiliate Division of Invertebrate Zoology Peabody Museum of Natural History Yale University BULLETIN OF THE PEABODY MUSEUM OF NATURAL HISTORY Curatorial Editor-in-Chief Jacques Gauthier Executive Editor Lawrence F. Gall Publications Editor Rosemary Volpe Publications Assistant Joyce Gherlone The Bulletin of the Peabody Museum of Natural History, Yale University, publishes in- dependent monographs in the natural sciences based on research in the collections of the Yale Peabody Museum. Volumes are numbered consecutively and appear at irreg- ular intervals. The Bulletin incorporates the Bulletin of the Bingham Oceanographic Collections, which ceased independent publication after Volume 19, Article 2 (1967). The series is now out of print. For submission guidelines and information on ordering Yale Peabody Museum publications, see our website or contact: Publications Office Peabody Museum of Natural History Yale University 170 Whitney Avenue P.O. Box 208118 New Haven, CT 06520-8118 U.S.A. Phone: (203) 432-3786 Fax: (203) 432-9816 E-mail: peabody.publications@yale.edu URL: http://www.peabody.yale.edu/publications/ ON THE Cover: Parahyalella kunkeli, new species, from Taiwan, named for Yale graduate B. W. Kunkel; YPM 8236, male paratype (above), and YPM 8237, female paratype (below). Copyright © 2001 Peabody Museum of Natural History, Yale University. All rights reserved. Photograph by William K. Sacco. FRONTISPIECE: Copyright © 2001 Peabody Museum of Natural History, Yale University. All rights reserved. Photograph by William K. Sacco. Copyright © 2001 Peabody Museum of Natural History, Yale University. All rights reserved. No part of this book, except brief quotations by reviewers, may be used or reproduced in any form or media, electronic or mechanical (including photocopying, recording, or by any information storage and retrieval system), without the written permission of the Peabody Museum of Natural History, Yale University, New Haven, Connecticut, U.S.A. ISBN 0-912532-58-0 ISSN 0079-032X Printed in the U.S.A. ©) This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). CONTENTS v1 Figures and Tables vul Abstracts vit Keywords 1x Acknowledgments 3. Introduction 5 ONE* Systematic Account of Parhyalella Kunkel 1910 5 Description of Genus 5 Species Descriptions 43 Remarks on Species of Parhyalella 49 key to the Species of Parhyalella 51 Two Systematic Account of Exhyalella 51 Description of Genus 51 Species Descriptions 65 Remarks on Exhyalella 65 Key to the Species of Exhyalella 67 THREE * Systematic Account of Marinohyalella 67. Description of Genus 67. Species Description 73 FOUR * Remarks on the Hyalidae 5 Placement of Insula 5 Ecological and Distributional Notes on Parhyalella, Exhyalella and Marinohyalella 77. key to the Genera of Hyalidae 79 References FIGURES AND TABLES Parhyalella batesoni Kunkel. YPM 8188, male holotype. Various appendages. 6 FIGURE 1. _ 8 Ficure2. Parhyalella batesoni Kunkel. YPM 8188, male holotype. Pereopods. 10 ~=FiGure3. — Parhyalella whelpleyi (Shoemaker). YPM 9398, male. AMNH 6831, male paratype. us Figure 4. Parhyalella whelpleyi (Shoemaker). YPM 9361, female. AMNH 6831, female paratype. 12 Figures. Parhyalella pietschmanni Shellenberg. NHMW 17926, male paralectotype. USNM 1000145, male. Parhyalella pietschmanni Shellenberg. NHMW 17926, female paralectotype. USNM 1000150, female. Parhyalella congoensis Ruffo. MBC 39376, MBC 39373, male holotype. 8. Parhyalella congoensis Ruffo. MBC 35636-35659, female allotype. 16 Ficgureg. Parhyalella barnardi Lazo-Wasem and Gable, new species. USNM 1000153, male holotype. USNM 1000154, female allotype. 18 =FiGure10. Parhyalella barnardi Lazo-Wasem and Gable, new species. USNM 1000153, male holotype. USNM 1000155, male paratype. 20 Figureu. Parhyalella barnardi Lazo-Wasem and Gable, new species. USNM 1000153, male holotype. Various appendages. SD 13. FIGURE 14 FIGURE 15 FIGURE 21. Figure 12. Parhyalella barnardi Lazo-Wasem and Gable, new species. USNM 1000153, male holotype. Pereopods. 22. Ficure 13. Parhyalella barnardi Lazo-Wasem and Gable, new species. USNM 1000154, female allotype. 24 Figure 14. Parhyalella kunkeli Lazo-Wasem and Gable, new species. YPM 8235, male holotype. Full body and various appendages 26 =Ficgure15. Parhyalella kunkeli Lazo-Wasem and Gable, new Spas YPM 8235, male holotype. Mouth parts. 27. Figure 16. Parhyalella kunkeli Lazo-Wasem and Gable, new species. YPM 8235, male holotype. Gnathopods and uropod 3. 29 Ficure17. Parhyalella kunkeli Lazo-Wasem and Gable, new species. YPM 8239, female allotype. 30 = Figure 18. Parhyalella nisbetae Lazo-Wasem and Gable, new species. YPM 9392, male holotype. YPM 9393, female allotype. 32. ~=Fricureig. Parhyalella nisbetae Lazo-Wasem and Gable, new species. YPM 9392, male holotype. 33. Figure 20. Parhyalella nisbetae Lazo-Wasem and Gable, new species. YPM 9390, male paratype. YPM 9392, male holotype. 34. Figure 21. Parhyalella nisbetae Lazo-Wasem and Gable, new species. YPM 9392, male holotype. 35. -Ficure 22. Parhyalella nisbetae Lazo-Wasem and Gable, new species. YPM 9393, female allotype. 36 = FiGure 23. Parhyalella ruffoi Lazo-Wasem and Gable, new species. USNM 1000158, male holotype. USNM 1000159, female allotype. A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) FIGURE 24. FIGURE 25. FIGURE 26. FIGURE 27. FIGURE 28. FIGURE 29. FIGURE 30. FIGURE 31. FIGURE 32. FIGURE 33. FIGURE 34. FIGURE 35. FIGURE 36. FIGURE 37. FIGURE 38. FIGURE 39. FIGURE 40. FIGURE 41. FIGURE 42. FIGURE 43. FIGURE 44. FIGURE 45. FIGURE 46. TABLE 1. TABLE 2. Parhyalella ruffoi Lazo-Wasem and Gable, new species. USNM 1000158, male holotype. Parhyalella ruffor Lazo-Wasem and Gable, new species. USNM 1000158, male holotype. USNM 1000160, male paratype. Parhyalella ruffo1 Lazo-Wasem and Gable, new species. USNM 1000158, male holotype. Parhyalella ruffor Lazo-Wasem and Gable, new species. USNM 1000159, female allotype. Parhyalella steelei Lazo-Wasem and Gable, new species. YPM 9381, male holotype. YPM 9382, female allotype. Parhyalella steele: Lazo-Wasem and Gable, new species. YPM 9381, male holotype. Parhyalella steelet Lazo-Wasem and Gable, new species. YPM 9381, male holotype. YPM 9382, female allotype. Parhyalella steelei Lazo-Wasem and Gable, new species. YPM 9381, male holotype. Exhyalella natalensis Stebbing. SAM A4574, male lectotype, female parallolectotype. Body, anterior, and uropods. Exhyalella natalensis Stebbing. SAM A4574, male lectotype. Exhyalella natalensis Stebbing. SAM A4574, male lectotype, female parallolectotype. Gnathopods. Exhyalella natalensis Stebbing. SAM A4574, male lectotype. Exhyalella indica K. H. Barnard. ZSIC 1728/1, male lectotype. ZSIC 1728/1, male paralectotype. Body, anterior, and various appendages. Exhyalella indica kK. H. Barnard. ZSIC 1728/1, male lectotype. Various appendages. Exhyalella hartmani Lazo-Wasem and Gable, new species. YPM 9280, male holotype. Full body and various appendages. Exhyalella hartmani Lazo-Wasem and Gable, new species. YPM 9280, male holotype. Mouthparts. Exhyalella hartmani Lazo-Wasem and Gable, new species. YPM 9280, male holotype. Gnathopods. Exhyalella hartmani Lazo-Wasem and Gable, new species. YPM 9284, female allotype. Marinohyalella richardi (Chevreux). MOM 37 0992, male paratype, female paratype. Marinohyalella richardi (Chevreux). MOM 37 0992, male paratype. Mouthparts. Marinohyalella richardi (Chevreux). MOM 37 0992, male paratype. Gnathopods and uropods. Marinohyalella richardi (Chevreux). MOM 37 0992, male paratype. Pereopods. Vil Geographic distribution of Parhyalella, Exhyalella and Marinohyalella. Key diagnostic characters distinguishing species of Parhyalella. Key characters distinguishing Parhyalella, Exhyalella and Marinohyalella. ABSTRACT The genus Parhyalella Kunkel, sensu lato, is revised, based on a review of type material for all previously described species, and on new material. Parhyalella, sensu stricto, is limited to four previously described species, P. batesoni Kunkel 1910, P. whelpleyi Shoemaker 1933, P. pietschmanni Schellenberg 1938, and P. congoensis Ruffo 1953; and to the new species P. barnardi, P. kunkeli, P. nisbetae, P. ruffoi and P. steelei. The genus Exhyalella Steb- bing is re-established as a valid taxon that includes E. natalensis Stebbing 1917, E. indica K. H. Barnard 