Contents October 15, 1994 Editorial. From our Readers The Amphipod Superfamily Dexaminoidea on the North American Pacific Coast; Families Atylidae and Dexaminidae: Systematics and Distributional Ecology. E. L. Bousfield and J. A. Kendall New Species of the Amphipod Crustacean Genera Photis and Gcim- maropsis (Corophioidea: Isaeidae) from California. K.E.Conlan ' 67. Vol. I, No. 2. Errata of subject matter. 74. The Phyletic Classification of Amphipod Crustaceans: Problems in Resolution. E. L. Bousfield and C.-t. Shih 76. National Library of Canada ISSN No. 1189-9905 AMPHIPACIFICA is an international jour- nal of invertebrate systematics, aimed primarily at publication of monographic treatments that are too large or bulky (50 - 100 printed pages including plates) for acceptance by standard taxonomic jour- nals. Initially* the contents will feature mono- graphic studies on crustaceans of the faunistically rich and geologically ancient North American Pacific coastal marine region. The scope of this new journal extends* geographically to other broadly Pacific regions* and faunistically to other arthropods, mollusks, annelids, to other regional invertebrate tax a, both aquatic and terrestrial* in- cluding parasites, and to aspects of vertebrate animals that may involve systematics* ecology and behaviour. The journal appears quarterly* or approximately so, with a run of 300-400 copies per issue, each of about 150 pages, and a volume (yearly) total of 600+ pages (approximately). The printed page size is 8,5 X 11 inches (22 X 27,5 cm). Paper quality accommodates line cuts and half tones at 400-600 d.p.i.* and a limited number of colour plates at author’s cost. Manuscripts are to be sub- mitted in “camera-ready” computerized format (IBM- or M AC-compatible diskettes), and also in hard copy, that have previously been refereed (name to be supplied) and text-edited at the au- thor’s instigation. Suitability of manuscripts, based on content and adherence to submission regula- tions will be decided by the Advisory Board of the Journal, The cost of printing and mailing of each issue is defrayed by institutional and individual sub- scriptions to the Journal of $50.00 Canadian funds ($40.00 US) per annum* and by page charges to the authors of $15.00 per printed page (including plates). For more detailed instructions and a sub- scription form, please see the inside back cover. Further information may be supplied on request. Publication. AMPHIPACIFICA is published by Am phipacifica Re search Publications, Registrar of Companies for the Province of British Columbia No. 0152988, 1993. Editorial Board. E. L. Bousfield, Managing Editor, Royal British Columbia Museum, Victoria. B, C, Canada. C. P. Staude, Associate Editor, Friday Harbor Laboratories, Friday Harbor, WA, USA. P. Lambert, Associate Editor, Royal British Columbia Mu- seum, Victoria* B. C„ Canada. Advisory Board. D. R. Calder, Royal Ontario Museum, Toronto, Ontario. D. E. McAllister* Canadian Museum of Nature, Ottawa, Ontario. Leo Margolis, Pacific Biological Station, Nanaimo* B. C. G. G. E. Scudder* University of British Columbia, Van- couver, B. C. C.-t. Shih, Canadian Museum of Nature, Ottawa, Ontario. Translation Services. Maijorie A. Bousfield, MSe* Montreal* Quebec. Sponsoring Agencies, Canadian Museum of Nature, Ottawa. Ontario, Canada. Friday Harbor Laboratories, University of Washington* Friday Harbor, WA, USA. Royal British Columbia Museum, Victoria, B. C., Canada, Royal Ontario Museum, Toronto* Ontario, Canada. Offices. (1) Subscriptions and Correspondence: Dr, E. L. Bousfield* Managing Editor, do Natural History Division* Royal B, C. Museum, Victoria, B. C„ Canada. V8V 1X4. (2) Mailing: Friday Harbor Laboratories, Friday Harbor, WA, USA 98250. Att: Dr. C. P. Staude. Authorization pending by the U. S. Postal Service for second class postage paid at Friday Harbor, WA., USA 98250. Registration. The journal AMPHIPACIFICA is registered at the National Library of Canada, Legal Deposit Office, 395 Wellington St, Ottawa, Canada, as ISSN Number 1 189-9905. Printing. Island Blue Print Co. Ltd., Victoria* B. C., Canada, AMPHIPACIFICA JOURNAL OF SYSTEMATIC BIOLOGY SUMMER ISSUE, 1994 AMPHIPACIFICA, Journal of Systematic Biology {ISSN No. 1 189-9905) is published quarterly by Ampldpadfica Research Publications, 611-545 Dallas Rd.. Victoria, B, C.. Canada V8V 1B3, Annual subscription rates are S40. US rands oar $50. Canadian Known office of Publication in the Untied Stales is Friday Harbor Laboratories. University of Washington. Friday Harbor, WA, h 98250-9218, USA, Application to mail ai Second-Class Postage Rates is pending at Friday Harbor, WA., alb Dr. Craig P. Staudc. Friday Harbor Laboratories, Friday Harbor. WA., 98250-9218. USA. DEDICATION The Journal AMPHIPACIFICA is dedicated to the promotion of syst- ematic biology and to the conservation of Earth's natural resources. Cover Design: Adapted from the rule page of S, J, Holmes (1904), "Amphipod Crustaceans of the Expedition, 1 ' Haniman Alaska Expedition, pages 233-244, From our Readers . , The second number of the new journal was mailed to subscribers in early June, and Uiis issue in October. We ack- nowledge slippage in planned publication dates, and our eff- orts to remove mechanical errors from the text have not yet been entirely successful. However. Ebe number of favour- able comments being received from the general readership concerning [he material and illustrations is encouraging. We are increasing the use of offers from colleagues for ref- eree services in their Gelds of specialization, and plan to in- clude t he ir eompreheusi ve re view ad idea on topi cs of aquatic biological concern. Continuing modification of the editorial terms of reference, outlined below, have expanded the scope of our service to both contributors and readership, Collabo- ration with the library exchange program of the Royal British Columbia Museum, and the steadily increasing numbers of new subscribers, have broadened the total to nearly 200 committed subscriptions, as of this date. Positive and helpful, commentary has been received from Jan Stock ( Amsterdam}. Niel Bruce (Copenhagen), Nina Tzvetkova (Leningrad), Horoshi Merino (Japan). Phil Gsbel (Chicago). Patsy McLaugb Jin ( Anacortes), Ann Quayle (Nanaimo), and several other colleagues An item in a recent issue of the Memorial University Alumnae “Gazetted by Carla Pomeroys concerns a new species of phosoeephaiid amptupod that, has been ranted Mandibufaphoxus mayi, Jarr- ett & Bousficld 1994 {see also p. 66 of this issue). In Pomeroy’ & article. Dr. May. President and Vice-Chancellor of the University, comments K Tve always wanted to spend my life on die beaches of British Columbia. 1 never made it, but no w P m there in name ! In all seriousness, w hen the La tin version of your name is used as the species designate for an animal or plant, you’re in quite good company, and we’re running out of animals to name. Usually this is reserved for those directly involved ir taxonomy". The editorial board of AMP HIPAC IFIC A encourages (he use of patronyms by contributors of manuscripts in recognizing those who ac- tively promote scientific investigation, and the conservation of the world's natural resources, A few readers have been concerned, understandably, about the possibility of competion for submissions between AMPHIPACIFICA and other journals of systematic biol- ogy. To a very limited extent, .such may be the case. However, AMPH1PACIFICA was designed initially to re- place former Canadian outlets for systematic biology -such as (he “Publications in . , T series {Canadian Museum of Nature, Ottawa), "Contribution s“ and "Technical Reports" (Royal British Columbia Museum), and to some extent the Canadian Journal of Zoology and die Journal of Fisheries and Aquatic Sciences (Ottawa). For many years, these journals had been publishing both long and short taxonomic papers, but recently these outlets have been discontinued and /or no longer accept long {>20 pp,) papers on taxonomy or systematic biology. AMPH (PACIFICA accepts mainly long papers on systematica (>30-40+ printed pp.) with em- phasis mainly on aquatic invertebrates, mainly of the North Paci fie region . Also, its page charges are modest* less than those of most other journals that do assess page charges, thereby hopefully stimulating publication by authors who may be without institutional backing or other major financial resources, and might otherwise be unable to publish good systematic work. In this issue we present further systematic and bio geo- graphical features of the North American Pacific fauna as revealed by the dexaminoidean amph ipods. Generic and species diversity of this group in tbe North Pacific is rela- tively low. hut remarkable in consisting mainly of very primitive and very advanced genera, with few phyietically intermediate representatives. In analyzing this fauna, in combination with that recorded elsewhere in the scattered literature, (he authors have been able to recognize, within the SLiperfamily, two dearly defined families, each with four morphologically and hiogeographteally distinctive sub- families, The North Pacific is the probable centre of origin and evolution of this relatively ancient and primitive group which has since spread mainly into the Indo-Pacific region, The more advanced members are now re -penetrating the North Pacific as commensals on lunteates, sponges, and other sessile colonial invertebrates, A second major paper outlines morphological and behavioural evidence for the development of new phyletic classificatary concepts within lire Amphipoda broadly. Am ph ipods may be grouped serni- phyletically and pragmatically into two main assemblages, the swimmers {"Natatitia'T, and the bottom crawlers r Rep lamia") based mainly on major differences in thdr reproductive morphology and life style. The authors recog- nize the subordinal distinctiveness and relatively primitive position of the small, hypogean. relict Ingnlfiellidea w.-a- viz the Ganunaridea, At the same tune, they provide evidence thai would derive the relatively advanced pelagic Hy peri idea from a common ancestor that is least far removed from the gammaridean superfamily Stegocephaloideai the benthic* clinging Caprellidea had earlier been derived, hy plhers. from podocerid members of superfamily Coruph- ioidea. Possible submergence of both the Hyperiidea and (he Caprellideaasirfraordmal taxa within SulwrderGammaridea therefore merits further consideration, Our editorial policy has been modified to accept review articles concerning aquatic biology, environmental protec- tion, and biological conservation, It has boon broadened to accept short articles, where publication of the new taxa is ur- gent, and where inclusion of the paper can be accommodated wilhin the total pagination of an issue at a required minimum of 140-150 pages, Ac cordin gly* we ha ve included here, on brief notice, a short paper by Kathleen Con Ian on new isacid ampbipods from off the coast of California (pp. 67*74), We conclude this editorial hy thanking the editors of "Deep Sea Research" and "The Canadian Field -Naturalist" for recent publicizing of AMPH1PACMCA in Those jour- nals. A notice concerning the history and aims of the Cam adian Field-Naturalist’' is provided in this issue (p. 75). AM PH IP AC IFIC A VOL. I NO. 3 OCTOBER 15, 1944 About Our Authors . . Jane Kendall, B.Sc., majored in biology, with minors in chemistry and mathematics, at Ottawa University. In 19S8 she commenced a 3 -year, part-time period of identification work on CMN collections of North Pacific amphipod crustaceans, with special attention to family groups within the Dexaminoideaand Lysianassoidea. She is currently a teacher of biology and mathematics at secondary schools of the Carleton Board of Education, She is married, with one child, and now resides in (he picturesque town of Burrin'* Rapids, Ontario, south of Ottawa, the nation’s capital. Kathleen E, ConJan, obtained an Honours B.Sc, in Biology at Queen's University, a Master's Degree in Marine Ecology at the University of Victoria, and a PhD in Systematic s and Evolution at Carleton University, Ottawa. Prom 1979 to 1989, Dr. Con lan worked with Dr. E. L. Bousfield, at the Canadian Museum of Nature, on systematic s of corophioidean amphipods. Since then she has expanded her research to ecology and behaviour. She has recently in it mated studies in Antarctica, the High Arctic, and the deep sea, both on questions concerning ampin pod*., and more broadly on benthic community response to anthropogenic disturbances. Chiang-tai Shih, PhD, is a Research Scientist (formerly Curator of Crustacea) at the Canadian Museum of Nature, Ottawa. Although broadly interested in aquatic biology, he specializes in the systematic* of crustaceans, with emphasis on planktonic amphipods and copepods, His research in biogeography of planktonic crustaceans led to de ve lopme nt of his t heory of speciatior in m arine wop} an klon that is termed planktopatric speeiation. Commentary on Dr. E. L. Bousfield w r as published in Amphipaciftea 1(1): 2, 1994. ANNOUNCEMENT: 7th Deep Sea Biology Symposium KNOSSOS ROYAL VILLAGE HOTEL CRETE Thursday, Sepi 2V - Tuesday, Oct J, im AM PH1P AC1F1C A VOL 1 NO. 3 OCTOBER 15, 1994 2 THE AMPHIPOI> SLPECKFAMILY DEXAM1NOIDEA ON THE NORTH AMERICAN PACIFIC COAST; FAMILIES AT YLIDAE AND DEXAM1NIDAE; SYSTEM ATf£& AND DISTRIBUTIONAL ECOLOGY* by E, L, B OHS Held 1 2 and J. A. Kendall* ABSTRACT Based mainly on study material from North American Pacific coastal marine waters (from the Bering Sea region to northern California h this study treats the systematic* and distributional ecology of regional family and subfamily members of the gammaridean amphipod SuperfatttSly Dexar inoidea. Cluster analysis of 22 component genera supports subdivision of this toxanoimcally clifficull group into two principal families* the primitive, thin bodied Atylidae, and the more advanced, broad bodied Dexsmunidac. Family Atylidae encompasses the subfamilies Atylinae Leacti IS 14 (revised status); Auatyltnae Bulycheva 1955 (revised status).. Lepechincllitiae Schellenfoerg.. 1926 (revised status); and NcslntmpUnae* new subfamily. Family Dcxamintdae here encompasses subfamily Dexgunininae Leach, 3 814 (revised status); Prophliantinae (Nieholls, revised Barnard. 1970): Dvxammociitmae, new subfamily, and PolydierUtiae, new subfamily. Material from the Study region contains representatives of both families and three component subfamilies Within subfamily Atylinae are newly described and figured AtylutgeorgkM us, new species and A bormtis, new species. Newly recorded from the study region and/or re figured are: Atylusatlasstivi (Guijanova, 1951), A. bruggeni (Gurjanovn, 1 93 S ), A. coltingi (Gmjanova, 1958), and A Uviden&us Barnard, 1954, A, tridens (Alderman. 1956), and the type species A, carmmusj, C, Fabridus, 1783. Rediagnosed and refigured from the western Pacific region arc Aiyttu ekrti&nl Gurjanova, 1938 and A. rylovi Bulycheva, 1952. Based on analysis of the literature and records from the western Pacific, subfamily Anatylmae here encompasses AnatyUts pffvtoVtifcii Bulycheva, 1955, an d Jfa meh a t\' las} upon icus (Nag a ta, 1961). Subfamily Nototropiinae contains the western Pacific Nototrvpis sp. (cf. gutmtv . f Costa) and the abyssal species, Ary/cts aberrantis (Barnard, 1962). here transferred from subfamily Lepediirel I inae in the monof ypic new genus AbmUykts. Within family Dexiiminidae. subfamily Folycheciinae is moderately well represented in amphi-Nonh Pacific waters. Newly described and figured are Pofycheria carinata, new species, and I 1 mhrffltie. new species, and Pohchma osborni Caiman, 1898 is redescribed. Subfamily Prophliantinae, is represented in Asiatic Pacific coastal waters by about a dozen described species, but in the North American study region, the sole species identified to date is Guemea retfwKQns fJ. L. Barnard, 1958). Subfamily Pexaitiininac is well represented in the western North Pacific by species of Paradexami/u* but is absent from (he eastern North Pacific, Biogeographically. within family Atylidae. members of the primitive subfamily Atylinae are endemic to the North Pacific region. From there, members of the more advanced Nototropiinae and AnatyUnae have apparently radiated into the Indo-Pacific and former lelhyan wain-water regions. Members of subfamily Lepechinellinae, Having a common ancestor with the Nototropiinae, have become abyssal. Nearly ail mem- bers of the more advanced family Dcxiuninidae arc warm -temperate and tropical bnta few species of the most advanced Subfamily, Polycheriinae, ha ve penetrated into the cold temperate North Pacific, apparently from two different wurces , The sole North American Pacific prophliantin species appears more closely related to counterparts in the North Atlantic region than to prophliantins of the western North Pacific. Estep! for (he abyssal Lepectiineilinae, mostdexaminoideans are littoral -subliuoral in depth range and all are exclusively marine, In life style, they mainly nestle on the bottom, in sea grass clumps* coral clusters, empty shells, or in pits excavated in the tests of colonial invertebrates, bu( dte Prophliantinae apparently burrow in soft sediments. Species densities are seldom highand total biomass is low, presumably with little significance in regional food energy cycles. The group may be considered a specialized relict offshoot of early gammarokfean ancestral types, from which may have also evolved the closely related, bul ecologically more successful, ampeliseoidean am ph ipods that are tube dwelling and deposit feeding in soft sediments, world-wide. 1 Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario KEP6F4 2 R. R W L Burrin' s Rapids, Ontario KOG 1B0 AMFHJFACIFECA VOL. I NO. 3 OCTOBER 15. 1994 3 INTRODUCTION The superfainily Dexaminoidea encompasses a group of benthic nestling ampbipoda that occur mainly on hand sab’ strata in tropical and warm lemperate regions of the world. The number of described species is relalivley small (< 200), and populations are generally of tow density. However, morphological diversity within the group is relatively high icf. Gammaroiden with 5 times the number of species), possibly reflecting the wide variety of solid (and some sedimentary) substrata on which various subgroups of Dexaminoidea have been modified tor existence. The animate arc characterized by variously carinated or dorsal Jy processiferous bodies, fused usosome .segments 2 &. 3, a tendency to piehen&ility (subcheJation)of peracopods 3- 7, and a reproductive life style- that involves mating freely in the water column. The gnathopods are weakly (or not) sexually dimorphic, but in the type genus Dexamine and dose relatives, the anterior margin ol the propod of gnathopod 1 tin the male) bears a characteristic notch or sharp excava- tion, of presumed (but unknown) reproductive function. Although basically free -living, with well developed perjeopodst pleopods, and (ail fan, the animals am typically slow -moving, even sedentary, in vegetative life style, Most species are deposit or trypton feeders, frequently employing specialized setae of the antennae and anterior peraeopods to rake in organic food material, from a nearly fixed position on the bottom. Members of the specialized genus Polycheria are commensal on the tests of colonial tunicates and sponges where they live ''upside down" in pits excavated in the surface of the host. However, unlike the closely related ampeliscoideans, of similar "upside-down" feeding style, dexamlnoldeans lack spinning glands in the anterior peraeopods and arc incapable of tube building. The classification of the group within suborder Gammaridea has gone through an early period of .stability, followed by a recent period of relative instability, Early workers (e.g Sars,1895; Stubbing, 1908) maintained the aty lids and dexaminids as separate families, and recognized the close similarity of alylids to the basic Gammandae' 1 amphipod type. Such stability was extended to the Lepech- ineltidae (Stebbing, 1 90S) and the ProphlianLidae CNichoils, 1939) by more recent major workers (e,g, Guijanova ? 1951 ; Barnard, 1969a), Soon afterwards, however, a developing trend to fusion of related higher taxis, led to submergence of al I dexamin idgroups within fan* it y Dexami nidae (e,g. Bel lan- San tuii, 19S2) or to formal creation of a new superfamily group, Dexaminoidea, in which family levels of distinction could be maintained (e.g. Bousiield, 1979, 1982). Dexaminoidean ampb ipods are among the few regional North Pacific garnmaridean groups that have received sig- nificant taxonomic attention. Within the Dex ami nidae proper, Polycheria osbowi was described from California by Calmand 898). Within family Atylidae, the genus Atylus had been unknown from the Pacific coast of North America prior to Alderman 1 * (1930) description of A. iridens from Califom ia. That record was c loscly followed by Gubanov a' s (1938) description of A collingi from eastern Siberia 3 Eld Bering sea regions, and by I, L, Barnard' s( 1956) deseripl ion of 4, fevidensus from California. Based on CMN material collected along the Pacific coast of Canada 1955 - 1959, Mills (1962) provided Ulus hated descriptions, keys, and distributional data on those three species from the coastal marine region of British Columbia. Further records from California were added by Barnard (1962, 1969 b andCadien (1991), Barnard { 1 975), and Stands ( 1987) included dex- aminkis in keyed and illustrated popular regional works and Austin ( 1985) summarized records from the cold temperate nor (beast Pacific region . Within family Prophliantidae, Gtterriea (Ptimssu^} redmems Barnard had been recorded widely along Californian coasts by Barnard (1958, 1969b, 1972) and Cadicn (1991), The biology of Polycheria osborni was studied in detail by Skogsberg & Vansell (1928), and some members of the Lepechinellidae were recorded from the eastern Pacific abyss by Barnard ( 1967, 1972). In tlie western North Pacific, the early work ofGinjanova ( 1938). and Stepheusen (1944) was encompassed by Gurf anova (1951), Subsequently Bulycheva (1952, 1955) pro- posed further species of Atylus, Polydie ria and Afiatylu^ and Biistein & Vinogradov (1955) recorded an abyssal I epoch in ell id. From Japanese waters Nagata (1961) de- scribed the aberrant Stylus japoniats. The more recent work of Game (1981) on Lepechwdki, of Hirayama (1984a, b, 1984, 1986) on species of Pamdexamine, Polyctieno , Guernea h and Atylus, and Ishimaru (1987) on Guenwa, and others, has been summarized most usefully in a catalogue of riexaminid amphipods of Japan by Ishimaru ( 1994), Dexaminid systematica and distribution have been treated comprehensively and most usefully by Barnard & Karaman (1991). However* the lumping of diverse sub- groups within one family, with recognition of only one additional subfamily and no superfamilies, and the use of loo few. or phyleticaJJy non significant, character states In diag- noses and keys, tends to create problems of iu consistency in taxonomic analysis and an unwieldiness of classification that may also apply elsewhere within gammaridean classifi- cation (e.g. within family Eusiridae). Our purpose here is to ( I ) develop new basic taxonomic information and analytical criteria from a study of the present North Pacific material; (2) incorporate this information with previous knowledge as a basis for numerical analysis of natural relationships be tween higher taxonomic categories, and (3) modify existing, classi- fications in a manner that more con si stent! y reflects diserthu - lionaJ, ecological* and behavioural, as well as taxonomic and phylede, differences between the subgroups. The authors have recently examined extensive new material in the amphipod collection of the Canadian Mu- seum of Nature (CMN), Ottawa, that supplements the earlier material of Mills (196 i), and material from (he Bering Sea region (Peter Slattery expeditions) and elsewhere. ■Station lists for CMN museum Enaterial, 1955 - 1980, are provided AMPHEFACU’ICA VOL 1 NO. 3 OCTOBER 15, IW4 4 by Milts (mi\ Bousfield (1 958, \%X 3 %$\ Bousiield and McAllister (1963), find Bousfield and Jarre tt (1981). This report provides an extensive review of the system alias, distribution al ecology of the dexiiininoideiin fauna from the North American Pacific coastal marine region and relates ft phyletically and biogetJgraphically to counterpart faunas of the western North Pacific and elsewhere in the world. ACKNOWLEDGEMENTS This study could not have been completed without [he help of many biological research agencies and interested colleagues, The field surveys were carried out with the full support of the National Museum of Natural Sciences (now Canadian Museum of Nature) in Ottawa, mid received vital technical assistance and ship-time from Canadian Pacific research centres and their staffs. These included the Pacific Biological Station, the Bamfield Marine Station, the Pacific Environmental Institute, the Royal British Columbia Mu- seum, the University of Victoria. University of British Co- lumbia, and the Institute of Ocean Sciences, Sidney, and. in the United States of America, the Friday Harbor Marine Laboratory and the College of Fisheries* University of Wash- ington. We are particularly indebted to Or. Peter Slattery, Moss Landing, CA.. for provision of material from the Bering sea region. Full acknowledgement to individuals of those agencies,, and to many others, are provided in the previously published station lists (above ), to whom we again express our deepest appreciation. W r e are grateful for the splendid published wort of our colleagues elsewhere on whoe illustrations we have drawn freely in developmentot' taxonomic and phyletie relationships. We thank especially Museum colleagues Ed Hendryek* and Judith C. Price for curatorial and cataloguing assistance . Preparation of the line illustrations was most capably assisted by Susan Laurie - B ourque, H ull, Quebec . We are grateful to Dr.. J , D . Thomas, USNM, for review of the text and kind pemtission to adapt illustrations from the pertinent research publications of the talc J, L, Barnard. Work by the senior author, and by the artist, was also Supported by operational grants from the Royal Ontario Museum, Toronto, and the Natural Sciences and Engineering Research Council, Ottawa. DEX A M IN OTDH A Leach { revised Bousfield) Pexaminoidca: Bousfield, 1979: 350. — Bousfield. 1982: 277.— Bousfield, I9S3: 2.63,— Sttam. 1986: 180. Dexaminidae Barnard, 1970; 161. -tabmtarite 1987: 141.2. — Barnard & Kanaman, 1991 : 260. Families; 1. Atylidae G, O. Sars, 1882; 26. Includes subfamilies Atylinae Sara (revised status): LcpechindJmae Scheilenberg 1926 (revised status); Nototropiinae, new sub- family: and An atylinae Bulycheva, 1955 (revised status), 2. Dexaminidae Leach, 1813/14:432. Includes subfamilies Dexamininae l-each (revised status); Polycheriinae, new subfamily: Dexamtnoculimae, new subfamily; and Pro- phliantinae Nicholls. 1939 (revised status, Barnard J970), Diagnosis (after Bousfield, 1982); Rostrum present, variable. Body (especially urosome) usually with mid- dorsal, and often dor stv lateral, carinations or teeth. Urosnme segment I dorsal ly carinate. Urosome segments 2 & 3 coalesced, often dorsally carinate, Sexual dimorphism pro- nounced in eyes* antennae, uropod 3, and coxal gilts, but weakly expressed in gnathopod 1, Eyes pigmented and multi -faceted, lacking in bathyal forms, Antennae variable; peduncles of flagella (male) armed with brush setae. An- tenna L peduncular segment 2 usually longer than 1; seg- ment 3 short; accessory flagellum minute or lacking. An- tenna 2 trending to shortening (female); flagellum elongate, nofi-calccolate (male), Mouthparts trending to modification. Upper lip entire. Lower lip, inner lobes variously developed. Mandibular molar tritunuive, but trending to reduction; leftlacinia basi- cally 5 ^dentate; palp various, weak or lacking. Maxilla l, inner plate 0-8 setose, outer ptale 7- M spinose; palp often I -segmented. Maxilla 2, inner plate the smaller, trending to loss of marginal setae. Maxilliped, outer plate large, inner plate and palp trending to reduction in size and loss of setae. Coxal plates 1-4 medium lo small, often notched or incised be low; coxa 5 strong, often a nJefo-lobafe. Guafeo pods small, weakly subchdate (palms convex), generally dissimilar in form. Gnathopod 1, pnoptxJ may be distinc- tively sexually dimorphic. Peraeopods 3 & 4 subequal, or peraeopod 4 smaller, trending to shortening of segment 5, Peraeopods 5-7 var- iable in form and size, bases unequally broad, trending to linearity; segment 5 variable; segment 6 and dactyl trend- ing to subchelation, Pleopods usually strong, especial ty inmate. Uropods I & 2, rami unequal, lanceolate, apically spinose, Uropod 3 aeqniramous; rami lanceolate, outer ramus 1 -segmented margins Setose in male, often so in female. Telson bilobaie. lobes variously fused basally, apices spinose, notched or finely crenulaied. Coxa t gi! I s sac- 1 i ke, on peraeopods 2-7(6) , often pleated or phyllofortn, especially in male. Brood lamellae medium broad or strap- 1 ike , trending to linearity. Reproductive Life Style; synchronous, mating freely in water column (presumed from morphology - nearly all members). Taxonomic and Biogtographk Commentary: Cluster analysis of all 22 generic-level laxa within superfamily Dexaminoidea recognixedJteft (p. 56 ) supports the validity of the family and subfamily components listed above, and detailed in (he following systematic accounts. The most primitive subfamily* Atylinae is endemic to Ehe twreaJ- su paretic North Pacific; others are components of mainly tndo-Paci fie faunas marginally present in this region. AMPHTPACIFICA VOL I NO 3 OCTOBER 15.1994 5 (a) ATYL1NAE (to) NQTQTRQPIINAE (c) LEPECHINELUNAE X? (d) ANATYL1NAE -0 , I FIG. 1. ATYLJDAE: TYPICAL CHARACTER STATES A - Rostrum; R - Urosome 1; C -Antenna 1; D-Coxa 1 ; E- Mandible; F - Gnathopod 1; G - Pcraeopotls 3*4; ft- Peraeopods 5-7; ^ - telson ; K - coxal gills 2 - 5 ; (from test plates) AMPHIPACIFICA VOL. I NO. 3 OCTOBER 15, 1994 ft (a) DEXAMININAE (b) DEXAMINOCULINAE (d) PROPHLIANTINAE (c) POLYCHERIINAE FIG. 2, DEXAMINIDAE: TYPICAL CHARACTER STATES A- Rostrum; B - Antenna 1; C -Coxa 1-4 : D ' Gnathopod 1 (male); £-peraeopods 3-4; F ■ peraeopods(F-7 bases) G ■ peraeopads5-7 (distal); H ■ abdomen dorsum; J - Pleon plates 2-3; K- telson (From test plates) AMPH1PACIFICA VOL. I NO. 3 OCTOBER 15, 199* key to world families of DEXAMINOIDEA I. Body slender: antennae medium-long: antenna 2 not reduced (female); mandible usually with palp; maxilla L palp 2-seg mealed; maxilliped almost normal; coxa 5 shallow, strongly an Eerodobate. Alylidae. —Body short, broad; antenna often short, A2 shortened (female): mandible lacking palp; maxilla 1, palp l- segmented; maxilliped palp distinctly reduced; coxa 5, broad, aequilobatc. usually deep . . Itexyminidae, Atylidae G. O, Sars Atylidae: G. O, Sars, 1 SS2: 26, — Stebbing, 1906: 327— Barnard, f%9a: 161— Bouslleld. 1982: 277. Anatylidae: Bulycheva, 1955: 204 — Bousl'idd, 1982: 277 Dcxaminidae (part); Barnard. 1970a: 164, — Bellau-Santinj. 1983: 212. — Barnard & Karaimin, 199] : 260, Type Genus: Asyius Leach. 18L5: 21, (Type A. carin- atus Fabricius 1793), Diagnosis: Body slender, laterally compressed, Poste- rior peraeofl&cdpleon var-iously carinate or smooth. Urosome 1 mid-dursallv carinate, Rostrum various, usually medium to strong Antennae not shod, antenna 2 the longer. Antenna I, peduncular segment 2 subequal to segment 1, often longer; accessory flagellum present minute. Lower lip, inner lobes usually lacking. Mandible with palp (few exceptions), trim rating molar, 5-dentate left larinia, and several blades in spine row, Maxilla 1. palp 2-seg- mented. Maxdla 2 normal, margins setose. Max illiped inner plaies normal, apex spinose; palp strong. Coxae 1-4 various, lower margins may be acute, occa- sionally incised, 2 & 3 deepest, 1 less deep. Gnathopods I & 2 weakly subchelate, weakly sexually dimorphic: caspus various, often slender. Pfcraeopods 3 A 4 + segment 5 usually much shorter than segments 4 & 6 and dactyls not elongate (except in Lepeehinellinae). Peraeopod s 5-7 not elongate, bases vari- ously expanded and lobate below, somewhat dissimilar in form; segmen I 5 various , Pleopuds various, usually strong. Pleon plates 1-3, hind corners squared or acuminate. Utopod 3, turn! lanceolate, margins setose (esp, male) or spinose. Telsoa lobes normal, short to medium, fused basally. Coxal gills often pleated or plaited. Brood plates broad, Taxonomic Remarks; The family Atylidae is here subcii vided into 4 subfamilies as diagnosed be! ow , They arc separated on character slates of the key (below) for which illustrations are provided in Fig. 1. and in pertinent sections of [he text. Subfamily itylirme Boeck (revised status] Atylinae Boeck, 1876: 320. Atylidae Stebbing, 1 906: 327. — Gurjanova, 1951:678. — Barnard. 1969: 163. Dcxaminidae (part) Bdl&n-Santini 3 982; 2 12. — Barnard & Katanian, 1991: 260, Type germs: Atylus Leach, 1815. Diagnosis: Generally medium to large atyiids (5-40 mm). Rostrum usually large. Posterior peraeom pleon, and KEY TO SUBFAMILIES OF ATYLIDAE 1. Peraeopods 3 and 4 closely subequal in size; peraeopods 5-7, segment 5 distinctly shorter than segment 4 &. 6: antenna 1 , peduncular segment I shorter than peduncular segment 2 (always, in female) Atylinae (p, 8) — Peraeopod 4 distinctly shorter than peraeopod 3 (in distal segments); peraeopods 5-7 segment 5 not dis- tinctly shorter than segments 4 & 5; antenna 1 peduncular segment 2 vamusly longer than segment l . 2, 2. Eyes lacking: anterior head lobe strongly bifid; peraeopods and daciyls very slender, elongate; teisnn lobe* short, diverging distaLly Leperfimellinae (p. 3 1 ) — Eyes present: anterior head lobe blunt or slightly emarginate; pemeopods and dactyls normal length and thickness; telson lobes normal, converging distal ly 3. 3. Picon segments 1-3 not carinate; tiresome \ with simple mid-dorsal tooth; mandible lacking palp; uro- pod 3, rami short, margins spinose; peraeopod 5, basis mil lobate below ; gills simple Anatylinae ip. 32) — Pleon segment 1-3 usually carinate mid-dof sally; urosome I wilh tooth and notch; mandible with palp (weak); uropod 3, rami lanceolate, margins setose (esp, male); peraeopod 5, basis with lower hind lobe; gills phylliform Nototmptirute (p 28) AMPHIPAClFlCA VOL. [ NO. 3 OCTOBER 1 5,. 1994 ft urosome usually dfimlly carinate. Antennae large, setose. Mouthparts basic. Mandibular molar, spine row, and palp well developed, Manilla Rinnerplate separate- Really setose, Max ilia 2, plate margins setose. MaxiBiped normal, plates and palp well developed. Coxal plates 1 -4 medium, lower margin* often weakly incised or subacute anter iorly. Gnaihopods medium, suhsiuiiiar, weakly sexually dimorphic. Peraeopods 3 & 4 subsitnilax in form and si?£; segment 5, distinctly shorter than segments 4 &. 6. Peraeopods 5-7, bases broadened; .segment 5 variously shorter than segments 4 & 6. Peraeopod 5. basis, hind lobe weak, not produced below, Peraeopod 7, basis very broad, posterior lobe present, acme or rounded below. Plcopods strong. Unopod 3, rami strong, tftargins spmose and usually setose (both sexes), Tetson lobe* medium to large, with apical spine(x). Anterior coxal gills usually pleated especially in males, Taxonomic and Distributional Commentary: The subfamily Atylinae presently con la ins a single genus, Ary/wr „ encompassing about a dozen species, almost all endemic to the boreal and .subarctic North Pacific region. The range of morphological variation ts sufficiently great that recognition of internal groupings (e.g. the collingi subgroup) may ever- lually justify subgeneric recognition. The princiapl features of subfamily Atylinae are eontgrasted with [hose of other subfamilies of Atylidae in Figure 1, In summary; (referred to in following text, where pertinent). Aiylus: Leach, I8i5 Atylus Leach, 1815; 2! .—Mills, 1961: 17 (key).— Barnard, 1956:38,— J.L, Barnard, 1969a: 163,— J.L, Barnard 1970a: 164.— Barnard & Karaman, 1991; 262 (part). Noiotroph Gurjanova, 1951: 680 + key. (part) non Anatylus Bulycheva, 1955: 205. -Tzvetkova, 1967: 391. non K am^lmtvtus Barnard. 1970b: 93. Type Species: Ccuntmrus catinalus J. C. Fabricius 1 793, monotypy. Species: A , dilmsovi (Gurjanova, 1951); A, borealis, new spec i A. bmggeni (Gurjanova, 1938); A. collingi (Gurjanova, 1938); A. ekmani (Gurjanova, 1938); A. georgiamts, new species; A ftvidettsus (J. L, Barnard, 1956): A. rylovi Bulycheva, 1952; A. widens (Alderman 1936); A. vittosus Bate 1862, (A. oriental!* Hirayatna, 19861- Diagnosis: Large atylids ( 10-30+ mm). Rostrum me- dium to large, Anterior head margin rounded, rarely bifid. Antennae moderately strongly sexually dimorphic. Ant- enna L peduncular segment 2 not longer than 1; accessory flagellum minutely I - segmented, Antenna 2, peduncular Segments 4 & 5 strong, often setose. Lower lip, inner lobes weaker lacking. Mandible: molar triturative; palp normal. 3-segmented, Maxilla 1, inner plate with 4-8 apical setae, Maxilla 2, inner plate with 1-8 proximal plumose marginal setae, Maxilliped, palp normal, 4-segmeuted. Coxae 1 4 medium deep, smooth or subacute below; coxa 3 antero-dis tally deepest. Coxa 5, anterior lobe broadly or sharply rounded below. Gnathopods 1 Sc 2 ordinary, very w r eakly or not sexually dimorphic; propod & carpus medium, usually subsimilar in length. Peraeopods 3-7 not elongate, dactyls relatively short. Peraeopods 3 & 4 , segment 5 markedly shorter than 4 & 6. Peraeopod 5 distinctly smaller than peraeopods 6 Sc 7; basis with small posiero-dislai lobe, Peraeopod 7, basis broad, po*tcro -distal lobe present, rounded or acute below. Peraeopods 5-7, segment 5 markedly shorter than segments 4 & 6, Pleopods regular plcon plates rounded below and be- hind, Urapod 2 short, ram i uneq ual , Uropod 3 strung, rami lanceolate, margins setose in male, setose and/or spirtose in female, Tel sort lobes not elongate, fused basal ly, not diverging dUtally. Coxal gill* sac-like, weakly to moderately pleated. Brood plates medium hruad, not slender. Variables: Rostrum long t type), medium (A, coliifigi. A, ge&rgiamisy, posterior peraeonites carinate (type), smooth (A. borealis, A. tyfovi A. trident); pleon carinate (type), smooth (A. hornets, A , ryiovi, A. indensK gnathopod 2, propod & carpus short, stout (type), slender (A, brtiggeni, A. ekmam. A, vittosus}: peraeopods 5 - 7, segment 5 only slightly shorter than segments 4 & 6 (A, vittosus). Taxonomic Cwnmtntary: Some species of the genus Atylus, as here define , overlap in some character slater with some specie* of Motor ropis, as def ined below (p.28), How- ever, the two genera are distinguished by the characters of the subfamily key (p. 8) and, in combination, by (be larger rostrum, heavier mandibular palp, the weak (or lacking) hind lobe of die basis of peraeopod 5, heavier uropod 3, and the pleated, raiher than phyllofonn (or dendritic) anterior coxal gills, especially in the male, Distributional Commentary: Most species are con- fined to subarctic and boreal coastal marine waters of the North Pactflcregion, and are mainly benthic. Arylus caritteaus is holarctk, bul A. viUosus has been recorded only front the southern oceans and may not be a natural member of the genus. Members of the genus Aiytus ( sens, sir , I are virtually non-overlapping distributionaily with members of ih&genu^ Noioiropls , as here defined. AMPHIPAC1FICA VOI., J NO. + OCTOUKR 15, L£M 9 kf.y to north pacific spkcif.s of atylus* (Characler stales illustrated mainly in Fig, I. p, 6) L Peraeon segments 6 & 7 and pleon mid dorsal] y toothed or carinated; urosome segment S with single (or bifid) mid-dorsal tooth; coxa 4 various, not crescent shaped posteriorly 2 . — All segments of peraeon and pleon mid-dorsaHy smooth; urosome segment ] with single mid-dorsal tooth and preceding notch; coxa 4 crescent- shaped posiero-venirally 8, 2. Rostrum large (1/2 head length); uropod 3 large, rami longer than twice peduncle, margins with spines and setae; telson lobes elongate. 1 1/2 - 2 X basal width . . . . 3. — Rostrum medium, 1/4- 1/3 head length; uropod 3 medium, rami shorter than twice peduncle, margins spinose; telson lobes short, length = width . . . .. . 7. 3. Gnathopod 2. propod and carpus stout, depth of each > 1/2 length: perafciipod 7, basts, posterior lobe targe, scute below; fused urosome segments 2 h 3 with low mid -dorsal carinaUon; mandihle, palp stout, segment 3 setose 4. — Gnathopod 2. propod and carpus slender, depth < 1/2 length; peraeopod 7. posterior lobe small, shallowly rounded below; fused urosome segments 2 & 3, dorsal process erect, projecting well above mid line * . 5, 4. Peraeopods 3 & 4 & peraeopod 6, segment 5 short, length < 1/2 segment 4; gnathopod 2, propod stout, length > carpus .... A. allassori (p. 11) — Peraeopods 3 & 4, and peraeopod 6, segment 5 medium, length > 1/2 segment 4; gnathopod 2, propod small, length < carpus A. cttrinatus (p. 11) 5. Eyes large; fused urosome 2 & 3 with bifid mid-dorsal tooth; uropod 3, margins of rami with spines and setae; peraeopod 7, basal lobe subacute A. bmggeni (p, 14) — Eyes small: fused urosome 2 & 3 with single mid-dorsal tooth; uropod 3 (female), ramal margins spin- ose; peraeopod 7, basal lobe rounded below 6. 6. Pleon segment 3 and urosome segment 1 with bifid mid-dorsal tooth; western Pacific, Sea of Japan , . , A, ekmuni (p. 16) — Pleon segment 3 and urosome J with single mid-dorsaJ tooth; endemic to the North American Pacific COdsI , . . .4. levidensux (p. 16) 7. Coxa 4 acute below: fused urosome 2 & 3 with low mid-dorsal ridge; gnathopod 1, propod, postero- disial angle with 5-6 transverse row of stout Spines . A, coUingi (p. 24) — Coxa 4 rounded beJow r ; urosome segments 2 & 3, with erect mid-dorsal tooth; gnathopod 1, postero- dorsai. angle of propod with 3 transverse rows of spines A, georgiamts (p. 26) 8. Peraeopod 6, segment 5 short {< 1/4 segment 4); telson large, length » width 9. - -Peraeopod 6, segment medium C- 1/2 segment 4); telson short, basal width about equal to length A , ryfavi (p, 18) 9. Antennae long, flagella with more than 20 segments; uropod 3 (female), apices of rami acute; telson elongate, length 1.5 X width A. tridens (p. 20) — Antennae shorter, flagella with fewer lhan 20 segments; uropod 3 (female) apices of rami rounded; telson medium, length 1.3 X width A. borealis (p. 22) * Aiylus orienmth Hirayama not included AMPHIPACIFKA VOL I NO. 3 OCTOBER 15. 1994 \Q Atytus carinatus ( Fabric! u s ) (Fig. 3} Gttmmarus carkiaius J. C, Fabricius, 1793: SI, 1 ', Atyhts ctttinafus Sars, 1895: 47 1, pi. 166, — Stubbing* 1906: 328,— Sh-remaker. 3920: 14 E. — Shoemaker, 1955: 45 — Gurjanova, 1951:679. — Dunbad 1954; 762.— Barnaul 1975, fig. 61. — Barnard & Karaman. 1991: 262. Material Examined: North-west Territories; Slidre fiord. Eliesmerc I T , Ait lie Biot. Sta., FRB, Canada, July 25, 1962 - 3 male (22,0 turn) (slide mount}: 1 female ov (28,0 mm) (slide mount). Many specimens in CMN Canadian arctic collections: none from immediate study region. Diagnosis Female (25.0-30,0 mm): Body large, laterally compressed. Per aeon and pi eon segments with mid-dorsal ridge, increasingly elevated as carinaiions on peraeon seg- ments 5-7, pleon 1-3, and urosome segment l. Fused uiosome segments l and 2 with low mid-dorsal and paired dom-lateral ridges. Head: rostrum large (> I /2head length);, anterior head Jobe bluitu, slightly cmarginate. Eye small, Antenna 1, peduncular segments I & 2 subequal in length, setose posteriorly; accessory flagellum small, 1 ■ segmented. Antenna 2 slightly the longer, peduncular segments heavily setose posteriorly. Lower lip lacking inner lobes. Mandible: molar large, Strong; spine tow with 5-7 blades: left lacinia 5 -dentate; palp normal, strong. Maxilla I, inner plate with 7 apical setae: palp large, Segmented, Maxilla 2, inner pi ate with 1 ^ stouL plumose inner marginal setae. Maxilliped normal* palp strong. Coxal plates 1-4 medium' deep; coxa l shortest, directed anteriorly; coxa 3, lower margin anteriorly subacute. Coxa 5, anterior lobe broadly rounded. Gnathopod* l & 2, very weakly sexually dimorphic; carpus and propod relatively short deep, suhequal in length, Peraeopods 3 & 4, segment 5 shorter than segment 4 & 6* dactyls stout, Peraeopods 5-7 (especially bases) somewhat dissimilar in form and sri.e; segment 5 slightly shone r than corresponding segments 4 & 6. Peraeopods 5 k 6. lower hind low very small, not produced. Peraeopod 7 t basis broad, poslero distal lobe rounded. Pleon plates i O broad, hind corners squared, Uropod 1 , rami lanceolate, subequal , uropod 2, outer ramus markedly shorter than inner. Uropod 3* rami narrowly lanceolate, > 3X peduncle; margins setose in female and male. Telson lobes deeply separated, not diverging* each with apical and subapical spines, Coxal gills large, sac-like, on peraeopods 2-7, anterior gills (male) weakly pleated basally. Distributional Ecology: Holarctic, in North America south to the Saguenay fiord in the east* and northern Bering Scat Kotzebue 5 on nd) a n the west (5 hoemak er, 1 955 ) mainly in shallow coastal waters (0*50 mk along mixed stony and silty shores. The species has. been recorded from the stomach contents of various arctic shallow- water fishes, and from eider ducks, and bearded and ringed seals (Dunbar, 1954). Taxonomic Commentary, An 1 tar carinatus is a distint live species of the genus that exhibits several plesiomorphic character states. These Include the accessory fiage Hum, strongly carinated dorsum of the posterior thoracic and abdominal 1 segments, and the strongly marginally setose rami of uropod 3 {both sexes), As the type of the genus Atvtax. a is combination of character slates separate it at gen us level from AnatyluS' pavkrvi Bulycheva, 1955, and from Nomropis smitii (Goes, 1866) with which A, airimtm overlaps distributional ly in high arctic and subarctic waters, A tytux atlas so Vi (Gurj anova) {Fig. 4) Nototwpis atlassovi Gubanova, 1951: 690. figs. 77A, B. Afyias fitfassovi Mills, 1961 : 19 (key only), — Barnard & Karamam 1991:262. Mute rial Examined: Bering Sea region: Amchitka Is* land, Constantine Harbor, C, E. O'Clair col I „ Qct. 5, 1969 - July 14, 1970: 4 lots with 15 specimens, including males* females, and immature*; female ov, (20 mm) (slide mount) fig'd. Si. Matthew Island, Walrus Cove* P. Slattery coll., June 29, 1983,- E male (28.0 tmnKfig.'d) CMN collections. Diagnosis. FemaEe (20.0 mm): Body large com- pressed. Per aeon segments each with shallowlly indented mid -dorsal ridge, slightly elevated to weak carnations on segments 6 & 7, Mid-dorsal carination weak on pleon segments 1-3, strong on urosome 1. medium on fused urosome segments 2 & 3, Head: rostrum targe (> 1/2 head length); anterior head lobe broad, shallowly iinarginate. Eye me- diu m , ve rti tally ovale . Antenna I , peduncular segment 2 not longer than I ; hind margin thickly short-setose; peduncle 3 short; accessory flagellum minute, Antenna 2, peduncular segments 4 & 5 stout* anterioi and posterior margins setose. Lower lip lacking inner lobes. Mandible; molar strong; spine row with 8-10 blades and accessory setae; left larinia 5 -dentate; palp stouh setose. Maxilla I, inner plate with 10- 12 apical setae; palp stout, 2 -segmented. Maxilla 2, inner plate with several inner marginal plumose setae. Maxi lliped stout, palp segment 2 short. Coxal plates 1 -4 relatively broad , lower margins nearly straight; coxa l about as deep as 2, weakly directed forward. Coxa 5, anterior lobe acute below, Gnathopod* 1 & 2 stout, 2 larger, moderately sexually dimorphic; propods relatively large, deep; carpus deep, shorter than respective propod. Gnathopod L propod with single distal now of pectinate setae; posterodislal angle with 3 rows (4-5 in male) of stout clasping spines. Gnathopod 2. propod* postero-distal angle with 2 rows (3 in male) of stout spines. Peraeopods 3 & 4 stout, margins spinose; segment 5 small, much shorter than segments 4 & 6; dactyls short. Peraeopod* 5-7, not markedly dissimilar; segment 5 much AMFHIFACIFICA VOL l NO. 3 OCTOBER 15, 1994 H PIG. 3. Atylus carinatus (Fabi\)« Female (28 A) mm), Male (22.0 mm) Slidre Fiord, Ellesmere I. A*4FH1FACIFICA VOL I NO. 3 OCTOBER 15. 1994 12 FIG. 4, Atylus atlassovi (Gurjanoval Female hr. Ill <20,0 mm) Constantine Harbor, Amchitka L Male ( (27*0 mm), St. Matthew L, Bering Sea, AMPHIPACIHCA VOL I NO. 3 OCTOBER 15, 1994 13 shorter than segments 4 &. 6; peraeopod 5, basis with weak lower hind cusp. Peraeopod 7, basrs. broad, lower TitncL lobe acute, PI con plates 1 -3 broad, hind corners acuminate. Uropod 3, rami large, broadly lanceolate. margins bluntly rounded and spinose apically t female); rami larger, nar rowly lanceo- late, inner margins setose and spinose. Tclson lobes, medium, fused basal ly, narrowing distally, apices nearly hare. Coxal gills sac- like, moderately pleated in males, on peraeon segments 2-7. Distributional Ecology: Sea of Okhotsk (Kamchatka !>CLiinsula) to Bering Sea and Sea of Okhotsk in subtidal shallows. Taxonomic commentary: Aiytun atlassovi is a distinc- tive but relatively primitive species. It clusters most closely with A, carfnatus, at less than 75% similarity level (p. 60). Atylus bruggetti (Gurjanova) (Fig, 5). Nolot ropis bmggeni Guijanova, 1938: 325, figs. 36, 37 — Gurjanova, 1951: 680, fig, 475, Atylus hruggeni Mills. 196 hOcey only), — Barnard & Kara ^ man, 1991: 263, Material Examined: BERING SEA; St Lawrence L, F. Slattery coll.. July 10, 1.980 * I male i female. Ibid. June. 1983 - 1 male. 1 female (hr. Ik 10 im. Paouk (, gravel. 5 m„ P, Slattery coll,, June 6, 1983, 3 lots - male (J 4.5 mm) (slide mt,); remale hr It (16 0 mm) (slide ml.); 200+ specimens including many males and some females ov. Si, Matthew L Walrus Cove, P. Slattery, June 27/83 - 3 lots (8 spins ), Ihad B ig B ite Bay , J nne 1 5/86 - male ( 1 5,0 tn m (slide ml,); female .with young (19,0 mm ) +40 other sped - mens, including mature males, females with hrood young. Pripilof L, St Paul 1., D, B. Quayle coll. Nov. 21. 1965 - 11 specimens, ALASKA MAINLAND; Oil ley Cape, 2 ft in depth, P. Slatlery call., June 24, 1984-4 specimens. Off Wat n wright, June 22/84 - 1 1 specimens, ALEUTIAN ISLANDS: Amchitka L Constantine Harbor. C, E.O’Gaircoli April 26. 1969- 1 male (22.0 mm > (slide mounrV Thld. Sept 27. 1969 ■ I male. 1 female. CMN col- lections. Diagnosis: Male (15.0mm). female (1 0,0 mm): Body large, strongly compressed,. Pefaeao and plcon with mid’ dorsal ridge, elevated to medium strong carina on peraeon segments 6 & 7, and pleoti segments 1-3. Urosome seg- ments 1, and fused 2 & 3, each with bifid mid-dorsal caiina. posterior tooth much die stronger.. Head; rostrum large (» 1/2 head length); anterior head lobe shallow, excised below eye. Eyes large, oval, larger in male. Antennae slender. Antenna 1 relatively short, peduncular segment 2 shorter than 1 (female), subequal (male), margins weakly setose (brush setae in male); segment 3 short, accessory flagellum mi nuic . An tenna 2. peduncular seginen 1 5 muc h longer | han 4 : margins spangly hcLosc; segments 3 &4 with brush setae (male). Lower lip lacking inner lobes. Mandible: molar strong; spine row with 5 blades and accessory setae; left 1 acini a 4(5) dentate; palp slender, weakly setose. Maxilla 1, inner plate with 8 apical setae; palp stout. Maxilla 2* inner plalc with several inner marginal pectinate setae, Maxilliped. palp slender, dactyl tong. Coxae 1-4 medium, lower margins various: coxa 3, anteriorly subacute below. Coxa 5. anterior lobe small subacute. Gnathopod 3 & 2 slender, very slightly sexually dimorphic, somewhat dissimilar, 2 die larger, Gnathopod 1, propod and carpus small, short; propod with antero-distal row of about 15-20 pectinate setae, and 3 clusters (2 in female) of longisb spines at the postero-distal angle, Gnathopod 2, propod and carpus more slender and longer; basis, margins lined with long setae, Peraeopods 3 & 4 medium strong; segment 5 small, much shorter than segments 4 & 6; dactyls short, Pemeopods 5-7 rather dissimilar In form; segment 5 shorter than segment 6 and much shorter than 4. Peraeopods 5 &. 6. bases lacking postero-distaJ lobes. Peraeopod 7, basis broad* subacuiely produced below, Pleopods strong. Pleon plates 1-3 broad, hind comers acuminate, Uropod 3 I & 2 strong, rami unequal. Uropod 3* rami lanceolate, margins setose (male), spinose and very weakly setose (fe-nti.c). Telson ordinary, lobes fused basally, not diverging distally; apices each with single small spine. Coxa! gills large, sac-like* simple (female): anterior gills moderately pleated (male) Distributional- Ecology: Bering Sea to Sea of Japan, in depths of 10 ■ 80 metres, mainly on sand, In North Amer- ica, from St. Lawrence Island and the Pribilof Islands to the Aleutian chain and mainland Alaska* from the shore line to depths or more than 10 m. Taxonomic -commentary : This species is distinguished by ils large body size and low bod y carinations, except on the urosoine where it is bicuspate on fused urosome segments 2- 3. Among other disiingu i shing features, the gnathopod s and uropod 3 (esp. in the male) are very setose, and the mandibu- lar palp is slender. This species evinces plesiomorphic character states such as the weakly subchelate and long wristed gnaibopods (both sexes) and strongly rostrate head. It also possesses apomorphic features such as Hie weakly 5- dentate mandibular left lacinia and weakly pleated coxal gills. AMPHIPACIFJCA VOL 1 MO. 3 OCTOBER 15.1994 [4 FIG. S. Atylus bruggeni (Gurjanava). St, Matthew L r Bering Sea, Female ov (19,0 mm) Male (15.0 mm). AMPHIPACEFICA VOL.. [ NO. 3 OCTOBER 15 , 1994 15 Atyfas ekmani (Gurjannva) (Fig. 6) Soiotropis ekmani Gurjanova T 1938: 323, fig, 35, — Gurjanova, 1951: 685, fig. 473. — Tzvetkova, 1968: 172, Atylus elmani Mills, (961: 19 (key),— Barnard & Kara- man. 1991: 264 (list). — Ofcadtx 1993: 7. Diagnosis. Female (20 mm): Body large strongly com- pressed laterally. Peraeon and pleon with mid-dorsal ridge elevated to low carinae posteriorly on personal segments (5) 6-7, and pleon segments 1-2, Pleosome segment 3 and uresome segment 1, each with strong bifid mid-dorsal tooth. Fused urosome segments 2 & 3 with single acute mid-dorsal earina. Head: rostrum arched, medium to large 1/2 head length); anterior head lobe narrow, angles rounded. Eyes small to medium, subovatc, Aniesmac slender, not elongate. Antenna L peduncular segment 2 shorter than 1, weakly setose behind: segment 3 short: accessory flagellum ves- tigial, Antenna 2, peduncular segments 4 & 5 weakly setose. Lower lip riot described (inner lobes probably lacking). Mandible: molar strong: spine row with 6-7 narrow blades and accessory setae; left lacinia 5 1/2 den late; right lacinia bifid, tips fiabe Hate; palp slender, weakly setose. Maxilla L inner plate with about 6 apical setae; palp strong. Maxilla 2, inner margin of inner plate with single stout plumose seta, Maxilliped, plates large, palp slender slightly shortened. Coxal plates 1 4 narrow, shallow, subacute betow; coxa 5, anterior lobe small, sharply rounded, Gnathopods 1. & 2 small, slender, unequal, little or not sexually dimorphic, bases not strongly setose behind, Gnathupod l, carpus medium depth, as long as propod; propod inner face anteriorly with 5-6 row's of pecti nate seiae, di stal 2 rows each with more than 20 setae. Gnathopod 2, carpus slender, longer than propod. Peraeoptxls 3 & 4 strong, spinose; segment 5 distinctly shorter than segments 4 & 6; dactyls med ium . Peracopods 5-7 dissimilar, segment 5 shorter than segment 6 and very much shorter than elongate segment 4. Peraeoptxls 5 & 6* bases with very small acme postero-distal lobes, Peraeopod 7 moderately broad, posiero-dislal lobe small* rounded be- low. Pleopods uudescribed, Pleon segments medium broad, hind comers rnucronate- Uropods 1 & 2 stout, rami unequal, margins spinose, Uropod 3, rami subequal, lanceolate. - 2X length of peduncle, margins spinnse. Telsoo lobes long, narrow, fused in basal 1/4, apices not diverging, each with notch and small spine. Coxal gills not described. Distribution: Western Nonh Pacific: Russian coast of ihe Japan Sea and southern Hokkaido, north to the Okhotsk and western coast of the Bering Sea, at subtidal depths. Taxonomic Commentary: Mills key to .species of Atylus includes A, ekmani erroneously in the group with 2 dorsal teeth on urosoane 5 & 6. This oversight, pointed out by Okada (1993), is collected in the present key (p. 10), The species clusters mast closely with P. bmggeni and P. ievi- deusm (p. 60). Features in common include the very thin body, carina led abdomen, weak gnathopods (propod of gnathopod 1 with heavy pectinate setae), unevenly scalloped lower margins of the anterior coxal plates, and the shorn spinose uropod rami in both sexes. Atylus kvideasus ).L. Barnard (Fig, 7) Atylus kvidensu i J, L. Barnard, 1956: 38. pis. !3„ 14. — Mills, 1961: 19, tig, I. — Barnard, 1969b: 94— Barnard, 1975 : 340, 359. fig. 133.— Austin, 1985: 604,— Slaude, 1987: 382, — Barnard & Kanrnm 1991; 264. Material Examined (CMN collections, Ottawa): S. E, .ALASKA: Prince William Sound {Kayak L) through outer coast (Siika region), to southern Alexander Archi- pelago (Bronson Bay), ELB Sms, June-Aug„ 1961 - 51 specimens in 8 lots, at: A3(l ), A6( I ), A22( 1 1 - [Including female hr 111 (10.5 nun) (slide mU, male (7,0 mm) (slide mUl, A 75(7), A8CK5)* A1 12(IL A1 5 1(5), A 175 (20). ELB Stns., Lisranski Strait to Sitka region, 1980 - 8 specimens in 5 lots ac $4B3(1), S4B4(1), S8BJ (2), S11B2(1), S19B113). BRITISH COLUMBIA; Queen Charlotte Islands: Graham L, north, outer, Emd inner coasts + M asset Inlet, ELB Sins, July- Aug,, 1957 - '■100 specimens in 13 lots deported upon by Mills, 1961), B. C. Mainland coast: Prince Rupert to Calvert Island, ELB Sms, July, 1964 - -200 specimens in 15 lots, at; Hl(19), H5(3), H7(50k H8U4), H12( 16), H26( 1), H33(l), H35(4), H39(50), H44(9X H47f6), H49(3), HSOdO), H530X H65 ( 6 ). North end Vancouver I., Cape Scott to Wickaninnisb Bay, ELB Sms* July, 1959 - 30 specimens in 6 lota (reported upon bv Mills. 1961 ). South end Vancouver 3 ., outer coast south to Victoria, surf coast locations, in Fhyilospadixc orms: July, 1955 - 7 speci- mens in 4 lots (reported upon by Mills, 1963 1. Barkiey Sound south to Sooke, ELB Stns, 1964-77 - 80 specimens in 17 lots, at; P702(2) > P7l9(5) > P710® t P711( l), P7l4(2);B3(52) [including female ov( 10.5 mm ) (stifle mt.), male (8.5 mini (slide mQJ, B4(5), B5(3): B8G), B 1 9(18), Strait of Georgia, English Bay, ELB eoll, - 1 female ov (12,0 mm) (slide mu); 1 male ( 10,0 mm) (slide ml.). WASHINGTON, OREGON. ELB Stns., Strait of Juan dc Fuca to Ottei Rock, July -August, I960 - -250 specimens in 7 lots, at; W30 (2),. W34{13), W36 (62). W4Q(50), W42(6), W58(65), W60(48). Coos Bay, Oregon, to Mendocino Co,, CA, KE Con tan Shis, July, 1986 -50 specimens in 5 lots, incuding 1 female br. HI (10,0 mm) (slide mt,); I male (7,0 mm) (slide mt). AMPHIPACIFICA VOL. I NO 3 OCTOBER 15,1994 16 B FIG. 6, Atylus: ekmani Gurjanova A. Female (20,0 mm )♦ Ja|janSea. (modified from Gurjanova, 1051) B. Female (20,0 mm) (modified From Ok ad a, 1993)* AMFHIP AC [FIC A VOL. I NO. 3 OCTOBER 35. 199 J 17 Diagnosis Female {1 2.0 mm); male (7-8 mm): Body medium, strongly compressed laterally. PcryeOrt arid pkon with mid-dorsal ridge, elevated to low carina on peraeon segment 6 &. 7 and pi eon segments 1-3- Urosomc segments 1 . and fused segments 2 & 3, each with acute elevated dorsal tooth. Head: rostrum large £“1/2 head length): anterior head lobe shallow, slightly emarginate. Eye small oval. An ten nae slender. Antenna 1, peduncular segment 2 shorter than 1 , not setose behind: accessory flagellum minute, Antenna 2. peduncular segments weakly setose. Lower lip lacking inner lobes. Mandible: molar strong; spine row with 4*5 blades and accessory setae; left lacinia 4 1/2 - dentate; palp slender* weakly setose. Maxilla 1 * inner plate with 5 apical setae, palp strong. Maxilla 2, inner plate with single large inner marginal plumose seta. MaxilEiped. palp slender, inner plate relatively short. Coxal plates 1-4 medium, Utile overlapping basal I y: coxa l not directed forward; coxa 3 anteriorly acute below. Coxa 5, anterior lobe narrowly acute below. Gnathopods 1 & 2 not discemibly sexually dimorphic; propod short, small, with distal row of numerous pectinate setae; carpus slender, little longer than propod. Peracopods 3 & 4 relatively short; segment 5 small much shorter than segments 4 6; dactyls short, Pceaoopods 5-7 somewhat dissimilar in size and form; segment 5 s mall shorter than segment 6 and much shorter than segment 4; bases moderately expanded lower hind lobes very small* not produced. Pleopods relatively short, weak. Pleon plates 1-3 me- dium broad, hind comers obtuse. Uropods 1 & 2 relatively short, outer ramus the shorter. Uropcd 3, rami short ( - 2X peduncle), thick, margins s p i nose (both sexes). Teison lobes narrow, slightly diverging distally, apices with single stout spine. Coxai gills on peraeopodx 2-7, medium large, weakly pleated in male, Distributional Ecology, North American Pacific; from Prince William Sound (S.E. Alaska) southward through British Columbia lo Central California (rare south of Monterrey) along open, high salinity, suit -exposed, bedrock shores, frequently among conns of Phyllospodix, in the lower intertidal zone, 1 1 was not taken in dredge hauls and is therefore ranked as a truly littoral zone species. It was also seldom collected in (he summer-warm, relatively brackish shallows of ihe Strait of Georgia, A. ievtdettstof , and A, tridens, were the only two species of Aiytar collected in modest abundance. Taxonomic Commentary; The species shows little variation in body size or morphology throughout Ms range, but is endemic to cold -temperate waters of the North Ameri- can Pacific coast, li dusters above the 75% similarity level with A, ekmani of western Pacific shores but only at the 65% level with A, bniggeni of the intervening Bering Sea region {Fig. 30, p. 60). Atyhts ryfo vi { B u I yehe . va) (Fig. S) Noioiroph rytow Bulycheva, 1952: 221, fig. 21. Afvfu.v rylavl Barnard & Karaman, 199 It 264, Ishimaru, 1994: 42. Diagnosis. Female ov, ( n .0 mm ): Body medium, corn- pressed. Ptraeon segments dorsally smooth, Pleon seg- ments J -3 with low ii 3 id -dorsal ridge that becomes a weak carina posteriorly. Urosomc l with posterior mid-dorsal car i nation and pre ceding notch. Fused urosome segments 2 & 3 with raised mid-dorsal tooth. Head: rast-rum arched medium - large f @ I f2 head length); anterior head lobe notched medially. Eyes medium, renifemn. Antenna medium. An- tenna 1* peduncular segment 2 not longer than 1 but “3 X segment 3; accessory flagellum vestigial? Antenna 2, peduncular segments 4 & 5 strong, moderately setose. Lower lip lacking inner lobes. Mandible; molar strong; spine row with 4*5 bladesffi; lacinia not described; palp medium, apicafly setose. Maxilla 1 inner plate with 3 apical setae: palp broad. Maxiila2. inner plate, inner marginal setae not des-cribed (several?). Maxibiped ordinary, plates and palp strong. Coxal plates 1-4 medium large, hind margins setose J ower margins gently con vex , Coxa 5 , a nteri or lobe bn>adl y rounded below. Gnathopods 1 Sl 2 medium slender, 2 the larger; margins of bases not strongly setose, Gnalbopod I, carpus not elongate, slightly shorter than propod; distal pectinate setae of propod not described. Gnalhopod 2, pro* pod and carpus longer and more slender than in gnatherpod I , Peraeopods 3 & 4. segment 5 small, much shorter than segments 4 & 6; dactyls short Pcraeopods 5-7 not markedly differing in length; segment 5 shorter than 6 and much shorter than segment 4 (especially in peraeopod 6); dactyls short. Feraeopods 5 & 6, hind lobes small, not produced be- low, Pertteopod 7, b ind lobe of basis sharply roc tided belo w . Pleon plates 1-3 broad, hind comers obtuse. Uropods l & 2 not clearly shown or described. Uropod 3, rami short { - 2X length of peduncle), broadly lanceolate, margins spinosc Teison short (width 3/4 length), lobes short, fused basal I y, apices narrowing abruptly, each with ! -2 short spines. Coxal gills and brood plates not described. Distributional Ecology; Pe ter-tfae-Great Ba y , R ussian coast of the Sea of Japan, in the littoral zone. Ovigerous females in September, Material of this species was not obtained at North American Pacific stations. Taxonomic Commentary: Although originally as- signed to the genus Noimropis (Bulychova, l o; r cj| .L rylovi is dearly referable to the genus Atyln $ in the form of its antennae, peraoopods, uropods and teison, Afy/KJ rylovi dusters with the A. widens group, including A. borealis. AMPHFPACIFICA VOL J NO. 3 OCTOBER 15, 1994 ]g FIG, 7* Atylus levidensus Barnard. English Bay, B, C + Female ov (12,0 mm) Male (10.0 mm) AMPHIPACIFICA VOL, ! NO- 3 OCTOBER l\ 1994 ]9 FIG. 8. Atylus ryiovi Bulycheva* 1952. Female ov, (1 LG mm), japan Sea. (modified from Bulycheva, 1952} Atylus {ride ns (Alderman) {Fig. 9) Noiotropis frtdens. Alderman, 1936; 58, figs 20-25, Afy/rts tndens Mills, 1961: 25, fig. 3 (parlim- cion-pelagic stage).— Barnard, 1975: 346, 359, Fig T 216 — Austin, 1985: 604— Staude, 1987: 382, figs, 1854, 18 . 63 — Barn art & Kanmian, 1991:265. Material Examined (CMN collections, Ottawa): SE ALASKA: None clearly separable from A. borealis in material taken &i ELB Stns in 1961 or 1980. BRITISH COLUMBIA: [Mills (1961 ) reported on 1955-59 collnsl. Queen Charlotte Islands, mostly Graham I. EL B Stns, July- Aug., 1957 - 20 specimens in 6 lots, at: W2(1), W8(3), W9(9), Wll(5X W 12(1) E17-1 8(1). B. C, Mainland. Prince Rupert ot Rivers IifleL ELB Stns, July, 1964 - ■'240 spins. in 1 1 lots, ai: H 1 (3), H4{5), H7( 15), H23(~8Q), H4H-85), H48(l), H49(2), H50(27) f H52(2), H57(~30) f H6KJL S, end Vancouver S. Wickanninish Bay and Barkley Sd. to Victoria and Nanaimo, ELB Stns, July- Aug., 1970-1977 - -200 specimens in 10 lots, at: P703 ( 1 male (1 1,5 mm) slide mU, P7 1 3( 1 ), P7 1 6(~50)* P7 1 7(47L B7 1 9(2); B4( 1 3), B 5(2), B9(2); 85(31), Bl la(~50- inch I male (1 1 , 0mm) (slide ml ) r I fern, ov ( 10,0 mm (slide mi). WASHINGTON, OREGON; Agate Beach, and Cape Flat- tery to Neskowin Beach, ELB Sms. July- Aug., 1966 - -400 specimens, mostly immature^, in 17 lots at; W33 (-200), W34(54h W 36(20), W39 (8). W40 (72), W42(ll+), W46 (2), W50f I ), W57 (24), W6I (5), Diagnosis; Female (10,0 inm), Male (9.0 mm) : Body small to medium, noi excep- tionaJly compressed. Peraeoit and pleon lacking dorsal cari nation. Urft&ome segment 1, and fused segments 2 & 3, each with medium tooth preceded by notch. Head: rostrum slender, medium (e 1/2 head length) ; anterior head lobe broad, slight! y emargtnale. Eye s very large, broad, subren i form (both sexes). Antennae long, medium strong. Antenna 1, peduncular segment ! longer AMPHJPACIFJCA VOL. J NO. 3 OCTOBER 15,1994 20 FIG. 9 + Atylus tridens (Alderman) Wickanninish Bay, R* C Fem, (10,0 nun) Male (11.0 mm). A MPH PACIFIC A VOL [ NO. 3 OCTOBER 15, J 994 21 than segment 2 tsubequal in male, with posterior marginal brush setae); segment 3 shod; accessory flagellum minute, Antenna 2, peduncular segments 4 & 5 long, moderately setose, much longer an darned anteriorly with brush setae in male; flagellum (female) with about 20 segments, each with short posterior spine. Lower lip with weak inner lobes, Mandible; molar strong; spine row with 5-6 large blades and accessory setae; left lacinia. 5-den Late, right lacinia bifid* tips flabellate; palp medium, distal segment setose. Maxilla 1 , inner plate with 5 apical setae; paip broad, strong. Maxilla 2, inner plate with single inner marginal plumose seta, Maxilliped ordinary. Coxae 1 4 broad, deep, lower margins variously convex; coxa 1 smallest, 4 largest and broadest. Coxa 5, anterior lobe small, subacute- Gnathopods 1 & 2 not grossly differing tn size but moderately sexually dimorphic; bases lined posteriorly with numerous long simple setae; propod and carpus slender, not elongate; propod of gnuthopodl armed anteio-distaJiy with 3-7 pectinate setae; postero-distal angle with 2-3 groups of sfrines. Peraeopods 3 &. 4 stout; segment 5 small, much shorter than segments 4 & 6 (in male, all armed posteriorly with plumose 11 swimming setae"); dactyls short. Ffcraeopods 5- 7 dissimilar in size and form; segment 5 small, much shorter than segments 4 & 6. Feraeopods 5 & 6, basis moderately broadened, lower hind lobes small, unproduced, Feraeopod 7, basis broad, hind lobe acute below, with notch. PEeopods strong, peduncles large. Pleon plates i -3 broad, hind comers acuminate. Uropods i & 2 SIOUL rami unequal. Uropod 3, rami strongly lanceolate, margins Setose (male) spino.se. and weakly setose (female), apex with 3 stout Setae, Telson ordinary; lobes not diverging, apices with small spine; in male, lobes more elongate and apices each armed with 3 strong setae, Coxal gills pleated, ba&ally lobaic fmalc). Dixirihiitkmal Ecology; Queen Charlotte Islands south along outer coasts of British Columbia (few inner) to Oregon and central California, in high salinities (mostly above 29%-:,) in surf exposed situations, mainly in or above sand. Range extends south of A. borealis, although the latter was not taken $. of Juan dc Fuc-a. Neither species was taken as far north as Prince William Sound, Taxonomic Commentary; The material examined by Mills (1961) has been re-examined and found to consist of two distinct species of which the large "pelagic stage” is the mature Form of A borealis . It tends to occur in deeper, colder, up-welling areas, from Juan de Fuca noith to SE Alaska. In mature male specimens, the pn.3xim.al flagellar segments were each armed poosteriorly with what appeared to be calyx-like protozoa, superficially resembling caleeoli- Atylus ttidens is more abundant at southerly locations, and in summer warm, brackish waters of the Strait of Georgia. Ajyt&x hnrefilis, new species (Fig. 10) A>'/kj f rirfnn Mills, 1961; 29 & Table 2 (pelagic stage). — Barnard & Karaman, 1991; 265 (part)? Material Examined (CMN collections, Ottawa): SOUTHEASTERN ALASKA; ELB Stn*J96l: A I ft, Mac Arthur Bay, Kuiu L June ft - 1 male (17 mm). 