1935, and Exhyalella hartmani, new species. Marinohyalella, new genus, is a monotypic genus erected for M. richardi (Chevreux 1902). The systematic position of the three genera within the traditional concept of the Hyalidae is discussed. Keys to marine hyalid genera of the world and to species within Parhyalella and Exhyalella are provided. Lectotypes for P. pietschmanni, E. natalensis and E. indica are also designated. RESUMEN Se hace una revision del género Parhyalella, sensu lato con base en el estudio de material tipo de todas las especies previamente descritas y de material recientemente recolectado. El género Parhyalella, sensu stricto queda restringido para cuatro especies anteriormente descritas P. batesoni Kunkel 1910, P. whelpleyi Shoemaker 1933, P. pietschmanni Schel- lenberg 1938, y P. congoensis Ruffo 1953, y para cinco especies nuevas, P. barnardi, P. kunkeli, P. nisbetae, P. ruffoi y P. steelei. Se reestablece, como un taxon valido el género Ex- hyalella Stebbing que incluye E. natalensis Stebbing 1917, E. indica K. H. Barnard 1935, y Exhyalella hartmani, especie nueva. El género Marinohyalella, género nuevo, constituye un género monotipico que se ha establecido para M. richardi (Chevreux 1902). Se discute la posicién sistematica de los tres géneros dentro del concepto tradicional de la familia Hyalidae. Se presentan claves para los géneros marinos de Hyalidae en el mundo y para las especies de los géneros Parhyalella y Exhyalella. Ademas se designaron lectotipos para las especies P. pietschmanni, E. natalensis y E. indica. KEYWORDS New species, new genus, talitroidean phylogeny, Insula, taxonomic keys, ecology, distrib- ution, Parhyalella, Exhyalella, Marinohyalella, status amendments, Hyalidae. ACKNOWLEDGMENTS From the onset of our study we hoped to examine specimens of all known species of Parhyalella sensu lato rather than rely on published accounts, as these contained too many inconsistencies. We were fortunate to receive cooperation from many individuals. C. Carpine, Musée de Monaco, provided us with type specimens from the Mediterranean and also outlined historical and geographical details not available in published accounts. Type specimens and copies of the original catalog data for specimens from South Africa were lent by M. G. van der Merwe, South African Museum, and final disposition of the specimens was handled by L. Hoensen. C. D. Quickelberge, Durban National History Museum, helped determine the depository for the type specimens. W. D. Hartman, Yale University, kindly arranged the loan, through S. C. Ray of the Indian Museum, for spec- imens from India. H. S. Feinberg, formerly of the American Museum of Natural History, lent the type specimens of Parhyalella whelpleyi. Specimens of P. whelpleyi were lent by the late J. L. Barnard, United States National Museum, and additional specimens of this species were donated by Y. Wakabara, Universidade de Sao Paulo, Brazil. C. O. Coleman, Zoologisches Museum, Berlin, allowed Gable to examine specimens of P. whelpleyi in that museum, and later re-examined these and provided further observations. V. Stagl, Naturhistorisches Museum Wien, lent type and nontype specimens of P. pietschmanni; questions were subsequently addressed by P. C. Dworschak of that institution. Additional specimens of P. pietschmanni were also lent by J. L. Barnard, United States National Mu- seum. Type specimens of P. congoensis and helpful correspondence were provided by R. Jocque, Koninlijkk Museum Voor Midden-Africa, Tervuren, Belgium. In addition to specimens of previously described species, several scientists gener- ously provided specimens that we have used in our descriptions of new species. The late J. L. Barnard, United States National Museum, provided specimens from the west coast of Mexico, and from Peru. D. H. Steele, Memorial University, Newfoundland, Canada, provided specimens from Madagascar (previously identified by him as P. pietschmanni in his population studies). Finally, C. F. Dai, Taipei University, generously donated a small collection of amphipods from Taiwan; these specimens prompted us to undertake the re- vision of Parhyalella. J. D. Thomas, Nova University, Florida, provided valuable insight into Parhyalella ecology that, along with information provided by D. H. Steele, led to the discovery of a new Caribbean species. E. Harrison-Nelson, United States National Museum, provided invaluable assis- tance in processing our many loan transactions and requests for specimen information. C. B. Kim generously provided both a copy of his dissertation and useful notes concern- ing Parhyalella in Korea. C. B. Robbins allowed us to use illustrations of P. batesoni from Lazo-Wasem and Gable (1987). A. Patnode III, formerly of Eastern Connecticut State University, helped translate papers written in French. O. Breedy, University of Costa Rica, kindly translated the abstract into Spanish. L. F. Gall and R. D. White, Peabody Museum of Natural History, Yale University, answered questions of type terminology. This paper benefited greatly from reviews by A. J. Baldinger, Museum of Comparative Zoology, Har- vard University, A. A. Myers, University College, Cork, Ireland, and W. Vader, University of Tromso, Norway; all errors are the sole responsibility of the authors. 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Hotei vs Mester wy i ‘saboq| ific (Ho08" Brrif A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) Introduction The genus Parhyalella Kunkel 1910 was erected for P. batesoni Kunkel, a species repre- sented by a single male specimen collected by A. E. Verrill in Bermuda (Lazo-Wasem and Gable 1987). Since Kunkel’s monograph (1910) , six other species have been assigned to the genus either through transfer or original descriptions. These are P. richardi (Chevreux) 1902, P. natalensis (Stebbing) 1917, P. whelpleyi (Shoemaker) 1933, P. indica (K. H. Barnard) 1935, P. pietschmanni Schellenberg 1938 and P. congoensis Ruffo 1953. Only three of these species, P. pietschmanni Schellenberg (Barnard 1970), P. batesoni (Lazo-Wasem and Gable 1987) and P. richardi (Krapp-Schickel 1993), have received any review beyond their original descriptions. The generic diagnosis of Parhyalella is based principally on the lack of a palp on maxilla 1. We examined the single specimen of P. batesoni deposited in Yale University’s Peabody Museum of Natural History. The mouthpart appears to have a slight indenta- tion where a palp would originate. This indentation seems natural but could be a site of microdamage or the site of origination of a broken-off palp on maxilla 1. The latter con- dition, if true, would be contradictory to the original diagnosis of the genus. To corroborate the existence of Parhyalella as a valid genus, we obtained type speci- mens of all named species. Through correspondence with colleagues we learned of unat- tributed specimens of Parhyalella and Parhyalella-like hyalids, and we included these in our examination. A thorough study of all the specimens available indicated that Parhyalella as described in the literature is actually a complex of three distinct genera: Parhyalella sensu stricto; Exhyalella Stebbing 1917, previously synonymized with Parhyalella and herein re- established as a valid genus (revised status); and Marinohyalella new genus. In addition, six new species were discovered and assigned to the corresponding genera. A diagnosis is presented for each genus, followed by diagnoses or redescriptions of component species and a section on morphological differences among congeneric species. We give geographic distributions and, when possible, ecological information for species in all three genera. Keys are provided for all species in Parhyalella and Exhyalella. We compare the three genera morphologically and address historical problems centered on synonymies. We also comment on phylogenetic affinities among the three genera and offer suggestions concerning their proper family assignment. A new key to the Hyalidae sensu lato, based on that of Barnard and Karaman (1991), incorporates the new and re- established genera. All body lengths are given in millimeters, measured from the tip of the rostrum to the base of the telson, with the amphipod in straightened (unflexed) po- sition. The gnathopods are considered equivalent to pereopods | and 2. Article 7 of the gnathopods and pereopods is referred to only as the dactyl. Institutional abbreviations used here are as follows: AMNH, American Museum of Natural History, New York, New York, U.S.A.; MBC, Museum of the Belgian Congo, Tervuren, Belgium; MOM, Musée Oceanographique, Monaco; NHMW, Naturhistorisches Museum, Vienna, Austria; SAM, South African Museum, Durban, Republic of South Africa; USNM, United States National Museum, Washington, D.C., U.S.A.; YPM, Peabody Museum of Natural History, Yale University, New Haven, Connecticut, U.S.A.; ZMB, Zoologisches Museum, Berlin, Germany; ZSI, Zoological Survey of India, Calcutta, India. ig tain 8 vk pies Todas ste Pogpeat) shan to se) Miced? st A of txt 2 aes nig is inal duel czy apps wha , (10 - 9 vin fea + A my. ea? 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NMegahil? 3 tae Lei rust wel ah nt i ioe | ; aan ; —: "i _ se. a Systematic Account of Parhyalella Kunkel 1910 Description of Genus PARHYALELLA KUNKEL 1910 Parhyalella Kunkel; 1910:74—76. Parhyalella Barnard and Karaman; 1991:372, fig. 70D, H. Parhyalella Krapp-Schickel; 1993:758. Parhyalella Gonzalez and Watling; 1998:140-141. Diagnosis: Male antenna 2 inflated, much larger than antenna 1, first article of flagellum conjointed. Female antenna 2, flagellum with aesthetascs. Right mandible molar with plumose accessory seta. Maxilla 1 lacking a palp, with distinct shelf on outer margin of outer lobe, distal margin of outer lobe bearing nine teeth. Maxilliped palp 4-articulate, article 4 unguiform, lacking whip-like seta. Male gnathopod 1 much smaller than gnathopod 2, palm of article 6 varying from transverse to oblique; gnathopod 2, article 6 broad, palm moderately oblique. Female gnathopod 1, palm of article 6 transverse; gnathopod 2 much larger than gnathopod 1, palm moderately oblique, of distinctly dif- ferent morphology than gnathopod 1. Female gnathopod 2 of similar morphology to that of male, but smaller. Uropods 1 and 2 (both sexes), outer rami spinose along mar- gin. Telson entire. Type species: Parhyalella batesoni Kunkel 1910. Distribution: Subtropical and tropical Atlantic, Pacific and Indian Oceans. Included species: P. batesoni Kunkel, P. whelpleyi (Shoemaker), P. pietschmanni Schellen- berg, P. congoensis Ruffo, P. barnardi new species, P. kunkeli new species, P. nisbetae new species, P. ruffoi new species, P. steelei new species. Species Descriptions PARHYALELLA BATESONI KUNKEL 1910 Figures 1, 2 Parhyalella batesoni Kunkel; 1910:74—76, fig. 28. Parhyalella batesoni Lazo-Wasem and Gable; 1987:328—331, figs. 5, 6. Parhyalella batesoni Barnard and Karaman; 1991:368, 372, fig. 70. Material examined: YPM 8188: male holotype, 6.8 mm, and microscope slides of ap- pendages; Bermuda; collector and date unknown. Diagnosis: Male. Eye large, oval. Antenna 1 longer than peduncle of antenna 2. Antenna 2 strongly inflated, peduncular article 4 distinctly thicker than peduncular article 1 of an- Graz hormmee Hy irr py ALL) Ppa ee, VA 2 2. Gnathopod 1, article 5, anterior margin with medial setae -------++-++++ essere eter eee g Gnathopod 1, article 5, anterior margin bare +++ +++ +++ seer eee e tee e eee eee é 3. Gnathopod 1, article 5, anterior margin with one seta +--+ + +++: sete tree reece eee 4 Gnathopod 1, article 5, anterior margin with more than one seta ------+++++++++++++++> 5 4. Antenna 2, article 1 of flagellum 5-conjointed -------+-+-++++ss+ees essere sees P. batesoni Antenna 2, article 1 of flagellum 3-conjointed -----++-+++++s ster t eee eee P. whelpleyi 5. Gnathopod 1, article 5, anterior margin with two setae ---------++++++++::- P. pietschmannt Gnathopod 1, article 5, anterior margin with four setae ----------+--+-+-++--- P. congoensis 6. Gnathopod Il palm concave CeCe Dew Cr CieCmiChO aCate Cr Oyu rat CF CheOmCmt 10 Stan) TIPO OPO OobeOed OO O16 elo OS OF O8O (Chat OPO c 7 Gnathopod I, palm straight (OLAV I\(D:eo UD UO OOO DID OO doo oOoGodpOooodD0oKK0OnG0G000G0000C 8 7. Antenna 2; article|1 of flagellum 4=conjointed ----+----=--s--*--)-- P. barnardi new species Antennal? article of flagellumm>-conjoimted= =< <> 2 --< Fe wii P. ruffoi new species Seantennad article lot flagellum 4-conjoimted! «<9 er i P. kunkeli new species Amtenna2vanticleveor fagelluma 5-comjomted! i-inr P. steelei new species ; Sia i ae els shee 1 a isa "ae eile Cao ja) 4 Maia his ¥ rr bee . wal ae " *h A yok. wy ar hc i, as : Sr 7 ~~ iT oo NP Oars it oS aed ober! WARiNg a 2A ne - nae bys we aber tin eye ; oe a te! } Gees a : - y a WY oy ee s\ aged ¥ ped aries, L vo hy d\n 2 ai) | is 7 og To pa rye & bd soe \ S =~Air “oe L 1 iat 7 » a ae PT be ea Barats Fo ee > h- egnbeter: 9) --4 Ooh ie fara tas aaa oe a ahs To Ailes pt: man Oe oie abn a Ee pica ligula Se ee ee. Le Lay halla he Ti, AA yen) a g = & 7 oy , f] ThA a J u i. BAN a (we (lew @ - a SF; © ' - - - ee) See = bem «vs live hianl Goreme ‘sh atl . . ine Gan is tomes SP a 7 q ie '; iS j mrs é | i “a TR a ic . Ti Mee cae poner DUO athe «lakh ea {ie eg gs . pow? act See (i) te. wT re ee - a a a Lore i ape te ate ea \ dipaetes 5 * ee) Fae? (ii tS : wes © oytele Lae et alg! La? elt ee OM gi pa? carstigt rien, Pa ve’ 70 GU, Ped i. ots ono Ant ate aay ala _ - 5. bs o..4 ; , =o ) nh ef ~muraie ied i y Pile “a te le! -_ a 7% Sty eeu apkhs ( A alg inti ghee — eel Ch. “2 eee o¢@ S Ore etev' \ (84 eae) dc mdr ae iJ = ie =a Ta. é _ ¢ 2 a ‘ ners Ai 2 7 Om - ' au ; a 9 i a rns j ihe Cy : ate re gs —_ Pe? F =e pee ti =) ae ori ee - i ae mas” «6 GU == ‘ “De er -_ ieee Cily py a) eS a] : Gn ru iy ae “Ao aay. Dea ae | 5 ot) o002 @Qagag? seherealeig opamp Sat i ) ot Sh 8 ee > dali (dine te eo ee err tit | ti Gage All _ ~ a a Systematic Account of Exhyalella Description of Genus EXHYALELLA STEBBING 1917 REVISED STATUS Exhyalella Stebbing; 1917:435. Exhyalella Stebbing; 1918:66. Type species: Exhyalella natalensis Stebbing 1917. Diagnosis: Mandibular molars without plumose accessory seta. Maxilla 1 lacking a palp but represented by a distinct small spine in shallow excavation. Maxilliped palp with 4-ar- ticulate palp, article 4 unguiform, lacking whip-like seta. Male antenna 2 peduncle equal in width to antenna | peduncle. Male gnathopod 1 much smaller than gnathopod 2, arti- cle 6 with transverse palm; gnathopod 2, article 6 palm strongly oblique. Female gnatho- pod 1, article 6 with transverse palm; gnathopod 2 much larger than gnathopod 1, article 6 with palm oblique, of distinctly different morphology than gnathopod 1. Female gnathopod 2 of similar morphology to that of male. Uropods | and 2, outer rami spinose along margin. Telson entire. Distribution: Indian Ocean. Included species: Exhyalella natalensis Stebbing, E. indica (K. H. Barnard), E. hartmant new species. Species Descriptions EXHYALELLA NATALENSIS STEBBING 1917 Figures 32-35 Exhyalella natalensis Stebbing; 1917:435. Exhyalella natalensis Stebbing; 1918:66—67, pl. 11. Parhyalella natalensis K. H. Barnard; 1925:359-360. Parhyalella natalensis K. H. Barnard; 1935:294—295, fig. 11f. Parhyalella natalensis Griffiths; 1974:253. Parhyalella natalensis Barnard and Karaman; 1991:372. Material examined: SAM A4574: male lectotype, 8.0 mm; female paralectotype, 10.6 mm; female paralectotype, 10.4 mm; male paralectotype, 7.8 mm; male paralectotype, 8.3 mm; 27 male and 19 female paralectotypes; four male and two ovigerous female paralectotypes separated out by C. L. Griffiths; South Africa, Durban; collector H. W. Bell Marley, 28 May PONT: Diagnosis: Male. Gnathopod 1, palm of article 6 transverse, posterior corner demarcated it} WN fm Nt HR ON } Wr Figure 32. Exhyalella natalensis Stebbing, SAM A4574. A. Male lectotype, 8.0 mm. B. Female parallolectotype, 10.6 mm. Abbreviation: U, uropod. 