9 hum.; A59, Dixon Hbr,. greet) Ling stomach contents. June 19 - i subadult male; A 1 40, McLeod Harbor, Montague L, June 1 3 - 4 males 1 female* 1 im, Chichagof 1. to KmofL* ELB Sms, 198(1: ■ -75 specimens in 8 lots at; S4B2(l), S4B3( 1 J, $4B4( -40), S4B5(2), S4B A48(l), AA63G), A 7 1(21 AS mi A8KU A84U), A133(l), Al4fl(-53), BRITISH COLUMBIA: Mainland Coast: ELB Stns, July, 1964: H13( 1 1 - including mule ( 1 1 .0 mm) (slide ml,), female ov (8.5 mm) (slide mi.), H16(1),H 17(21). Diagnosis, Ferny le hr. Ill (17.0 mm), male {19.0 min): Body large, strongly compressed, Peraeou and pleon with mid-dorsal ridge, elevated to low carina posteriorly on peraeon segments 6 & 7 and pi eon segments 1-3. Utosome segment J with strong mid-dorsal crest. Fused unosome segments I k 2 with mid-dorsal crest, and weak dorso- lateral ridges. Head: rostrum medium, nearly straight (- \l 3 bead Length); anterior head lobe broad, slightly emaiginate: upper angle subacute, Eyes small, lateral. Antennae rela- tively short, stout; flagella short. Antenna 1, peduncular segment 2 shorter than 1 (both sexes), posteriorly moder- ately .setose (brush-setose in male); segment 3 medium: flagellum 1 2-segtnen ted: accessory flagellum minute. An- tenna 2, peduncular segments 4 & 5 stout surfaced with numerous clusters of short setae; posterior margin with short setae (both sexes); peduncular segments 4 & 5 stouter and more elongate in male. Lower lip. inner lobes very weak, not well de lined. Mandible: molar suiting; .spine row with 5-6 narrow blades and accessory setae; left laeinia 5-dcniate, right I acini a bi- fid. apices 3-5 dentate; palp strong, segment 3 distally set- ose. Maxilla L inner plate with 6-7 long apical setae; palp stout, proximal segment short. Maxilla 2, itfrrerplatc with 6- 7 pectinate inner marginal setae, MaxiBiped normal, inner plate relatively tail Coxae 1-4 large, deep, lower margins of 1-3 convex, of 4 subacute. Coxa 5 deep, anterior lobe small rounded, Gnathopods 1 & 2 stout, subsimiiar (2 larger), weakly sexually dimorphic, bases posteriorly strongly setose. Gnathopod l, carpus short, hind lobe deep; propod with an loro -distal group of 4- 5 pectinate setae, and 6-7 rows of short stout spines (5 rows in female) at postero-distal angle; dactyls basally thick. Gnathppod 2, propod and carpus slightly larger and more elongate than in gnathopod 1 . Peraeopods 3 & 4 stout, spinose, 4 slightly heavier than 3; segment 5 small shorter than segment 6 and much shorter than 4: dactyls mediums 1/2 length segment 6V. Peraeo- pods 5 - 7 dissimilar; segment 5 small, shorter than segment 6 and much shorter 1 ^n segment 4, Peraeopods 5 &. 6, bases moderately broadened, hind lobes moderate, not produced. Peraeopod7, basis, posicro-dislal lobe rounded helnw, AMPHIPAOFICA VOL. T NO. 3 OCTOBER 15, 1994 24 FIG, 11. Atylus colltotgi (Giirjanova) St. Lawrence t Bering Sea, Male (19.0 nun), Femalebr, III (17,0 mm). AIWPHIPACIFICA VOL. ] NO. 3 OCTOBER 15, L994 25 Pteopods stout. Picon plates 1 -3 broad, hind corners acuini- nate= Uiopod ] stouf rami suhequa]. Uropod 2, rami relatively long, unequal. Uropod 3, rami short (< 2X peduncle), apices acute, margins weakly spinose. Telson lobes short (shorter in male), fused 1/3 basally. converging di stall y. apices each with single slender spine Coxal gills on pmcopods 2-7. weakly pleated anteriorly, smaller, simple posteriorly, in males and females, Brood plates broadly strap- Like. Distribution: Japan Sea U> (he Chukchi Sea, in deplhsof 3- 1 0 m, North American Pacific region: from the Bering Sea and Aleutian Islands south to the Strait of Juan de Fuca and Puget Sound. Taxonomic Common tary: The western Pad tic material figured by Guijanova (1951) differ from North American material in its somewhat smaller, more acute coxal plates f ■ 4, deeper hind lobe of the hasi* of peraeopod 7, and inner plate of maxilla 2 that has only 3 apical setae. Atylus gtorgianus, new Specie? (Fig. 12) Arytea collingi Mills, 1961: 23, figs. 2, 4B A, — SUiude. 1987: 382(part?),— Barnard & Karaman, 1991; 263 (part). Material Bxamimd (CMN collections, Ottawa): BRITISH COLUMBIA; ELB material (1955, 1957. 1959) from the Queen Charlotte Islands and Vancouver I„ reported upon by Mills (1961). has been reexamined, and proves to consist entirely of this species. Additional material, from S, Vancouver I. tnludcs: ELB Stn. H43 (Witty's lagoon) July 28. 1964 ■ 17 imm, Satuma T, JFL Hart call,, Aug. 26, 1955 - female ov (8.0 uun). Holotype (slide mU, CMN Cat. No. NMCC 1994-0387; ] male (7.5 mm). Allotype (slide mU, CMN Cal. No, NMCCI994-038B; 10 other specimens, Paratypes CMN Cat. No. NMCt 1994-0389. Head of Departure Bay, JFL Hart oolL Aug, 25, 1938 - 2 males (8.0 mm). 1 female hr. IN (9,0 mm). WASHINGTON: North of Columbia estuary, ELB Stns, J td y J 966 - 6 small specimens in 4 lots at: W26b( 1 ) . W3 5(2) , W4(KT), W46(2). Diagnosis; Male (7.5 - 8.0 mm); female (8 ~ 9 mm); Body medium, laterally compressed. Peraeor and pleon witli nud-doml ridge increasingly elevated to weak poste- rior carina on peraeon segments 6-7, and pleon segments ] - 3, Urosome Segment I , and fused segments 2 & 3, each with single elevated rounded tooth. Head: rostrum short, extend- ing little beyond weakly acute anierior head lobe. Antennae medium, much as in A. colfingi, but shorter and less setose. Lower lip, inner lobes small, Mandible: molar medium; spine row with 6^7 slender blades and accessory setae; left lacmia strongly 5 -dentate; right lacmia simply bifid; palp slender, segment 3 setose apical ly. Maxilla J, inner plate with 5 apical setae: palp medium slender. Maxilla 2> inner plate with single inner marginal plumose seta. Maxilliped slender, basal segment with long distal faeiaJ setae . Coxae I -4 large, deep, overlapping, rounded below. Coxa 5 deep, anterior lobe small rounded, Gnathopods 1 & 2 medium (less strong than in A, collingiY* slightly sexually dimorphic; bases setose posteriorly. Gnathopod 1, carpus very short, lobe deep: propod with antero-distaJ group of 3- 5 pectinate seiae, and 4 groups of slender spines ai postero distal angle. Gnathopod 2, propod and dactyl larger, heavier than in gnathopod 1 . Pcraeopods 3 & 4 medium strong, margins s pi nose; segment 5 small much shorter than segments 4 & 6. dactyls medium. Peraeopods 3-7 medium, less spinose, dissimilar: segment 5 much shorter than segments 4 & 6. Pemeopods 5 &6, hind lobes of ba;is small or lacking. Peraeopod7.basis, hind lobe rounded below. Pleopods medium. Picon plates I hind comers squar- ish. not acuminate. Uropod Lramisubequal. Uropod 2. rami unequal. Uropod 3 short rami ~2X length of peduncle, margins spinose (both sexes), Telson lobes short, fused in basal 1/3, converging di stall y, apex of each with slender spine, Coxal gills sac-likc. anterior gills pleated In male, simple in female. Etymology: The trivial name gevrgianus alludes to the Strait of Georgia where the species is commonly encoun- tered. Distribution: Endemic to the North American Pacific coast: Queen Charlotte Islands and central B, C.. Strait of Georgia, to Washington-Qregon coast, frequently in beds of eel grass, in sandy shallows, Tuximomk Commentary: Atylusgeorgiamis is closely related to A. coiling} Guijanova but differs in mandibular palp, annature of plates of the maxillae, size of gnathopods. and shape of the uix^omal carinae. The subequal size of the mature male and female is distinctive. Atylui occidentals Hirayama Atvtus occidental! s Hirayama, 1986: 4, figs, 1-4. -- Ishim&iu 1994: 42. Taxonomic Commentary, The original material was from Otsuchi Bay, Japan. We tentatively accept the designation of this species by Hirayama and Ishimaru floe, cit) . as ei member of the genus Atvltts (seas. str.T Regretahly, how- ever. we have seen no material of this species, and the literature is noL available to us. The species is therefore not included (0 the regional key (p, 10) or analysts of species relationships (p. 58-59), AMPHIPACIFICA VOL. I NO. .1 OCTOBER 15. 1994 26 FIG. 12. georgianus, new species. Saturna T„ B. C. Fem. ov. (8.0 mm) Male (7,5 mm) AMFHIPACIFICA VOL. I NO. 3 OCTOBER 15, L 994 27 KEY TO GENERA OF NGTOTROPIINAE E. Pigmented eyes well developed; anterior head lobe, inlum or slightly emarginaie; peraeopods 3 & 4 n seg- rnent 4 small, much shorter than segments 4 & 6; coxal plates 1-4 large. deep, strongly overlapping . . . , . , . , ft'ototroph (p. 28) — Pigmented eyes lacking; anterior head lobe hi fid; peraeopods 3 & 4, segmenl 4 slightly shorter than segments 4 & 6,; coxal plates I small, shallow, basal] y overlapping only AbemtfyJus (p, 30) NolotropHnae. new subfamily {see Fig. 1(b)) Aly Lite (purl) S [ebbing, 1906:329. -Bamaid. !969a: 161 — Gurjanova, 1951; 327, Atylidae Lincoln, 1979: 438. Dexammsdae (Dexamininae) (part) BeUan-Santim, 1982: 212.— Barnard & Karaman, 1991: 260, Type genus: Nottitrapis Costa, 1 SS3: 170, Genera: Aberraiylu i, new genus ip. 30). Diagnosis: Similar to AtyMnae (p. 8) with the following differences: Body medium, occasional ty large. Peraeon, posterior segments often not Jtt id-dorsal !y toothed or carinate , Pleon various, often smooth above, Urosomc l usually with mid-dorsal tooth and preceding sharp notch. Rostrum short to medium. Eyes large (when present). Antennae slender; antenna I, peduncular segment 2 longer than 1. Mou&hparts haste. Lower lip. inner lobes variously developed, or lacking. Mandible, molar strong* palp slen- der, 2-3 segmented, weakly setose. Coxal plates 1 -4 large, lower margins smooth or rounded. Gnathopods 1 & 2 subsunikm moderately to strongly subchelate (esp, in male), variously sexually dimorphic* or not. Peraeopods 3 & 4 distinctly unequal in size, peracopod 4 the smaller, shorter in segments 2, 5 & 6; segment 5 variable, but typically small, much shorter than segments 4 & 6; segments 2,4, and 6 (male) often posteriorly armed with "swimming setae' 1 . Peraeopod 5, basis, posterior lobe usu- ally produced below, Peraeopods 5-7, segment 5 not short- ened, 5 & 6 subequal; segment 4 often elongate, Pleopods powerfully developed, stronger in die male, Pleon plates 1-3. hind corners mueronate Uropod 3, rami long* lanceolate, margins variously setose (both sexes), Telson, lobes ordinary, dee ply separated, apices sputose. Coxal gills 2-5 strongly phylloform or dendritic (espec- ially in male), simple on peraeopods 6 & 7. Brood plates medium to strap- like. Taxonemix Comit ternary: The subfamily overlaps with subfamily Atylinae in a number of character states, but can be distinguished reliably by the combination of character states illustrated in Fig. HbHp, 6), Nototropis Costa (sec Fig. 13) Noroiropis Costa, 1853; 170. — Slebbing, 1906; 329, — fiuijanova* 1951: 680 (most). Atylus Lincoln, 1979; 438 ( pan),— Barnard. 1969: 163 (part), — Bcllan-Santini, 1982: 212 tall).- — Barnard & Karaman, 1991: 262 (part). Paraiylus G. O, Sarx, 1895: 462. Type species; Nototropis gutratus Costa 1953 (,-Noto- i topis spimilicmuki Costa), Species; Nototwpis brtvUarsus Ledoyer; 1979; N. comes Giles, 1888; JV. deFUflntfSehellenberg > 1931;^.jfe/c^f(ry (Metzger. 1871); N. $rtmufo$tti Walker, 1904;#, hcrtwchir (Has well, 1885): N. mass Hens is Bellan-Santini, 1975; N. mevalop-'f (Moore, 3 984) ; jV, mfkuwps Oldevig, 1959; N. minikoi W alker, 1905; .V, twrdkmdicus Boeck, 1871; JV. re ductus K, H. Barnard. 1930; M serratus Scbellenberg, 1925; A'. smini Goes, 1866; A( swammerdamei Milne- Ed wards, 1830; N. taupe* J.L. Barnard. 1972;A r . ur&carinatus McKinney, 1980; .V, vedlomemis Bate & Westwood, 1863; NQf&trapis sp, (-N. gmmm s Irie> 1965)7 Diagnosis; Small to medium (occasionally large) aty lids. Rostrum short to medium. Eyes often very large, especially in males. Peraeonal segments 5-7 and pleon segments L3 dors-ally smooth, occasionally mueronate Urosome 1 dorsal ly with carina and preceding notch; fused urosome segments 2 &. 3, median dorsal carina variously developed Or lacking. Antenna 1 , peduncular segment 2 not shorter than segment 1; accessory flagellum minute or scale-like. An- tenna 2, peduncular segments 4 & 5 strong, weakly margin- ally setose. Lower lip, inner lobes various, occasionally lacking. Mandible, palp slender, (2)3-segmenied, Maxilla l, inner plate with 3-8 apical seiac; palp (l)2-$egmeiued; Maxilla 2, inner plate with stout inner marginal plumose seta, Maxilli- ped, palp normal slender. Coxal plates 1 -4regular, medium, lower margins rounded or straight, not acme. Coxa 5 antemlobate, lobes rounded below, Gnathopods 1 & 2 variously sexually dimorphic; caipus and propod relatively short, subequat in length; propod of gnatbopod 1 with aruero-distal dusters of pecti- nate setae, Peraeopod 4 distinctly smallerorsborter than peraeopod AMFH1PACIFICA VOL 1 NO 3 OCTOBER 15, 1994 28 FIG, 13, Notolropis guttatus Costa, 1853. Female (9*11 mm) Mediterranean Sea. (modified from Bella n-Santim, 1982) 3; segment 5 (in both) usually much shorter than segments 4 & 6; in male, anterior and posterior margins of segments 4-6 and distal portion of segment 2 often lined with ‘'swim- ming" setae.. Peraeopods 5- 7 n ot strongly dissimilar in size and form; bases, bird lobes strong, often acute below; segment 5 large, usually longer than segments 4 and/or 6; dactyls medium, Pleopods strong . Pleon si-Ele platen hind comers acumi- nate, not produced- Uropod I rami subequal in length . Uropod 2, outer ramus the shorter. Uropod 3, rami strong., lanceolate, subequal, margins setose in male, spinulosc and or setose in female, TeLson regular, lobes medium, apices obliquely truncate- Brood plates medium to broad, margins simple- setose, Anterior (pertteopods 2-5) coxal gills strongly dendritic or pbyllifonii, especially in the male. Variables; Rostrum large (.V. anterior personal segments dorsally carinate {N. homochir ); gnathopod 2, propod and dactyl elongate (A/, tempo* N. smitti); peraeopod 7, basis, postero-distal lobe weak or lacking {N. korruKf ifr, vV. me lamps, N, smith i] urosome weakly or not carinate OV. m e galops); tel son lobes short (AT, Further generic and/or subgencric categories may yet be required to reflect the taxonomic significance of these variables. Distributional Commentary: Component Species of Nototroplf are strongly Lelhyan in distribution, occurring mainly in tropical and warm temperate coastal waters of the Mediterranean -Caribbean Atlantic and Indian oceans, w ith a few morphologically aberrant outliers in arctic and austral" regions. To date, one species, implausibly identified as the Mediterranean species jV. guiMlus Costa by Me (1965) represents a questionable record of this genua and subfamily from Japanese waters. None was identified in present sludy materia! from the North American Pacific region. AMFHl PACIFICA VOL. I NO. 3 OCTOBER 15. 1994 29 FIG. 14 Aberratvlus aberrantis (J. L, Barnard). Female (48 mm > Male 6.1 mm) (Modified from Barnard, 1973) Aberraiylus , new genus (see Fig. 14 ) Aiytus J. L, Barnard. 1962; 69 (part,); Lepechmelia J. L, BamEird ; 1973; 7 (port). — Barnard & Karamari. 1991: 26L(part>- Type species: Atylus aberraniis J, L, Barnard* 1962: 69. figs. 66. 67, — Barnard, 1964: 40. fig. 32. ij=Lepechin- elk r aberramis J. L. Barnard, 1973: 7. figs. ). Diagnosis: Rostrum medium. Peraeon segments 1-6 smooth dorsal ly. Feraeoti segment 7 and pleon segments 1- 3 posterodorsally mucronate. Urosoine segment I mid- dorsally with two teeth and intermediate notch; fused wtjsomc segment 2 & 3 with prominent earina. Anterior head lobe weakly bifid. Pigmented eyes lacking. Antenna 1. peduncular segment 2 slender, elongate; accessary flagellum I -seg- mented, Antenna 2 peduncular segments slender, elongate, weakly setose, Lower lip, inner lobes present Mandible, palp weak, segment 3 short; axilla] , palp broad. 2-segmentcd; inner plate with 2 apical setae. Maxilla 2, inner plate with single inner marginal plumose seta. Myxilliped, palp and plates normal. Coxal plates 1 4 small, basally contiguous or overlap* ping, !ow r er margins entire, denticulate, not acute or processiienoos. Cosa 5 anlcrolobate, anterior lobe various . Gnathopods 1 & 2 subsimilar (2 larger), very weakly sexm ally dimorphic; carpus and propod medium, palms very oblique. Peraeopods 3 & 4 slender, but relatively short; segment 5 slightly shorter than segments 4 & 6: dactyls medium. Peraeopods 5-7 dissimilar in size; bases little broadened, lower hind lobes small, not acute; segments 4 & 5 subequal in length, boih shorter Lhan 6; dactyls medium (= segment 6) Pleon pi ales 1 - 3 broad, hind comers in ucronate , Uropods slender; umpod 1, rami subequal; uropod 2, outer ramus (he shorter. Uropod 3, rami slender lanceolate, inner margins weakly setose, Tel son ordinary, lobes medium length, not diverging, apices with single spine. Coxal gills not described (probably pleated), Brood plates not described. AMPHIPACIFICA VOL. 1 NO. .3 OCTOBER 15, 1994 30 FIG, IS, LepechirteMa itfhu J. L. Barnard Male [7 6 mm) Female ( 8,5 mm) i mod Hied fmm Barnard, 1973) Lepcchrndttnae Schellenbcrg (revised status) [see Figs. 1(c); 15) Lepechinellidac: Sehellcnberg, 1926: 344. -Gurjanova, 1951: 6-74,— Barnard 1969: 236.— Bousfield, 1982; 273. Dexaminidae (part): Barnard, 1973a: 5. — Bellan-SantinL 1982: 212,— Barnard & Karaman, 1991: 260 Diagnosis; Peraeon (variously) and pleor segments miid-domily proecR&iferous and/or densely covered with small setae and spines. Urosome I with single mid-dorsal process. Urosome 2 & 3 not carinate. Head, nostrum spike - like; anterolateral head margin acutely bifid. Pigmented eyes lacking. Antennae long, slender (both sexes); antenna I the shorten; peduncular segment 2 long; accessory flagellum present 1 -segmented. Lower lip, inner lobes welt developed. Mandible, palp slender, reduced. Maxilla I palp 2 -segmented, distal segment broadened. Maxilla 2, inner plate narrow. MaxillTped, outer plate large; inner plate arched disto-medb ally; palp 4-scgmen ted. Coxa plates 1-7 narrow , small, separated basal I y. Coxae 1-4 incised or acute, often bilobate- below; coxa 1 distinctly deepest; coxa 5 small, antcrolobate. Gnathopods slender, weakly or not snbchelate: carpus (especially in gnathopod 2) longer than propod. Peraeopods 3 -7 slender, elongate, Peraeopods 4 sligh tly shorter than 3„ mainly in basis; segment 5 Utile (or not) shortened; dactyls elongate (often > segment 6) . Peraeopods 5-7 subsimilar in form and size; bases subliflcar. PleopoUs slender, elongate. Uropods 1 & 2 slender. Uropod I , outer rEnnus enlarged. Uropod 2; outer ramus not shortened. Uropod 3, rami sublinear, rod-Uke. margins sparsely (or not) setose, apices spinose. Telson lobes short to medium, fused basally by more than 1/3; apices usually diverging, distall y narrowing. Coxal gills pleated , Guvkerat LepectanellQ (Lepechiwtta} Stubbing* 190S: 19 1; Pvralrpechinetki Pirlot, 1933; 161; Leptchinelloides Thurston, 1980: 81; Lepechinett&psis Ledoyer, 1982: 365. Taxonomic and Biogeographk Commentary: <‘Upechine lla ” abert&rtiis J, L. Barnard. 1964, is basically an at y lid that exhibits a very few “IcpcChineliid" character states (of head, peraeopods 3 & 4 , and uropod 3). Accord- ingly, the species is here reassigned within family AtyUdae AMPH1PACIFICA VOL I NO 3 OCTOBER 15, I W 31 KEY TO GENERA AND SUBGENERA OF LEPECHINELL1NAE K Mandibular palp servient 3 elongate, telson lobes not diverging Paralepecktnella — -Mandibular |>alp segment 3 short or lacking. lelson lobes diverging r T r 2. 2. Cephalic projections lacking; mandibular palp 1 -segmented tepechinelfoutes — Cephalic projections prominent; mandibular palp 3-segmented 3. 3. Outer ramus of uropods i-3 reduced LepechineOopsis — Outer ramus of u reports 1-3 normal tepee kind fa to subfamily Nototropiinae with which ii appears to have closest morphological affinities (p. 28), A new genus, Abenwyhu (p, 30) is here erected to accommodate its uni- que combination of character state s. The pbyietic and artificial keys to Lepecftinelfa devel- oped by Barnard (I ^73) suggest further internal subgroupings that might merit formal subgeneric recognition, Thus, a group contajniQglc/jfcWrtcl^Jducd, L c&cHi, L cetmta, and L httaco exhibits plesiomorphie (atylinid) character states including a lade of mid-dorsal teeth on three or more peraeonal segments, coxae 1 -4 weekly proccssiferous below, and per- aeopod dactyls less markedly elongate than in other Icpech- rneilid species groups. About 35 described species, in 4 genera, can be assigned to subfamily Lepechinelllnae, all abyssal and bathy pelagic - epibenthic. At least two species arc known from abyssal depths off Japan (Gamo, 198 1 ). None was recorded front the Cascadia Abyssal Plain off liic coast of Oregon by Dick- inson and Carey { 1978), at least not in significant numbers, and none was found in CMN amphipod material from other North American Pacific deep-water sites. Subfamily Aunty I ferae Bulycheva (Revised status) (Figs. Ltd); 16) Anatylidae Bulycheva. 1955: 205, Dexaminidae (Anatybnae) Barnard, 1969a: 202 . Dexaminidae (part) Barnard &. Etaraman ; 1991; 260. Type Genus: Anaiylus Bulycheva 1955: monoiypy. Genera; Kamehatylus Barnard. J970b; 93 (revised sta- tus). Diagnosis: Smallalylids(3-6mm). Body thin. Peraeon segments 5-7 and pleoo segments J-3 variously carinate or smooth tnid-dorsally. Rostrum weak. Anterior head lobe shallowly excavate. Pigmented eye small, Antennae 1 &2 short; flagella short. 4-5 segmented. Antenna 1 , peduncular segments l & 2 subequal; accessory flagellum vestigial. Antenna 2, peduncular segments weakly setose Lower lip, inner lobes very weak. Mandible: molar trending to reduction; left tacinia 4-demaie; palp lacking. Maxilla L inner plate with 2 apical setae; palp slender, Maxilla 2, inner plate slender. Maxilliped normal; palp strong. 4 -segmented. Coxae 2-4 relatively shallow, narrow, lower margins gently excavate. Coxa I tapering, subacute below. Coxa 5 shallow Gnathopods I & 2 slender, dissimilar in size; propod, palms very oblique. GnaUiopod L propod and carpus relatively short. Gnatttopod 2, carpus elongate, Rcracopods 4 distinctly shorter than 3. mainly in basis and segments 5 & 6: segment 5 short. Fcraeopods 5-7 bases dissimilar, lower lobes very small or lacking; segment 5 not shortened, longer than segment 6 . Pleon plate 1-3 deep, hind comers obtuse or rounded. Uropods L & 2, rami medium, unequal. Uropod 3 short, rami stout, margins spi nose. Telson medium short, lobes deeply separated, converg- ing di Stall y, apices with single spine Coxal gills undescribed, but probably sac-like, unmodi- fied Brood plates undescribed, probably strap-like, Male unde scribed, Taxonomic and Distributional Commentary: To date, the subfamily contains but 5 described species in two closely similar genera, of Indo-Pacillc and western Pacific affinities, as detai led be low . The present study restores [he group to the subfamily status proposed initially by Barnard (1969a). Anafylus Bulycheva Aiimylus Bulycheva : 1955; 205, original designation. — Barnard, 1969a: 202 tin Dexaminidae) (part) Barnard & Karaman, 1991: 262, Type Species: Anatylus pavt&vski Bulycheva, 1955. Diagnosis; Bodv medium, thin. Peraeon, segments 5-7 and pfeon segments I -3 carinate along dorsal margin (of, Atylus levidensus). Rostrum medium strong. Anterior head lobe shallowly excavate, Pigmented eye small+ round. Antennae I & 2 short flagella 4-5 segmented. Antennal, peduncular segments 1 & 2 subequal; accessory flagellum vestigial. Antenna 2, peduncular segment 5 longest. Lower lip n inner lobes present, moderate. Mandible: palp lacking; moLar reduced, weakly triturative. Maxilla 1 . inner ptate fused 10 base of outer plate, w ith 2 apical setae; palp 2-segmented. Maxilla 2, inner plate small, lacking strong plumose inner marginal seta. Maxilliped normal, palp AMPHIPACIFICA VOL. 1 NO. .* OCTOBER 15, 1994 3 2 FIG. Ifi. Anatylus pavUmkii Bulycheva, 1955. Female (6-8 mm) japan Sea (modified from Bulycheva, 1955) strong. 4-segmented. Coxae 2-4 relatively shallow, hevtow, s^i^tiUy cmarginaie below. Coxa 1 tapering, subacute below, almost as deep as coxa 2. Coxa 5 broadly antero-lobate. Gnathopods 1 & 2 slender, dissimilar in size (2 larger); carpus longer than propod, palms very oblique. Gnathopod 1, propod and dactyl relatively short. GnailK>pNl 2 t propod and dactyl rel- atively long, slender Peraeopods 3 & 4, segment 5 shortened. Peraeopods 5- 7 slightly dissimilar, segment 5 not described. Peraeopod 7, basis lacking distinct posts ru-distal process, Pleopods not described, (not powerful?), Picon plate 3 deep, rounded below. Unopods 1 & 2 not described, Uropod 3 short, rami heavy, lanceolate, margins sparsely spinose( female). Telson medium short, lobes deepl y sepa- rated, converging distal ly, apices each with single spine. Coxal gills and brood plates undesciibed. A natylua pa vlovstcii Bu lyche Va (Fig. 16) Anaiytus pavtovskii Bulycheva, 19 55: 206, Fig, 6. — Buly- cheva, 1957: i 04 —Tzvetkova. 1967: 173, Atylfispavlovskii Barnard &. Kataman, 1991: 262, fig.SOA, Diagnosis: With the characters of the genus Distribution? The monotypic species A. pavlovskii is known only from the Russian portion of the Japan Sea (Peter- I he -Great Bay), in medium depths (Bulycheva, 1955), Taxonomic Commentary^ As figured and described by Bulycheva (1955) and refigured by Barnard & Karaman (1991), this species bears a combination of character states A MPHEP ACIFIC A VOL l NO. 3 OCTOBER 15, L 994 33 that are remarkably similar to those of Kamehatylus, origi- Dally diagnosed as a submenus of Afvto tawd on the Hawaiian species K. nani (below). Regretably, Bulycheva did not fully describe or figure the diagnostic character states ofpeiaeopods3-7. Until further material cad be studied. the diagnostic subfamily character states are assumed to be similar to those of Kematylus japonicus which occurs a( other localities in the Sea of Japan, The two genera appear closely similar in described character states, although the type of Kamehanius is based on a species with all three UTOSOittiteS fused. Whatever Tut me studies reveal in this regard, the name Anatylus Bulycheva 1 955 would be a senior synon yrrt and is therefore retained here as a valid full genus, KfWiekatylas J. L. Barnard, revised status (see Figs, 1(d); I7A.B) Atyfus (Kamehatyiua) J. L r Barnard,. 1970b- 93.— Ledoyer, 1979 b: 157.- -Barnard & Karaman, 1991: 262. Type Sptc its: Afy/n.r (KamchcnyiiiS )iWni J. L. Barnard. J 970b: 93. figs. 43,49. Species: Kamehatylus japonic u s (Nagata, 196 3); K. processicer (Siviprakasam, 1970); A", mlearensis (Ledoyer, 1984)? Diagnosis; Small, morphologically modified atylids. Rostrum short. Eyes small. Peraeon and pleon dorsally weakly carinate or nearly smooth, Urosome segments Land fused 2-3 dorsal! y toothed; all three urosome segments fused in the type species, Antennae short, slender, flagella few- segmented; accessory flagellum lacking. Antenna 1 , pedun- cle 1 with posierodistal tooth or process. Antenna 2, peduncular segments 4 At 5, margins nearly smooth. Lower lip tacking inner lobes. Mandible: palp absent: molar process medium; spine row with 2-3 blades and accessory setae; Left lacinia 4-demale, right lacinia bifid- fiabellate, Maxilla 3 , inner plate with 2*3 apical setae; outer place w r ith 10 apical spines; palp slender, 2- segmented. Maxilla 2, inner plate, inner margin subapically with single large plumose seta. Maxilliped, palp slender, shortened, Coxae 1-4 short, shallow, lower margins rounded or slightly incised. Coxa 1 subacute below. Coxa 5, anterior lobe small, Gnalhopods l & 2 slender, dissimilar, probably little or not sexually dimorphic. Gnatiiopod I , propod shorter than carpus, with antero-distal median facial clusters of pectinate setae. Gnalhopod 2, carpus slender, longer than in gnathopod 1. Peraeopod 4 distinctly smaller in size than peraeopod 3: segment 5 (of both) small, much shorter than segments 4 8l 6; dactyls short. Peraeopods 5-7 subsimilar in size* bases not broadly expanded, lower bind lobes small or lacking; scg. mem 5 not shortened, longer than segment 6, but not mark- edly longer than segment 4: dactyls short, Pleon plates 1-3 regular hind corners mucronate. Fleopods not described. Uropods 1 & 2 slender. rm\ un- equal, Uropod 3 rami short, subequal, margins spinosc. Telson lobes deeply separated . d i verging d isial ly, apice s singly spinoac, outer margins bare. Coxal gills sac-like, simple. Brood plates strap- like, not broad. Mature male (Leftover, 1979b): Eye slightly larger, antennal flagella longer, than in female. Taxonomies I and Distributional Commentary. The tew described species of this genus are essentially ludo- Pacitlc in distribution, northwards in the Pacific to southern Japan, but not yet recorded trout the North American Pacific coast. The species appear morphologically specialized for a cryptic life style on coral reefs, in association with large, sessile invertebrates such as sea lilies (Siviprakasam* 1970). Kamekatylus japonic us (Nagata) (Fig. 17 A) Aiylus japonicus Nagata, 1961: 216, figs. 1, 2, — Nagata, 1965a: 202. fig. 19 - Barnard & Karaman, 1991: 263. jl£H Atvlus ( j Katn chai vhts) japonicus — Ledoyer, 1 97% : 1 56 fig. 7(H). TaxonomR Commentary : The species has been well described and figured by Nagata 1 961 4 965, Idl cit) whose figures are partly reproduced here (Fig, 1 7 A), Nagatn's species conforms closely will! the subgeneric diagnosis of Barnaul ( 1970b) that was based on (he Hawaiian species, K. n am. Ho wever, in the Japanese species, the posterior per- aeon and pleon are more strongly carinatcd, urosome seg- ment 1 is not fused with segments 2 & 3, and the gnathopods arc more slender. Despite these and other miotF differences, the authors consider Nagata's material from Japan conge- neric with that of Bamard, and have broadened the generic diagnosis to accommodate both species. Ledoyer ( 1 979b, local) described a very similar species from the Moluccas Islands, Indian Ocean, to ivhich he had perceptively assigned the name Aiylus i Kamehatylus ) japonicus Nagata, Ledoyer’s figures, reproduced here (Fig. 17B), do show remarkable similarities to those of Nagata, including the relatively small eye and excavate anterior head lobe* the postero-dista! process of peduncular segment 1 of amentia l , and the unfused urosome segment 1. However, on close inspection, his Moluccas material i.s seen to differ in a n umber of spec ific feat ures such as its weaker body earinaiion, shorter carpus of gnat ho pod 2, and more acute apices of the rami of uropod 3, Ledoyer’s materia) is therefore regarded here as a species different from A. japonic us Nagata* and awaits Icrmai designation as a possible new taxon. Distributional Commentary. Kamehotyius jiij>on.icuS has been recorded £rom Japanese waters mainly from Honshu and more southerly localities (see summary of pcrimmeni literature by l.sliimaru, 1994). AMPHIPACfPICA VOL 1 NO 3 OCTOBER 15 , 1994 34 FIG- 17. Kamehatyluj japonic us. A. K< japonicus Nagata Female (3 - 5 mm) Seto Intend Sea (from Magata, l^WO) B. K. japonicus Ledoyer Male 13.4 mm) Moluccas Ids* (from Lcdoyer, 1979), DEXAMINIDAE Leach Dexaminidae Leach i8]3/34; 432,— Stebbing, 188& 573, — Guijanova, 195 1:788 — Lincoln, 1979:448.— Bousfteld, 1982: 212, Dexaminidae (part) Barnard. 1969: 21)0, — Barnard 1970: 163. — BeUah-SanUrtl 1982: 277. — Barnard & Karaman. 1991:260. Subfamilies: Pcxam i ni nae Leach ; Pe xam inooi iinae, new subfamily; Foiycheriinae, new subfamily: Proph I laminae Nicholls. Diagnosis: Body small stow, compact, not compressed, Sexual dimorphism expressed in eyes, antenna, uropod 3. typically in gnathopod 1 , pleopods. arid telson PeraeOn seg- ments 5-7 usually smooth above, occasionally with mid- dorsal teeth and/or darso-lateral mucronations. Pleosome and urosome, less often posterior peraeon, armed dorsal! y and occasionally dorso- laterally with leeth or spines, Ros- trum short. Antcrio r head margin rounded or acute; may be produced strongly as ocular lobe, Eyes medium to large, Antennae shorn (female), Antenna 2 often reduced, not longer than l Antenna 1, peduncular segment 2 various; accessory flagellum minute or lacking. Lower lip. inner lobes usually strong. Mandible, molar usually strong, triturating; spine row weak; left lacinia often 4-dentate; palp lacking, Maxilla I, palp 1 -segmented (rarely 2); outer plate with 7-1 1 apical spines; inner plate with 0-2 apical setae. Maxilla 2. plates variously reduced, often weakly setose, Maxilhped, inner plate reduced, with apical setae only: outer plate large, broad, palp variously shortened, dactyl reduced or tacking ( ^segmented). Coxae 1-4 deep, shortest anteriorly, little (or not) in- dented below. Coxa 5, broad often deep. Gnathopods, un- equal subchfilatc (rarely chelate); gnathopod l the smaller, with short carpus, propod (male) strikingly notched or ex- cavate anteriorly, Gnathopod 2^ carpus usally longer than propod. Peraeopods 3 & 4 subequal various, segments 5 & 6 trending to reduction in length, and subchelation, Ptracopods 5-7 sulxqual in length; bases typically unequally expanded, trending to linearity; segment 5 normal, occasionally short- ened, segment 6 & dactyl often shortened, Pleopods short to medium. Pleon plates 1-3. hind comers acuminate, often produced, Uropod 1. rami subequal, tips Spinose. Uropod 2 much shorter than I, outer ramus the shorter, Uropod 3, rami lanceolate (often broadly), margins variously plumose -setose, especially in male, Tdson deeply bilobate, lobes not diverging, apices subtruncaic, variously armed. Coxal gills on j^aeqpods 2-7 (6), variously pleated, not phyllifoitn. Taxonomic Commentary; As noted previously, Barnard (I970a< lac, cit) combined a number of dexaminoidean families (including Atylidae, Analylidac, Lepcchinellidae, Frophliantidae) within family Dexaminidae, His decision was based on the presence of one or more species deemed intermediate in form (often on single character stales only) AMPHPAC3FICA VOL. 1 NO 3 OCTOBER 15. 1994 35 KEY TO SUBFAMILIES OF DKXAMESIDAE LPfcrawpods fundamental Ey simply not subcheliform; body fesp, pleosome) varitnisly can rated dr pro- eessiferous; - . . , , 2. — Peraeopods variously subebdiform; body (estcepjt urosome) smooth Pol^cheHinav (p. 37 ) 2 ► Eyes enormous located at end of interantennal lobe: coxa 3 short; antenna 2 very short in female Itexaminoculmue fp. 49) — Eyes normal, not at tip of interantennal lobe; coxa 3 normal deep; antenna 2 little shorter than antenna J (female) - 3. 