1 (Yo A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) by distinct protuberance and stout, recurved spine. Female. Gnathopod 2, article 6, palm strongly oblique, very densely lined with long, fine, distally curved setae, posterior corner with broad excavation, marked by three large spines at point of dactyl closure. Type locality: Durban, South Africa. Description: Male. Eye large, oval, pale yellow in alcohol-preserved specimens. Coxal plates 1 to 3 deeper than broad, coxa 1 anteriorly produced, coxa 4 as broad as deep, nearly twice as broad as 3; distal margins of coxae smooth. Antenna | extending beyond peduncle of antenna 2, peduncle slightly thicker than that of antenna 2, less than one- half length of flagellum, flagellum of 16 articles. Antenna 2, peduncle articles 4 and 5 subequal in length, flagellum slightly longer than peduncle, with 16 articles. Left mandible, incisor with five teeth, lacinia with three to four proximal teeth, distally with bifurcate blade; right mandible, incisor process with four teeth, lacinia 5-toothed; molars of both left and right mandible strong, triturative. Maxilla 1, inner plate with two distal plumose setae, outer plate with eight distally serrate teeth, lacking palp but with single small spine on outer margin. Maxilla 2, inner and outer plates of equal width, inner plate with row of stout, plumose setae along apical margin and distal half of inner margin, outer plate with distal setae only. Maxilliped palp much larger than outer plate, 4-artic- ulate, article 4 unguiform, with large distal nail. Gnathopod | much smaller than gnatho- pod 2, article 5 slightly longer than article 6, posterior lobe broad, spinose; article 6, broad, subquadrate, palm with strong recurved spine arising at base of bulbous protu- berance of posterior corner, hind margin with three setal clusters, dactyl stout, distally bi- furcate, directed inward. Gnathopod 2, article 5 short, strongly produced between articles 4 and 6, article 6 twice as long as deep, palm strongly oblique, lined with spines and setae, hind margin 30% length of palm. Pereopods 3 and 4 slender, article 2, anterior margin with distal setae, articles 4 to 6, anterior margins nearly bare; pereopod 5 stout, spinose, shorter than pereopods 6 and 7, articles 2 and 4 with posterior lobes; pereopods 6 and 7, articles 2 and 4 with posterior lobes, pereopod 7 longest. Uropods 1 and 2 slender, spi- nose; uropod 1, rami subequal in length, of same length as peduncle; uropod 2, rami longer than peduncle, inner ramus longer than outer, four times longer than broad. Uro- pod 3, peduncle with two distal spines; ramus 50% length of peduncle, distally with one spine and a few setae. Telson apically tapered to acute angle. Female. Antennae long and thin; antenna 1 flagellum, 14 to 20 articles. Coxal plates deeper and broader than in males. Gnathopod 1, article 5 subequal in length to article 6, article 6 twice as long as wide, palm transverse, slightly convex, two spines defining pos- terior corner, hind margin setose along distal half. Gnathopod 2, article 5 lobe very deep, as wide as length of article; article 6, width about 60% of length, palm strongly oblique, convex, very densely lined with fine, curly tipped setae, three large spines at point of dactyl closure. Remarks: The type material consists of approximately 50 specimens and one slide of dis- sected appendages from one male and one female. From the records available, no speci- men was specifically designated as the holotype. The data in the original catalog at the South African Museum that correspond with this lot indicate the original manuscript name of “Parhyalella retundata” and make no reference to type status. One 8.3 mm male and one 10.4 mm female included in the specimen lot had been previously dissected, with the missing parts partially corresponding to those on the microscopic slide prepa- ration. Unfortunately, the poor quality of Stebbing’s figures do not allow detailed com- parisons with those of the dissected specimens and microscope slide. We consider the 5 NS SES e SS a creeds SSS eran ae —_ Ste ea Ss Figure 33. Exhyalella natalensis Stebbing, SAM A4574, male lectotype, 8.0 mm. Abbreviations: LL, lower lip; Md, mandible; Mx, maxilla; Mxpd, maxilliped; plp, palp; 1, left; 1, right. Figure 34. Exhyalella natalensis Stebbing, SAM A4574. A. Male lectotype, 8.0 mm. B. Female parallolectotype, 10.6 mm. Abbreviation: Gn, gnathopod. SS ee ee Figure 35. Exhyalella natalensis Stebbing, SAM A4574, male lectotype, 8.0 mm. Abbreviations: P, pereopod; T, telson. | A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) 5) two dissected specimens to be figured syntypes from among a large series of syntypes, and so, to clarify application of the name, we hereby designate the 8.0 mm male speci- men from lot SAM A4574 as lectotype for Parhyalella natalensis. EXHYALELLA INDICA (K. H. BARNARD 1935) NEW COMBINATION Figures 36, 37 Parhyalella indica K. H. Barnard; 1935:294-295, fig. 1 la—e. Parhyalella indica Sivaprakasam; 1969:299-301, fig. 2 Parhyalella indica Barnard and Karaman; 1991:372. Material examined: ZS] 1728: male lectotype, 7.2 mm; four male paralectotypes. India, Tuticorin Harbor, shore; collector H. S. Rao, Feb—Mar 1926. Diagnosis: Gnathopod 1, palm convex, posterior corner of palm rounded, defined by two stout spines projecting distally; gnathopod 2, pala sinuous. Type locality: Tuticorin Harbor, India. Description: Male. Eyes medium, black in alcohol-preserved specimens. Coxal plates 1 to 3 deeper than broad, coxa | anteriorly produced, coxa 4 as broad as deep, and one and one-half times as broad as coxa 3. Antenna 1, peduncle article 1 slightly wider than pe- duncle of antenna 2, flagellum extending 50% length of antenna 2, composed of 14 arti- cles. Antenna 2, flagellum longer than peduncle, composed of 14 articles. Gnathopod 1 much smaller than gnathopod 2, article 5 slightly longer than article 6, posterior lobe broad, with relatively few setae; article 6 subquadrate, slightly longer than broad, palm transverse, convex, lined with setae, posterior corner rounded, marked by two stout spines directed distally, inner spine one-third larger than outer, hind margin with short row of single setae, dactyl simple, annulate. Gnathopod 2, article 5 short, produced be- tween articles 4 and 6; article 6, length nearly twice the width, palm strongly oblique, dis- tinctly sinuous, moderately lined with spines and setae, posterior corner with excavation and four spines. Uropods | and 2 normal, spinose. Uropod 3, peduncle nearly three times wider than ramus, distally with two stout spines and a few setae; ramus 50% length of pe- duncle, distally with one stout spine and a few setae. Telson quadrate, width equal to length, posterior margin bracket-shaped. Female. None examined. We had an opportunity to examine only male specimens and therefore must rely on the original description of K. H. Barnard (1935) and the short re- description by Sivaprakasam (1969) for female morphology. Although neither author produced greatly detailed figures of the gnathopods, both refer to a dense brush of setae on the palm of gnathopod 2. In comparing the gnathopod 2 setal brush of female E. na- talensis and E. indica, K. H. Barnard (1935) noted it was more dense in the latter. His fig- ure of the female gnathopod 2 of E. indica is poorly drawn (for example, the hind margin of article 6; Barnard 1935:295, fig. 11b). Gnathopod 2, as illustrated, seems produced posteriorly (ironically, he comments on the poor quality of Stebbing’s [1918, pl. xi] fig- ures of E. natalensis). Sivaprakasam’s (1969:302, fig. 2) figure of female gnathopod 2 is substantially better and does not show a posterior projection as illustrated in K. H. Barnard’s (1935) figure. We believe that Barnard’s illustration is incongruent with hyalid female gnathopod 2 morphology, and that Sivaprakasam’s (1969) illustration, therefore, likely represents the true state of this character. Remarks: Barnard did not specifically assign type status in his original description. The five specimens we received from the Zoological Survey of India were labeled “types” > ELT pea! =m, = Figure 36. Exhyalella indica K. H. Barnard. A. ZSIC 1728/1, male lectotype, 7.2 mm. B. ZSIC 1728/1, male paralecto- type, 7.0 mm. Abbreviations: Md, mandible; Mx, maxilla; U, uropod; |, left; 1, right. eee NG ie es 2 Se ee ee Figure 37. Exhyalella indica K. H. Barnard, ZSIC 1728/1, male lectotype, 7.2 mm. Abbreviations: Gn, gnathopod; T, tel- son; U, uropod. Figure 38. Exhyalella hartmani Lazo-Wasem and Gable, new species, YPM 9280, male holotype, 6.8 mm. Abbreviations: T, telson; U, uropod. Figure 39. Exhyalella hartmani Lazo-Wasem and Gable, new species, YPM 9280, male holotype, 6.8 mm. Abbreviations: LL, lower lip; Md, mandible; Mx, maxilla; Mxpd, maxilliped; UL, upper lip; |, left; r, right. Figure 40. Exhyalella