3. Body carinated on urosome; peraeopod 7, segments 4 & 5 broadened, strongly setose; gnathopod 1 propod no I sexually dimorphic ....... PropbUantinae Cp. 53) — Body earinated on pi eon and urosome; peraeopod 7, segments 4 & 5 not broadned or heavily setose gnathopod 1, ptopod lypcially sexually dimorphic . , , , r De Kami u lnae (p. 36 ) between the families in question. As noted else where, this philosophy of taxonomic fusion dties not recognize die Dar- winian evolutionary thesis that predicts “intermediate 1 morpbotypex existing, at one time or other, between all ex - tarn and past organisms. Thus, we agree with fshimaru < 1 993) thai the pre sence of si ngle taxa that appear to "bridge" otherwise morphologically discontinuous higher taxa docs not, alone, constitute a valid basis for merging of the perti- nent higher taxa. The Barnard ian classification is therefore not followed here- in this study, numerical taxonomic anal ysis(p, 56) strongly supports recognition of just, two family -level d$ xaminoidcan subgroups, the Atylidae Cp. 8) and the Dexaminidae (above). The analysis further supports recognition of four distinct subfamily groupings within family Dexaminidae, as listed and keyed above. Subfamily Dexaminuiae (revised) (see Fig. 2(a)) Dexamininae (part); Barnard & Karaman 1991; 260. Dcxamininae Is him aru, 1987: 1412, Type genus: Dexamine Leach, 1813/14. Genera; Demminc Leach, 1814: 432; DexamineUa, Scbellenberg, 1928: 654; Parade xamine, Slabbing, 3899; 210; Sebadexius Ledoyer, 1984: 5& t Syndexarnine Chilton, 1914; 332, Diagnosis; Body generally toothed or piocessiferous above, not strongly compressed, Rostrum medium. Eye normal. Antennae regular. Motiihparls typical of family; Lower lip, inner lobes variously developed, Mandible, spine row weak, Maxilla l, outer plate with 10- tl apical spines. Maxilliped, outer plate large; aimer plate distinct; palp variously reduced, segments 3 & 4 shortened or vestigial. Coxae l *4 regular deep. 3 smallest Coxa 5 medium, Gnathopods typically subehelatc, occasionally chelate; car- pus not elongate. Gnathopod I. propod sexually dimorphic. Feraeopods normal, not subefoeliform nor elongate; seg- ment 5 not unusually lengthened or shortened; dactyls me- dium; peraeopods 5-7 subequal in length, bases dissimilar in form, variously broadened; segment 5 normal, PI eon segments dorsad y and dorso-iateraily carinate. Heon plates 2-3, hind comers variously acuminate or pro- duced. Telson elongate, lobes deeply separated, not diverging. Brood plates sublinear. Taxonomic and Biog^ugraphlcCoinmencary: As here defined: the subfamily Dcxamminac encompasses five gen- era and about 60 species that occur mainly in southern oceans. Pamdexamhie, with more than 40 described spe- cies, is essentially Indo-P&cific, with outliers extending to the Mediterranean South America and Japan, The Japa’ nese fauna comprises -8 described species (Lshimain, 1994), all confined to Kynshu and the southern archipelagos; none reaches northern Honshu, and no member of the genus reaches the Pacific coast of North America. Setwdexws Is monotypic in New Caledonia, Syndexamine contains 6 Species, in littoral waters Of New Zealand and sou them Australia. Demminelfa, containing 3 species, is confined to the northwestern Indian Ocean and Red Sea. However, Demmine* with only 3 recognized species (Barnard & Karuiiian, 1991) Is confined to the boreal and temperate North Atlantic region, extending southward along eastern shores to the Mediterranean and Senegal, and along western shores to (he Middle Atlantic States and Chesapeake Bay. Members of this subfamily have yet to be recorded autben- tically from the North American Pacific region and are not treated further in this study. AMPHIPACIFICA VOL I NO 3 OCTOBEE 15,1994 36 KEY TO WORLD GENERA OF DEXAMIMNAE 1 . Gnaihopods eheliform; maxilliped palp various* usually small lo vestigial . , - Sebadexitm Ledoyer, — Gnathopods subcheliforttt* ittaxillipcU palp 3-4 seg melted , • , 2. 2 . Pleon segments distinctly carinate mid-dorsally and/or dorso-Jaterally, incc^ument normal 3, — Picon segments indistinctly or not carinaicd; integument often thick, heavy 5. 3, Picon segments 13 carinate laterally and dorsal] y . Pwvd&utmuu Stebbing — Pleon segments earinaled doraafly only , . . 4- 4. Maxilliped palp 3 -segmented DexammeUa Scheltenberg — Maxilliped pttlp 4 -segmented tlexamine Leach 3, Uropods 1 & 2t inner rami reduced; peraeopod 6 massive Delkaryle J. L. Barnard — Uropods 1 & 2 normal; peraeopods 5-7 subequak 6 rot massive Syndexamine J, L. Barnard Species of both Tritaeta and Polycheria are commensal mainly on sponges and colonial tunicate* (Vade* 1969), POLY CHERUNA E* new subfamily (See Fig. 2(c)) Dexaminidae Cpa^LJ Stebbing, 190 *: 514 , — Barnard* 1969 : 200 ,— Lincoln, 1979 ; 448 — Bellan-Samini, 1982 : 212 ,— Bfimard & Karaman. 1991: 260, Type Genus: Polycheria Halved, [879:345. Generic Content: Tritaeta Boeek, 1876:317, Diagnosis: Body smooth* carinate (weakly) only on urosome, Head; rostrum very weak or absent, Anterior head lobe variously rounded. Byes pigmented, large. Antennae 1 & 2 medium. subequaL flagella usually setose. Antenna 2, peduncular segment 4 longer than 5. Accessory flagellum lacking. Upper iip, epistome weakly produced anteriorly Man’ dibular molar, left andrighL sides unequal. Maxilla L outer plate with 7-9 apical spines, Maxilliped palp 3-, or weakly 4- segmented. Coxa 1-7 shallow, variously bifid or acute below. Gnai hoped* slender, dissimilar in length; weakly subcbelate, Peraeopods 3-7 delicately prehensile (subchelate, or pseudo -carpochel ate): segment 4 elongate; segments 6 and/ or 5 shortened, Ptraeopods 5-7 subsintilar, bases subbnear. segment 7 and dactyl often reversed, Pleopods medium, peduncle and rami not powerful. Picon plates 1-3 , hind comers m ucronate Uropod l, rami subequai, Uropod 2 short, rami unequal. Uropod 3. rami lanceolate, margins setose (mate), Telso-n lobes elongate, deeply separated* marginally spinose. Coxal gills weakly pleated, on |>eiaeapotls 2-7, Brood plates stibhnear* strap-like. clinging upside dow n in small pits excavated in surface test of host, and feeding in the fashion of ampeliscoideans, Taxonomic and Distributional Commentary: The subfamily presently contains two genera, Polycheria and Tritaeta, not very closely related (p. 57), characterized by a trend to prehen si lily (subchdation) of peraeopods 3-7. The peraeopod* of Tritaeta are caipochelaie (fig. 28). About 20 species of Polycheria are known, most from tropical and warm temperate lndo- Pacific regions. Three species were previously described from temperate waters of the Asiatic Pacific coast (Bulycheva, 1952; Hirayama, 19.84) and one from the Pacific coast of North America (Caiman 1898; Barnard. 1969b). Triiaeta contains only two species (many synonymies), both in the northeastern Atlantic and Mediter- ranean regions (Lincoln, 1979; BeMan-Santinb 1982), The phylchc relationships of subfamily Polycheriinae are with (he Dexaminmae { p . 36; fig; 2(a) ). Thus* males of I he more primitive genus Tritaeta retain (be distinctive dex- aminid dorsal ly notched form of the propod of gnathupod 1 Polycheria Has well Polycheria Has well, 1879:345. — Stebbing, 1906 ;519.— Holman & Wading, 1983; 221, — Thurston. 1974: 18. — Barnard & Karaman, 1991: 271. Typt Spedes. Polycheria temipea Has well 1879. Species (North Pacific region). Polycheria osbomi Caiman 1898; P, carinata, new species (p. 42): P. mixillae, new species (p. 44); P. atmkusaensis Hirayama 1984a; F, orientaUs Hirayarn* 1984a: P.japomcus Bulycheva* 1952. AMPHIPACIF1CA VOL, 1 NO, 3 OCTOBER 15,1994 37 KEY TO GENERA OF SUBFAMILY HOLY CHERIINAE [. Ferueopods 3-7 pse utk^earpCKdtelate (carpus expanding and strongly spinose distally, propod lacking palm); antenna! flagella smooth; gnalhopod 1 T propod markedly sexually dimorphic Triiaeta . — Peraeopods 3-7 distinctly subdielate (propod with distal palm, carpus not expanding distal ly); antennal flagella strongly setose: gnathopod 1, propod not markedly sexually dimorphic. , . ,PoIycheria (p. 37). Diagnosis: Body stout, broadest at peraeon segments 4 &. 5 h mkd’doisally canny ted on urosoroe segment I; paired dorsolateral ridges or small spines usually present on fused urosome segments 2 & 3, Head: rostrum very weak: anterior head lobe variously rounded; eyes 1 urge . se xual 1 y d tm oiphi c . Antenna 1, flagella usually strongly setose. Lower lip, inner kibes well developed. Mandible, left and right molars dissimilar in size. Maxilla 1 , outer plate with 7-9 apical spines. Maxilla % apical setae weak. MsKdtiped, palp 4-segmented. Gnaihopods very weakly subchelate, Gnathopod i, propod not striking !y sexually dimorphic; palmar margin short to obsolescent. Perticopods 3-7 del icate I y su be he late ; dactyl short + clos- ing on short Fixed finger: segment 5 short, not expanded or strongly spinose distally, variously shorter or longer I him segment 6, Peraeopods 5-7, bases. subli near (inay he slightly broadened in peraebpods 5 & 6). Uropod 2. outer ramus usually the shorter. Uropod 3 t female), rami variously unequal. Tel son lobes variously fused basal ly. margins spino.se. Sexual ditnoiphism strongly expressed in eyes, antennae, pleopods.. and uropod 3 r Taxonomic and Distributional Commentary: North American Pacific species differ from Asiatic Pacific species in several character states. mostly apomoipliically (pp.6Ct52 and key below ). Both groups d r ffe r from the genera! I y more primitive species of the southern hemisphere as exemplified by the P, antarctica complex of species ( Holman & Watling, loc. cit 'i. Species of I lie Nort h American study region are characterized by: maxilla I , outer plate with 7 (vs. 9) apical spines: maxilliped palp short (vs. medium) ; coxa 1 acute (vs. rounded) below; gnathopod palmar margins distinct (vs, obsolete); peraeopods 3-7, segment 5 shorter (vs, longer) than segment 6; uropod 2 inner ramus (vs, outer ramus) the shorter; uropod 3 (female), rami subequal (vs, unequal); and telson lobes more strongly fused basally. These differences point to the need for an extend ve revision of the gen us, based On re-examination of species world-wide, that is beyond the scope of the present study Palyc herfoosbomi Caiman {Figs. 18, 19, 20) Polycherto osbvmi Caiman, 1898: 268, pi, 32,. fig 2.— Skogslierg & Vanselh 192S: 268, figs. 1-26, — Barnard, 1975: 363, key 4 fig. 53. — Barnard, 1 969a: 103. —Barnard. 1969b: 200, fig. 25g— Staude, 1987: 382 4 key.— BarnanL 1979b: 33.— Barnard & Karamam 1991; 272 (list), Polycftetia antarcrica (Stubbing, 1875): Stebbing, 1906: 520 (part). — Alderman, 1936: 63. — Barnard, 1954a: 21. Material Examined (CMN collections, Ottawa); SE ALASKA: Stilka region. Slocum Pt., ELB Stn S4B4, under boulders, July 27, 1980 - l female ov (slide uil). BRITISH COLUMBIA: Queen Charlotte Islands: none raken at outer coast sites, Noith Cental coast: Oval Bay, surf shore at LW. ELB Sin H10, July 12, l%4 1 female hr, IT (slide ml.), 2 other females, 5, end Vancouver I; Uclnelet, outer coast. L Macnun coll,, July, 1909, (identified initially as P. tenuipes HasweU)- 1 lot dried specimens, Barkley Sd. region, ELB Sms, 1975-76: Taylor I. b Trevor Ch.annel. ELB Sm, P5b c.onascidiansaud sponges, LW, July 25, 1975 - \ female hr 11 (4.5 mm) (slide mt); 1 female ov (5.2 mm) (slide mt.); 2 female ov. (4.5, 4,8 mm) (slide- mts,); [ male many specimens, Kirby Ft., Diana 1 , ELB Stn, Pl7d, Oh sponges and ytunicates from rocky walls of surge channels, LW and subtidaL Aug, 6, 1975 - 1 female ov, (5,8 nun) (slide mt - fig'd specimen); 1 mate (3.7 mm ) (slide mL- fig'd specimen), 2 subad, malts (4,3* 4.5 mm); 3 subad, female (4,2 mm) (slide ml.); several other specimens, Bordelais Islets* mouilh ofTrevorGi.,, ELB Sin. F20c, from sponges and tunicate* on rocky walls of surge channels, Aug. 9 + 3 975 - J fem ale ov. (6,0 mm) (slide mt>2 subad, males (5,0 mm, 3,8 mm) Edward King J. /Taylor L,ELB Sin B28a* under boulders at LW, July 10, 1976 I female ov, (5.0 mm) (slide mt.); 1 female br. M (5,3 mm) (slide ml,); several other speci- mens, mostly subad. females, WASHINGTON -OREGON: No specimens were found in (collections from apparently suitable habitats at localities along the outer coast (see Bousfield & Jarretk 1981). Diagnosis, Female ov. (5.8 mm); EJrosome 1, mid- dorsal carina low. weakly toothed behind. Eye medium, covering anterior l 3X basal width, tinathopod 1 , carpus and propod subeqtml in length, carpus proximally deepest; dactyl slender, projecting :> 50% of its length beyond short palm. Gnalhopod 2 V propod more, slender, shorter than carpus, palm short but distinct, slightly exceeded by closed dactyl. Peraeopods 5-7, segment 5 shorter than segment 6, Peraeopods 3 & 4, basis slightly broader than distal seg- ments. Peraeopods 5-7, bases sublinear, not broadened; segment 6 shorter than in peraeopods 3 & 4. A MFH! PACIFIC A VOL. I NO. 3 OCTOBER 15,1994 39 KEY TO NORTH PACIFIC &PEC1ES OK POLYCHERJA 1. Urosome segment I postMoriy extended. partially concealing fused urosome segments 2 & 3; peraco-pods 3-7, segment 5 not shorter than 6; uiopod 2, outer ramus shorter than inner; maxiJlEt 3, outer plates with 9 apical spines: in as i (la % inner pi ale, inner margin setose (Asiatic Pacific) . . . 2. — Urosome segment 1 not extended posteriorly, based of uro&ome segments 2 & 3 open; peraeopods 3-7. segment 5 shorter than 6; urepod 2, inner ramus the shorter; maxilla 1. outer plate with 1 apical spines; maxilla 2, inner plate with weak apma! setae only (North American Pacific) 4. 2. Gnathopcds 1 & 2 subchelate, palm distinct; peraeopod 5, basis expanded, length < 2X width . . , . . . P.japoaieas ( p. 44) — Gnatliopods 1 & 2 nearly simple, propod palmar margins very short or obsolete; peraeopod 5, basis sub- linear, length > 2X width , . 3. 3. Peraeopods 6 & 7, segment 6 distinctly shorter than segment 5; pleom pftite I, hind corner rounded . r i , » . i ............................. L x . .... R ianakanaenais (p r 46 ) — Peraeopods 6 & 7. segments 5 & 6 subequal in length; pleon plate I, hind comer acuminate .......... - P \ orientalis (p.47 ) 4. Eye medium, covering anterior ha If of head; gnathopod 1 . dactyl long, extending >50^. of its length beyond palm; coxa 3, anterior process strong, length >3 X basal width; tel son, lateral margins with 7-8 spines , . , , , P, osbomi (p. 38) — Eye large, covering 3/4 width of head; gnathopod 1, dactyl medium, extending 50% of its length beyond palm; coxa 3, anterior process medium length 2-3 X basal width; telson, lateral margins with 5 6 spines T - 5. 5, Antenna ] strongly setose posteriorly on flagellum and peduncular segment 2; gnathopod l, propod distinctly shorter than carpus, dactyl basaMy broad, thick; coxa 3, anterior process medium, length > 2X basal width P. carirmut (p. 42) — Antenna 1, flagellum and peduncular segment 2 moderately to weakly setose posteriorly; gnathopod I; pnopod and carpus subequal in length, dactyl basally slender; coxa 3, anterior process short, length < 2X basal width T . ... P. mix Ufa \e (p. 44) Pleopods medium, rami - 1 2 -segmented; pleon plates I - 3* hind comers squarish or obtuse, Uropods I . peduncle, anterior (outer) margin richly setose, apical spines of rami elongate, Uropod 2, inner ramus the sborL inner margin with 2 medial long spines, Uiopod 3, outer margin shorter, outer margin 4-5 spinose, Teison, lobes slender, basal 1/4 fused margins with 7-8 short spines, apices acute. Mate- (5,0 mm): Eye very large, broadly reverse- reniform, covering 5/6 head width, Atitennae 2 longer than antenna 1, brush setae present on the posterior margin of peduncular segment 2, antenna I s and the anterior margin of peduncular segment 3 & 4 of antenna 2; flagellum lacking feeding setae, Gnathopod I , pnopod mo re slender and palm virtually lacking; gnathopod 2, propod longer and more slender, and palm very much shorter, than in female. Pleopods, peduncles strong, massive, nearly 2X longer than in female; split- tipped clothespin spines on 5-6 proxi- mal segments of inner ramus, tiresome, mid-dorsal carina elevated, not mueron&te behind; fused urosome segments 2 & 3 with mid-dorsal notch. Uropod 2. inner margin of peduncle with a few plumose setae; inner margin of inner ramus with 3 slender spines. Uropod 3 T outer ramus slightly the shorter, outer margin with a few spines, all other margins (of both rami) heavily plumose-setose. Tblsoti relatively shorter, broadest medially: lobes more deeply separated, irtergins less spinose than in female. Distribution: Commonly encountered in teste of Amaroudum (Skogsberg & Van sell, 1928), from Central California north to Eri lish C olumhia and southeastern Alaska, questionably southward to the Gulf of California and Galapagos, The probability is high that P. osborni is a complex of sibling species over such a broad geographical range. Taxonomic Commentary: The female ol the present materia I compares closely w r i th the original figu res of Caiman (fig. 19, above) based on material from Puget Sound. Par- ticularly diagnostic of the species is the small palm of gnathopod 1, greatly exceeded by the dactyl. The species Folycheria antarciica (Stebbing. 1888), described origi- nally from sponges in the Antarctic mad ANZAC region^ is not a true synonym of P. osbomi, but is a distinctive species that exhibits general ly more plesiomorphic characters states (p, 49, fig, 25). AMPHiPACIHCA VOL I NO 3 OCTOBER 15. 1994 40 FIG, 19, Pvlycheria osborni Caiman, Kirby Pt T Diana I T Barkley Sound, Male 0,7 mm) AMFfflPACIFICA VOL- 1 NO. 3 OCTOBER 15< 1994 41 UROS MXPD FIG. 20, Polycheria osborni Caiman* Female ov. (7.0 mm) Puget Sound (modified from Caiman, 1S5>S) Polycheria cannot*, new species (Fig- 21 ) Materia] Examined: BRITISH COLUMBIA: Mainland eo®s£ Athlone 1., ELB Stn, H53 T under boulders, LW, Aug, 7, 1964- 1 female ov (5,8 mm) Pa*aiype( slide mL) CMN Cat. No. NMCC1994'0392; 2 additional females, S. end Vancouver I,: Taylor L Trevor Channel ELB 3tn, P5c, from ascidi^s and sponges beccaft boulders, LW, July 25. 1975 - 1 female br. II (4.0 tnm) (slide mt X MoCaulay PL Victoria, B, C„ GW O'Connel dive coll., Aug, 26. 1976 - 1 female ov. (4.0 mm) Holotype (slide ml.) CMN Cite, No, NMCC 1994-0390; 6 female, 1 subaduk male speci- mens, Paralyses, CMN Cat No, NMCCI994 0391, Diagnosis Female br. II (4.0 mm). Urosome segment 1 and fused segments 2 & 3 dorsal ly anddorso-laterally sharp- ly ridged or keeled, not acuminate behind. Eye large, red or black (in alcohol), covering anterior 374 of bead width. Anterior head lobe very broadly rounded. Anlennaesube^ual. flagella and distal peduncular segments richly armed with iongish food-gathering (feeding) setae. Mouthpans typical of N. American generic subgroup. Maxilla 1, outer plate, apical spines relatively long, palp short. Max illy 2, Outer plate, apex subtruncate, weakly setose. Maxilliped, palp very short, dactyl small; outer plate with 10 inner marginal spires. Coxa 1 acutely produced anteriorly; coxa 3 moderately produced, tength> 2X basal width; coxa 4 blunt, rounded in front. Gnathopod 1. propod relatively short and deep, lower margin with several stiff setae; palm very short, dactyl normal slender (in paratype), large, heavy, basally thick or broad, apparently abnormally developed in holotype. Gnathopod 2 more slender, carpus and propod subequal in length, palm very short. Peraeopods 3-7, segment 5 shorter than segment 6. Peraeopods 3 & 4 S basis relatively heavy, broader than distal segments, Feraeopods 5 - 7, bases narrow, slightly broad- ened in 5; segmeni 6 with re l a lively strong antero-distal cluster of setae. Pleopods medium, rami 12- 14 segmented Ficon plates 2-3 1 hind camera squarish, noi acuminate; pleon 3 setose below. Uropod I. peduncular anteriorly line with setae; rami clsely subequal apical spines not elongate* Uropod X rami much longer than peduncle, apical spines short, Uropod 3 f outer ramus slender, length about §0% inner ramus, outer margin with 2-3 short spinet olhe rroargins spinosc. Telson lobes nairowing distal ly. fused in basal 1/4. outer margins w f ith 5-6 small spines. Distribution Know n from Southern Vancouver 1, north to Athlone I central B. C. coast. Host unknown. Taxonomic Commentary:. The species Is closest to P. mixillae in most characler$tale$, but is distinguished mainly by features of the key (p. 40), AMPHIPACIFICA VOL. t NO. 3 OCTOBER 13.1994 42 FiG. 21. Polycheria carinata, new species, McCaulay Point, B. C. Female Br, II (4.fl mm). AMPHIPACIFICA VOL L NO. 3 OCTOBER 15.1994 43 Polycheria f tin: iliac, new species (Fig. 22) ' Material Fxatnmed (CMN collections, Ottawa): BRITISH COLUMBIA; S. end Vancouver L: Diana I., Kirby Pu ft. Anderson coIL from sponge (Mmlla inemstam), June 25, 3976- 1 female hr II (4.0 mm) HoLotype (nlatic mt), CMN Cal. No. NMCC 1994-0393; 9 Other females, Paratypes. CMN Cat No. NMCC1994-0395. Bordelais Islets, entrance to Trevor C ha nnel. ELB Stn, P20c, LW, in sponges and tunicate* tundci. ) collected from rocky walls of surge chaniids, Aug, 9; 1975 ■■ 1 Female hr, IT (4,0 mmHslide mt,). Diagnosis. Female ov, (5,0 mm). Urosoine 3, dorsal carina low, not produced posteriorly. Urosotlle segments 2 & 3, carinac or ridges inconspicuous. Head relatively shallow, anterior head lobe strongly rounded. Eyes very large, ovate, weakly faceted, covering anterior 3/4 of head Antennae subequal, slender. Antenna 1, segment 2 poslero- d is tally with longish setae; segment 3 shod, flagellum -16* segmented, moderately strongly setose, setae long. Antenna 2, flagellum 3 -segmented, Lower lip broad, inner lobe* large Mandible, spine it™ with 2-3 blades. Maxilla J, inner plate with 1 apical seia: outer plate with 7 slender apical spines; palp short Maxilla 2, inner plate small* weakly setose apical! y; outer plate, apex subacute. Maxilliped palp short, dactyl sloul; outer plate, i nner margin with 7-tf weak masticatory spines, Coxa 1 , anterior process short, with 2 apical setae. Coxa .3, anterior process relatively short, with single apical seta; coxa 4, anterior lobe rounded, Gnathopod l, basis lacking hind, marginal setae; propod shorter thaw carpus, lower mar- gin dislaJJy with 5-6 stout seme; palm short, (acceded by nearly 50% of slender dactyl when closed. Gnathopod 2 slender, propod much shorter than carpus, palm distinct, barely exceeded by simple dactyl. Peraeopods 5-7, segment 5 shorter (or not longer] (hart segment 6, Peraeopods 3 & 4, basis heavy, broader than distal segments, Peraeopods 5-7, bases subl inear, very slightly broader in peraeopods 5 & 6; segment 5 shorter than in peraeopods 3 & 4; coxa 7 produced posteriorly, subacute. Pleopods medium, rami - 1 3-1 f egmejit&d Pleon ptatcs 1- 3 broad, hind comer* squarish or obtuse. Uropod 1. peduncle, anterior margin strongly setose; mmi slender, suhecjual, apical Spines elongate. Uropod 2, rami longer than peduncle,, inner minus short, inner margin with 2 longish slender spines. Uropod 3, inner ramus with inner marginal spines and a few setae; outer ramus shorter, outer margin lined distally with 3-4 short spines. TeJson lobes ha sally one* fourth fused, narrowing dist- ally T margins distal ty with 4-6 short spine?, apices acute. Coxal gills large, sac* like, weakly pleated, on peraeopods 2- 5, smaller on peraeopods 6 & 7. Brood plates sub linear. Mature male Luide.scribcd, Etymology; The root name refers to the genus of sponges, Mlxithi w i|h which the amphipod species appears to be oommensally associated. Distribution. Known only from the Barkley Sound region of Vancouver E. Commensal on Demospongia (Mix ilia in crust anO Taxonomic Commentary: The species is closely related toP. catittam within the North American taxonomic complex of species. P, nuxiHae is distinguished from it by Characters provided in the key (p. 40), by the somewhat less strongly reduced palp of the maxLLliped, and by ihc mom setose inner ramus of uropod 3, WESTERN PACIFIC SPECIFY OF POLYCHERIA. Tile principal character states of the three specie* of Patycheria, previously described and figured from die west- ern Pacific region, are here summarized for inclusion in analysis of relationships of the North American Pacific fauna (see al*o Table III, and Fig r 31f Polycfteritt japonica Bulycheva (Fig, 23) Polyeheria japofflea Bulycheva, 1952 ; 233.— Barnard & Karaman, 1991: 272. Taxonomic commentary: The original description and figures were based on a male specimen, but pertinent nor sexual character stales arc here summarised; Fused urosome segment* 2 & 3 bearing small dorsal spines and paired lateral ridges. mHcro* laterally masked by posterior projection of urosome segmeni 1. Antenna 1, peduncular segment 3 longer than adjacent flagellar seg- ments. Mandible, left and right molars unequally reduced. Maxilla L outer pi ale with 9 apical spines; palp large. Maxilla 2, inner plate strongly setose. Max Wiped, palp medium, slightly exceeding tall outer plate. Coxae l it 2 anteriorly rounded below. Coxa 3 lacking anterior process. Gnathopod 1 . propod relatively short deep; palm large, not exceeded by dactyl, Gnathopod 2. propt.xl slender, subequal in length to carpus, palm distinct, Peraeopods 3-7, segment 5 larger (noi smaller) than segment 6; bases stout somewhat broadened, Pleon plate* 2-3. hind comers acuminate. Uropod 2, outer rani us the Shorter, Uropod 3, outer ramus the shorter, outer margin spmose. Telson lobes narrowing distal I y, fused in hasal one-sixth, margins weakly spinose. AMPHIPACIFICa VOT.. 1 NO- 3 OCTOBER (5, 1994 44 FIG. 22, Polycheria mixilUie* new species. Diana L, Barkley Sound, Female ov (5.0 mm) AMPHIPAC1F1CA VOL. I NO 3 OCTOBER 15, 1994 45 FIG. 23, Polyeheria japonica Bulycheva. Mate (5.0 ttifri), Peter-the-Greal Bay. Poiyeherkt amakasaettsis Hirayama (Fig, 24B ) Potycheria amnkuxaertsis Hirayama, 1984a: 194, figs. 1Q<5- 108, — Barnard & Karaman, 199 J: 27 L— Ishimam, 1994; 43. Taxonomic Commentary: Hirayama' & descriptions and figures f ]j)g--C kj pertain essentially to a male specimen, but pertinent non -sexual character states are here summarized: Fused uRisome segments I & 2 with paired lateral ridges, basally masked by posterior projection of uro&otne segment I, Antenna 1, peduncular segment 3 Linger than adjacent flagellar segment; flagellar setation probably as in P. oriental: s. Mandible, left and right molars unequally reduced, Maxilla I, outer plate with 9 apical spines; palp long, Maxilla 2, inner plate with strong medial setae. Maxilliped palp medium, about as tall as outer plate. Coxae 1 & 2 rounded below. Coxa 3 rounded anteriorly, Gnathopod I, propod subovate, lacking palm; dactyl short, strongly curved, Gnaihopnd 2. propod slender, shorter than carpus, palm and dactyl short. Peraeopods 3-7. segment 5 larger (not smaller) than segment 6; bases little broader than distal segments except in peraeopod 5, AMPHIPACIFICA VOL. I NO. .1 OCTOBER 15, 1994 46 KTG* 24, Polychena species . West Ky us hu> Japan* (after Hir ay ama, 1984)* A. P. orienialis Female (4*5 ram)* IS* P> amakusaensis Male (4*5 mm), Pleon plates 2-3. hind comers acuminate. Uropod 1, rami subequal. Uropod 2, outer ramus the shorter, Uropod 3, outer rajnus slightly the shorter, outer margin weakly spinose. Telson lobes of female not described (probably as In P. orients Its), Polyckeria orientalis Hirayatna (revised status) { Fig. 24A ) PofocheritiQtoliiorfcnHilis Hirayama 1934a: 1 £7, figs. 10 J, 103-1 05 . — Barnard Sl Karaman, 199!: 272, — Ishiniaru. IW; 43. Taxonomic Commentary: : The pertinent taxonomic charae ter slates of H irayama' s description and figure s, based on a female specimen, are summarized here: Fused nroaome segments I & 2 (one illustration shows an inter- segmental line ! 1 with small spines and paired lateral ridges, based parti y masked by posteriorprojection of tiresome I . Antenna 1, peduncular segment 3 longer than adjacent flagellar segment; flagellum richly anned with feeding setae. Mandible, left and right molars not shown, probably as in R amakusaensis Maxilla U outer- plate with 9 apical spines; palp large. Maxilla 2- inner plate marginally setose, AMFHIPAClFlCA VOL.1 NO. 3 OCTOBER 15,1994 47 FIG. 25. Polycheria anturctica species complex 1. acanthopoda Thurston . 2. dentata Schell . 3. gracilipes Schell . 4. nudus Holman & Walling, (modified from FloJman & Wat ling, 1983} AMFHIPACIFJCA VOL. I NO. 3 OCTOBER IS, J994 48 Maxilliped, palp slighily exceeding tail outer plate. Coxae L 2, & 3 rounded ante ro veolrally Gnathopod !. propod and carpus stibequah palm short, barely exceeded by dactyl. Gnuihopud 2, propod shorter than carpus; palm small; dactyl very small, hoot-like. Peraeopods 3-7, segment 5 Lillie shortened* distinctly longer than segment 6; bases suM inear but broader than in P. arnnkusaemti' Pieort plate 2 -3. hi ml comers acuminate. Uropud l . ram i subequal. Uropod 2, outer ramus the shorter, llropod 3, outer ramus the shorter, outer margin weakly spinose, Tel son Lobes long, narrowing distal! y, fused in basaJ ore-eighth, margins weakly spinose. EXTRALEVUTAL SPECIES Potycheria Antarctica fSlebbingl (Fig. 25.) Dexamtne Antarctica Siebbing 1875; 184. Triiaeto aruar&kti Stubbing 1888:451 Polyc tiffin antarctica Stebbing, 1906: 520* figs. 90, 91,- - Schcllenberg, 1931:214. — Thurston, 1974: IS.— Holman & Wading, 1 983: 22 L figs. 6-9 (including forms ncanthopoda Thurston; demata Sebel lenberg \gracUip?s Sehe Elenberg; nud- us (Holman & Wading),— Barnard & Karaman, 1991: 271. T&xonnmlc Commentary: Pertinent taxonomic char- acter slates from an assemblage of Tormae"of P. amantka (cf. Holman and Wailing, 1983), restored as distinct species of die antaretka complex by Bam-anl& Karaman (Inr. riri , provide broader perspective to the analysis of North Pacific species relationships fp. 61. Eg, 31). Fused urosotne segments 2 & 3 dorsal ly with 4 spines, and paired lateral ridges. Urosome I with low dorsal Carina, not produced postern- laterally to conceal base of urosome 2. Antenna I, peduncular segment 3 slightly longer than adja- ceni flagellar segment: antennal flagella setose. Mandibular molars probably unequally reduced (cf, illustration of Stebbing, 1906). Maxilla t, auierplaic with 9 apical spines; palp medium, slightly shorter Lhan outer plate. Maxilla 2, inner plate with sparse inner marginal setae. Maxilliped, palp little reduced, exceeding tall outer plate. Coxae l & 2 rounded below. Coxa 3 with strong aniero- ventral process, Gnathopod I, propod slender shorter than carpus; palm medium, little exceeded by dactyl. Gnathopod 2, pro pod shorter than carpus, palm relatively large, not exceeded by dactyl, Feraeopod 3-7, segment 5 reduced, shorter than 6: bases sublinear. little broader than distal segments, Fleon plate 2 &. 3, hinti comers weakly acuminate, Uropod 1 Jnnei ram ucdisii tic Ely the shorte r. Uropod 2, ram subequal. Uropod 3, outer ramus much the shorter, outer margin nearly bare. Telson, lobes elongate, separated nearly to base, margins distal ly bare or weakly spinose, apices each with spine. DEXAMINOCIJLJNAEi new subfamily (see Fig. 2(b):26) lilccrLic- scdls , Barnard. 1969a: 480* fig, 1 7 3a. De xa mi nidae ( part ) Leftover, 1979:65.— Lowry, 1981; 190. Prophlianiinae Barnard & Karaman, 1991; 273 (key) (part). Type genus: Dexaminocuiu^ Lowry 1981: 191. (Spiiaer- vpfithalmus Spandl. 1923). Diagnosis: An Indo-Pacific monoiypic group, of unu- sual morphology, about w hich little is known except for the studies of Lowry fl oe, cit. ). Body smooth or weakly toothed on peraeon. Pleon segments and urosome I, each wilh mid-dorsal cari nation and postero-lateral marginal teeth or cusps. Urosome seg- menis I & 2 ridged mid -dorsal I y and mid-lateral ly. Rostrum medium, slender. Eye large, on produced lateral cephalic lobe. Antenna 1 elongate (boih sexes), accessory flagellum vestigia L Antenna 2 very short, flagellum vestigial (female), elongate, with peduncular brush setae (male). Mouthparts nearly regularly dex&ninid. Mandibular molar irituraiJve. blades few, Maxilla I , outer plaic with 1 1 apical spines. Maxilla 2, ptates not slenderized. Max- ill iped, inner plate small; palp 3-scgnaeoted (female). Coxae 1-4 medium, unequal. 3 smallest (allowing for respiratory current exit?), lower margins cumulate and/or setose. Coxa 5 large, anierolobaie. Gnaihupods dissimilar in size and form, distinctly subchelate. Gnathopod I . propod sexually dimorphic, somewhat as in the typical dexamirtid. but with the dorsal notch reduced to a shallow depression,, and Ehe palm deep! y excavate, rather than con vex, Peracopods 3-7 slender, regular (noi subeheJate); peraeopod 5 slightly the longe.sE, Peraetipods 5-7, bases dissimilar, variously broadened and lobatc below: segment 5 not shortened; dactyls slender. Fieon plates large; pleon plates 2 & X postern lateral margin uw>Ehcd, hind comers acuminated, hooked. Pleopods roc described. Uropods I & 2 large, regular; uropod 2 short. Uropod 3, rami large, broadly lanceolate. Tdson large, elongate, lobes not diverging apical ly. Coxal gills and brand plates not described. Species: D-extiminoculusai' utipei Ledoyer, 1979 (Mada- gascar); D. cavimamtx Ledoyer, 1982 (Madagascar); and D. groebbetii (Spandl, 1923) (Lowry, 1981) (Madagascar to Australia), Taxonomic and ftiugwigrsiphfc Commentary: The genus Dt>xitmmacuhn was first described as Sphaer~ ophrhatnum by Spandl (1923) and placed iu taxonomic cat- egory irtccrta icdifc by B arnaid ( 1 969a), Two further species were described* both from Madagascar, by Ledoyer ( 1 979, 1982). The genus is narrowly IndoFacifiCi not yet known from Japan and the North Pacific region, but might be anti- cipated at the northern limit of coralline substrata. AMPHIPACIFJCA VUL ] NH .1 OCTOBER 15, 1994 49 FIG. 26* Dexaminocutus grobbetii (Spaiidl). Female (3.6 mm) Male (3 t 9 mm) Great Barrier Reef, (after Lowry, 1981)* AMPHJPACIFICA VOL. r NO 3 OCTOBER 15, 1994 The genus was renamed and fully redescribed by Lowry t. Iny £jj), based on more complete material from the Great Barrier Reef of Australia (Fig. 26), He likened ii most closely to the germs Dexiimiwlta Schellenberg (1928), On questionable grounds, Barnard & Karaman (1991) placed the genus within their realigned subfamily Pnophliantirtac. However, as Lowry (R ^ L _£il. ) and Ishimam (1987) con- cluded, the balance of character states of Dexaminocuktx are closer to the true dex am in ins . Dexa m in? , Pa radexamin e an d espee tally Dexamine tfa (Figs-, 2 (c ); 29). Parti cularlv sigoifi- cam is the form of the coxal plates, pi eon carnation, and the sexually dimorphic gnathopod L as well as mouthpart mor- phology. However, the extreme location of the eye is non dexamiiun. and die lack of subchelae on the perueopeds is non polycherijn. The author therefore propose the new subfamily Dexanrinoculinae to facilitate recognition of its distinctive, major, taxonomic differences. PropMantinae Nicbolls (see Fig. 2(d)) Prophliantidae: NiebolLs, 1939:312. — Barnard, 1969a: 432, — Bousfidd, 1982: 278.— Ishimam, 1994:43. Dexaminidae (part): Barnard, 1970a: 163- — BeUan-vS&ntini, 1982: 212. Dexaminidae (Prophliantinae) Barnard, 1970: 161: Ishimani, 19&7: 1413 — Barnard & Karaman, 1991: 273. Type genus; Prophlkis NicholU, 1939:312. Genera: Guernea Chevron*, 1887: 302 {=Fma&sus, -Dexamonica ) ; Hmsio rftfflsis Sche llenberg , 1938:12, Diagnosis: Body small, short, broad, surface often with rugose integument. Feraeon with low tttid-dorsaJ carina (part or all), but no dorsal processes. Urosome segment I may be fused with fused segments 2 & 3. Rostrum very short. Anterior bead lobe mainly rounded. Eyes pigmented, me- dium. Antenna 1 (female) short, peduncular segment 2 shorter than 1. Accessory flagellum minute or lacking. Antenna 2 (female) short: in male, peduncle short, segment 4 broad, flagellum elongate. Mouthparts modified. Lower lip, inner lobes distinct. Mandible; molar variously reduced or modified: spine row lacking. Maxilla 1 * palp I (2) segmented: outer plate with 7-9 apical spiles. Maxilla 2. plates modified, reduced, MaxilUped, outer plate large, inner plate small, palp short- ened Coxae 1-4 slender, deep: coxa l shortest. Coxa 5 very large, Gnathopods slender, weakly subehelaie; carpus usu- ally longer than propod; palmar margins small, distinct,. Gnathopod l, propod not sexually dimorphic. Peraeopods 3 & 4 simple, not subehelaie, segment 5 not strongly shortened. Peraeopods 5-7 short, generally dissimi- lar in form hut little in size (petaeopod 7 shortest); bases variously broadened, unlike; segment 5 little shortened often broadened; dactyls simple, short to medium. Pleopods small: peduncle broadened, rami short. Urupods I & 2 short rami usually unequal in length. Uropod 3 short, margins spinose (weakly setose in male), Tel son lobes medium, separated nearly to base, not diverging, apices truncate, spinose. Coxal gills simple, not strongly pleated or 1 abate, on peraeopods 2-6 only. Brood plates small, linear, with apical setae. Taxonomic Commentary : The authors concur with the decision of Barnard ( 1970aj followed by Hirayama ( i 984, 1986), to transfer Guemeu from family Dexaminidae to the Prophlismlisiae. Cluster analysis (p. 56, Fig. 29) further confirms Us relatively dose morphological similarity to Praphlias and tiausioriopsis. Guemea is a complex of diverse species groupings* some of which have been giver formal generic and/or subgeneric status (Prinassus in the N, Pacific region and Guernea elsewhere)* However* the a uthors aho agree with the decision of Bdlan-S antlni (19§3) and Ishimaru ( 1987) to resubmerge the names Prinassus. and Demmomca in the syncmoiiiy of Giiemea Chevereux, 1887, Barnard and Karaman (1991, Inc. eh .) reduced the Frophlianlidae to subfamily status within the Dexaminidae. This decision Ls supported by the present analysis (p, 56). As noted by Tshimaru \ 1 987), those two authors also relegated the genus Demminoculus to the Prophlianiinae on dubious grounds, arid as noted here, without suitable concordance with their own subfamily diagnoses* The coral-dwelling Dexaminocuhis is here considered distinctive at subfamily level (above). In balance, its phyletic affinities Eire; closest to the primitive, nestling Dexamintnae, and rather remote from the l'ossorially specialized and apomorphic PinuphlLamirtae. Guerttea Chevreux Guemea Chevreux, 1887b: 302,— Slcbbing, 1906: 521 (part). — Barnard, 1970a: 11, figs. — Hirayaimi, 1985; 395,— Bcllati-Santini, 1982: 225. — Ishimaru, 1987: 1395.— Ishimaru, 1994: 43, Guemeu {Guemea) J.L, Barnard, 1970a; J 69 h — Hirayama, 1985:1. — Hirdyttma- 1986:488 Barnard & Karaman, 3991: 274. Prtnqs&us Hansen, 1888: 82. Guemea (Prinasm^ T, L, Barnard 1970a: 169. — Hiraymna, 1985: 8 — Hirayama, 1986a: 493. — Barnard & Karaman. 1991: 275. Dexamonica J.L. Barnard. 1958: 1 30, pis, 26-27. — Barnard, 1969a: 203. Type Species: Helleria coalite Norman, 1868, AMPHfPACIFICA VOL. I MO. 3 OCTOBER 15. 1994 51 Species: About 24 described species and subspecies world -wide (Barnard & Karaman* 1991, updated). The following 1 1 species ace recorded from the North Pacific region: G. ezomxis Ishknaru, 1987: G iongirfaciyia Hirayama, 1 986a; G, mackiei Hi ray are a, l9S6a: G, mag- naphUostomci- Hirayama, 3985: G. minor Ishinwu 1987; G. nuliispina Hirayama* 1885; G. quadrispitwsa Stephenson. 1944; G. rectocephalux Hirayama, 1 935: G, redunaina J.L. Barnard, 1958; G. sombaii Hirayama, 1986a: G. lereftimina Hirayama, 1935; G. tomiokmnsix Hirayama, 1985. Diagnosis: Posterior peraeon and all pteon segments weakly carinated and/or posteriorly mucranate. Urosomlte 1 separate, with mid-dorsal keel or hump (both sexes), tfeoSOTdtes 2 & 3 coalesced, variously with small dorsal spines. Rostrum very short; anterior head lobe sharply rounded. Eyes medium, rounded, weakly faceted. Antennae (female) short. Antenna 1, flagellum 4-8 segmented; acces- sory flagellum minute or lacking. Lower tip large, outer lobes with prom bent shoulder cones. Mandibular molar variously triturulive* often com- plexly divided; left lacinia 4(5) dentate. Maxilla 1, palp l(2)-segmentedt outer plate with 7-9 apical spines, inner plate 0( D-setose. Maxilla 2. inner plate small: 2-5 setose. Maxilliped, inner plate very short, apex with 2-5 long setae; outer plate large, palp 4‘segmented, dactyl short. Coxae 1-4 medium, narrow, strongly overlapping, rounded below, Coxa 5 very large, deep, ijosterodobaie. Gnathopods I slightly smaller than 2, basis with distinct proximal “buccal bend"; carpus relatively short and deep, little longer than propod; palm destine L Peraeopods 3 & 4. segment 5 shorter than 4 & 6, posterior margin spinose; daclyls medium. Peraeopods 5-6 subsiiniiar in form and length; segment 5 not shortened dactyls various, usually reversed. Pcraeopod 7, basts very broad; segment* 4 & 5 broadened (not greatly, and/or y symmetrically, as jn fltiuiioriopyis), margins strongly setose; dactyl short. Pleon plates 1-3, hind corners rounded* or squared, Uropods 1 Si 2, outer rainus be longer {usually ). apices with long apical spine. Uropod 3, rami short, subequal* inner margins spinose (setose in male). Telson lobes medium, not diverging, outer margin and apex variously armed with setae and/or spines, Distribution; Mainly tropical and warm -temperate (Indo-Padfic and tethvan) coastal shallows; fossorial in fine sediments. Of the 24 species and subspecies described to date wor Id-wide, 12 (one half) have been recorded from the North Pacific region, but only one of these from the North American Pacific coast. Taxonomic Commentary: In balance of character states. Guernm appears more closely related to the type genus Pro- phiias than to the more highly specialized genus Hauti- oriopsis. It differs from ProphUas , however* in iKskon ger gnaihopods. unexpanded segment 4 of peraeopod 5* and its dorsal Jy carinated, unfilled urosome segment I . Guzrrtm Fedurtcttn# (J. L. Barnard) (Fig. 27) Dexatnotlicd redunctuw J, L, Barnard, 1958: 130* pis, 26, 27,— Staude, 1987: 382. GuemeafPriaassus) redunegtu Barns'Ll 1970a; 1 '73-. figs. 3-3 — Barnard Si Kamman, I99J: 275. Guentea reduncans Austin, 1985: 604, Material Lxu mined: BRITISH COLUMBIA: Queen Charlotte Islands, ELB Sins, 1957: H4a mouth oi'Yakoun Bay* July 19- 1 female with juveniles); W 11. Head of G tidal Bay, Graham I., July 28 ■ I imm, Vancouver 1.. ELE Sim, B27, Dodger Channel, SW end Diana I. r July 8* 1976- l mate*! female ov. OflMcCauley* Pl* Victoria* B. C.> G. W. O’Connell Stns.* Aug, 28, 1976: WJOB - I male; WI56B - 1 male t 3 females (ov) Cfig’d. specimens). Off Victoria, C. Low colL Aug., 1981-3 males, 5 females, 10 imm. Diagnosis: Female ov, (2,4 mm); mate (2.5 mm). Pcraeori segments 6 & 7, and pteon segment 1-3 will] low mid-dorsal ridge, slightly acute behind, Urosotnite 2 with recurved mid-dorsal canny. Fused urosomltes 2 & 3 some- whal humped nud-dor sally, with 2-4 small spines. Eye medium, subovate* about 25- faceted. Antenna I. flagellum 5-segmentcd; peduncular segonents 1-3 (male), anterior margins minutely ere nutated; segment 1 deep, posterior margi n dis tal ty w i th c I usters of long isli hntsh setae. An te nna 2 1 flagellum 3-segmented: flagellum (male) elongate (20+ segments), peduncular segments 4 & 5 enlarged, anterior margin of 4 wilh clusters of short brush setae, Mandible, grinding surface of molar modified but en- tire. distal plumose sola short; left lacinia 4 -den tale. Maxilla 9, outer plate with 7 apical spines; palp I segmented* apex with 2 setae. Maxilla 2. inner plate narrow* with 5 marginal setae, MaxillipciL inner plate with 3 long apical setae; palp segment 3 and short dactyl exceeding lall outer plate. Coxae 1-4* lower margins finely crenulate and weakly setose. Coxa 5, anterior lobe small rounded, hind lobe very large, deeply rounded, Gualhopod 3, carpus and propod relatively short, deep, subequal in length; fotspod widening distally to convex palm, with 3-4 poslero-dlstal spines. Gnathopod 2 slightly larger than gnathopod i; carpus and propod slightly more slender and elongate: palm of propod with 3 poxtertvdislal spines. Peraeopods 3 &. 4, .segment 5 distinctly shorter than 4 & 5, hind margin with 3 stout spines increasing distal ly; dactyls medium. Pcraeopods 5 & 6, segments 5 , 6, and dactyls re- versed; basis Of peraeopod 5* hind lobe not strongly prod- uced below; segment 4 normally broadened, Peraeopod 6. AMPHIPACIP1CA VOL. I NO. 3 OCTOBER 15. 1994 52 FIG* 27* Guernea reduncans (Barnard)* OIT Clover Pt*, B. C Fern* (2*3 mm) Male (2*0 mm) AMPHIPACIFICA VOL. I NO- 3 OCTQRFR 15,1994 53 basis narrowing dis tally. hind margin nearly .straight, nol markedly concave. Peiaeopod 7, segments 4 & 5 not exceptionally broadened, length of each greater than width; dactyl slender* medium. Pleon plates 2 & % hind comers squarish or rounded, lower margins weakly seto.se. Uropod J, tips of rami exceed- ing uropod 2 but not uropod 3; peduncle with 34 proximal outer facial setae. Uropod 2, outer ramus Hie longer, apical spine about 2/3 its length. Uropod 3, rami about 50% longer than peduncle, margins with a few stout spines; in male, inner margin or both rami are pJuraose-sdhm Telson not Longer than wide, lobes fused basally. submarginaJJy with pemcillale setae, apices each with single spine. Distribution; Southern British Columbia, Washington and Oregon, to .southern California, subtidalJy to about 100 in. in deplh, in fine sand and muddy sand, The present records are the first authentically from British Columbia- Taxonomic Commentary: The species apparently varies somewhat throughout its range. Material from Cali- fornia, illustrated by Barnard (1970a, cif) . exhibits distinct, posteriorly mucronaie, personal and pleonal carmatioiis, and more elevated dorsal tooth on urosomtte I . Uro somites 2 & 3 bear 6 (vs. 24) dorsal spines, and apical spines of the uropod rami are longer. In southern material, the eye of the female is smaller, the flagellum of antenna I is 6- (vs. 5-) segmented, the posterior spines of .segment 5 of pci aeopods 3 & 4 arc longer, the posterior lobe of the basis of peraeopod 5 is deeper and, in peraeopod 7, segment 5 is shorter and broader. Moreover, in maxilla ! of Californian material the palp has a weak suture dividing it into two segments, the outer plate bears 8 apical spines, and (he inner plate a single apical seta. In males, the eye of northern material is larger with more numerous onirnatidia, Guernea reduncans appears more ciOSdly similar to G, coaika and G, nordmskioldi of the North Atlantic region than to species of the western Pacific described and figured by Hirayama (I9S5, 1986a) and Ishimaru (1987) (see be- low ), WESTERN N. PACIFIC SPECIES OF GUERNEA To date eleven species of Gitenieo have been receded and/or newly described from Astatic North Pacific localities as follows: L Sea of Japan Sea. Russian Coast, 1. Guemeti species {identified as G, nordeftskiotdi by Bulycheva, 1955). II. Coast of Hokkaido i material of Ishimaru, 1987), 2, Guenxea ezoemis (males, females) - Otsucbi, Noisufce peninsula. 3, G mnor (males* females) * Sbirahania, II, West Kyushu coast, Japan (material of Hirayama, 1985): 4, Guernea ttt&g nop h i to .v t&ma. \ males, females) - Ariake Sea, 5, G. wreiamim (female) - Shijiki Bay. 6, G, lomlakaertsis (females, mates) Tomioka Bay. 7, G, iwllispin# (male. im matures) - Tomioka Bay. 8, G. reemcephah (females) - Tomioka Bay, III. China Sea Coast (material of Slephensen. 1944), 9, Guernea quadrifipinosa (male) - Liao-tung peninsula. TV. Hong Kong (material of Hirayama, 1986), 10, G.uernva wmbati (male, female), 1 1, G. lotigidoayta (male). 12, G, tftfccfcrei (males, females). Taxonomic Commentary: The above species from (he Japan and China Seas exhibit a considerable range of mor- phological diversity. However, a reasonably close perusal of illustrated charactei states did not reveal well-defined sub- groups but rather a series of morphological specializations that presumably adapt each species for a particular niche Eind life style. The species range phyletically from the relatively primitive G. emerisis t in which most chEiracter states are plesiomorphie. to t be highly specialized minute species. G. minor, in which most character stales are apomoiphic, None closely resembles the type species. G. ctmtiia (Norman) from the North Atlantic region* but differs especially in the form of the gna thopods, and in the shape of the mid-dorsal process of urosome l. Barnard (1970a) has previously commented on differences belween the material of Bulycheva ( 1955) from the Japan Sea (No, 1* above), and his material of G, mrdenskioldt from North Atlantic coastal regions, and of G- rvduncamrnns from California, The last two species were boih fully illustrated in his extensive generic revision (Barnard, 1970a), A Jce y to North Fau die species is beyond die scope of th i s study, However. G, re dune tins was found to differ from Species Nos. 2, 6, and 10 in which the outer plate of maxilla 1 has 9 apical spines; from Nos, 3. 5, 7. 8, and 12 in which the outer ramus of uropod 3 lacks plumose swimming setae iu the male; and from No, 9 m which the apical spines of (he rami of uropods 1 & 2 are extremely long. G, redmeana differs perhaps least from Nos. 4 &. 11 (above) but both the latter species have relatively slender gnathopods, and telsou lobes that are marginally and/or apically setose. Hopefully, this study may stimulate a thorough revision Of this challenge mg assemblage of western Pacific prophlianlids. AMPHIPATMCA VOL. 1 NO. 3 OCTOBER 1994 54 P4 PJ& FIG. 28- Prehensile Peraeopods in Dexaminoktea {modified partly from Vader, 1983) A. Nvtotrupis fa f coins B, Delkarlye ensamcUkt C. Tritacta gibhnsv D. Polycheria ubtusa Discussion and Conclusions* Tikis study treats the sys tenuities and distributional ecology of some 12 species of dexaminoidean amphipod crustaceans occurring in North American Pacific coastal marine waters, from the Bering Sea to Northern California. This fauna is small and relatively minor in contrast to several Large and diverse regional ganmuiridean superfaiiijlies pre- viously treated (e.g. Garnmuroidea (Bousfield* 1979); Ampeliscoidea (Dickinson, 1982, Coropbioidea (Conlan, 1983); Phoxoeephaloidea (Jarre tl & BoiiS field. 1994), and others of this series now in preparation (e.g. Talilxoidea, Eusiroidea. Had 2 iodeatB 0 usl 1 etdS 1 aiJ.de, 1994), Moreover, regional dexamiuoadeans include only aboui 7% of the —200 species described to date, world-wide, How- ever. tins small fauna is remarkable in containing: (Ha large component of the single most primitive subgroup, the sub- family Alylinae; (2) significant representation from the most advanced subfamily, the Polychcriinae; (3) only one species from the Other sixpbyletieally intermediate subfamily groups. Thus, in combination wilh oourilcipart feaminoidean groups from the Asiatic North Pacific coastal marine region, this modest North American assemblage makes up in taxonomic and ptiylcUc quality what it lacks in species numbers, and thereby provides a basis for review and rectessi fieation of the entire world fauna not previously realised. Natural relationships among species and generic groups are here tented more critically by means of a modification of the pbtocUc UPGMA {cluster analysis) system of Sneath and Sokal (1973), The modified but relatively unsophisti- cated system employs an overall criterion of phyletic simi- larity termed the Pletf o-Apomorphic (TV A } Index in which low numbers signify phyieiically primitive, and high num- bers advanced, taxonomic groups, The system has been utilized effectively in similar studies by Conlan (1983), Staude (198b) and Jarrell and Bousfield (1994). Within the superfamily Dexami noidea, analysis of generic similarities is based on 21 characters and corresponding 42 character slates given in Table 1 (p. 57). The lepechindiids arc here represented pragmatically by one gztn& t L?pechirtrit&, mainly hecaust it contains more than 90% of the species, and the three other described genera do not show differences (from it) in the character states utilized in this analysis. The resulting phenogrem (Fig. 29) ‘"clusters out' 1 two main subgroupings at Jess than 50% similarity, viz, a primb live, thin-bodied, atylid family group (with P. A. indices of 9-24) on the left, and a relatively advanced, broad-bodied, dexami md family group (with P,- A. indices of 1 5-29) on the right. The atylids are especially primitive in relaming a number of presumed ancestral features (c.g.Bousfieid, 1983) such as basic body cari nation, peraeopods, pleopods, mouth - parts, and pleated gill .structure, whereas tbedtexaminoideans lend more strongly to reduction or loss of body carination, moutbpart armature, and modification of the peraeopods towards "prehensiliiy” on the one hand (Fig.28, above) or fossorial l ife style on the other (Fig. 2 (d), p. 7), Within the Atylidae, four subgroups "cluster out” with paired character slate similarities between 60 and 75%, that are here recognized at subfamily level. These include the very primitive large-bodied Atylinae (P,-A, Index of 9) on the one hand, and the advanced small -bodied AuatyUnae (P,- A, Index of 22-24) on the other. The other two groups, Notolropiinae and LepechtnelLmae, intermediate in body size and phyletic positioning (P. A. indices of 16-19), trend to a more free-living, epihentbic and pelagic life style, with strong deep-water and abyssal components. The hiogeo- graphicai significance of these phyletic relationships is noted below (p. 60), Within family Dexaminidae, containing nearly twice the number of genera, four subfamily groupings are similarly recognized. These “cluster ouf at slightly higher levels of character stele Similarity (60-77%). These subfamily group- ings include the relatively primitive Dexamininae contain- ing six relatively similar genera (p. a. indices of 15-24) on the .left, and the advanced, highly specialized and commensal pah of genera comprising the Polydieriinue (F. A. indices of 27-29) on the right, The iwo phyietically iniennediaie subgroups (P. A. Indices of 19-21) encompass two sub- families of widely differing morphologies and life styles, viz. the monolyplc, coral -dwelling Dexaminoculinae. on the AMPHIPAC1FICA VOL. 1 NO 3 OCTOBER 15, 1994 55 TABLE 1. GENERA OF DEX AMIN O H) K A : CHARACTERS AND CHARACTER STATICS CHARACTER CHARACTER STATE VALUE plesiomorphic Intermediate ApumOrphic 0 1 2 1 . Rostrum long medium short 2. Body form very slender short and stout 3, Thorax, dorsum spinose not spinose 4. Accessory flagellum 1 -segmented minute lacking 5, Sexual dimorphism of strong weak or none antennae, gnathopds 6. Mandibular palp present, strong weak lacking 7, Mandibular molar large, triturative non-triturative 8. Lower Up, inner lobes lacking weak well developed 9. Maxilla L palp 2-segmented 1 ^segmented 10, Maxilliped palp 4-segmented 3 -segmented 1L Coxal plates M smallest anteriorly deepest anteriorly 12. Coxal plate 5 shallow deep (about = 41 13, Gnathopods 1 & 2, elongate short & deep propod & carpus 14, Peraeopods 3 & 4 simplidactylate. suheheiate 15. Peraeopods 3 & 4 , . > segment 4 < segment 4 « segment 4 length of segment 5 16, Peraeopods 5-7, broad, guborbicular narrow width basis 17, Peraeopods 5-7, similar in size and unlike in size unlike in size similarity form or form and form IS. Pteon, dorso- lateral toothed smooth armature (or nearly so) 19, Urosome 5 & 6 - present weak lacking dorsodatenit “wings" 20, Uropod 3. rami lanceolate; margins 1 inear: margins plumose-setose spinose 21, Telson lobes separate, converging basal! y fused one hand, and a complex of three small bodied, fossorial genera within the Prophliantmae on the other. The Dex- juninoeulmae and Polycheriinae are linked (laterally to Hie Dexamioinae by greater overall character state similarity of the peraeopods and most other body appendages, including similar sexual dimorphism of the propod of gnalhopod 1. apparently unique to this family within all gammaridean amphipod superfamilies (Fig, 2, p. 7). Close comparison ol individual character states suggests that the Prophliantinae differ from the Dexamminae somewhat more strongly than semi-phyletic numerical taxonomic methodology actually reveals, This methodology may be arguably more suscep- ti hie to homppMous or convergent sim Parities than c lad i s tic analytical methodology A broader cladistic analysis, rot AMPHTPACTICA VOL. I NO 3 OCTOBER 15, 1994 56 FIG, 29 ♦ DEXAMTNOmEA; PHE NOGRAM OF GENERA, ABODEFGHJKLMNOPQRS >- l- tt < _i W LU I- < I- c n cc LU I- o < < x o Q LU DC < CL 42 40 33 36 34 32 30 28 26 24 22 20 13 16 14 12 10 “i — i — i— i — i— i — i — i — j — r — i — r — i — r — i — i — i — i xn _ 9+ 19 1 6 1 7 22 24 16 17 19 17 19 2* 21 19 20 1 9 27 29 P.-A. INDEX G E N E Ft A AT Y L 1 D A E DEXAHINIDAE A Mym$ K - Dexaminetta a LepGG hiftatte L - Syndexainine c Abetratyfus M - Deikaryle D Not Off op ts H " Datarti^ocu^us E Anatyjtis O - Gu«msa F Kamehatylus P - Proptilias <3 D&xamina q Hauatorlnpafa H Paradexnmine Ft - Tritaetfi J SehBdsxius S Pofycharia £ 50 _ Cfl 25 al tempted in this regional study, may show greater phylelk significance to the differences, especially in gnat hoped structure, and perhaps justify restoration of the Pruphliantins to family level of recognition. Within thcmonotypic genus Afyfa.* (subfamily Aty linae). an amphi-Narth Pacific near-total assemblage of I 0 species may be phenetlcally analyzed, based on 20 characters and character states outlined in Table If The resulting phenogram (Fig. 30, p. 60) encompasses two not very closely similar subgroups, a primitive large bodied cariri&ius-leiidftisus assemblage (P. A, Indices of 10-2 1 ) on the left, and a more advanced, generally smaller bodied coHrngi-tridens assem- blage (P, A. indices of 20-33) on the right. The most primitive members of the carinatus subgroup, A. carinam and aitmsovu appear not far removed in basic morphology from large regionally occurring gammaroidean am ph ipods (e.g. various Anisogammaridae, and the Gammarm mosus -wilkUzkU complex within family Gammaridae: see Bouslield, 1979), Within the A, levidenms subcomplex. including A. ekmant and A. bmggeni , some reduction of inouthparts (e,g. mandibular palp) and specialisation of body appendages (e.g. pectinate setation of gnathopod AMPHIFACEHCA VOL. I NO .1 OCTOBER 15. 199 * 57 TABLE IT SPECIES OF ATYLUS: CHARACTERS AND CHARACTER STATES. CHARACTER CHARACTER STATE VALUE Pltfsiunwrahic f fiterjiuidiaU Atwtnomhic 0 i 2 1 . Body form very slender short and stout 2. Thorax, dorsum spinose not spinose 3. Accessory flagellum 1 -segmented, minute lacking 4. Sexual dimorphism of antennae, gnalhpftds strong weak or none 5. Mandibular palp present, strong weak lacking 6, Mandibular molar large* iri curative TJOil'lriluraiivC 7, Lower lip. inner lobes lacking weak well developed 8. Maxilla L palp 2-segmented 1 -segmented 9, Maxi lliped palp 4-segmenied 3 -segmented 10. Coxal plates 1 -4 smallest anteriorly deepest anteriorly J 1 . Coxal place 5 shallow. deep (about - 4) + 12. Gnathopods 1 & 2, propod & carpus 13. Peraeopods 3 & 4 elongate simplidactylate. short & deep subchelate 14. Peraeopods 3 & 4, length of segment 5 15. Peraeopods 5-7, width basis 16. Peraeopods 5-7, Similarity > segment 4 broad, suborbicular similar in size and. form < segment 4 unlike in size or form « segment 4 narrow unlike in size and form 17, Pleon, dorso- lateral armature 18, Urosome 5 & 6* dorso-latera] "wings" J9. U roped 3, rami 20, Telson lobes toothed present lanceolate; margins plumose-setose separate* converg- ing or straight weak smooth (or nearly so) lacking ] inear; margins spinose basal ly fused, spreading propods) i s ev idem (Fig. 7). Wi Lh in the collingi grou p. the more advanced tridem subgroup exhibits weakest body cut inahom and most strongly modified peraeopods in which character states trend, probably ecmvergenUy, with compa- rable features of the Nototropiinae (Fig. 1(b)), The North Pacific species of the highly specialised genus Polycheria (dexamini d subfamily Polycheriinae) may also be analyzed numerically on the basis of 20 characters and corresponding character states outlined in Table ITT (p, 61). Character states of Che P. ammftca complex of species of southern oceans is included here for broader perspective on morphological relationships within the genus, The re- sulting phenogram (Fig. 3Lp. 62) "clusters out" ‘ two major subgroups^ a primitive japonica subgrouping of three west- AMPHIPACIRCA VOL. I NO. 3 OCTOBER 15 J 994 5g FIG. 30, PHENOGRAM OF SPECIES OF ATYLUS cc < (fi UJ < cc UJ 46 46 44 42 40 38 36 34 32 V> 30 28 Q 26 < I < X Q Q UJ E < OL 24 22 20 18 16 14 12 10 ABCDEFGHJK to 1 T" 14 19 "T 1 1 r- 19 21 20 22 T 27 31 33 R~A. INDEX SPECIES LEGEND A e c D E carinatvs F coltingt atlassovi G - cjeon^Jrtus bruggenr H rylovi levidensus J tridens ekmani K borealis £ 75 GZ < -J G) 60 25 em Pacific species (P. A, indices of 8- 19) on the Left, and a highly advanced osbomi subgroup (P. A. Indices of 26-28) on the left. The osborrii subgroup exhibits signifieanily greater reduction ofmouihparts and specials /ation of tmai plates, peraeopods. and uropods. differences perhaps related 10 differing life styles in association with differing host organisms, Although the combined North Pacific and Potyeheria amarctica assemblages, in nm represent only aboil one- third of the world- wide fauna, some tentative inferences may he drawn. The North American and Asiatic subgroups differ very signifi candy froFn each other, clustering at less than 50% similarity, and perhaps meriting separate generic (certainly suhgcncrie) reeogniiion of (he North American assemblage Such would seem further justified by the fact that I he antarctica subgroup, closer to the generic type P r tenuipes Haswell from southern oceans, clusters much more closely with the Asiatic than with the North American Pacific osbomi group . Hopefully, this limited study will point ihe way to a more broadly based solution io phylchc relationships and formal classification within subfamily Polychcriinae. AMPHIPAOHCA VOL I NO i OCTOBER 15, I W 59 TABLE 1IL SPECIES OF PQLYCHERfA : CHARACTERS AN|> CHARACTER STATES CHARACTER character state value PfesJouorphlc Intermediate Apomorpliit 0 1 2 i. Antenna i, segment 3 length > flagellar segm' i - flag, segm't < flag, segm't 2. Mandible: number of blades in spine row 4 3 1-2 3 t Maxilla 1, number of outer plate spines 9 7 4. Maxilla l T length exceeding outer = outer plate shorter than of palp plate outer plate 5, Maxilla 2 + inner plate. No. marginal setae many C>10) 3-5 0-2 6. Maxilliped. length of exceeding outer = outer plate shorter than palp plate outer plate 7. Coxae 1 & 2, lower margin rounded squared front acute 8. Coxa 3, lower front comer rounded process small process large 9. Gnathopod l, palm of propod long medium short 10. Gnathopod 2, palm of propod long medium short 1 1 . Peracopods 3 & 4, length of segm't 5 > segm’t 6 - segm’ 1 6 < segm H i 6 12, Peraeopods 5-7 length of segnTt 5 > segm't 6 - segm’t 6 < segmT 6 13. Peraeopod 7, width of basis (segm't 2) broadened si. broad subi inear 14. Peraeopods 5-7, length of seizin’ t 5 >segm't 6 - Segm ' l 6 outer ramus - Outer ramus < outer ramus 18. Uropod 3, length of rami (female) subequal slightly unequal markedly unequal 19, Telson lobes, basal fusion little (1/6) intermediate much (1/3- 1/4) 20- Tclson lobes, number of lateral spines many 7-8 intermed.(4-6) few (0-3 Biogeograph ic Considerations The limited occurrence of Dexaminoidean amphipods in the North Pacific region a Slows for few broad conclusions concerning regional biogeography of ihe group. However, [he regional and world-wide distributional record of compo- nent families and subfamilies, including (be more diverse western North Pacific dexaminoidcan fauna, is more helpful (see Table IV, p. 62), Less than 200 world species are encompassed by 22 genera and 8 subfamilies (columns 1 , 2), The low species/genus ratio provides a relatively high index of morphological diversity within the super family and, by inference, a relatively long or ancient evolutionary history of the group as a whole. Within family Atylidac,. 2 subfamilies are mainly litto- ral and sublittoral (Column 7), whereas (he Lepechineiiinae AMPHJPACtHCA VOL. I NO. 3 OCTOBER 15, L 994 60 FIG. 31. PHENOGRAM OF SPECIES OF POLYCHERJA A B C D E F G > h- CC < CO Lii H < h- CO cc LU H O < CC < X o Q LU X < CL 27 28 R-A, INDEX [containing nearly half the known aty I id species) is abyssal, mainly mlndo-Pacific and Atlantic regions, The Li species of subfamily Atylinae {Atylus) are endemic to the North Pacific region, with a single outlier in the Atlantic and one possibly in the Antarctic, By contrast* the 20 species of Nototropi inae are mainly Indo-Pacific and Atlantic* with a few outliers reaching the western Pacific. The little known subfamily Anafytinae is also mainly Indo-Pacific* with 2 species reaching the Sea of Japan but none attain the North American Pacific coast. Within the more diverse and species- rich family Dexaminidae* all four subfamilies are primarily Indo-Pa- cific* and the few described species within subfamily Dexaminocnlinae are endemic there. A few members of the primitive subfamily Dexaminmae penetrate into temperate waters of the North Atlantic and southern Japan (Kyushu), Subfamily Prophliantinae is also Indo-Paeifie and southern, but with stronger representation in the North Atlantic and western Pacific regions. By contrast, the phyletieally most advanced subfamily, Polycheriinae, penetrates fairly strongly AMFHIP ACIFIC A VOL I NO. 3 OCTOBER 1 5, 1994 6 | TABLE IV. GEOGRAPHICAL DISTRIBUTION OF DEXAMINOIDEA* TAXON DIVE! RSITY DISTRIBUTION DEPTH ZONE- NO. GEN. NO, SPP. NORTH PACIFIC N, ATL. INDO- PACIFIC ASIATIC N. AMER. AT Y LIN At 1 ! 1 11 X X x X? L-SL NOTOTROFIINAE 2 ~20 X 0 X X L(A) LEPECH.’INAF 4 ~34 X X? X X A ANATYUNAE 2 4 X 0 0 X L DEXAM1NINAE 7 ™55 X 0 X X L-SL DEXAMINOCTNAE 1 1 0 0 0 X 1. POLYCHERIINAE 2 ~24 X X X X L-SL PROFHL1ANT1NAE 3 -40 X X X X L-SE Totals 11 -190 AMPELI SCOIDEA 4 -230 X X X X L*A * Data updated from Karimrd & Keraniatt (IWl), + LEGEND: L- Littoral; SL • SuhlirtoraL; A - Abyssal.. X- common; * - Spwtefc few; 0- absent. northwards along both Asiatic and North American Pacific coasts, with its most primitive members (within genus Tritaeta) confined to the Mediterranean and eastern North Atlantic regions. With respect to local distribution, the North American Pacific coastal marine fauna here consists of 8 alyl Ins, 3 pclycberiing, and one prophliantin, Three species of lepechinellins occur at abyssal depths off tine eastern Pacific continental slope, from Cental! America north to Baja and southern California but, to dale, none has been recorded from off Oregon or points northward (Barnard, 1973; Barnard & Katanian, 1991 >. As noted previou sly in this text, of the 8 regional species of Atyius, three species within the more primitive carmatus-ievidensus subgroup (i.e., A, carinam, A. mlassovl and A. bruggeni) do not extend south of the Bering Sea, and only A. levidensas reaches California. Within the advanced cottingl-iridens subgroup, all four species occur in the central region of British Columbia. However. A, Tridens and A. georgianus do not extend north to the Bering Sea. tar occur southward to central California. Of seven alyl in species recorded from coastal western Pa- cific waters. A, ekituirih A. rytr>vi< and A. octidtmtatLss; (advanced morphological counterparts of A. levider isus t A. iridens, and A. coding!) also extend furthest southwards. The more southerly occurrence, in North American Pacific waters, of pbylebcaUy advanced members of major taxo- nomic units has been noted previously within subfamilies of IheFtiOxocephaloidea, especially subfamily Metharpiniinae (Jairetland Boustield, 1994) and subfamily FicusUnae with- in the Leucoihoidea (Bousfield & Hendrycks, 1984), The evolutionary significance of this phenomenon is yet incon- clusive, but possibly reflects the overall depressant effect of low temperatures on rates of evolution, all other factors being equal (Ekman, 1 93 3), A MPH I PACIFIC A VOL I NO, 3 OCTOBER 15, JW 62 The distribution ofampelistoide^n am ph ipods. consid- ered to be close but mo ft highly advanced and specialized phyietie counterparts of d&xaminoidws, stands in marked contrast (Table IV), Through mod tlications of peraeopods 3-7, ampeliscoi deans are able to construct and live (in the “upside down” fashion of polyeberiins) within protective vertical tubes of their own construction. They thereby ex- ploit, in vast numbers, the rich tryptonic and deposit food resources on and above sedimentary substrata. Ajnpeliscoideans occur abundantly along ah marine coast- lines. including the arctic and antarctic but relatively few have penetrated the deep sea (Table IV, columns 3 -Oh However , relali ve to the de xami rtoid&aos, the larger n umber of described species (column 2) is encompassed by only 4 genera and one subfamily (column l h three- fourths in the essentially liltorabsubiitlaraJ genus Ampelisca. TTiis high specics/genus ratio implies a relatively low index ofmorpb- o logical diversity and a relatively recent evolutionary his- tory. This di fference would suggest that the Dex amino idea is, paiaeohistoncally, an older superfamily group than the Ampeliscoidca, The most primitive members (e,g, of Ary ) now exist in phyleticaHy relict or semi-relict fashion, still occupying marine "nestling" niches that gamitiaroideann and other more euryiopie and more successful ecological coun- terparts have apparently not yet penetrated, REFERENCES Alderman, A, L„ 1936, Some new and little known amphi- pcxls of California, Univ, Calif. Publ. Zook 41: 53-74, 51 figs, Austin, W. C., 1985, An Annotated Checklist of Marine Invertebrates of the Cold Temperate Northeast Pacific, Kh votan Marine Laboratory, Cowichan Bay, B. C. Vols. l-HL 682 pp, Barnard. J. L„ 1 954. Marine Ampbipoda of Oregon, Oregon State Monogr. Studies in Zoology. 8: 1-103, 33 pis. ■, 1956. Two rare amphipods from California with notes on the genus Atylus. Bull, So, Calif, Acad. Set, 55: 35-43. — 1958. ,A new genus of'dexuininid amphipod (marine Crustacea) from Califommia. Bull. So. Cal. Acad. Sci. 57: 85-90. — , 1962, South Atlantic abyssal amphipods col- lected by R.V, Verna, Abyssal Crustacea. Verna Re- search Series I: 1-78, 79 figs, , 1964. Deep- sea Ampbipoda (Crustacea) col- lected by the R7V "Verna” in the eastern Pacific Ocean and the Caribbean and Mediterranean Seas, Bull. Amer Mus Nat. Hist. 127( l):l -45, 33 figs, , l%9a. The families and genera of marine gammaridean Amphipoda, Bull. U„ S. NaU- Mus, 271: 1-535, 173 figs. . 