hartmani Lazo-Wasem and Gable, new species, YPM 9280, male holotype, 6.8 mm. Abbreviation: Gn, gnathopod. —————————————— eee A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) 63 (=syntypes). To clarify application of the name, we hereby designate the 7.2 mm male specimen from lot ZSI 1728 as lectotype for Parhyalella indica. EXHYALELLA HARTMANI LAZO-WASEM AND GABLE NEW SPECIES Figures 38—41 Material examined: YPM 9280: male holotype, 6.8 mm. YPM 9284: female allotype, 6.2 mm. YPM 9282-9283: two male paratypes, each on SEM stub. YPM 9286: 11 male para- types. YPM 9287: male paratype, 6.2 mm. YPM 9835: male paratype, 6.2 mm. YPM 9836: male paratype, 6.2 mm. YPM 9837: male paratype, 5.1 mm. YPM 9840: male paratype, 6.3 mm. YPM 9281: female paratype, 5.1 mm. YPM 9285: four female paratypes. Sey- chelles, Beau Vallon, Mahé Island, Yale Seychelles Expedition Station 33, lat 4°37’S, long 55°26’E; collector A. J. Kohn, 24 Jan 1958. Diagnosis: Male. Gnathopod 1, article 6, proximoposterior corner of palm demarcated by a distinct protuberance and stout, curved spine; gnathopod 2, article 6, hind margin with three bifurcate spines distinctly set away from posterior corner of palm. Female. Gnathopod 2, article 6, palm strongly oblique, weakly sinuous, densely lined with long thin setae, posterior corner with three large spines and slight excavation. Type locality: Mahé Island, Beau Vallon, Seychelles. Description: Male. Eye large, oval, black in alcohol-preserved specimens, coxa | anteriorly produced, coxae 2 and 3 longer than broad, coxa 4 broader than deep, nearly twice as broad as coxa 3. Antenna 1, article 1 of peduncle slightly thicker than peduncular articles 4 and 5 of antenna 2, flagellum composed of 16 articles, extending beyond peduncle of an- tenna 2. Antenna 2, peduncle slightly shorter than flagellum, flagellum composed of 13 ar- ticles. Upper lip broadly rounded, distally setose. Mandibular molar triturative, incisor 6 toothed, left lacinia mobilis with five teeth, right accessory plate bifurcate, one margin un- evenly toothed and serrate. Maxilla 1 palp lacking, but represented by a small spine on a distinct prominence on exterior margin. Maxilla 2, inner plate slightly narrower than outer plate, distal half of inner margin lined with row of plumose and simple setae, prox- imally defined by one stout plumose seta; outer plate distally setose with submarginal setae and one plumose seta. Maxilliped palp unguiform, composed of four articles, article 4 with distal nail. Gnathopod 1, article 6 nearly one-third longer than wide, palm trans- verse, nearly straight, and lined with setae and small facial spines, posterior corner with stout, slightly curved spine, hind margin weakly concave at distal end, dactyl bifurcate. Gnathopod 2, article 5 strongly produced between articles 4 and 6, margin very spinose, length of article 6 twice the width, palm strongly oblique, spinose, sinuous, proximally de- fined by four spines somewhat stouter than those lining palm; hind margin proximal to palm one-third length of palm, nearly straight, with three marginal spines set off from the posterior corner. Pereopods 3 and 4 slender, article 2, anterior margin with distal setae, posterior margin with medial and distal setae, articles 4 to 6, anterior margins nearly bare; pereopod 5 stout, spinose, shorter than pereopods 6 and 7, articles 2 and 4 with posterior lobes; pereopods 6 and 7, articles 2 and 4 with posterior lobes, pereopod 7 longest, 66% of body length in largest males, spines on anterior margin of article 6 in groups of three. Uropod 1 extending beyond uropod 2, peduncle subequal in length to rami; uropod 2, inner ramus longer than outer; uropod 3, peduncle with many small facial spines and two (rarely, three) stout distal spines; ramus nearly 75% length of peduncle. Telson longer than wide, tapering to a weak point, lateral margins with setae, and apex with two pairs of setae. Female. Antenna 1, flagellum with aesthetascs. Gnathopod 1, article 6 rectangular, length Figure 41. Exhyalella hartmani Lazo-Wasem and Gable, new species, YPM 9284, female allotype, 6.2 mm. Abbreviation: Gn, gnathopod. A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) 65 of article 6 twice the width, palm transverse, posterior corner rounded, defined by two stout spines, hind margin weakly setose. Gnathopod 2, posterior margin of article 4 se- tose, length article 6 nearly twice the width, palm weakly sinuous, densely lined with setae (some plumose), with three stout spines at point of dactyl closure. Habitat: Shallow water, among coralline rubble. Etymology: This species is named in honor of Willard D. Hartman of Yale University, the senior invertebrate zoologist of the Yale Seychelles Expedition (1957-1958). Remarks on Exhyalella The three species of Exhyalella are primarily distinguished by characters of gnathopod morphology. Unfortunately, unless both males and females are collected together and ex- amined, specific identification can be difficult. Males of Exhyalella indica lack the prom- inent tubercle and stout recurved spine at the posterior corner of article 6, gnathopod 1, found in both E. hartmani and E. natalensis. Unfortunately, because of considerable mor- phologic and allometric variation, no single character definitively distinguishes the males of E. hartmani and E. natalensis. However, several characters in combination will sepa- rate these two species. The relative size of the eye is greater in E. natalensis than in E. hart- mani: in similar-sized individuals the eyes of the former appear 20% to 25% larger than those of the latter. In E. natalensis, the length of the ramus of uropod 3 rarely exceeds one-half the length of the peduncle, and often is much shorter, especially in larger indi- viduals. In E. hartmani the ramus of uropod 3 is often 75% or more of the length of the peduncle. Finally, the hind margin of article 6 of gnathopod 2 is usually much shorter relative to the palm in E. natalensis than in E. hartmani. Aside from the characters that help to distinguish males of E. hartmani from males of E. natalensis, there are clear morphological characters that separate the females of these two species. Females of E. natalensis and E. indica have a brush-like appearance of the setae lining the palm of gnathopod 2, which is lacking in E. hartmani. In E. hartmani, although the palm of gnathopod 2 of the females is densely lined with setae, the setae are much stouter and do not present a brush-like appearance. Females of E. natalensis and E. indica, however, are virtually indistinguishable. Key to the Species of Exhyalella (males and females) 1. Female gnathopod 2, article 6 palm with dense, brush-like setae --------+-++++++++++++: 2 Female gnathopod 2, article 6 palm with fine spines and setae, ILO fare (OC AGHA LAU S INU OR h =e Pasian a eho 2) 2 = 2-22) 0c inne ein ei E. hartmani new species 2. Male gnathopod 1, posterior corner of article 6 with distinct protuberance ----- E. natalensis Male gnathopod 1, posterior corner of article 6 broadly rounded, lacking a protuberance [0 le! ie) 6),6@:.6, 101 0) .@) 1e)6) 0) Je)e) (0, .e; (e) (w/a Velo! 0116! 6) 40).18) 0) 01 "e) 16) 16 7:0 e) 18). ele] (a kee) le eels see nes ee E. indica ie 6 7 4 vd py Td ery r i ibe a : ne : . wiling Aline gill tp Nya a» eee) dh By ng a ye GO| ‘soot ols ; aula . 