1969b, Ganunaridean Amphipoda of the Rocky Intertidal of California: Monterrey Bay to La Jolla, U. S. Natl, Mus, Bull. 258:1-230, 173 figs. , 1970a, The identity of Dexamonica and Prin- sjjffj, with a revision of Dexaminidae (Amphipoda). Crustaceans 19: 161-180, 1 970b. S ubl ittoral Gammaridea o f the Ha wai- ian Islands Smiths. Coutr. Zool, 34: 1-286, 1 80 figs, , J972a. The Marine Fauna of new Zealand: Algae-I lying Liuoral Gammaridea (Crustacea Amphi- poda). Mem. N. Z. Goeanogr, Inst, No. 62: 1-216. 100 figi. , 1972b, Gammaridcan Amphipoda of Austra- lia, Part 1. Smiths. C’ontr, Zool. No. 103: 1-333, 194 figs, . 1973. Deep-sea Amphipoda of the genus L^ptduneiki (C r istacea). Smiths, Contr, Zool, No, 133: 1-31, 12 figs. — — , 1974. Giutmi andean Amphipoda of Austra- lia. Pan II, Sm!tl)$. Contr.ZocL No. 139: 1-148, 83 figs, -, 1 975, Amphipoda Gammaridea. pp 31 3 ■ 366, pis. 70-85. in R. I, Smith & J, T, Car Lon (eds).Light's Manual. Intertidal Invertebrates of the Central Califor- nia Coast. 3rd edition, Univ. Calif, pres*, Berkeley. CA. 716 pp. , 1979. Li ttoral gammaridean Amphipoda from the Gulf of California and the Galapagos Islands, Smiths. Contr, Zool. 271: 1-149, 74 figs, & G. S. Karaman, 1991. The Families and Genera of Marine Gammaridean Amphipoda (Except Marine Gammaroidsk Rec. Austral, Mu$„ Suppl. 13, Pts, 1 & 2: 1 866. 133 figs. Barnard, K. H.. 1930. Amphipoda. British Antarctic ('Terra Nova 11 ) Expedition. 1910. Natural history Reports, Zool- ogy 8: 307-454. 63 figs, Bate, C, S., 1 862, Catalogue of the specimens of amphipodous Crustacea in the collection of the British Museum, Lon- don, 399 pp„ 58 pis. Rale, C, S. k J. 0. Westwood, 1863, A history of the British Scssile-Eved Crustacea, London. I: 1-507, illustr. Belkin -SantiTii, D,, 1975. Au sujet d une nouvelle espcce d'ArWwv (Amphipoda, Dexaminidae) de Mcditerranee: Atyius tmssitietisb n. sp, Boll, Mus. Civ. Stor Natnr. Verona 1 : 437-479, 2 pis. , 1982, Family Dexaminidae. pp. 212- 232. in S. Ruffo ed. The Amphipoda of the Mediter- ranean. Mem. Inst, Oceanogr. Pan 1. Gammaridea (Aeanthonotozomatidae toGammaridae). No. 13:212- 232. Birstcln, J, A. s & F4, E. Vinogradov, 1955- Pelagicheskei gammaridy (Aniphipoda-Gammaridiey) Kuril o-Kam- ehatxkoi Vpadiny, Akad, Nauk SSSR, lust, Okeanol. Trud, 12:210-287, 35 figs. Boeck, A., 1971, Bidrag til Californiens amphipodenfauna. Forhandl. Vidcnsk. Selsk. Christiana 1 87 1 : 32-5 L I pi. — — , 1876. DeSknndinaviskcogAriiske Amphipoder, 712 pp, 32 pis. Christiana: A, W, BrOgger, Bousiicld, E. L„ 1958, Ecological Investigations on sea shore id vertebrates of the Pacific Coast of Canada. Bull, Natl. Mus. Canada 147: 104-115, AMPHIPAC1PICA VOL S NO 3 OCTOBER i5 .1994 63 , 1963, Investigations on sea-shore inverte- brates of the Pacific coast of Canada, 1957 and 1 959, 1, Station List. Bull. Nad, Mus. Can. 185: 72-89. , 1968. Studies or Mttond marine inverte- brates of the Pacific coast of Cana, 1964, 1, Station Hat. Natl Mus, Can, Bull. 223: 49-57. , 1979a. A re vised cl ass i fi cation and phy logen y of amphipod crustaceans. Trans, Roy. Soc. Canada 4: 343-390, Bousfidd, E. L.. 1979b- The amphipod superfamily Gam- maroidea in the northeastern Pacific region: systemal- ics and distributional ecology, Bull, Bioi. Soc. Wash. 3: 297-359. 12 figs. „ 1982. Amphipoda, Gammaridea. jn Synopsis and Classification of Li y ing Orga n isms. McGra w -Hi] i , New York, vol. 2; 254-285. > 1983. An updated phyletic classifies lion and palacohistory of the Amphipoda, Crustacean Issues , 3 ; 257-278, & E, A. Hendry eta, 1994 The Amphipod Superfamilv Leucothoidea on the Pacific Coast of North America, Family Pleustidae: Subfamily Pleustinae, 5y si- emaUcs and Biogeography. Amphipacifica 1(2 ): 3-69, 38 figs. and D, E. McAllister, 1963. Station List of ihc National Museum marine biological expedition to south eastern Alaska and Prince William Sound, Natl. Mus. Can, Bull. 183:76-103. and N. E, Janett J98 1 , Station lists of marine biological expeditions of the National Museum of N aE- ural Sciences in the North American Pacific coastal re- gions, 1966 to 1980. SyllogeusNo. 34, I -66, &C. P, Staude, 1994. The Impact of J, L. Barn- ard on North American Pacific Amphipod Research: A Tribute, Amphipacifica 1(1): 3-16. Bulycheva, A. f T 1952, Novye vidy tKteopfcivov (Amptvi- poda: Gammaridea) i? Japonskovo Morei. II, Akad. Nauk SSSR, Trud, Zool, Inst. 12; 195-250., 39 figs. , 1955. Novyc vidy bokoplavov (Amphi- poda, Gammaridea) i z Japanskovo Morei. If Trud. Zool, Inst. Akad. Nauk SSSR.2J: 193-207. (ki Russian), , 1957. Amfipody (Amphipoda) severo- zapadnoi chasti Japansfcovo Morei. Akad. Nuuk S5SR, Issled. Dalhev. Morei, 4: 85-126, 3 figs, Cadien, D, B.. 1991 , List of the Marine Amphipod faunas of the temperate and boreal northeastern Pacific Ocean, including literature rc cords oloccun e nee between Bahia San Quintrn, Baja California. and the south side of the Aleutian Islands, incorporating tinmenclatural changes listed in Barmaid & Karaman, 1991, SCAM IT Tech. Pub l,, Los Angeles, California, Sept,, 1991, 21 pp., list. Caiman, W, T,. 3 898, On a collection of Crustacea from Puget Sound. Ann. N. Y. Acad. Sci. XI (13): 259-292, pis. 31-34 Chilton, C„ 1914. A new ainphipodan genus ami species (Family Dcxaminidae) from new Zealand, Jour. Linn, Soc. London, Zool, 32: 331-336, 2 pis. Conlan, K. E„ 3983. The amphipod superfamily Corophioidea in the northeastern Pacific region, 3 , Family Isaeidae: systematics and distributional ecology. PubJ, Nat. Sci., Nad. Mas. Nat, Sd., Canada 4: 1-75, 36 figs. Dunbar, M , J„ 1 954 . The amphipod Crustacea of Unga va Bay .Canadian eastern arctic. Jour, Fish, Res, Bd. Canada 11:709-798, 42 figs. Dickinson, J. J., & A. G, Carey, Jr„ 1978, Distribution of gamniand Amphipoda (Crustacea) on Cascadia Abyssal Plain (Oregon). Deep-Sea Research 25: 97-106, 2 figs, Ekman, 5,, 1953, Zoogeography of the Sea, Sidgwick Sl Jackson, London: 437 pp.. illnstr. Fabrici us, J. C„ 1783, Entomologies systematica emends ia et aucta 2: 5 1 9 pp Hafniae: Christ. Gotti. Proft, Game, 1981. A new deep sea Amphipoda, Lepechinella $agamien$i$ sp. tiov, from Sa garni Bay. Proc, Jap. Soc. Syst. Zool. 20: 16-20, 2 figs, Giles, G. M,. 1888. Further notes on the Amphipoda of Indian waters, Natural history notes from H. M.'s Indian marine survey steamer "Investigator' 1 , commander Al- fred Carpenter, R,N., D.S.O, commanding. No, 1 5. lour Asiatic Soc. Bengal 57: 220-255, pis. 6-12. Goes, A. . 1866. Crustacea Amphipoda mans Spetzbergiam alluentis, cum speciebus aJiis arcticis e numeral. Ofv. Kong I, Veutcnsk, Akad.Furh. IS65:517-536 T pLs. 364] , Has well, W. A., 1879. On some additional new genera and species of amphipod oils crustaceans. Proc, Linn, Soc. N, S, Wales 4: 319-350+ pis. 18-24. - - . 1885. Notes on the Australian Amphipoda, Proc. Linn. Soc. N. S. Wales 9: 993-1000, pis, 48-49. Guijanov.a, E. F.. 1938, Amphipoda Gammaridea of Siauku Bay and Sudzukhe Bay (Japan Sea). Rep, Japan Sea Hydrobioi. Explor. Zool, Inst. Acad, Sci. USSR in 1 934, 1:241-404, (in Russian) — , 1951, Amphipoda of the Seas of the US SR , Opted. Fau nc SS SR . Akad. Nauk 41:1-1 029, 29 5 tigs. (In Russian) H tray am a. A., 1984a, Taxonomic Studies on the shallow- waler ga mmEiridcan Am ph ipoda of West Ky usfe u , Japan. Ill- Dexairtmidae ( Polycfw ria a nd Pamdexatnine I . Publ, Seto Mar, Biol. Lab. XXIX, Nos. 4/6: 187-230. , ]yx4b. Taxonomic Studies on the Shallow W r aier Gammaridcan Amphipoda of West Kyu- shu J apan IV Dcx am i n idae (Guemeu } f Eoph 1 iati L idae, Eu si ridae. Halt stori idae , H yal idae „ Isch yroceridae . Pu hi . Seto Mar. Biot. Lab. voL 30, Nos, 1/3:1-53. „ 1986a. Marine Gatnmandear Amphi- poda (Crustacea) from Hong Kong 11 The Family Dcx yimnidae, Proc. Second Ini, Mar, Biol. Workshop: The Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong, 1986, (B. Morion, edj, Hong Kong Univ. Press: 487-501, 9 figs, . 1986b. A new alyl id species (Crusta- cea: Ainphiptwla) from Gteuchi bay, Northeast Japan, Pme Japan Soc. 5y*(. Zool, 33: 4-10. AMPHIPA0MCA VOL, I NO. 3 OCTOBER LI 3994 fi4 Holman, H., and L, WatJing, 1983, Amphipoda from (he southern ocean: families Colomastigidae, Dexammidae, Leuoothoidae* Liljeborgiidae, and Sebidae. Biology of the Antarctic Seas XML Antarctic Research Series 38; 2 15-262, 35 figs. Erie, H., i960. Amphipoda, m Uchida et al, i960: Encycl. Zoologiea Illustrated in Colours, VoL IV, 1-247, Hot' uryukan, Tokyo. Ishimam, S.-l,. 19S7, Description of two new species of GttetHea (Crustacea, Amphipoda, Dexaminidael from Japan, with tentative revision of subfamily Prophliant- idae, Jour. Nat, Hist, 21: 1395-1414. , 1994, A Catalogue of Gammaridean and Ingol lie! I- idcan Amphipoda Recorded from the Vicinity of Japan. Kept, Sado Mai', Biol, Sta„ Niigata Univ. 24; 29-86, Jarrell, N. E. h & E, L. Bousfield, 1994, The Amphipod Superfamily Phoxocephaloidea on the Pacific Coast of North America, Family Phoxocephadidae, Fart L Mel- harpiniinae. New Subfamily, Amphipaeilica 1(1): 58- 140. 31 figs, Leach, A., 1814, Cruslaceology. Appendix, The Edinburgh Encyclopaedia 7: 29434. 3 1815, A tabular view of the external character of four classes of animals which Li nine arranged under Insccta; with I he distribution of the genera composing three of these classes into orders &e. and descriptions of several new genera and species, Trans. Linn. $oc. Lorn don 1 1: 306-400. Ledoyer, M. n 1979a, Les gamniaricns de lapente externc du Grande R^elf de Tulwar (Madagascar) (Crustacea: Amphipoda). Mem, Mus, Civ, Stor, Nai. Verona 2: F 150, 1979b, Expedition Rumphius 11(1975), Cmsiacds parasites, commensaux, etc, (Th. Monodet R, Serene, M), VI, Crusiact Amphipodes Gammariens. Bull, Mus. Nat. Hist., Pains 4° ser. I, 1979, .sect. A, n" 1: 137- 18 1, , 1982. Cmstacts amphipodes gammariens, Families dcs Acamhonotoioinatidae a Gammaridae. Paris. Centre National de la Recherche* Seientificiuc: Faune de Madagascar, 59(1): 1-598. — -,1984. Les Gammaricns (Crustacea, Amplii- poda) des herbiers des phanerogames marines dc Nouvd le Caledonie (region de Noumea). Mein, Mus. Natl, d’Hist. Nat,, new series, ser. A. zoologie. 129; 1 13 pp., 48 Tigs, Lincoln, R, J„ 1979, British Marine Amphipoda: Gamnw- idea. Bril. Mus. (Nat. Hist.) PubL No. SIS: 1-658, 280 figs, Lowry, J. K., 198 L A rede&cTiption of Sphaentphthctmus: grohbem Spandi based on type material from (he Red Sea and new material from the Great Barrier Reef. { Am- phipoda, Dexaminidae), Crustaetana 41(2): 190- 1 98. McKinney; L.D,, 1980. Four new and unusual amphfpods from the Gulf of Mexico and Caribbean Sea. FroC. Biol. Soc. Wash. 93: 83-103, Metzger, A.* 1871. Die wtrbullhsen Mcciesthiere der ost- friesischen Kuste. Jahrsb, Naturh, Gesellsh , Hannover 21:20-34, Mills, E. L., 1961, Amphipod crustaceans of (he Pacific Coast of Canada. I, Family Atylidae. Nall, Mus. Can. Bull. 172; 13-33, Milne-Ed wards, H„ 1830. Ex Emil de reoberches pour scrvii a I 'histoire nature lie des crustaces amphipodes, Ann. Sei. Nairn, 20: 353 399. pis 10, 11. Moore, P, G„ 1984. Cammaridean Amphipoda (Crustacea) collected by the yacht Tulip from surface waters of the Arabian Sea, Jour, Nut. Hist. 18: 369-380. Nagata, K„ 1961, A new atylid amphipod from Japan, Annol. Zool. Japan, 34: 216-218, - -. 1965. Amphipoda Gammaridea. JpM, Iwasa & K. Niigata's Illustrated Encyclopedia of the Fauna of Japan: 559-572. (in Japanese) Ni cholls. 1939, The Prophlianlidae, A proposed new fam- ily of Amphipoda, with description of a new genus and four new species Ree, S, Austral. Mus. 6: 309-334, 10 figs. Norman. A.. 1868, On Crustacea Amphipoda new to sci- ence or to Britain. Ann. Mag, Nat. Hist. ser. 4. 2: 4 1 1- 421, pis. 21, 22, & pi 23, figs, 1-11. Okado, M,, 1993, The occurrence of Atyivs ekmmi (Crust aeea. Amphipoda, Gammaridea) from Southern Hok- kaido, Japan, ip Bull. Fac. Fish. h Hokkaido LIniv. , 44 (J ); 6-14. ; figs. l-4 h Oldevig, IL. 1959, Arctic , subarctic, and Scandinavian amphipod s on the collections of the Swedish Natural History Museum in Stockholm. Goteborgia Kunglia Vete nskaps- V i Uerticvts-SamlLal les HandUnger(6B)8 (2): 1-132, Pi riot, 1933, Les Amphipodes dc I'cxpedition du Siboga. Deuxiemc par tie. Les amphipodes gammarides II, - Les amphipodes de la mer profondc, 1 (Lysianassidse, StegoeephaJidae, Slcnothoidae, Plena tidae, Lepechin- eilidac). Siboga -Expedition, Monogr. 33: 115-167, figs. 35-60. Sars, G, 0, ? 1895, An Account of the Crustacea of Nor- way. J. Amphipoda, Christiana & Copenhagen, 711 pp,, 240 pls„ 8 suppl. Sehdlenbcrg, A„ 1925- Crustacea VIIL Amphipoda, vol, X pp, 1 1 1 -204, 27 figs, la W. Michael son (ed.). Beitr. Kennt. Meenesf. Westafrikas. Hamburg: L, Friedrich - sohn & Co, ,1926. Die gammariden der Deutscben S udpolar-E x pediti on 1901-1 903 . Deutsches S ndpotar- Expedition 18: 2354214.68 figs, 1928, Zoological Results of the Cambridge Expedition to the Sue? Canal, 1924. 35- Report on the Amphipoda. Trans, Zool Soc. London, 22(5): 693- 692, figs, 198-209. AMRHIPACIFICA VOL, I NO, 3 OCTOBER 15,1994 65 — — * 1931, Gammariden und Caprdltffcn de* Magdlangebietes, Sudgeorgiens iind dcr Westantark- tis. Further Zoological Results of the Swedish Antarct- ic Expedition 1 90 M 903, 2(6): 290 pp. 1 pL, 136 figs. , 1938. morale Amphipoden des tropisch- en Pazifiks. Kungl. Svenska Veteask, Haudl. (3) 16(6): 105 pp. n 48 figs, Schrant, F, R,. 1986, Crustacea, Oxford University Press, Mew York. 606 pp., illustr. Shoemaker, C. R-, 1920, The am phi pods of the Canadian Arctic Expedition, 1 913- J 918. Report of the Canadian Arctic Expedition, J 91 3d 918, 7E: 30 pp, 6 Egs,. app > 1955. Amphipoda collected at the Arctic Laboratory. Office afHaval Research, Faint Barrow. Alaska, by G, E, McGinitie. Smiths. ML&c, Coll. 128 (1): 1-78, 20 figs. Sivapnikasam, T. E., 1970, Description oiAtyfas {K#nieh- atylu$)proce$$ket sp> nov, { Amphipoda: Detain in idae) from the Gull of Mannar (sic), India. Jour. Mar. Biol. Assoc, India 10: 93-9fr I fig. Skogsberg, T., and G, H, Vansell, 1928. Structure and Be- haviour of the amphipod, Polychem osbornL Proc. Cal. Acad. Sci., ser. 4, 17: 267-295, 26 figs. Sparadl, H„ 1923. Amphipoden tier 'Pola 1 - Expedition in das Rote Meer, Akad, Wissensch. Wien, An?, 60 Jafir- gang, 1923 (in 3 parts): 17-20. 87-89, 111-112. Staude, C, R, 1986. System sties and Behavioural Ecology of the amphipod genus Pummoera (Gammaridea: Eu- siroidea: Pontogenriictee) in the eastern North Pacific. PhD thesis. Univ, Washington Press,, Seattle, 511 pp. , 1987, Amphipoda: Gammaridea, 346- 39| r ja_E. Kozloff (ed.). Marine Invertebrates of the Pacific Northwest. Univ. Wash. Press. Seattle. 5n pp. Stubbing, T. R. R.. 1875, On some rtciv exotic sessile eyed crustaceans. Ann. Mag. Nat, hist., ser. 415: 184-188, pi 15 A. 1888. Report on the Amphipoda collected by H.M.S, Challenger during the years 1873-76. Rt^ port on the Scicnlific Results of the Voyage of H.M.5. Challenger During the Years 1873-1876, Zoology 29: 1737 pp., 210 pis, London, Eyre & Spottiswoodie. , 1899. Revision of Amphipoda. Ann. Mag. Nut. Hist. sec. 7 3: 350, — , 1906. Amphipoda T. Gammaridea. Das Tiemeich, Berlin. 21; 806 pp., 127 figs. - T J 90S. On two new species of northern Amphipoda. Jour. Linn. Soc. London* Zool, 30: 191- 197, pis. 27-28, Stephen sen, K., 1944. Some Japanese aniphipods Vid- ensk. Medd, Dansk Natuib. For. 108: 25-88, 33 figs. Thurston, M,. 1974. Crustacea Amphipoda from Graham Land and the Scotia Arc, collected by Operation Tab- ari n and I he Falkland Islands Dependencies Survey, 1944-59. British Antarctic Survey Scientific Reports 85: 89 pp r , 28 figs. , 1980. Abyssal benthic Amphipoda (Crust- acea) from the east ice tend basin 2. Lepechinelia and an allied new genus. Bull, firit. Mus, Nat Hist, (Zuolj 38: 69-87, 12 fir;, Tzvetkova, N, L.. 1967. On the fauna and ecology of am- phipods (Amphipoda, Gainmaridea) of Possjet Bay, (Sea of Japan). Acad. Sd. USSR. Zool. Inst, , Issled, Pauny Morci 5: 160-195, 7 figs. Vader. W., 1969. Herkerming en bioloop van de West- Europese Dexammittee (Crustacea- Amphipoda). Zool. Bidj, H, Bidj. Faun. Nederland I: 59-67 , 2 figs. , 1983. Prehensile pereopotis in gaiiim andean Amphipoda. Sarsia 68: 139-148, Walker, A, 0„ 1954, Report on the .Amphipoda collected by Professor Hodman* ai Ceylon, in 1902. Report to the Government of Ceylon on the Pearl Oyster Fisher- ies of the Gulf of Mannar, Suppl. RpL, 17: 229-300, , 1905, Marine Crustaceans XVI. /Amphipoda. Fauna and Geography of the Maidive and Laccadive Archipelagos, 2, Suppl. 1:923-932. 2 figs., I pi. LEGEND for figures Al - antenna 1; A2 - antenna 2; ABD - abdomen; ACC FL - accessory flagellum; BR - branchia (coxal gill); CX - coxa; BP - epimera (pleon plate) ; GN - gnatbopod, HD - head; JV - juvenile; LL - lower lip; LFT - left; MD - mandible; MX - maxilla; MXPD maxilliped; PL - pleopod; PLP - palp; RT right; T - tdson; UL - upper lip; U - uropod; UR OS ■ urosome ; X - enlarged; O - male; O - female. AMPHIPACIFICA VOT, I NO. 3 OCTOBER 15,1994 66 NEW SPECIES Of THE AMPHIPOD CRUSTACEAN GENERA PHOTIS AND GAMMA ROFSIS (COROPHIOIDEA: ISAEIDAE) PROM CALIFORNIA. by Kathleen E. Conlan T ABSTRACT Three new species of the amphtpod supertamlly Corophioidea have been found m depths of 92 10 2005 meters off the Pacific coast of California, Phorh (Photis) typhl&ps t new species, Photis {Phans) Imeanmnus, new species, and Gammaropsis {Podocempsis) ocellata, new species, are here described and illustrated* and their morphological relationships with oilier regional species are -discussed. INTRODUCTION Three new species ofCorophioidca have been found in benthic collections from offshore waters of the const of California. These ait Photis (Pltotis) typhlops , new species, an eyeless form recorded from depths of 8 12 to 2005 meters, Photis (Photis) tinearmiintts, new species* an eyed taxon from a single collection at a depth of 92 meters, and Gamma ropsis (Podot^rttpsis) acethita, new species, an un- usual podoceiopsid having Faceted hut un pigmented eyes* taken at a depth of 590 meters. The most recent review of regional species of these genera i s Conlan ( 1983 k L ists and nurrtbe rs of region al gen - era and species can also be Found in overview treatments by Austin (1985), Cadien ( 1991), anti Bousficld and Staude ( 1994), The new species are here described and compared with their local relatives, The present study raises the number of North Pacific species of Photis (Photis) to32, and of Gammaropsis (Podocewpsis) to 9, The coraphioidean genera Photis and Gammampsfs are here assigned <0 family Isaeidac. Families Isaeidae and Aoridae had been merged within family Corophiidae by Barnard and Karaman (1991). However, continued recog- nition of the Isueidae as a distinct coiophioidean family is in keeping with the recent work of Myers (1988)* and with North Pacific regional comprehensive listings of Ishimaru [ 1994) and Bousfidd and Staude ( 1 994}. acknowledgements This work was conducted as pail of a contract with Science Applications International Corporation for produc- tion of a guide to the Corophioldea in the Taxonomic Adas of the Macryi n vertebrate Fauna of Lhe S anta Maria Basin and the Western Santa Barbara Channel. Jim Thomas. Hans Kuck, Joe) Martin, Paul Scott, Terry Gos liner, and Judith Price provided specimens and data, C-t. Shih and E. L, BousOekt reviewed the manuscript. Susan Laurie-BouTKiue illustrated the three new species fully treated here. METHODS The amphipod specimens were pan of a large collection of Cotophi oidea that was examined for preparaiion of a taxonomic atlas to the ben (hie invertebrates of tee Santa Maria Basin and (he western Santa Barbara Channel, Right appendages and mouth parts were ill ustrated from slide moun ts in polyvinyl lactepbenot. Body length was measured from the tip of the rostrum to iheba.se of the telsor. Material was deposited at the Canadian Museum of Nature (CMN), (he California Academy of Sciences (CAS), the Smithsonian Institution. L 5 .S, Museum of Natural History (USNM), the Los Angeles Coun Ly M use urn of Natural History (NHML AC), and tee Sania Barbara Museum of Natural History (5BMNH). J Canadian Museum of Nature. P.O. Box 3443, Siaiion D, Ottawa, Ontario, Canada K IP 6P4. AM PH IP AC1F1C A VOL.I NO. 3 OCTOBER \5, 1994 67 SYSTEMATIC^ Phoris (Phatis) lyphiapx, new species D Material examined TYPE MATERIAL: Holotype: adult male (USNM, catalogue no, 260403)1, U.S.A,: Califor- nia: off San Francisco ( 37° 22.3 1 'N, 123° 19.24* W), station 3-1 8. 26.829. 9-9 1 . EPA site 1 02. ! 990 m, 1 5 Sepi, 1 99 1 , J, A, Blake, collector. Allotype, adult female (USNM. cata- logue no. 266404). same location. Paratype^ about 300 individuals (USNM, catalogue no, 266405 (adult mates), 266406 (adult females), 266407 (unseeable juveniles}): 6 males, 8 females, 30 juveniles (CAS, catalogue no. CASIZ 085729); 6 males, 8 females, 30 juveniles (NHMLAC, catalogue no. LACM 9 1 -190,1); 6 males, 8 females, 30 juveniles (CMN, catalogue no, NMCC 1 993-000 1 ). a FI same location. OTHER MATERIAL (excluded from the lype scries); l j u veni le from U.5 , A.; Cal iforo la; off Pt. B ueb on (35° 15,72' N, 121°04,68"W), 396 in, California Phase II Monitoring Pro- gram, Minerals Management Service, Pacific OCS Office. Santa Maria Basin Project, station 020-BSS-01 -TX; 2 juve- niles from same area, off Pi. San Luts (35 o 05.07'NL 121900.75 WL 390m, station 025-BSR-OI TX; 5 juveniles from same location as preceding. 390 m, station 025-BSR- 02 -TX; 2 juveniles from same location its preceding. 390 m. station 025 -B SR *03 TX; 2 adult females, 4 juveniles from same area, off Purisima Pt, (34°37.SO*N T I21°01.66 W). 591 m, 6 Jan 1984 ; station 050 BSS-pl -TX; I adult female, 7 juveniles from same area, off Pt. Argue Elo (34°33.66*N. 1 20°56.3 1' W), 590 m, station 05 5 - B SS -0 1 TX (all of above SBMNH); ] juvenile from Eel River Basin (4l°S6.33’N, l24°38 t 0Q'W), 552 tn, dissolved oxygen 1 .03 ml/1, bottom water temp, 5,86°t\ sill-clay sediment, % organic carbon 1,473, 22 Nov, 1987, station SB-4 (NHMLAC); l juvenile from same area as preceding (4]°3Q,77‘N., 124°29,33'W), 524/549 m, dissolved oxygen 1,6 ml/1, bottom water temp. 5,92 0 C, sand-silt-day sediment, % organic carbon 0,859/ 0.782, 22 Nov, 1987, station SB - 12 (NHMLAC); 1 juvenile from same area as preceding (40^57, OG’N, 1 24°23.42‘WL 188 m, dissolved oxygen 4,35 ml/3, bottom water temp, 8 j67°C t silt-clay sedimen t h % organic carbon 0,924, 1 9 Nov . 1 987, station SB* 14 (NHMLAC); 1 juvenile from same area a s preceding (40°57 .20' N , 1 24°33,20 : ' W ), 555 m, dissolved oxygen 2,51 ml/l, bottom water temp, 6.56°C silt-clay sediment, % organic carbon not recorded, 18 Nov. 1987. station SB 16 (NHMLAC) (all Eel River Basin samples collected by MEC Analytical Systems lnu, Carlsbad, Cali - forma). Diagnusis, Eye lacking. Antennae half length of the body, with long setae. Coxae 1-5. ventral margins with 0-4 long setae, Gnattiopod l, carpus longer than propod us. propodus slender, palm convex or shallowly excavate Gnathopod 2 of male, basis with row of xtiidulaiioti ridges angled across lateral face; palm of propodus transverse, with tooth and long spine at palmar defining comer and two small teeth in palm, Description, Adult male (3,2 mm) Holotype: Head lobe triangular. Eye lacking. Antennae 1 and 2 about equal in length. Antenna I weakly setose, article 3 longer than article I ; accessory flagellum microscopic button. Antenna 2 moderately setose, flagellum not pedilonn, longer than article 5, festally spmose. Upper lip, epistome triangular . Mandible with 34 raker spines; molar Hake present; palp strong, article 3 hardly wider dis tally than proximal I y, both articles 2 and 3 wife numerous setae, article 2 longer than article 3, Maxilla l, inner plate without setae; palp narrower than outer plate. Maxilla 2, inner plate about same width as outer, with facial setae. Maxilliped, inner plate riot reaching end of article 4; outer plate not reaching end of article 5; unguis (article 8) about equal in length to article 7. Peraeopods 1-5, ventral margins of coxae with 0-2 long setae each. Goalhopod I, coxa more anterodistally produced than coxa 2; basis inserted mid-proxitnally on inner face, not densely setose; carpus about equal in length to propod us, anterior margin distal I y setose; propodus, palm shallowly excavate, defined by single spine; dactyl longer than palm of propodus, posterior (inner) margin with few short setae and cusps, Gnathopod 2. basis, lateral face with row of stridula- lion ridges: carpus shorter than propodus; propodus, width 1 .5 times width of propodus of gnathopod I , palm transverse, with 2 excavations and tooth and spine at defining comer, setae at dactyl hinge less than half length of propodus; dactyl overlapping palm hf length of unguis, without tooth, with spine and selal duster proximal of unguis. Peraeopod 3, coxa with row of stridulatlon ridges on ventral margin, Peraeopod 4, coxa, posterior margin not ex- cavate. Peraeopods 3 and 4, basis not expanded, uterus wid- er than carpus and produced anteriorly over less than 1/4 of carpus, dactyl shorter than propodus. Peraeopod 5, coxa similar in depth to coxa 4; basis broad, not posteriorly excavate; menis not posteriorly excavate; merus and carpus not spinose; propodus with only single long spine ai anterotlistal comer; dacty] with 2 pronounced cusps at junc- tion of unguis, Peraeoptxls 6 and 7, coxae smaller than coxa 5; otherwise articles similar in shape to peraeopod 5 T al- though base* narrower and dactyl not eusped, Pleon plates 1 -3 not posterodistatiy notched. Picon and urosomc without dorsally erect setae or cusps, Uropod 1, peduncle without lateral eedysial spine pro* imally or tooth - like process extending ventrafly below rami; rami tipped by 1-2 short spines. Uiopod 3> peduncle not spinose; outer ramus nearly as long as peduncle and tipped by 1 -2 long setae, inner ramus about 1/4 length of outer, lipped by single short spine, Tclson apices marked by single long seta and small knob, AMPHIPACIFICA VOL. 1 NO. 3 OCTOBER 15, 1994 FIG. 1 . Photis typhiopsi new species. Adult male, (3.2 mm) HOLOTYPE: whole body, distal articles of gnathopods I and 2 (setae omitted), mouthparts, uropod 3, and telson, Adult male 1 (3.4 mm) PARATYPE: distal articles of gnathopods 1 and 2 (setae omitted). Juvenile male (2.9 mm): distal articles of gnathopod 2 (setae omitted). Adult female br. III. (3.4 mm) ALLOTYPE: gnathopods \ and 2. Adult female I br. HI. (3.3 mm) PARATYPE; gnathopods 1 and 2. magnifications (setae omitted), Lateral views: whole body, maxilla 1, uropod 3, and telson: other views medial Scale OT mm. AMPHIPACIF1CA VOL. 1 NO. 3 OCTOBER 15, 1994 69 Condition. With all appendages. Right appendages, moutbparts, and telson slide mounted. Adult female, Body length 3.4 mm. Gnathopod 1, carpus and propodus similar to but slightly slenderer than in male. Gnathopod 2, basis without stridulariou ridges; propodus, palm convex. Brood plates moderately wide, setae without hook at each (ip, Other features as in rnafe, Condition. Wild all appendages. Right appendages, mouthparts, and telson slide mounted, Variation. The narrowness and amount ofindeutation of the propodus of gnathopod 1 of the male varies, becoming narrower and more excavate in larger males. Tooth length in the palm of gnathopod 2 is also greater in larger animals. The number of long setae on the ventral margin of coxae I -5 may l^e as much as 4 . I mrna lure fcina Its bear ase lose brood plate s or lack then] altogether. Etymology. From the Greek tvphfops, meaning blind, alluding to the absence of pigmented eyes tn this species. Distribution and ecology. Collected from 188-2005 m off Santa Barbara, San Francisco. and Eureka-CresceOtCUy, in the San Francisco collections, Photis typhfops was found from 812 m to 2005 m depth, with density peaking at 9500 individuals/m^ at 1770 to 1090 m depth. At this density the ai n ph ipods were dearly visible as a thick mat concentrated at the sediment surface. This is the first known record of a deep water amphipod mat (J. A. Blake, peas. comm., 7 Dec, 1992). Evidently Photis typhfops can occur in areas of low dissolved oxygen, judging from (be Eureka-Cresdent City collections. Taxonomic Commentary, Two Other blind species of Photis arc known to occur in the North Pacific: Photis {Ph&its) kurilica Gurianova and Photis {Cedrophotis) maUftako J. L. Barnard. Photis kuriticu differs from P typhlops in the following respects: head lobe rounded ventraily; antenna 3. flagellum 8 articles, slightly longer than peduncle; antenna 2, peduncle article 4 3 times as long as article 3; gnathopod 1 of male, basis, anterior and posterior margins covered with abundant plumose setae, carpus equal in Length to propodus. propodus, palm concave; gnathopod 2 of male, coxa with 9 long setae on ventral margin, basis with abundant short, stout setae anteriorly and long, slender setae posteriorly, propodus. palm concave, without tooth; gnathopod 2 of female, propodus, palm shallowly concave; uropod 1 with 8-11 lateral spines on peduncle and rami; uropod 2 with 2d 2 lateral spines on peduncle and rami Photis kuriiicu has only been recorded from the east coast of Russia (Gmjanova, 19553. Photis tmUnako, from the Ccdr os Trench, Baja Califor- nia. has a much longer inner ramus on the third uropod (half the length of the outer - a defining character of the submenus), more slender propodus of gnathopods I and 2. broader coxa 1, and less spinose uropnd.s I and 2 (J. L. Barnard. 1967k Another blind .species of Photis is the South Atlantic abyssal Phot i.v caeca J . L. Barnard. This species differs from P- typhtops as follows: antenna 1, article 3 only slightly longer than article I ; gnathopod 1 of female, coxa square, basis with 3 long setae anteriorly and 1 posteriorly; gnathopod 2 of female- coxa square, propodus much narrower than wide; peraeopodlk coxa covering only 1/3 of basis; peraeopod 5, basis, width 3/4 of length; uropod I rami, outer ramus with 1 spine, inner ramus with 0 (J. L. BairtarcL 1962). All four blind spscte&of Photis possess distinctly longer antennae than in eyed species of Photis . a characteristic which apparently correlates with lack of visual sensory organs, Photis i Photis ? finettrmanux, new species (Fig, 2) .Material examined. TYPE MATERIAL; Bolotype: adult male (USNM. Catalogue no. 239498), Ll.S.A,: Califor- nia; off Purisima Point |34 o 43 0'N, 12(P47.4’W>, 92 m, May 1 987, California Phase II Monitoring Program. Miner- als Management Service, Pacific OCS Office, Santa Maria Basin Project, cruise L3, station R-4. replicate 1, Battel le, collector. Diagnosis. Eyes small, pigmented. Coxae 1-5, ventral margins with 2-1 1 long setae, Gnathopod I, carpus shorter than propodus; propodus broad; palm strongly excavate in male, GnathoptMl 2 of male, basis with lew giridulation ridges on anterodjstal margin; palm of propodus oblique, linear, defining corner not marked by spine or change of angle, with 2 small teeth in palm. Description. Adul ( male ( 3 .4 mm ) Hdot ype : Head lobe triangular. Eye black, oval. Upper lip. cpistorne triangular. Mandible with 3 raker spines; molar flake present: palp strong, article 3 hardly wider dis tally than proximal))-, both articles 2 and 3 with numerous setae, article 2 longer than article 3. Maxilla I, inner plidc without setae; palp narrower than outer plate Maxilla 2. inner pl^e about same width as outer, with facial ^etae, Mhxilliped, inner plate not reaching end of articled; rioter plate not reaching end of article 5; unguis (article 8) about equal La length to article 7. Peraeopods 1-4, ventral margin of coxa 2 with 1 1 long setae; coxae 1. 3, and 4 with 2-5 sclEte. Gnathopod ] : coxa different in shape from coxa 2, narrowed distal ly; basis inserted midway on inner face, not deadly setose; carpus shorter than propodus, anterior margin serose only at anterior junction with propodus; propodus, palm concave, defined by single small spine; dactyl only as long as palm of propodus, posterior (inner) margin, with few short setae and cusps, Gnathopod 2, basis, anierodistal margin with few striduta- tion ridges; carpus shorter than propodus; propodus, width AMPH1PACIF1CA VOL, 1 NO. 3 OCTOBER 15, 1994 70 FIG. 2. Photis ttnearmanus, new species. Adult male (3,4 mm) HOLOTYPE. Lateral views: whole body, mandibles, and maxilla 1; other views medial Scale 0.1 mm. J.7 times width of ptopoto ofgnathopod 1, palm oblique, with shelf at dactyl and shallow protuberance ir idway* setae at dactyl hinge nearly as long as propod us; dactyl scarcely overlapping palm, inner margin evenly curved, with spine and seta! cluster proximal of unguis, Peraeopod 3, coxa with row of stridulation ridges- on ventral margin, Peraeopod 4, coxa, posterior margin not excavate. Peraeopods 3 and 4, basis not expanded; mcrus wider than carpus and produced anteriorly over about 1/4 of carpus; dactyl shorter than propod us. Peraeopod 5, coxa similar in depth to coxa 4. Peraeopod* 6 and 7, coxae smaller than coxa 5; other articles of peracopods 5-7 lacking. Pleon plates 1-3 not posierodistally notched. Picon and urosome without dorsally erect setae or cusps, Llropod 1, peduncle without lateral eedysial spine proximally oriooth- likc process extending ventially below rami; rami tipped by 1 short spine, Uropnd 3, peduncle with single spine ventrally at origin of rami- outer ramus 273 length of peduncle and tipped by I long sc La; inner ramus about 1/4 length of outer ramus* ripped by single short spine. Telson apices marked by jingle long seta and small knob. Condition, Without antennae, right peraeopods4-7 t and left peraeopod* 5-7/ Adult female. Unknown, Etymology. From the Latin, linearis, meaning linear, and meaning hand, referring to the oblique, nearly I inear palm of the propod of gnathopod 2 of the mature male. Distribution* Known on ly f rum Ui issingle collection in Santa Maria Basil], at 92 in in depth. Taxonomic Commentary. This is (he only species on the northeastern Pacific coast whose adult male has an oblique palm on the propod us of the second grialhopod. The relative sparsity of setae on the ventral margins of the coxae, the cluster of long setae at the origin of the dacty l on the male s gnathopod 2, and the concave palm of the male's gnat hoped I are also distinctive, although not unique among regional species. AMPHIFAC IF! C A VOL. I MO, 3 OCTOBER 1x1994 7| Gammaropsis l Podaceropsis) ocettau j, new species (Fig, 3) Material examined* TYPE MATERIAL: Holotype: adult male (USNM, catalogue no. 239495), U.S. A*: Califor- nia: off Pt. Argue No 134°33,66'N, 120°56.3 1 ' W), 590 m. 5 January 1984. California Phase II Monitoring Program, Minerals Management Service, Pacific OCS Office. Santa Maria Basin Project , station t)55, B5S-0J -TX,MBC Applied Environmental Sciences. colleeior. Allotype, aduli female (USNM. catalogue no. 2394961, same location. Paratypes: I adult female, 3 juveniles (US MM, catalogue no, 239497); 1 adult females, ZjuvcnilesfNHMLAC, catalogue na LACM 84-285, 3); 2 adult females. 2 juveniles (SBMNH. catalogue no. 35646); 2 adult females. 2 juveniles (CMN, catalogue no. NMCC 19934X53), all from the same location. Diagnosis, Eyes weakly faceted, unpig menlcd. Anten- nae, setae maximally as long as Iasi peduncular article. U roped J, peduncular ventral spinous process less than half length of shortest ramus. Gnat hoped 2 of male, propodus, palm nearly transverse, centrally notched, and defined by I spine and change in angle; dactyl not longer than palm. Peraeopod 5 ol male, basis shallowly excavate on posterior margin. Gnathopod 2 of female, propodus, palm shallowly excavate, Description. Adult male (3.8 mm) Holotype: Head lobe triangular, but not anteriorly acute. Eye oval, with about 12 unpigmented facets Antennae 1 and 2 equal in length. Antenna 1 moderately setose with long setae posteriorly, article 3 longer than ankle 1 ; accessory flagellum micro- scopic bulton. Antenna 2 moderately setose, with long setae alsOj flagellum not pediftimi* longer than article 5, distall y spinose. Upper lip* episiome acutely produced. Mandible with 5 rater spines; molar flake present; palp strong, article 3 hardly wider d is tally than proximal ly. both articles 2 and 3 with numerous setae* article 2 longer than article 3. Maxilla 1, inner plate with single long seta; palp somewhat narrower than outer plate. Maxilla 2, inner plate nearly as wide as outer, wi tli row of facia I setae. Max ill iped. inner plate nearl y reaching end of article 4; outer plate not reaching end of article 5; unguis (article 8) as long as article 7. Peraeopods 1-4, coxae., ventral margins with minute setae only. Gnathopod I, coxa similar in shape, to and not shallower than coxa 2; basis inserted tiiid-distally on inner face, not setose an tenodistally; carpus longer than propodus, with anterodistal cluster of setae; propodus nearly simple, palm indistinct, defined by single long spine: dactyl much longer than palm of propodus. posterior (inner) margin with several short setae and cusps. Gnathopod 2, basis without stri dotation ridges; carpus shorter than propodus: propodus, width 2 r 5 times width of propodus of gnathopod L palm nearly transverse, with protuberance near origin of dactyl followed by oval incision, spine at palmar corner, setae at dactyl hinge about 1/2 length of propodus; dactyl not toothed, only as long as palm. Peraeopod 3, coxa without stridulation ridges art ventral margin. Peraeopod 4, coxa, posterior margin not excavate. Peraeopods 3 and 4 basis not expanded, merus wider than but hardly produced over carpus; dactyl not elongate, much shorter than propodus, Peraeopod 5, coxa as deep as coxa 4; basis moderately broad- shallowly posteriorly excavate in aduli male; merits shallowly concave posteriorly; carpus with cluster of spines at posterior junction of propodus: propodus with few- spines along anterior margin; dactyl not cusped. Peraeopod 7, coxa noi expanded. Peraeopods 6 and 7 simitar in shape lo peraeopod 5, although bases narrower. Picon plates 1-3 with few minute setae posterodistally hut without cusps or ridges, Urosomc segments 1 and 2 with pair of dorsal I y erect setae but without cusps. Uropod I, peduncle without lateral ecdysial spines, hut with spinous process extending ventraliy below rami about 1/3 length of outer ramus; rami tipped by 2-3 spines. Uropod 3, peduncle spinose dorsal! y at origin of rami; outer ramus nearly as long as peduncle and tipped by 1-2 long setae, inner ramus as long as outer, tipped by l spine. Telson apices marked by nipple and setal cluster. Condition, Without peraeopods 5-7. Adult female ov. {4.4 mm) Allotype; Gnathopod 2, propodus, palm shallowly excavate. Brood plates moder- alely wide, setae with book at each (ip. Other features as in male. Condition, Without righi peraeopods 4,5* and 7, and left peraeopods band 7, Etymology, From the Latin oCfilata, referring to the relatively small unpigmented eyes of this species. Distribution, Known only from this single location in l he Santa Maria Basin, at 590 in in depth. Taxonomic Commentary, The faceted but unpigmented eye distinguishes Gammaropsis ocellatu from other mem- bers of the subgenus on the North American Pacific coast. Another deepwater species* Gammaropsis {Podoceropxis} kemadea Stebbing, also lacks pigmented eyes* but differs considerably from G. ocdldta in having a much broader propodus of gnathopods 1 and 2, a more enlarged and transverse palm on the propodus of the male gnathopod 2, and a longer carpus re I alive to the merus on peraeopods 3 and 4. The body is also dorsal ly setose, which is not the case in G. oceltiita. Gammaropsis acelhta most closely resembles; G- {P). bamardi Kudryashov and Tzvetkova, which has been described from southern and western Sakhalin, Russia (50°N, 145°W) and Vancouver Island, British Columbia (48°4S , N 1 1 25° 12.5’ W) (Kudryashov and Tzvetkova, 1975; Coulan, 1983). Gammaropsis octMafa differs from G. bamtirdi in having an unpigmented eye, more transverse AM PH I PAG F! C A VOL. i NO. 3 OCTOBER 15, 1994 72 FIG. 3. Gammaropsis ocellata. new species. Adult male (3 8 mm) HOLOTYPE: whole body, distal articles of guathopods 1 and 2 (setae omitted), mouth parts, and tel son; Adult female (4.4 mm) ALLOTYPE: guathopods I and 2. Lateral views: whole body, mandibles, and maxilla 1 ; other views medial. Scale 0, 1 mm, palm of gnathopod 2 in the male* more concave palm of gnathopod 2 in the female, and leas excavate peraeopod 5 basis in the male . DISCUSSION Ph&tte typhfaps arid G. oceliam demonstrate the ten- dency of deepwater or caveniicolons ampin ipods to lose eye pigmentation and/or Facets and lengthen their antennae. Since no phyleiic treatment has been developed for either genus, it cannot be determined whether these species bear other apomorpbic features* Phot is typhiops and F. hneartmnus belong to the poorly setose group of photids which lack a dense fringe of setae on the ventral margin of the coxae. Males of both species are stridulators and the second g rta thopods are moderate! y sex ually dimorph ic . S Lridu h \ im is the norm for photids, and is presumably of value for com- municating mating intent, particularly in the close commu- n ity con lac t that was fou nd for Photis typhfops< Gammaropsis oceU$%i shows the same sort of sexual traits as other tnetro hers of the subgenus. The subgcmis Is very conservative in its range of sexua l dimorphism. Gammaropsis oceitata show's less exaggerated alteration of the second gnathopod and fifth peraeopod than in some other species, suggesting that the specimen described here may not have reached fully mature size. However the toss of eye pigmentation is significant and unique in the genus. REFERENCES Austin, W. C. H 1985. An annotated checklist of marine im vertebrates of the cold temperate northeast Pacific. Khyotan Marine Laboratory, Cowichan Bay. B.C. Vols, I III: 682 pp. Barnard, J. L.. 1962. South Atlantic abyssal ampli ipods collected by R. V, Verna. Abyssal Crustacea, Verna Research Series 1: 1-7S, AMPH IPACIFIC A VOL.i NO. 3 OCTOBER 15, 1994 73 f 1967. Bathyal and abyssal gaminarideafl Amphipoda of Cedros Trench, Baja California. Bull. U.S.Natl.Mus. 260: 1-205. & G- S, Karaman, 1991. The families and genera of marine gam maridc an Amphipoda (except marine gammaroids). Rec. Austral. Mus, Suppl- 1 3, 866 pp. Bousfiehf E. L., & C P. Staude. 1994. The impact of J. L, Barnard cm North American Pacific ampbipod research: A Tribute. Amphtpacifica I (1): 3-16. Cadien, D., 1991. List of the marine amphipod faunas of [he Temperate and Boreal Northeastern Pacific Ocean, in- eluding literature records of occurrence between Bahia San Quiniin, Baja California, and ihe south side of the Aleutian islands, incorporating nomcrtdattirai changes listed in Barnard k Katanian, 1991. SCAM1T Tech. Pnhl., Sept. 1991. 21 pp. Con Ian. K. E. , 1 983 . The amphi pod superfam il y CortJjtfuoidea in die northeastern Pacific region, 3. Family Isaeidae: systematic s and distributional ecology. Nail. Mus. Nat. Sci. (Ottawa), Publ. Nat, Sci. 4: 1-75. Guijanova.E F. , 1955. Novyc vidy bnkoplavov (Amphipoda, Gammaridea) iz sevemoi chasti Tixogo Okcana. Trud, Zool. Inst, Akad. Nauk SSSR IS: 166-218, Eshimaru, S. t 1994. A catalogue of gammaridean and ingol- fieilidear Amphipoda recorded from the vicinity of Japan, Re-pc, Sado Mar. Bio. Sta., Niigata Univ. No, 24: 29 - 86 . Kudryashov , V . A , &. N . L, Tzvetkova, 1 975 . New' ami rare species of Amphipoda (Gainmaridea) front the coastal waters of the South Sakhalin, Zool. Zhur. 54: 1306- 1315. (In Russian), Myers, A, A., 1988, A el ad i Stic and biogeographic analysis of the Atirinae Subfamily nov. Crustaceana, Suppl. 13: 167 - 192 . LEGEND FOR FIGURES GN - gnathopod; JV - juvenile; LL - lower lip; LFT - left; MD - mandible; MX - maxilla; MXPD - maxilliped, PLP - palp; RT - right; T - telson; UL - upper lip; U ■ Limped. Vol I, No. 2. ERRATA OF SUBJECT MATTER Editorial. p. L . The correct address for Roy J, Kropp is: Battel le Ocean Sciences, Duxbury, MA 02332 USA, Paper No, I: Pleustinae p. 42 et sequ. Apologies are extended to readers who may have encountered difficulties in connecting labels with fig- ures. Paper No. 2: Phoxocephalidac, p. 83. The tribute to Dr. Arthur May should read: "The species Ma fid ih u/o p h avu .% mayi Jarreti & Boustield. 1994, is named in honour of Dr. Arthur W. May, President and Vice-Chancellor of Memorial University, St. John’s, Newfoundland, and former President, Natural Sciences and Engineering Research Council of Canada”, p. 77 et sequ, Apologies arc extended for die heavy over- printing of the plates, esp, figs, 8, 17, 28, 29, p, 102, et sequ, Berkeley Sd, - corrected to Barkley Sd, p. 120, Fig. 27. Reversed curved line on pcracon segment 2 should he removed. p. 125. Add to the bottom ofihe page: M. videns is similar to H, conlame and H. ellisi ", p r 147- All references following Austin, W, C,. and before Barnard, K. H„ are attributable to Barnard, J, L, AMPHIPAGRCA VOL r I NO, 3 OCTOBER 15.1994 74 The Canadian Field- Naturalist A continuation of 115 years of publication on northern North American biodiversity — 1880-1994. The Canadian Fie Id- Naturalist is the official publication of The Ottawa Field-Naturalists 1 Club and features both articles and notes on original research and observations on the natural history of not them North America (including distribution* faunal analyses, taxonomy, ecology, and behaviour). Issues include news and comment (Club annual meetings and aw aids, tributes and review papers) and book review and new title sections. Since 1 984, it has presented edited Status Re pons for individual species designated by the Convention on Species of Endangered Wildlife in Canada (COSEW 1C). It is entirely supported through club membership and subscriptions, page and reprint charges. The Ottawa Field Naturalists 1 Club has 1041 members and The Canadian Fieid- Namralm an additional 248 individual and 573 institutional subscribers* for a distribution of 1 862 copies. The Ottawa Field-Naturalists' Club was formed in 1879 by scientists from embryonic federal departments including the Geological Survey and the Dominion Experimental Farm and leading amateurs; this type of mix remains its strength to this day. The Club quickly emphasized publication, and for seven years beginning in 1880, it annually issued the Trans act ions of the Ottawa Field- Naturalists *■ Club, With volume 3 in 1 887, the Transactions became a subtitle of Volume 1 of The Ottawa Naturalist, a monthly publication. With Volume 3 of The Ottawa Naturalist in 1889 the emphasis changed from local members reports to national ones, and in 1919 the journal was renamed The Canadian Field-Naturalist (starting with Volume 33 which was Volume 35 of the Trans- actions but this subtitle was subsequently dropped). The issues per year were gradually reduced from 1 2 to 9 to 6 and eventually to 4, the latter beginning with Volume 67 in 1953, However, the annual number of pages increased, in 1988 (volume 1 02) reaching a record of 798 with a the largest single issue of 2 16 pages in 102(2). Since 1967, the Club has separately published a local (Ottawa area) natural history journal, Trail & Landscape i now also issued 4 times a year. Submissions to The Canadian Field-Naturalist and predecessors have long been peer reviewed, first through a ' Publishing Committee", later "Sub- editors”, and then “Assistant Editors” until the designation “Associate Editors” was adopted in 1885 and maintained ever since, Currently, most submissions also go to at least one (anti often more) additional neviewer(s). Associate Editors are listed in every issue and since 1 982 additional reviewers been acknowledged annually in the Editor's Report. A formal publication policy was published in The Canadian Field-Naturalist 97(2): 231-234, “Advice to Contributors 11 is published in one or more issues annually. The current subscription rate is $23 for individuals and $38 for institutions. Postage outside Canada is $5.00 additional. Subscriptions should be sent to The Canadian Field-Naturalist, Box 35069 Westgate P.O., Ottawa. Ontario, Canada KIZ 1A2. Manuscripts for considera- tion should be addressed to Dr, Francis R. Cook, Editor, Canadian Field- Naturalist, RR 3 t North Augusta, Ontario, Canada K0G 1R0 Francis R. Cook AMPHIPACTFICA VQL r I NO. 3 OCTOBER 15,1994 75 THE PHYLETIC CLASSIFICATION OF AMPHIPOD CRUSTACEANS: PROBLEMS IN RESOLUTION* by L. L. Bouslicld 1 and C.-t $hth J ABSTRACT Tlii: phyletic classification of amphipod enk-itaceans has been a major source of disagreement among principal recent workers. The disagreement results; at leat>n partly from the marking effevls of converger! or bomoplaaioui morphology not oit3v on snpeffamijy and subordmai recognition, but also on the determination of closest phyletic lister groups to (he Ampbipoda within theMalacostraca. The most recent attempts eil phyldic classification of amphipods. Based on new evident*, partly from recent behavioural work of CVIN colleague K. E. Cnnlan. this study takes a morphological -behavioural approach to solving such problems at al] levels of classifies! ion- Among malacoslracan potential sister groups, die Amphipoda appears phylclically least remote from die Mysidacea, hut more remote from the Henhetmdea and Lite [sopoda. Within the Amphipoda, two' natural subordinal groups are reccigtiized. vi 2-, (he primitive, relict Ingolfielhdea. and the more advanced, dominant Gammandea, both with extant members itt marine and freshwater habitats. Within the Gam mar idea, two exclusively marine, infraordmal groups, the Hyporiidea andtheCaprelhdea, have possibly arisen from stegocephalid- and potiocerid-hke ancestor respectively The mfraorders and super families within (he Gammaridea may be oraLinized broadly and semi - pby 1 etic all y iruo "Amplnpoda Nataima" and “Amphipoda Reptan ti a 1 1 , Arttdogotis to categories formerly employed within the malacostracan Decapods- The former category includes reproductive]} 1 free-swimming groups, with direct mating (usually Lacking pre- amplexus ) mostly freely in the water column. Typically here. J he male in sexually specialized in the antennal sensory organs fe.g. possesses callynophore.mlceoli and brush setae i. eyes, and tail fan, but seldom in the gnaihopods, The mature male stage is also smaller than the female and is a terminal life stage (non -moulting, often non*feeding}, Cotnponen is of ihe second c a Legary are mostly ben tli ic or in faunal in all life stages, m atiisg occurs ort/in the bottom, with preamplejuis (precopulatory grasping of the female and/or agonistic behaviour toward other mates). Here also, the male is usually the larger, is usually sexually specialized in the gnathopods but not markedly in sensory organs or tail fan, and is indeterminate iu growth (mates during two or more life stages). The very few anomalies within Ibis classification are Variously attributable to delayed loss of plesiomorphic structures or to convergent morphology and behaviour, in specialized forms. INTRODUCTION The phyletic classification of amphipods has long been trough! with difficulties m<\ much controversy among prin- cipal workers. Their views lend to lie "colored” by their experiences with various taxonomic and ecological subgroupings, particularly within the Gammaridea (e.g, Bousfield(1979, 1982a 1983); Barnard and Karaman (1980); Holsingcr (1989); Stock (1985); Ruffo (1989); Lincoln (1979); Schram( 1986 k Contributing to thi&diffieulty isthe relatively large size of this; crustacean ordinal group (more than 7000 described species in 4 suborders and more than 1 25 families), and the large number of external morphologi- cal characters 1100 +) employed variously at higher levels of classification. The current stale of the problem of classifi- cation within the Amphipoda seems analogous io (he tale of the three blind men who were asked to describe an elephant based on the part of the beast that each happened to be touching — trunk, leg, onail — with three widely differing rational piclure therefore seems possible only by character' izing all body pans, of all component groups, simultane- ously. Faced with these difficulties and limitations, some authors (e.g, Ruffo cl a I (1990), and Barnard and Karaman ( 199 1) have expediently adopted a simple, pragmatic, alpha- betical listing of families within suborder*, as is widely accepted for classifying genera within families and species within genera, However, a useful phyletic 'lead ' has been provided by major workers within suborder Hyperiidca (c.g. Bowman & Gniner, 1973) and Caprelhdea (e.g. McCain (1970); Laubiiz (19701k Also* in order io avoid being overwhelmed by unwieldy numbers of names and volume of taxonomic detail within the much larger suborder Gammaridea, others (e.g, Lincoln (1979); Schram (1986); and the writer (Sons Feld. 1979. 1983) have attempted to reduce the classificatory problem to a manageable li eompro- mise" by utilizing a pbylelicaliy defined super family concept. Within the Gammaridea, this method reduces a results. An overall, comprehensive, and phylctically ^BaKtd orTthe Pfenary lecture, Fir*fEurope;m Crustacean Conference:. Paris. August 3], 1 9*52 ' Researcher Emeritus, Canadian Museum of Nature, Oiiiiwa, Canada, K t A 6P4 ? Research Division. Canadian Museum of Nature, Ottawa, Canada, Kl A 6P4 AMPbllPACIFIC A VOL ] NO 1 OCTOBER 15, 1994 iiumnomicaliy unwieldy 90+ families to less than two dozen readily conceptualized and readily manageable super tami- lics. ALSO, t tie number ofstatistkally significant characters of in ajar taxonomic value is. reduced to less than 50. thereby facilitating numerical taxonomic analysis. The need for a well founded, widely acceptable phyletic classification of amphtpods, especially within the Gam- maridca, is of Increasing concern. Owing to new taxonomic discoveries and revisions of older laxa. species diversify within this subordinal group is increasing at the rate ofaboul l -2% per ye ar. Correct fare i I y and superlamrJ y placement of laxasuchas Aetiopedes Moore & Myers (1 98 8) remains highly subjective and increasingly difficult Without a Confirmed phytogeny, character states cannot be “ordered" or polarized at appropriate taxonomic levels, nor can family-level units be properly defined in relation to one another. Lack of a recognized, phytogeny severely handicaps students of amphipod behaviour and physiology who require stable ancestral reference points in formulating their conclusions. Today, the Amphipoda reclaim one of the few major animal groups in which alphabetical classifi- cations appear more widely utilized than pbylctic arrange- ments. Such lack of consensus constitutes an impediment to systematic work within this suhdlscipline of crustacean sy stem atics, l c our view, this problem merits f urthe r close .scrutiny and. hopefully, early resolution. The phylelic position of Line other broadly recognized suborders of Amphipoda. the Hyperiidea< Caprdlidra* and Ingoifiellidea viz>a-viz the Gamin aridea, has been unevenly examined by previous workers. In the '' pre-ingolfi-ei I i<3 " classical arrangements of Stebbing 0 888) and Sai's { 1&95), the hyped ids were considered among the most primitive, and the caprelli ds among the most advanced higher categories of amp bipod crustaceans. Although recent literature on hyperiids contains little ' outgroup" phylelic conjecture (e.g. Bowman & Corner, 1973k the early status quo has appar- ently been rnaintaiued. With respect to the eaprellids, die more recent ' in-depth' 1 studies of Laubii* (1993) and Tak- euehi (1993) confirm, widely acceptably, the highly prob- able corophioidean origins of (he capnellids. whether mcno- ot poly- phyletically. The morphologically advanced post- lion of caprellids is maintained in cl tiding, by inference, that of their relatively recently evolved cetacean-parasitic cy amid con fees, ‘Hie small relict group of highly modified in faunal and hypogean ingolfiellid amphipotte is generally consid- ered to be phylciically very old and worthy of maintenance at subordinal level (e.g. Ruffo, 1969; Stock 1977), a view that is ampil fied here- (pp, 120). Bowman and Abele ( 1 982), however, would include the ingolfkllids within the Gant’ maridea, close to family Gammaritiae. Schnun (1986) has provided one of the most recent comprehensive reviews ofamphipod classification. Whereas he has acknowledged the relatively primitive phylettc posi- tion of the ingolridlids and hyperiids. and followed phyletic arrangements of supecfamilies and families within the Gam- uiaridea proposed earlier (e.g. Bousfield, 1979, 1982a, 1983), he lias placed the caprellids in a primitive position, close to the ingolfiellids. He has advocated the use of rigid dadi sric techniques (e,g. a Wagner 78 program) in produc- ing a natural classification. However,, In agreement with Ridley (1983), we find many bask or "obvious " as sump- tions about character states to be often flawedby homoplasies; resulting cladograms in which- these are not recognized are thus less realistic than phenngrams in which homoplasious tendencies are selected out or otherwise minimized In this essay, we propose to treat the classification of ampbipods phyleiically, but with a somewhat semi-prag’ malic approach, Aiter the fashion of D. H. Steele (1988, et scq r ) who noted dial ampbipods were primarily swimmers and dingey and Secondarily crawlers and burro wers, we Lave borrowed from older decapod crustacean classification the terms ‘Natan Lia'" (for the reproduetively swimming and pelagic types) and “Replant ia” (for the reproductive)# bot- tom-crawling and benthic categories). This approach uti- lizes reproductive (mating) morphology and behaviour, in both sexes, as its principal phyletic basis. Whatever the nature of the morphology and life style of mature Females and immature stages of both sexes, reproductive morphol- ogy tends to be displayed most diagnostically in die mature male stage. Of partcular significance there is the form and armature of Ihc antennae, gnathopods, and uropod 3 and, to some extent, in the mechanical coupling organelles of peraeopods,p]copisi$.andurof>ods. The approach also fac- ilitates the solution, or neat -solution, of some longstanding problems of natural ordering of character states, and their application at proper levels of phyletic classification. ACKN OW leix; EMENTS The work was conducted mainly at the National Mu- seum of Natural Sciences iia Ottawa over the past five years, and collated in research association with the Royal British Columbia Museum in Victoria, Officers of the following institutions have contributed significantly to the refinement ot ideas in this work; Dr. K, E. Cunlan, , Canadian Museum of Nature, Ottawa, Canada; Df.J.R, Holsinger, Old Domin- ion University. Norfolk, Virginia: Dr, D r H. Steele Memorial University, Si. John's, Newfoundland; Dr. Pierre Brunei, University de Montreal, Quebec; Dr, Roger Lincoln, British Museum, London; Dr. J. K. Low ry. Australian Museum, Sydney ; and Dr, Les Walling, Darling Marine Center, Mai ne, US A. We are especial I y grateful to Dr , D. R, Calder, Royal Ontario Museum, Toronto, who .supplied material of a new species of Vulettbpsis , and to Moira Gal-braith, Victoria, who supplied material of pelagic eusiroideans and lysianassids used in these comparative morphological stud- ies, Helpful editorial commentary was provided by Dr. C. P. Siaudc, Friday Harbor Laboratories, and by Philip Lambert. Royal British Columbia Museum. Preparation of the line illustrations was greatly facilitated by Susan Laurie Bourque, Hull. Que,. and Floy F,. Zinin, Cupertino, Califor- nia, Basie work leading to compilation of the presen I results was carried out partly under government of Canada NS ERC grants (1987-90), AMPHIPACIF1CA VOL. I NO 3 OCTOBER 15, 1994 77 External Morphology of the A mp hi pod Crustacean. By way of review, the general external morphology of 'a g&ammaridean amphipod has been diagrammed previously in Bousfield (1973), Barnard &. Karman (1991). and in several other popular arid semi-popular works (e.g. Staudc. 1987). In Figs. I A and ] B, the principal features of repre- sentative member of Amphipoda ''Natantia" and "Repiantia' respectively, are outlined. Amphipods ate similar to most members of the subclass MaJacostraca (large crustaceans) in having a finitely taeina- tifced body; head with 5 pairs of appendages; thorax with 8 pairs (first, pair fused to head as maxillfpeds); abdomen with 6 pairs, and terminating; in a small supra-anal flap or telson. The order .Amphipoda is- superficially similar to most other orders within supraordcr Peraearida in which the carapace is much reduced or tacking; in having eyes that are sessile or near-sessile; mouthparts that am concentrated in a buccal mass beneath the head; thoracic legs that arc uniramous (or nearly so); and lecithotropk (nonplanktonlc) development of eggs within a thoracic brood pouch of the female. Amphipods differ from all other malacottracans in hav- ing ambulatory thoracic (peril eonal) legs arranged m two dislinct groups; the first four pairs are directed forwards, wutti the dactyls (claws) backwards, and the Iasi three pairs are directed backwards, the dactyls forwards, hence the name. “amphi'V’pod” (both kinds of feet}. This contrasts with the “tan- wise” or radiating position of the thoracic legs in isopod crustaceans, A second distinctive feature, unique 10 amphipods, is the arrangement of abdominal limbs: due first three pairs are biramous swimming legs (pleopods) and the hind three pairs are thrusting legs (uropods), This arrangement of abdominal limbs contrasts with that , which consists of fi ve pairs of pleopods and one pair of uropods in all other cumalacostracan crustaceans. In amphipods, tail thrust drives [be animal forwards. whereas in decapods the tail thrust is typically a rearwards 'escape reaction ", The diagnostic features of amphipods that male freely in the water column (Natan(ia) are shown in Figure l A. and are descibed in detail elsewhere. Tire body is slender, often toothed or carinate above, with large- powerful abdomen., large pleopods, and lanceolate, serially spinose uropods. The head is generally short and deep, with rostrum , and eyes variously pigmented or lacking (abyssal forms). The antenna are slender and elongate. Antenna b peduncle stout; basal segments of flagellum often fused and strongly armed with aesthctascs ( eh e mo- sensory filaments), forming a callynophore; accessory flagellum short or lacking (in byperiidsj. Antenna 2, peduncular segments 3-5 slender, anterior margin (of male) lined with fine filaments ( brush setae} and often calccoli; flagellum elongate (esp. in males), often with calceoU. Mouthparts basic, mandibular and maxillipedal palps usually projecting anteriorly. Coxa) plates 14 various, usually shallow, similar but often unlike. Gnaihopods I & 2 usually slender, weakly subchelate, with slender carpus and propod, seldom sexually dimorphic. Feraeopods 5-7 usually slender, usually subsimtlsr (homopodousL bill peraeopod 6 is often longest; coxae postcrolobate (hind lobe larger). Telson usually large, and bilobate (fused and plate-like in hy peri ids). Coxal gills large, often pleated on peraeopods 2-7. Diagnostic features of benthic amphipods* the Replan ha, that mate on or in me bottom substrata, are shown in figure I B . The body tends to he short and compact, often flattened dorsvenually. seldom with dorsal teeth or eari nations. The head is usually long and shallow, lacking rostrum* eyes usually small. The antenna tend to be short, with stout peduncular segments, especially in males; callynophore and brush setae never present and calceoli rare. Monthparts variable, mandibular and maxillipedal palps usually visible. Coxal plates 14 various, from large, deep, overlapping, to small and basal I y separated- Gnaihopods often large, strongly subchelate, stToogly sex uad y dimorphic. Peraeopods with relative short stout segments, and anierolobate coxae (front lobe the larger). Abdomen short; plcopods medium to reduced or highly modified, Uropods short, stout, rami Linear, with apical spines, Uropod 3. rami usually shore margins spinose. or highly modified, seldom sexually di- morphic, Telson lobes variously fused, plate-like. Coxal gills plate-1 ike or sac-like, never pleated, often lacking on peraeopod 7. These diagnoses arc intended as a generalised guide to basic amphipod motophotypes. They do not apply to any part icu Ear species, nor to immature stages, Within each group are eKCeptioiMf cases that resemble species of the other group. S ueb encounters provide one of the fmslrat ing 'joys n of attempting to classify amphLpod crustaceans. The phytogeny of the Amphipoda as a group within Ihe Peracaride The phyletic positioning of the Amphipoda has also been the subject of considerable controversy. The most wide ly he Ld (cl assical ) v iew , tha l arnph tpods are most closely related to isopods, is held by a number of modem workers iodl uding Bowman and Abe le ( 1 982), Stock (pers . comm u n , ) and Schrem 1984, 1986). Other workers including Dahl (1963), Walling (1981), and Bousfldd (3988) have pre- sented evidence that the natural sister group among the Pertcarida is the My sidacc a (sens, fat , ), A few others (e ,g. D. H. Steele, and recently War! ing (pers.communic.)) have looked for an ancestry outside the Feracarida, and do no! rule out the Syncarida as the closest natural outgroup among ike EumaJacostraca, A basis for a possible mysidacean common ancestry is depicted in Figure 2, A typical giimmarideaD amphipod is represented by the frftoxocephaloidean (lower right), At first glance, it appears to have Li Hie in common, at least exter- nally, with the- various forms of Mysitfacea in the upper fig ures- The Mysidacean s are much more plesiomorphic in possession of a distinct maxillary carapace, and fully btramou s thoracic limbs, among other differences. However, the re tab vel y primili ve i ngol fiel lidea n amphipod ( lower m iddle AMPMPACMCA VOL. 1 NO. 3 OCTOBER 15, 1994 7 # Rostrum Ranis , papilla Teleori Uropod Brush Setae Calcooli Teiaon Gnathopods Ur o pods Poraeopods Ploon Plates i Tela an Gnathopods Uropods Antennaa Pleopods FIG, L Ik^ic Morphology of the Amp hi pod Crustacean. A. Natan tirt (Hyperiop&idaeJ tJ. Keptantia (Meiitidae) AMPHIPACJFJCA VOL. I NO. J OCTOBER 15, 1994 79 L PttSJWCEA i t'ETALDPHTH^LUT DA£ (Hansenomysts falkiandica) C L0PH0GA3TR1M (Gnathophausia) f, MlfSlDftCEA % (Hansenomysls an t arctics) DL MV5] DACF-A J HY3IWE (My sis reiicta) H. AWPHIWOA IKOLFI'ELLIOU G. AITHIPDCA : GftlflAVDEA J. JhhffHIPCDA: PflMOCEPHAi.tME FIG. 2 EXTERNAL AND INTERNAL ANATOMICAL RELATIONSHIPS! EUPHAUSI ACEA, LOPHOGASTR1DA, MYSIDA, AMP HI POD A. (After Watlin^, J.9SL, and other eg orresj AMPHIPACIF1CA YOU] NQ- 3 OCTOBER 15, 1994 $Q A. CUWCEA r CTNURIM r AHBSmxtEA B. MfflOSBtiJIACtt G. SfflWRrM : JINKPIQWEA C, SPEUEMBIPHACEA H. CUMACEA D HICTACEA J. UNM DACCA E. KiCTtfCA K. ISOPOCW, PIC 3. EXTERNAL AM) INTERNAL ANATOMICAL RELATIONSHIPS OF MALACOSTRACAN SUPERORDERS SYNCARIDA, CUMACEA, BRACHYCARIDA, MICTACEA AND ISOPODA {After Walling, 19M, Bowman & Tllife, 19SS„ and other sources) AMPHIPACtFICA VOL. I NO. .3 OCTOBER 15, 1994 gl right, and also Fig, 27) shows (1) vestigial stalked eyes, (2) partly Cleft maxilliped basal segment, and (3) urtipod 2 much larger and stronger than uropod 1, both with serially setose rami, as in pleopod* elsewhere. All of 111 esc features tine more prominently and function ally present tnmy&idaceans, esp. in family Petal opthth-almidae Fig. 2£, Th us, the enlarged uropod 2 of the ingolfiellidean may be homologous pleopod 5s anomalously longer than die anterior plcopods in this mysidacean family. Members of this family also dem- onstrate. a trend to "fore and aft" subdivision of the thoracic legs, as in the Amphipoda. Also, the internal anatomy of the mysidacean e inconsistent with the overall morphological evidence developed by other work- ers, Although other* soon adopted the alphabetical system (c,g, Ruffo el al(l983, 1990 ), his phyletic thesis received little published support elsewhere* As pointed out by Scbram (1994), his co-author (in Barnard & Karaman, 1983) wrote a dissenting opinion in a separate appendix to that paper. Despite these informal phyletic proposals, the -classification system of Barnard's subsequent collated works (e.g. Barnard &. Barnard* 1983; Barnard & Karaman, 1991) continued to be essentially alphabets cai. Meanwhile, the need to develop a broadly acceptable basis for natural classification of related higher taxa was be- i ng more widely recogn ized. Bui ycheva (1957) aebi e ved a phyletic "breakthrough 1 by introducing the soperfamily con- cept. TaJitroidea, that combined, all terrestrial talitrid and aquatic byalid-like families. The success of this move was soon followed by i. L. Barnard's grouping of all "llcshy- telson ' tube-building amphipods within newly proposed supeifamily Corophioidea (1973). Similarly, the families of shallow -water gammaridean amphipods of the N, Ameri- can Atlantic region were presented mainly in closely related groupings (e.g. Pojitogeneisdac-Bateidaie-Olliopiidac- Eusiildae, and Dexaminidae-Atylidae-Ampeliscidae) each equivalent to an informal superfamily, by Bousfield (1973), During the mid- 1970's, however, the need to group related families was matched by an equally strong need to separate out obviously unrelated major taxa that hud long been, submerged as informal subgroups within an "um- brella" higher category, Titus* several distinctive free-swim- ming or free-crawling, marine, freshwater, and hypogean groups had previously been "dumped' within an increas- ingly large and and unwieldy liclerogeneous family concept long know n as ’good old Gammaridae 1 , Similarly, several families of free -burrowing but phyletically disparate amphipods (e.g. Phoxocephalidae, Haustoriidae (Ponto- poreiidae), Argissidao. the urotholds. and even ihe Dogiel- inotidae) had long been listed in close phyletic or semi- phyletic proximity (e.g by Sars (1 895), Stebbing ( 1906), and Guijanova (1951, 1962). The gammaioidcans were soon broken up into several new superfainUies.indudirig the Crangonyctoidea. Mclphidippoidta, Melttoidea (later Had- ziodea), Bog idle] loidea; with various family allocations (e.g. Gammarellidae) to Eusiroidea* etc. (BousfiekJ, 1977), With the su perfanu ly concept thus broade ned, a phy lelic arrange - mem of all gammaridean amphipods was then formally attempted (Botisfield, 1979a), Encouraged by the accept- ance of several of these linkages by Lincoln ( 1979), Holsinger (1992a) and others, the supeifamily and family concepts were further refined (Bousfield. 