7 | App § tas) divAal? « mein 4A Aue ; cpt eriatle. Large Systematic Account of Marinohyalella Description of Genus MARINOHYALELLA LAZO-WASEM AND GABLE NEW GENUS Diagnosis: Mandibular molar (left and right) with plumose accessory seta. Maxilla 1 with minute, |-articulate palp. Maxilliped with 4-articulate palp, article 4 unguiform and lack- ing whip-like seta. Male antenna 2, peduncle not greatly thicker than peduncle of antenna 1. Male gnathopod 1 much smaller than gnathopod 2, palm transverse; gnathopod 2, palm strongly oblique. Female gnathopods subequal in size, palms transverse, of similar mor- phology. Female gnathopod 2 of distinctly different morphology than that of male. Uropods 1 and 2, outer rami lacking marginal spines. Uropod 3 uniramous. Telson entire. Type species: Hyalella richardi Chevreux 1902 (by monotypy). Distribution: Mediterranean Sea, off Isle of Alboran, Spain; Banyuls, France; and Genoa, Italy. Included species: Marinohyalella richard. Remarks: Chevreux placed his species in the freshwater genus Hyalella (his fig. 1 erro- neously indicates the genus as Hyale), undoubtedly because the characteristics of the gnathopods and maxilla match those for Hyalella. Possibly because it occupies a fully marine habitat, Charniaux Legrand (1951) removed this species to the genus Parhyalella, but gave no specific reasons for doing so. Species Description MARINOHYALELLA RICHARDI (CHEVREUX) 1902 NEW COMBINATION Figures 42—45 Hyalella richardi Chevreux; 1902:223—227, figs. 1, 2. Hyalella richardi Chevreux; 1935:121, pl. 16, figs. 1, 2. Hyalella richardi Brian; 1940:5, figs. 1—4. Parhyalella richardi Charniaux Legrand; 1951:377. Parhyalella richardi Barnard and Karaman; 1991:372. Parhyalella richardi Krapp-Schickel; 1993:758—759, fig. 519. Material examined: MOM 37-0992: male holotype, female allotype, male paratype, 8.6 mm; female paratype, 5.5 mm; six paratypes. Mediterranean Sea, off Isle of Alboran, Spain, “Princess Alice” Station 643; collector Richard and Neuville, 1896. ZMB 25333: four males, one ovigerous female. Italy, Genoa, on beach or near shore, in sea grass; col- lector [L.] Brian, 7 Apr 1940. Type locality: Mediterranean Sea, off Isle of Alboran, Spain. Figure 42. Martnohyalella richardi (Chevreux), MOM 37 0992. A. Male paratype, 8.6 mm. B. Female paratype, 5.5 mm. A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) 69 Diagnosis: As for the genus. Description: Male. Eyes small, nearly round. Coxae | to 4 large, broad and deep, lower margins bare. Antenna | slender, extending well beyond peduncle of antenna 2, flagellum with 12 to 13 articles. Antenna 2 slender, flagellum with 13 to 16 articles. Mandibles lack- ing a palp, with 5-toothed incisor, left mandible with 5-toothed accessory plate (lacinia), right mandible with serrate, bifurcate accessory plate, proximally with two teeth. Maxilla 1 with minute, 1-articulate palp with distal spine, outer plate distally with eight (nine?) serrate spines, inner plate distally with two large, plumose setae. Maxilla 2, inner plate inner margin with a few small setae, distal half with row of setae and a single proximal plumose seta, outer plate with long setae distally, outer margin finely setose. Maxilliped inner plate with setose inner margin and three stout apical teeth, palp 4-articulate, arti- cle 3 with facial setal row, article 4 unguiform with distal nail. Coxal gills from gnatho- pod 2 through pereopod 6; pereopods 5 and 6 with accessory gills. Gnathopod 1 much smaller than gnathopod 2, posterior lobe of article 5 broad, lined with small marginal setae; article 6, palm transverse, posterior corner with stout spine on inner and outer face, dactyl stout, closing tightly against palm. Gnathopod 2, article 6, palm strongly oblique, nearly full length of article, densely lined with fine setae, inner margin lined with several rows of small spines, dactyl long and curving, closing against excavation and small spine at posterior corner of palm. Pereopods 3 and 4 similar, weakly setose, article 2 narrow; pereopods 5 and 6, article 2 broad, articles 4 to 6 strongly spinose; pereopod 7 spinose, article 2 broadly expanded, posterior margin moderately serrate. Uropods | and 2, inner rami slightly wider than outer, two spines along inner margin; outer rami with terminal spines only; uropod 3, peduncle and ramus with two distal spines each. Telson subquadrate, posterior margin slightly pointed. Female. Eyes relatively larger than those of male and slightly oblong. Antennae 1| and 2 slender, with 8 to 10 flagellar articles. Gnathopod 1, article 5 broad, article 6 twice as long as wide, palm transverse; gnathopod 2 of similar size and form, article 6 relatively longer than that of gnathopod 1. Brood lamellae from gnathopod 2 to pereopod 5, thin, con- secutively smaller from 2 to 5, densely lined with short setae. Uropods | and 2, outer rami with terminal spines only. Remarks: Chevreux indicated a type series that included 11 specimens; we found only 10 specimens. Krapp-Schickel (1993) states in her generic diagnosis that Parhyalella lacks a palp on maxilla 1, but she clearly illustrates and describes a rudimentary palp on that mouth- part in her figures of P. richardi (1993:759, fig. 519). Our figures agree with hers except for a few minor differences in setation; most notably, her figure of male gnathopod 2 (1993:759, fig. 519) lacks setae on the posterior margin of article 2, which are evident in the type material examined. Figure 43. Marinohyalella richardi (Chevreux), MOM 37 0992, male paratype, 10.4 mm. Abbreviations: Md, mandible; Mx, maxilla; Mxpd, maxilliped; 1, left; r, right. L Gn2 iF) ap j sugagetle™ i ANG | Te ie ‘oa Why ye Wy Sig Figure 44. Marinohyalella richardi (Chevreux), MOM 37 0992, male paratype, 10.4 mm. Abbreviations: Gn, gnathopod; U, uropod. Figure 45. Marinohyalella richardi (Chevreux), MOM 37 0992, male paratype, 10.4 mm. Abbreviation: P, pereopod. Remarks on the Hyalidae Historically, the generic diagnoses within the hyalid—hyalellid talitroideans have been de- fined as in Bulycheva (1957). The currently accepted and expanded diagnoses are those of Barnard and Barnard (1983) and Barnard and Karaman (1991). Although the core of nontalitrid genera has been realigned several times with respect to familial or subfamily placement (Hyalidae/Hyalinae or Hyalellidae/Hyalellinae), the generic composition has been generally stable. Characteristics of the telson (cleft or entire), maxilla 1 palp (lack- ing, 1-, 2- or 3-segmented), uropod 3 inner ramus (present or lacking), and basic gnatho- pod architecture have served to distinguish the various hyalid genera. It is somewhat surprising, therefore, that Marinohyalella richardi could have been assigned to the genus Parhyalella. Although Charniaux Legrand (1951) did not provide justification for doing so, the change has been virtually unquestioned for more than 50 years. Similarly, Ex- hyalella natalensis Stebbing and E. indica were incorrectly subsumed under Parhyalella without justification. In light of the clear and accepted morphological distinctions sepa- rating the genera now assigned to the Hyalidae by Barnard and Karaman (1991), the transfer of Hyalella richardi to the new genus Marinohyalella is definitely warranted. By similar reasoning, both of the previously known species of Exhyalella must be removed from Parhyalella, and Exhyalella re-established as a valid taxon. The principle diagnostic features distinguishing the genera are summarized in Table 2. The genus Parhyalella has been placed in various families, although most often it has been aligned with the Hyalidae. Bulycheva (1957) placed Parhyalella in the Hyalidae after devising the concept of the superfamily Talitroidea, an arrangement accepted and followed by Barnard (1969). Bousfield (1982) removed Parhyalella from the Hyalidae and placed it within the Hyalellidae. Zeidler (1991) also recommended aligning Parhya- lella and Allorchestes with the Hyalellidae. Barnard and Karaman (1991) repositioned Parhyalella again into the Hyalidae, an arrangement recently followed by Ruffo and Vader (1998) and Wakabara and Serejo (1998). Previously, Krapp-Schickel (1993) had placed Parhyalella and Hyale within the family Talitridae. Bousfield (1996) again assigned Par- hyalella and Allorchestes to the Hyalellidae; in all previous classifications other than Zei- dler’s (1991) Allorchestes had been placed within the Hyalidae. Gonzalez and Watling (1998) also place Parhyalella within the Hyalellidae but did not address family level clas- sification. In most of the taxonomic assignments described above, authors have not justified their deviations from Bulycheva’s (1957) original arrangement of talitroidean genera. We have adhered to the classification of Barnard and Karaman (1991) and place Parhyalella in the Hyalidae. Our justification for doing so is pragmatic: most authors have associated Parhyalella (and Allorchestes) with marine and estuarine hyalids, rather than with fresh- water hyalellids, and generally identify taxa according to this arrangement. Clearly, rig- orous cladistic and molecular analyses are needed to resolve talitroidean classification more fully. In the case of the former, variability of potentially useful characters within the 74 PEABODY MuSEUM BULLETIN 46 Table 2. Key characters distinguishing Parhyalella, Exhyalella and Marinohyalella. Mandibular Gnl A2, A2, molar, and peduncle, flagellum accessory Maxilla 1, Gn2, males article | seta(e) palp females Parhyalella Tumescent Conjointed On right Absent, margin Dissimilar mandible often finely setose Exhyalella Normal Normal Absent Absent, margin Dissimilar with a single spine Marinohyalella Normal Normal On left Present, Similar and right 1 articulate shape mandible and size three genera must be assessed if an analysis is to be based on stable character states. As a taxonomic aid, we offer here a revised version of Barnard and Karaman’s (1991:366) “Key to Genera of Hyalidae,” but have included new couplets to account for the inclusion of Exhyalella and Marinohyalella and have arranged for the placement of the questionable genus /nsula at the end of the key. In Barnard and Karaman’s (1991:718) “Key to Families of Talitroidea,’ couplet 6 separates Hyalidae from the Hyalellidae by the possession of a cleft telson (Hyalidae) rather than an uncleft telson (Hyalellidae). How- ever, Barnard and Karaman (1991:366) include Parhyalella (which possesses an entire tel- son) in their key to the hyalid genera and recognize both cleft and entire telsons in their description of the family Hyalidae. Within the context of the talitroidean genera (Bulycheva 1957, Barnard 1969) or the current arrangement of Barnard and Karaman (1991), the distinctions among hyalid, hyalellid and talitrid genera remain clear even with the addition of Marinohyalella and Exhyalella to the talitroidean complex. Ruffo (1953) was the first to observe that three species groups occur in Parhyalella, noting that P. batesoni, P. pietschmanni and P. congoensis are morphologically allied, and distinct from other congeners. Furthermore, he noted the close morphological relation- ship between Exhyalella (then Parhyalella) natalensis and Exhyalella (then Parhyalella) indica and their combined distinctiveness from Marinohyalella (then Parhyalella) richardi. Barnard (1972) suggested that Parhyalella represents a composite genus, and he discussed its morphological, ecological and evolutionary relationships with other tal- itroids. Although he clearly disagreed with Bulycheva’s synonymy of Parallorchestes with Parhyale on the basis of mouthpart morphology, he did not extend this reasoning to the morphological incongruencies of the Parhyalella complex. Parallorchestes, a marine genus with a 2-articulate maxilla 1 palp and a minutely bi- ramous uropod 3, has been treated as one of the most primitive hyalids (Barnard 1979; Bousfield 1981; Barnard and Karaman 1991). With an adaptation to the terrestrial habi- tat in talitroids, originating from a Parallorchestes-like ancestor, there is a clear morpho- logical trend toward reduction of the maxillary palp and the disappearance of the A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) 75 biramous condition of uropod 3. Parhyalella, Exhyalella and Marinohyalella richardi with their reduced or absent maxilla 1 and uniramous uropod 3, occupy a position between Parallorchestes and the advanced, land-dwelling talitrids. However, the relationships among these three genera is unclear. If Exhyalella and Marinohyalella are excluded, the remaining genus, Parhyalella, apparently represents a morphological and ecological link between the fully marine hyalids and the terrestrial talitrids. Although Parhyalella pos- sesses typical preadaptations to terrestriality (for example, the incrassate condition of male antenna 2), Barnard (1972) has pointed out that the entire telson of Parhyalella sensu lato represents an improbable reversal in the progression from marine hyalids to the terrestrial talitrids, both of which have a fully cleft telson. It is possible that Parhya- lella represents a sister group, along with Exhyalella and Marinohyalella, to an antecedent of the terrestrial taxa. Demonstrating this would require a detailed phylogenetic analysis of all genera within the talitroidean complex. Placement of Insula The genus Insula was erected by Kunkel (1910) for a single species, Insula antennullela from Bermuda. Barnard (1969) speculated that I. antennullela was probably incorrectly described, in part because of the improbability of a hyalid possessing a 3-articulate (rather than 4-articulate) maxilliped palp. Barnard (1969) suggested that the single spec- imen used by Kunkel for his description was actually a juvenile Hyale. Strangely, Barnard (1979) later synonymized all three species of Hyale reported by Kunkel (1910) as occur- ring in Bermuda with Parhyale hawatiensis (Dana). Recent collections, however, conclu- sively show that Hyale spp. occur in Bermuda (Baldinger and Gable 1995), and therefore ironically support Barnard’s (1969) original suggestion concerning Insula. Several years ago, Lazo-Wasem located the microscope slide apparently used by Kunkel for his illustration of the maxilliped of Insula. Unfortunately, the slide is dried out and the features of the maxilliped are obliterated. It is therefore impossible to determine the status of Kunkel’s Insula based on original material. Furthermore, an examination of newly collected material from Bermuda over the last decade has failed to produce any specimens conforming to Kunkel’s description of Insula. We agree, therefore, with Barnard’s opinion, and believe that disregarding the genus /nsula is warranted. It is pro- visionally included in our key for comparative purposes, but we have not made reference to the questionable feature of the maxilliped. We believe Insula should be suppressed after a careful review of all Bermudian hyalids. Ecological and Distributional Notes on Parhyalella, Exhyalella and Marinohyalella The distribution and ecology of Marinohyalella richardi and Exhyalella spp. can be sum- marized only in general terms, because little detailed information exists beyond the orig- inal collection data associated with type specimens and a few subsequent collection records for some species. At present, Marinohyalella is known only from shallow waters of the Mediterranean Sea, where it sometimes occurs in large numbers. Exhyalella, re- stricted to the Indian Ocean, is found intertidally near sandy beaches. Two species, E. in- dica and E. natalensis, have been found nestled among algae. For species of Parhyalella more detailed distribution and habitat information is known, although again, provided only from original collection data for most species. ‘saidads vyjajvAysvg ‘dsq s12j2a1s gq ISd ‘Hoffn4 _ Jad sMagsiu q ‘sud syayuny q “Ud ‘Ipsvusng d AQ ‘sisua08uo0) gq “udg suuviuyssjaid gq dg ‘1Aajdjaym q [Md ‘1u0sajKq viJalvAYysvg 4qd ‘ubiiivy “y AY ‘voipur “gq “UTA ‘sisua]vJvU vpyjajvAyx] QUA IpsvyI14 vijapV. (YOULIDIY DAY ‘SUOLIDIAILQQY “v]Ja[vAYOULIDPY pur vIja[vAYxX ‘VIJa]vAYAvg JO VOLNGLYSIP sydeisoary ‘Op AINSI A REVISION OF PARHYALELLA KUNKEL (CRUSTACEA: AMPHIPODA: GAMMARIDEA) Wh Parhyalella has a circumglobal distribution, and is widely distributed in the Atlantic, Pa- cific and Indian Oceans (see Figure 46). Half of the species of Parhyalella have been recorded from beaches adjacent to near-shore coral reefs. Parhyalella whelpleyi is the only species that has been recorded from salt marshes with estuarine salinity (da Cunha Lana and Guiss 1992; Wakabara, personal communication). Although Parhyalella spp. are typ- ically found intertidally, P. kunkeli has been recorded from an offshore locality in rela- tively deep water (20 m). An unidentified species of Parhyalella is found associated with mangroves (predominantly Rhizophora stylosa) in Australia (Lee 1997). Several species (e.g., P. nisbetae, P. steelei and P. pietschmanni) have been collected at the ocean-shore interface, often among algae or turtle grass washed onshore by pre- vailing winds. J. D. Thomas (personal communication) has reported occasionally en- countering large numbers of an unidentified species of Parhyalella in the Florida Keys (United States) after periods of extended onshore winds; at other times in the same places he never encountered Parhyalella. It is possible that some species of Parhyalella normally inhabit subtidal algal or grass beds, and wash inshore when wind patterns are favorable. If this is true, it may explain our inability to recollect P. batesoni in Bermuda. In the years we collected in Bermuda, prevailing wind patterns did not carry large amounts of plant material inshore, as often happens in Bermuda (Gable, personal ob- servation). If P. batesont is associated with offshore subtidal plants and algae, then wind and storm conditions would have created unfavorable collecting conditions during our stays. In sum, the species of Parhyalella occupy a surprisingly diverse and perplexing array of habitats, and detailed field work is needed before even a basic understanding of their ecology can be obtained. Key to the Genera of Hyalidae (modified from Barnard and Karaman 1991) 1. Dactyl of maxilliped short and blunt On UEopodss)lackine TamniSi-yei< F-12i- een hak ans eos heels ee oes Ceinidae, especially Hyachelia Dactyl of maxilliped unguiform ©. (6) (6) (0. 6fe a) hn). s) (a (aoe) je 2) ese se fene, 0, ,9rle) eo! a) ¢e) Siiewenelese slislielécemels pela) omens eles D) De Uropod 3 with minute inner ramus 0) fe (6, “e) @) a) (6) (0, <0).0fe) 0! @) (e! (0) 0) (a) a) (opie; (0) 0) (0.0) .0 (eye) ese) ars) enag sel ie) Jee le kedeliel die 3 Uropod 3 lacking inner MNNAlUIG oO OOO OOO ObIDIO nono Ooo duo o oC oOD DOOD oD ooo oG Oooo OGo OK 4 3). Maxilla ] palp l-articulate COoOMOODDOOOO UC DOOoOOoDD ODOC ODO OHH ODDO OA GO OOD OOOO SD Parhyale Maxilla ] palp 2-articulate SOouUAoDODOODOODOODOCOU ODD Ob dG Un HOO o nH o ADO GON aC Parallorchestes Ay Telson cleft TOO AOUOCUOMmEOD OOO OOOO OU OOD AMO DOOD OOO OUD OOO ODA dodseUEOoOoSODdUbOS 5 Telson entire Sse SORA A OOOnMO OO Oo DOD CO DODO ODdUoDO Doar dT Dona aOC OUR OO dO OOOO oO Ss 8 SDacal otmealliped lackine dons: wiip-like Setai- "om 717 -ia ee hak Soy eos 6 WactylLofinaxilliped bearme lone whip-like seta. <>< -- <7 -~ 44 - io < ie he 7 6. Male gnathopod 2, article 5 produced into a lobe between articles 4 and 6; palp of maxilla 1 short, NOPE RrCMe In eat OucHGrOmOUel plates = chic tery a-ha ote gs eens ree ened ae Allorchestes Male gnathopod 2, article 5 not produced; palprotmaxila Wong jextendinsto.end of outer plate --- 2° << -- Hyale 78 PEABODY MUSEUM BULLETIN 46 7. Male and female gnathopods diverse, male gnathopod 2 enlarged ---++-++++++++-- Lelehua Gnathopods of both sexesismalland temaleshkev" =~ 72-7 >So ese et Micropythia 8. Male gnathopod 2 enlarged, different from gnathopod 1 -+--+ +++ seers rtt etter tees: 2) Male onathopod aired 2:of mrediumiisizes simmilanirs er elctiis 1) - oe oem eho iid 10 9. Male antenna 2 peduncle tumescent, palp of maxilla 1 absent -+--+++++++++++- Parhyalella Male antenna 2 peduncle not thickened, similar in diameter toranternal; palp ofmiarallanlvalsingle:setaty iF ior 2) ele ae ae Exhyalella 10. Maxilla 1 palp 1-articulate, vestigial, length equal to width ------+-++++++-- Marinohyalella Maxilla 1 palp 1-articulate, length three times longer than width ------++++++++++-- Insula REBERENCES ANDRES, H. G. 1975. 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A new genus and species of phreatic amphipod (crustacea: Amphipoda) belong- ing in the “Chiltonia” generic group, from Dalhousie Springs, South Australia. Trans. R. Soc. S. Aust. 115(4):177-187. a. Pairs ‘ ; ne Ors ieee 5 i or ete es’ - ws a ices SE - - 7 a a -- > tor ane a ove age | _ : > Sy ' (~—e oy ve 7 : Bey vies a we ae = yah » > leat a le Va “pioe camuimnal a "ee e¢2 -« . ley iin > ee ay) = 2 r= a ‘ ‘ : aa = We ies Ici) BAK ps -% eDi) (ee 7. il = vt f > ‘ aie ot Puree i.e He ; ? a iatien a sf - - - ia mu , ie ——— ' - ~ . : hd a: — — os an | a ae, Toss - nea nyt be, at pane re ame § ee ae ; @ . 7 eam > > ~ ¢ ‘ f _ : Ory ieee \ me - ya, 2s 7 & ee pier 2 “te a : Te; ivivte Wasdale 7 7 PAI ie oe ek a} 11 5, petytes “eepatre - fee noe & | 7 year Wea) “ Aubee 2s) 04 a + a (Ric naan Sek Eee GP 2s se> 2h. culttion Any ive nt} otal Ht ie oa/en pa Tye Drive iT ae ag aa eek tics. Comite a Net te, bert, ot Meinl _ ere. 7? F Vakpes | sty Ae ° & Chikts! GQatiis ie fax... ee aes 4" Vv ie Tae; : - a Rina. ¥ m THe, bs OC a roe. oe sethe: ~ re eas ees o ie he i ¥' 4 ; ; —% uv m3 (ge ww mame | ae hy ’ i. @ ton P| a y i : oo on ; = > , , 4 7 Sf ae ' ug We 5? \ +” ti, Ry » 7 as t i a _ we ¢ , 6 7 dl 4 A } ~ on Na v ’ p *) 26°14 (2c ae 1949 oer * | - hte. 5) , mt 7 \ aw es » : 7 ec = 2 } aa Gul a. a 4 oy Ge as = = y¥ }o 9 7 r4 } - Dy > e ai o a | = _ ' > - uy eé rae a him ; . ( & 7 j a. anes! 1 Mae sna < pul oa : : a i 4 _ otis, (22. hdotiesiateney tat ease: te at ( ship 121 1 sen Weh tiie : <= \ Ger Licegé @) GE 2) >. ing : - ’ 7 al (At wry Vv YVR peaihactyan ad vat . 5, eet erate eh eres YALE PEABODY MUSEUM PUBLICATIONS The Yale Peabody Museum Publications Office supports the communication mandate of the Yale Peabody Mu- seum’s mission by making the Peabody’s research, collections, exhibits and educational activities accessible to the academic community and to the general public. An important part of the Publications Office’s responsibil- ities is publication of the work of curators, staff and research associates, and their colleagues, with an emphasis on research based on the collections of the Yale Peabody Museum. The Publications Office administers the Yale Peabody Museum's scholarly publishing program, and produces the following titles: BULLETIN OF THE PEABODY MUSEUM oF NATURAL HIsToRY In publication since the opening of the Yale Peabody Museum building in 1925, the Bulletin publishes monographs of original research in the natural science disciplines represented by the collections of the Pea- body Museum’s curatorial divisions. Volumes are numbered consecutively and appear at irregular inter- vals. The Bulletin incorporates the Bulletin of the Bingham Oceanographic Collection, which ceased inde- pendent publication after Volume 19, Article 2 (1967). POSTILLA Published at frequent intervals since 1950, Postilla pre- sents short papers based on original scientific research in the natural science disciplines represented by the col- lections of the Peabody Museum’s curatorial divisions. YALE UNIVERSITY PUBLICATIONS IN ANTHROPOLOGY YUPA embodies the results of research in anthropol- ogy directly conducted or sponsored by the Yale Uni- versity Department of Anthropology and the Peabody Museum Division of Anthropology. Occasionally, other manuscripts of outstanding quality on subjects of special interest to the Faculty of the Department of Anthropology may also be included. Titles range in size from brief papers to extensive monographs, are numbered consecutively as independent contribu- tions, and appear at irregular intervals. SPECIAL PUBLICATIONS The Peabody Museum on occasion produces special publications based on exhibitions, conferences and other Museum projects as appropriate. Publications available in each series, along with ordering information and forms, are available from our website at http://www.peabody.yale.edu/publications/, or contact the Publications Office at the address below. (GUIDELINES FOR AUTHORS Priority is given to manuscripts by Yale Peabody Museum curators and staff, and to research associates using the Museum’s collections or research facilities. For the YUPA series, members of the Department of Anthropology are given priority over outside submissions. All manuscripts are sent out for external peer review by one or more referees, and are also reviewed internally at the Peabody Museum by the Executive Editor in consultation with the Curatorial Editor-in-Chief for the series, and with one or more members of the appropriate curatorial divi- sion. Manuscripts may go through a single or several revision cycles; revision and major changes are not per- mitted once page proofs are set. At present there are no page charges for accepted manuscripts. Detailed submission guidelines with manuscript preparation instructions are available on our website at http:// www.peabody.yale.edu/publications/Guide.html, or contact the Publications Office at the address below. Interested authors should contact the Executive Editor before submission of manuscripts for possible publica- tion. Address correspondence to: Lawrence F. Gall, Executive Editor Peabody Museum of Natural History Yale University P.O. Box 208118 170 Whitney Avenue New Haven, CT 06520-8118 U.S.A. Phone: (203) 432-9892 Fax: (203) 432-9816 E-mail: lawrence.gall@yale.edu URL: http://www.peabody.yale.edu/publications/