1982a, 1983). These included a phyletic sorting out of the major sand-burrowing taxa, a reclassification still in progress (e.g. Bousfield, 1 989), in support of the initial formal phyletic arrangment of superfamilies; Bousfield (1979) developed a phylogenetic tree of relationships I hat is examined again in this study {p. 1 25), Trees provide a quick ” visual" of basic relationships between groups of organisms, and have been widely ac- cepted in cumalatx>siracan classification (e.g. Slewing, l%3) r By employing numerical taxonomic methodology modified from Sheath and Sokal (1973), these relationships became more widely acceptable (Bousfield, 1983). The classifica- tion was recognized in principle in Mark Ridley's (1983) exptanat ion of organ ic d iversi ty, a nd i n corporate*) in Lowry's ( 1986) analysis of callynophorc distribution and, with some reserv ations, in Scbram s coot preheat ve book on Crustacea (1986), The chart of Bousfield (1983), summarizing the range of plesioaporphy in selected characterxtates within 22 AMPHIFACMCA VOL.] NO. 3 OCTOBER L5, 1994 34 Suborde Coropkiidea Corophiida Corophioidea Ischyroceridae Corophiidae Caprogammari dae Caprellida Caprello idea | Caprellidae Cyamoidea Suborder Gammaridea Gammarida Gammaroidea Gammaridae [etc.] Urothoidea Talitrida [etc.] Suborder Hyperiidea Pkysosomata Physocephalata Ancestors FIG. 5. PHY LET 1C RELATIONSHIPS WITHIN THE AMPH1PODA PROPOSED BY J. L. BARNARD (1969). sub- ordinal andsttperfamily categories within the Amphipoda is provided in Fig. 6 (p. S6). The character states are ordered, with ptcsio-apoiiKirphic values of 0, 1 , and 2, and the values apply to component families of the almond-shaped enve- lopes for each superfamily and xu bord inal taxon . An i ndex of plesto-apomorphy (P./A index) was derived by adding the values across the 12 characters for each taxon and expre ssing them as a percentage of 24, the highest total possible , High EVA values denote advanced, and low values primitive* taxa. In terms of present classification orientation, we may note C ha i the envelopes for superfami I ies of Natan t la range main! y below, and those of the Replan tia mainly above, the 50% PI A level A certain degree of overlap is not unexpected, where the more advanced groups of Natan lia (e.g, Oedicerotidea, Hyperiidea, Ponluporeioidea) range above, and the more primitive groups of Reptantia (e.g, Crangohyoioidca, Gammaroidea) range below, the 50% level. Recently, computer-based methodology has been more widely employed and the results more widely accepted. However, these results may not necessarily correspond to the actual route through which a group of organisms evolved. Thus* using a Wagner 78 program, Schram anti Brusca had (by 19S6 t above) produced a cladogram of relationships among amphipod taxa that was "quite at odds with anything (then) currently its the literature 1 - Although apparently yet unpublished, such a result would command respect. Brusca and Wilson (199 1) obtained highly credible results in reclas- sifying the Isopoda, using a number of cladistic analysis packages that included HENN3G86 and FAUP (version 3.0). On the other hand, by means of a Wagner 78 program, Schram (1984) had employed 31 paired character states in developing 4 clatkgrams of relationships of major taxa within the Eumalacoshaca, all of which placed the l&opoda as the phyleiicaJly closest outgroup to the Amphipoda, However, the character slates found to be phyle heady ' sytiapoinorphic 1 in these two taxa (nos, 13, 14 , 2 1 , 22, and 31 - i.e, uniramcHJS iboracopods, pleopods lost or reduced, presence of thoracic coxal plates, eyes sessile, carapace absent) are features that are especially vulnerable to broadly eumalaeostracan convergent evolution. In our view* the AMFHIPAC1F1CA VOL. I NO..? OCTOBER 15. ]9M g$ FIG. b. RANGE OF PLESIO-AFOMORPHY IN SUBORDINAL AND SUPERFAMILIES OF AMPHIPODA (AFTER BOUSFIELD, 19S3; basic differences between isopod£ and ampliipods (c.g, t in embryonic development in inouthpan morphology, and in annulate vs. flabellatc pleopods) are more significant and less subject to homopl&Sy; moreover, such character states of the Amphipod a find much closer parallels within the Mysidacea and Lophogaslrida. as noted in the analysis of Brusca and Wilson (1991). A recent analysis of amphipod classification, using the PAUP Version 3.0k program, has produced 5 eladograms of phylogentic relationships of amphipod families and subor- ders considerably at variance within anything previously published (Kim & Kim (1993). However.the validity of these results has been questioned by Schram (1994) t since the anlysis of the entire amphipod taxonomic assemblage considered only 20 families (about 1 5% of the total land only 10 characters (of more than 50 that could be deemed useful). A further review of that study also reveals that 10 (62%) of I he setocled characters concern only mouthparts, uropods, and pleopods, of essentially non- reproductive orientation, and thus of probable lesser phyletic significance. Investigations elsewhere contribute usefully to the so- lution of problems of amphipod phyletic classification, Conlan (1990. 1991a) is continuing studies on the signifi- cance of sexual dimorphism of the gftathopods and of matt’ guarding strategies in the phyletic relationships of corophiodean am ph ipods, As we find in the present study „ her work applies more broadly across the superfamilies of Replan (ia and across the Amphipoda generally. Other major workers in amphipod phytogeny are inves- tigating potential ainphipod-syncarid relationships (D. Steele, L. Wading, personal communication). In present studies, we have yet found little evidence for such a relation- ship, but applaud I heir wide and stimulating interest in class- ificaloiy aspects of amphipod crustaceans, AMPHIPACIF1CA VOL I NO, 3 OCTOBER 15, 1994 ^6 A New Approach to Amp hi pod Phylelic Classification As outlined previously (p. 77), the current status of phylelic classification ol the Amphipoda finds no single system universally accepted or satisfactorily inciting alt major problems of natural relationship The following semi-phyletk approach to classification of amphipod crustaceans is based primarily on reproductive morphology and behaviour, as outlined recently by Conlan (1991 a, b : Fig, 7, here), In summary, amphipods that search out and mate freely, usually in the water column, tend to he closely related physically, and may he collectively ter med Amphipoda NalafUia. Those that mate on or in bottom sub- strata, following a period of male- guarding" proximity between males ami Females, are less closely related to each Other phylclically. but exhibit such similarity of life style us to be conveniently and pragmatically termed Amphipoda Reptantia. The primary features that distinguish these two principal categories are given in Table , and treated in greater detail in the following lest. Although this semi- phylelic approach covers all major groups of amphipods. at subordinal and superfamily levels, it does not pretend to solve all problemns of natural classifi- cation, at ah taxonomic levels. In this essay we have attempted to tackle some of the more vexing problems, using [be Naiiamia-Reptantia approach in a manner that may point to ultimately correct phylelic solutions. Many problems remain unresolved and await input from yet undiscovered ram and broader input from more recent and more basic taxonomic toots such as ultrastructuial analysis, electro- phoretic serology, and eventually DNA-DNA hybridization. Especially vexing to gross morphological analysis are those iaxa whose immediate characteristics are "replant" ( at fam ily and generic level) but which prove more or less closely related to groups that are primarily "natanf. We conclude that the problem of convergence is encountered in virtually every facet of phylelic in vest iga lion, and allowances lor this phenomenon must be made accordingly. In the following sections we consider the phylelic sig- nificance of sexually dimorphic characters and character states, as evidenced in both Ehe Natantiaand Reptantia. In the first part of the analysis, we consider the antennal sensory organelles, reproductively significant features of the gnathopodb, and phylelic trends cxihited by uiopod 3 and the tel son. In the second pan, we examine dassifiealory prob- lems posed by the fire sent status of hyperiid-gamrnarid and ingolfiellid-gani tnand morphological relationships, and the difficulties encountered in the study of fossorial amph ipods, and enigmatic hypogean taxa, In our concluding section we present, in tabular form, a broadly revised listing of subordinate super family, and fam- ily level laxa within the umbrella concept of Nataniia- Reptantia. Because the concept concerning Reptantia is essentially pragmatic, and because dadistic taxonomic analy- sis is especially difficult to apply within the Amphipoda, our concept of higher level phylelic relationships is presented in the form of a phylelic tree, revised from previous studies, FlG. T- Nut uni Jim! kfpiani Amphtpoda k*pruluftlYt St til he iidtrCoiilBn, 1991). In a more complete study, we might have included analysis of other major groups of appendages, especially the moulhparts, peraeopodw, and p looped s. The significance of moulhpart morphology in the phylelic classification of amphipod crustaceans has been outlined previously for gamimirideans by BouslTcId {1979. 1982a f 1983, etc,) and Barnard (3 969* etc,), for caprelhdeans by McCain (1970) and others, and forhyperiids by Bowman and Gmtier ( 1973). ]n general, mnuthpart morphology is a direct reflection of food preference and feeding methodology and is Significant mainly at the family level of classification. Although their character states seldom mirror reproductive behaviour, cer- tain features, especially of the mandible, are considered basic top phylelic class if ical ion , However, for development of more credible phylelic results we would advise caution In utilizing mouth part morphology to the exclusion of reproductively significant character stales. AMPHIFACIFTCA VOT 1 NCJ 3 OCTOBER 15, 1994 #7 The Natatitia-Reptanliii SemM’tjylctic Concept of Amphlpod Class iffeiit kin Jn a recent Slu<3y of the enigmatic new gammaridean genius AcUopede Moore and Myers (1988) opined that amphipod classification lacks a "soundly based analysis" of higher taxa or a "well founded" phytogeny, Such a comment may technically be true in a etadistical analytical sense. However, it apparently overlooks the long period of systematic stability during the first half of this century when the most widely accepted classification of amphipods was based on ihe semi-phyletic arrangements of Sars (1895), Stubbing (1 906) and other major workers, The lack of cladistic analyses in no way prevented development of universally accepted natural classifications within other major animal groups, e.g. Mammalia, Aves, Reptiles, to name a tew, lit this study, [he new higher class ill calory concepts are based on what might be termed "first principles" that may be tested eladistkally ai a later stage, and are diagnosed and described as Follows: AMPHIPODA "NAT ANTI A" 1. Primarily strong swimmers during reproductive behaviour, even where the vegetative life style is benthic or infaunal; 2. Sexes mate freely (usually synchronously ( in water column, or on/in the substratum, 3. Sexual dimorphism: in mate- seeking mates, the body form, antennal size and army lure, eye size, and structure of the pi copods, uropods and telson differ, usually strongly, from those of the female. Sexual dimorphism in gn&thopods is weak or lacking. The male is typcially smaller than the female. 4. Mate morph has a determinate moult cycle (6-8 stages); the adult stage is terminal and the male dies after mating. Females are usually seinclparous, 5 . Th e male antenna I i s near I y al wa y s equipped w ith a cal lynophore ; pedune u tar segtn cn I s 3 -5 of an (e nda 2, bear anterior marginal brush setae. Calceoli are frequently and variably present on one or both anten- nae. The flagellum of antenna 2 is frequently elongate in the male. 6. Reproductive behaviour typically docs not involve pre -ample ix us, except in in some Gedieerotoidea, and a few other phytetieally advanced taxa. 7. Almost all taxa are exclusively marine, often with strong represenEatior an the deep sea (Lysianassoidea, Phoxoeephaloidca, Stegoeephaloidea, Hypcriidea, Synopinidea, Pambmseoidea, Dexaminioidea, Ampeliscoidea, Melphidippoidea. A few eusiroideans, melphidippoi deans and allied groups (e,g., Phreut o gamma ru s, Sensomtor), and some oedicerotoideans inhabit fresh water and pontoporeioideans inhabit mainly fresh or brackish waters. The vegetative life style is free-living or commensal: a few lystauassoKJeansand p;trdaliscoideans; are ccto-paiasitic. Some eusiroidcan genera (within Pontogeneiidae and Cailiopiidae) and a few melphidippoideans ( Phreaiogammanti and Sensottator} are bypogean in fresh water. AM PH IPOD A M REPTANTIA |P 1 . Primarily mate-guar ders during reproductive behaviour. Free living forms tend to be casTiers : an Litiliie gnathopods in pre-amplexus with the female until her ovulating moult, Tube builders and semi-sessile groups are mate a (tenders. 2. Sexes mate on nr in [lie bottom, rarely in water column. 3. Sexual dimorphism of gnathopods is usually strong. The male is typical huger than the female but otherwise not markedly different in form. The antennae may differ in size sexually,. 4. Mate morph growth stages are lndeterminale(8+).wiih two or more sexual lusters; continues to leed and mates continuously alter maturity. Females are usual! uv (tern parous.. 5. Male antennae lack cal lynophore and brush setae and are seldom rarely equipped with calceoli, except in some primitive taxa. The flagellum of ahntenna 2 is not elongated. 6. Mating behaviour involves involves pre-amplexus and/or mate-a (tending agonistic displays by males, often of lengthy duration. 7. Most groups arc marine (Lencotboidea, Capreilidea) or mainly .so (HixJ/ioidea, Lilje-borgioidea, IngoJfiellidea, Corophioidea) but wilh relatively limited representation in the deep sea. Nearly all have freshwater representatives. The vegetative life style is fre-living or commensal, fossorial or domicolou*, and occasional ty parasitic (ex temal) ► The Crangonyc teddea, Gammaroidea, Bogid re U oidea and Talitroidea are primarily (or nearly exclusively) freshwater and/or terrestrial. Alt groups except the Lenoothoidc^ Corophiioidca. and Caprelfidea contain one or more hypogean species, and the BogldicLlOidea and Ingolftellidea are exclusively so. AMPHIPACMCA VOL T NO. 3 OCTOBER 15. J994 The CallvnopJiore The passible significance of Ihcciillynophtirc in phyteiic classification of amph ipods was first introduced by Lincoln and Lowry ( ! 984) ami amplified formally by Lowry r 1 986). This structure consists of a bundle of generally drwe-sel aesthetascs on the posterior, tr postcro-mediaL margin of the fused (or conjoi nt ) basal segments of (he flagellum of anten- na I. The callynophore is distributed across a wide spectrum of amphipod (ax a, including all Hypcriidea, hut is character- istic of Huperfamlly groups within the Natan tia (Fig, 8), It a Iso occurs widely across pelagic marine Malacostraca such as the Mysidacea, Lophag&slrida, Eupbausiacea, and Decs- ptida Natatilia (e.g,, Dendnobranehiata, emitted) (Lowry. 1.986). The structure almost certainly occurred in extinct presumably pelagic malawstraciin groups such as the Pygo- cephalornorpha (Mysidaoea) and various 'Eocaridacea' and WatcTStonellideic but present interpretation of fossil specie mens does not dearly demonstrate this feature (e.g, in Schram, 1986). However, (he callynophore occurs only sparsely in re productively pelagic males of the infaunal Cumacea, and is rare (perhaps secondarily developed?) tn isopods, It is apparently lack Eng in slomatopods, svucaridi, and all other essentially benthic, replant, or freshwater iml- acostrucans. With respect to function, since the callynophore con- sists oracMhetascs of various sizes and densities, its primary role is almost certainly chemosensory K but in some decapods may also be tactile or mechanical. In most am phi pod groups the callynophore ts developed only in the final adult male instar, and would seem to be of direct reproductive signifancc in the detection of females within (tie water column. How- ever, in some generic groups tc.g,, within Lysianassoidea. S ynopioidca h cal 3 ynop here- like si ruciures ma y also be pres- ent in mature females and subadult stages, perhaps indicat- ing a possible secondary' role in detection of food resources. Representative forms of cal lynopbores. within the Am- phipoda, are illustrated m Fig. 8, Lowry (1986) has de- scribed a one-field arrangement of the caJJynophorc within families Platyislinopidae. Urothoidue and Phoxocepablidae (Ptipjtoccphaloidea), a condition he considers primitive, and in some hyperiidste.g. Archaeo&cimdae), perhaps converg- ent Jy. Ln all other taxa the arrangmenl is two-field. The callynophore is especially strongly developed in pelagic carnivores and nevrophages, often where ealceoli arc weak or lacking, such as within the Lysianassoidea, Synopioidea. Pardaliscoidea, Siegocephaloidcu. and Hypcriidea, How- ever, with few exceptions, the callynophore is weak or lacking in reprodudively pelagic btil vegcla lively benthic groups such as the nestling Dex amino idea and tube -building Ampeliscoidea, and the fossorial Phoxocephaloidea and Pontoporeioidea, It is also weak or lacking in several subgroups within Natantia where the total life cycle is essent- ially benthic and infaunal (e,g. Hattslociidae ) + orcommensaJ- parasitic (e.g. some Lysianassoidea) and/or where preamplexmg reproductive behaviour has secondarily and con verge nil y developed ft.g. in Paracalliopiidac and Exoedicerotidae within Gediccrotoktea), Curiously, the callynophore is surprisingly infrequent, or weakly devel- oped, in the mainly marine, but main3y acaiceolate family Oedicembdae and, within superfamily Eusiroidea, is appar- ently restricted lo the pelagic, primitive family Eusiridae, The callynophore is almost totally lucking in the reproductive!} benthic Repiantia. including the CaprtUidea and Ingolfieilidetu even in those that have apparently be- come secondarily pelagic (e.g.„ Ma croheciopus : Gammar- oidcaj. However, callynophonde-ltkc structures have been reported from a few Amphiloctiidae (^g.Amtropheonoides, Peitocom) and C ressidae (Cressa cristate) within the primi- tive subgroups of Lcucothoidea (Lowry, 1986). We may reasonably conclude, therefore, that the callynophore land iis character stales) offers one of the potentially most n.seful criteria of reproductive life style within the Aniphipoda. Although Its occurrence across the spectrum of amphipod superfamilies is subject to some homoplasious tendencies, such abernmeies may be cor- related with nnri- reproductive life style and are thua predict- able, In broader perspective, the presence of a callynophore is a plessomorphic. or basic feature of makuiostracan repro- ductive morphology, and in our vie w provides a primary basis for development of a pltyletk classification within the Amphipoda Antennal Brush Setae The term ‘"brush setae" was first applied by the author (Bousfield, 1979a) to describe the dense tufts or clusters of short brush-like setae thai variously line the anterior margins of peduncular segments X 4, and 5, of antenna 2. A more refined term "caJlymosctae" might he coined from the Greek root employed by Lowry < 1986) in naming lire callynophore. Brush setae may occur also on ihe posterior (lower) margins of peduncular segments 1-3 of antenna l (e.g., in Dexamin- oidea). To dale, brush setae have hecn found only in the terminal male stage of pdagtcally reproductive amphipod superfamilies, and not yet m subadult males, females and/or iirunjumes. They also tXiVur in pelagic males of other pern' earidan tax a such as the Cumacea and MyswJacsa, Brush setae iire weakly lo moderately developed in calceolaie am- phipod taxa such as the Phoxocephaloidea, Poruoporeioidea, Eusiraidea, Oedieeroioidca. and Lysiariassoidea. They are almost invariably present. and most strongly developed, in non-caleeotaie superiorities of Natan tia such as the Partial - iscoidea, Synopioidea. Dexaminoidea, Ampeliscoidea, and Mflphidippoidca, but Eire less well developed or even rare within the Stegocephaioidea and Hy peri idea (Figs, 8, 30), The function of brush setae is yet unknown and conjec- tural. Although they have not yet been studied in ultra - structural detail, in gross morphology they appear as modi- fied setae, rather ihyn thin-walled as in aesthetics Their role may be tactile, during the process of copulation, when the mate is briefly in close contact w ith the female. The presence of brush seiae only in males and only in plesiomor- phtc la*a (with in the Natantia) suggests strongly lhai their function is of reproductive significance .and thus potentially of primary value in phylelic classification. AMPHfPACIFICA VOt, [ NO .1 OCTOBER L5 T FIG* 8 . TYPES OF ANTENNULAIl CALLYNOPHORES [after Barnard (1969), Bowman (1973) and other sources] AM PH1P ACIFIC A VOJ [ NO. 1 OCTOBER 15, 199-t 9[j Tilt Calceohis: Occurrence within the Amptiiptkiit. The possible significance of antennal calceoii in the phyletic clitssi fication of the Amph ipods has been alluded to variously by Bousfield <1979a, 1953), Lincoln and Hurley 0951), Lincoln fl984) and more recently by Godfrey, Hol-singer &. Carson ( 1 9S8), .Staple ion, Williams & Barnard 1 19S8) r Holsinger £ 1992)* and Steele &. Steele (1993). The principal features of these anieimal inicrostnictures have been, outlined bv Godfreyiml ( 1988), with special ref- erence to those of genera within Lfoe primitive superfamilies Cmngonyctoidea and Gamin aro idea of the R epttntia. The ealceolus is a slipper shaped membranous miooslrueture attached variously to the anteromedial segmental margins of the flagella and peduncles of berth antenna I (antcnnule) and antenna 2, The combination of its structural form (in advanced ibntis: similar to that of a parabolic radar "dish"), and its anterior antennal location, may indicate that it func- tions primarily as a rnechanorecepEor for detection of aquatic vibrations. Howe ver its ennervalion ami connection to the brain has nm yet been ascertained, nor havemicio-acousiicaJ studies yet confirmed its true function, The ealceolus is not to be confused with the aestbetasc, a sublincar thin -wailed microstnicrure of mainly chcmoscnsoty function, found only on flagellar segments of antenna I in nearly all species of Amphipotia. The aesthetic also occurs widely across malacOatracun ordinal subgroups, including the Dccapoda, The ealceolus is also readily distinguished from brash setae and other seta- like structures co-occurring on antennal peduncular and flagellar .segments. Representative types of amphipod calceoii are illus- trated here (Figs. 9 &tQ). Cal ecoli -like structures are found on the proximal flagellar segments of antenna 1 (male! of a few othermalacostmcars, notably within the Synoiritia ( An- aspidacea: Komuttga cursor ) and the Mysidaeey (Mysidaj Xerwcanthomysis pseudomacropsis). Such structures are not considered calceoii by Lincoln (pets, commimic.) and may be of different function, or convergent in form. How- ever, they arc included here as of possible phyleiie signifi- cance within theMalaeostraea and, in our view, merit further detailed comparative micro-anatomical and behavioural study. Within fire Ampbipoda, the ealceolus of the Crangonyctoidea (Ffg8 9»W) appears to be The most simpli- fied, and probably most plesiomorphic in form (category 9, of Lincoln and Hurley, 1981). It consists only of a basal stalk and elongate (usually narrow, occasionally distal ty broadened) fwdy that bears numerous (20+) elements of similar simple structure, Holsinger ( 1 992 ) has distinguished two subtypes of calceoii within the CratigonycLoidca. The cakeolus of northern Crangonyctidae is slender and elon- gate, with an simple branched internal ^brcHxirak'' configu- radon. Some separation of basal elements in Crtmgonyx rkhmondensis (illustrated by Godfrey et ah 1988) are suggestive of pftHri-^ceptacies . By contract the cakeolus of I he austral Slcrciophysingidae and Paramelitidae is typ- ically broad, paddle-shaped, and its internal tree-trunk con- figuration has more njmemus indistinct branches, a seemingly more primitive condition. Tn slightly more advanced types of calceoii (Fig. ID: Fhoxocephaloito), the elements arc fewer (ID- IS in Platyischnopidae; 4-6 in Fhoxocephalidac ); and the body may be short and spam I ate. or barrel -shaped as in same Phoxoeephalidae. In more advanced types of calceoii, the basal dement is broadened and modified into a receptacle (weakly developed in Pontoporcioideaand Gainmaioidea, strongly so in Eusir- oideah and the stalk is dis tally expanded into a bulla or resonator, weakly and more strongly in those same groups, respectively. In some Pmtoporeioidea (Bathyporeiidae). finger-like processes protrude over the proximal elements. In the most advanced types of calceoii (viz., in some Eusimidea: Gamnwcllidae, Eusiridae; Fag.9 , ) t and in some pelagic Lyrianassoidc-a (e.g. iclwopus «,pp., Lowry and Sioddare 1992), the distal elements are few and widely separated from one or more large, cup- shaped receptacles, and the bulla is prominent. With respect to the Eusiroidea, Steele & Steele ( 1 992 \ found t wo types of ca Iced i in Gamma re Mtt$ ang ulosus, viz, a large "porlogeneiid" type and a smaller, but more com- plex "gammarel lid" type. The former occurred singly only on flagellar segments of fir st and second antennae of mature males. The latter were found encircling the flagellar seg- ments of larger immature* and females as well as mature males. Although Steele & Steele Hoc. ciL l have urged caution in the use of calceofi in higher classification, their work may be interpreted as directly supportive of such use. Thus, the basic pomogeneiid type, in males only, would directly link the Gam marc tlidae to Other families with simi- lar mate-only calceoii. now placed within superfamily Eusiroidea. The smaller, more specialised calceoii of all sexes and stages of Gammarelhi^ are almost certainly not reproductive!) significant. Instead, these may assist in the detection of pelagic prey Organ isms by all life stages of these raptorial predators. The evolutionary morphological sequence within the calceoii portrayed here is believed to match more closely the phytogeny of corresponding superfamily groups, based on other character stales [see below), than does the somewhat pragmatic sequence originally provided by Lincoln and Hurley (1981). A graphical plot of the type* of calceoii and their distribution by antennal site, sex, and higher taxon, can be linked by means of a branching arrangement w ith relation- ships that, in part are remarkably similar to phyletie ar- rangement derived elsewhere from analysts of other char- acter slates (Figure 11 1. In (he first two categories this arrange mem goes somewhat beyond the relationships pro- posed by Lincoln (1984) on the basis of the taxonomic ( classi fi calory > distri bution of calceoii , In the present chart, the positions of the jnajor taxa in the various "boxes" are correlated primarily with the distribution (or lack) of calceoii on one or other (or both) antennae, along the horizontal axis and with the morphological type and its sexual occurrence* AMEWACIFfCA VOL, I NO. 3 EXTTOBER IS. 1994 $[ A. SYNCARIDA B. M YS I DACE A Xena c^r thorny sis pS£ udo mi crop S is B PONTOPORBQIDEA Pontoparniidae Dtpof&a hoyi Battiyp&refB sarer D. brew corn is Amphiporeia vfrpinlan* C. GAMMA FODEA MesogammaridaB Gammaridae Paramesogammams Mn&ieanisB Gamrnarus ocaantcus Puntog*n&iida* D EUSIROIDEA Wvyprechtia ptnguis Psaudomoara *P P sraCeptamphopi do& Pftitf- tnfiitarf dottn P&r&leptatriffh&piia Sp. cuapidata Gammara/Ius sp Gammaraffps sp. FIG* 9. TYPES OF CALCEOLI IN' GAMMARIDEAN AMPHIPODA AND POSITIONALLY SIMILAR ORGANELLES IN OTHER MALACQSTR AC ANS [modified from Lincoln & Hurley (1981 } and other sources! AMFHIFAOFICA VOL, I NO. 3 OCTOBER 15, !994 §2 PHOXOCE PH A LO IDEA P HO XOCEPH A L ID AE Tipimeginae Waft art gi sp. Pontharpiniinae MonditiviaptiQxue sp UROTHOIDAE Urothoinae UratTfO# ap. FIG. 10. PLESIOMORPHIC CALCEOLI: REPRESENTATIVE SUPERFAMILIES AN1J FAMILIES [after Jarrctt & Ekiusfirlrt, (1994 a* b), Godfrey ct ul (1988), and other sources} on the vertical axis. The vertical and horizontal axes also simulate, fan wise, an approximate evolutionary lime scale for the probable first appearance of the ancestral type of each major taxonomic group. In tliis tentative scheme, the arrangement is rooted in a presumed mysiddikeuul-group in which cafecol us-like st rue lures were possible (cf Jtenattmthoiii$tLr t Fig. 9 ), at least on antenna 1 of I he male. Such structures very probably occurred in presumed former epigean and pelagic marine ancestors of Ibe now bypogean relict suborder Ingollieliidea. and of the continental (resh water-endemic Crangonyctoidea. Such epigean and marine ancestral types have not yet been found extant, or in the fossil record, but are predicted from this study and Irani earlier considerations (e.g. Bou^rteld 1 9K2b) , t n this two-d irnens ional scheme, al 3 members of the seven calculate superfamilres, and the enigmatic imelpbidippoidean?) hypogean calceolate Sensonator valentiensis Note nboom ( 1 986k cannot t>e confined e lean ly with* n any given graphical box. Such variance is attributable to parallel development, diversification, and subsequent loss of calccoh from Ihe antenna of both sexes, presumably in response to changing life Styles within the various taxonomic subgroups. Notably, the more strongly cakeolate super- Luii ily groups (eaJccoli on both A I and A2> left column} are those in which members are primarily pelagic and/or mate freely in the water column. These include most of the PtloxOccph a Inidea. Pontopurcinidea, Lysianassoidcii, Eusiroidea, and Gediceiotoidea, The less strongly calculate superfamibes (with rare exceptions calceoli on A2 only, righi column} are found in the most primitive members of benthic superfamiliesof the Replan tia, such as the Crangonyctoidea. and Gammaroidea. The position of acalceolate superfamilies is tentative, but is guided partly by the presence or absence of an antennal caliynophore and other presumably p~ l mitive T often vestigial characters such as male antennal brush setae (see below! With respect to the sexes, the more primitive types of calceoli occur (with very few exceptions} in the 'males only' category of presumed most primitive superfamily taxa such as the Crangony cloidfca, PhoxocephaJoidea, Pon toporekridea, and most of the Ly sianassoidea ( upper two rows), Calceolate females are frequent in pelagic (especially raptorial) mem- bers of Eusiroidea (e.g. Eusiridae and GammarellidaeK in some large hypogean predators in more primitve groups (e.g. Crangonyx packardL Sensonator, p, 123), but rare in the fossorial Oedicerotidac, and benthic Gammaroidea. With respect to calceolus morphology, the more ad- vanced types occur mainly in ihe carnivorous family sub- groups, of tb e pelagic -mating Eusirioidea and Oedicerotoidea, and in the primitive be mhic Gammaroidea Gower two rows). AMPHIP ACIFIC A VOL I NO 3 CKTOBER 15, 1994 93 MYSIDA OUT-GROUP * DISTRIBUTION OF CALCEOLI A1 ONLY A2 ONLY LACKING PROTO-AMPHSPOD ingolfieliidea EKflnciepIgean ancestors PR QTQ-C RAN G ON Y CTI D Exfinci marina ancestors Perthta Ufungonyx g yrwreita CHEIDAE CON DU KN DAE ZOBRACHOINAE UFlOHAUSTOFaiNAE PHQXQCEPH ALOPSJDAE Other exUnci marine amphipods PHOXOCEP H ALOlDE A~PLAT W SC HI NOF1 □ AE UROTHQINAE’ PHO moc eph ali d ae PrtscSltini 7 HVPERIOPSIDAE \ VALETTlfOAE -"V • LYSIAMA&SOIOEA PONTOPOR EIO I D E A — ^ PONTOPGREIIDAE B ATH YPQREJ I D A E HAUSTORltDAE LJRlSTinAE MESQGAM1UARIDAE TYPH LOG AM MARI DAE MACROHECTOPIDAE CYPNOCARIDA, GAMMAROIDEA Mcrutopus GAMMABGPOREIIDAE V GAMMARIDAE AN l$OG AMM ARJ D AE EUSIROIDEA NIPM ARGIDAE PM REATOGAMMARI DAE MELPWDIPPQIDEA EUSlfltDAE OEDLCEHOTOIDEA BOGIDIELLOIDEA HADZIOIDEA PAR ALEPTAMPH DPI D AE Ex oetfc erodes PONTOG EM El I □ A E PARACALUOPKDAE COR OPH 10 IDE A CAPRELLIDEA TALfTROIDEA LEUCOTWOIDEA EXDEDlCEnQTlDAE BitiftyQ 0 r# 3 tpLi$ Kieloedlcefos ATEJDAE OEOfCEftCOlDAE Ncto&f ceres Rami. DEXAMINOIDEA \ \ SYNOPODEA • • HYPER IIDE A p Afl D ALISCO ID E A S\ AMPEUSCOIOEA L1UE&ORGI0IDEA • STF GOCEPH ALO IDE A CALLYNOPHORE: • Strongly developed, or present in most members * Moderately developed , or present frequently * Weakly developed, or in few members FIG. II. PHYLOGENETIC RELATIONSHIPS WITHIN THE AMPHIPODA ACCORDING TO ANTENNAL DISTRIBUTION AND SEXUAL OCCURRENCE OF CALCEOLI, AMPWPAQpIC A VOL L NO. 3 OCTOBER 15, 1994 94 These di&iri butions suggest that cal cool i developed ini- tially fin males only) as a device presumably lor delecting species-specific swimming vibrations of females at mating time. Tbecalccoli became have become secondarily adapted, and more complex structurally {in free-swimming raptors), for detection of escape vibrations of frce swimmmg prey, and Phis developed in females Etnd immature^, as we f[ as in reproductive males. As mating styles changed from pelagic lo benthic and/or hypogean. lode to Italic, marine to fresb- waicr, involving pre -ample xus (see below J. the primary role ofcalceoli correspondingly diminished or disappeared. The of reduction and dtsappearanceol ealceoli from male anten- nae was apparently first from antenna \ , and then antenna 2: in the latter, the sequence was first from the peduncles and finally from the tlagelltmi. However, as noted above, ealc- eoli persist (or become secondarily developed) in both male* and females of someepigeanle.g., tn some Anisogaimnandae and Gammaridac ) and/or cave pool amphipods Ce.g. in Crangtmyx packardi and Stemophysitix caiceala Of Crangonyetoidea; Sensonctor valemiensis (Melphidip- poidea'/X and some large paraleplamphopid etisiroideans of Mew Zealand) (Bousfield, 1980) where life styles presum- ably remain free -swimming and raptorial, Gnathoti Structure and Ph vie tie Significance Of all morphological characters of amphipod crusta- ceans, Ihc gnatbofxxJs (peraeopods l & 2 of formal malao □stracan terminology) have previously been considered one of the most significant and fundamental indicators of high level phylelic relationships;, ai least within the Suborders Gammaridea r.S tabbing. 1 90b; Barnard & Karaman. 199 ] ); andCaprellidea (Laubitz, 1993: Takeuehi, 1993). Initially, and based on early taxonomic studies on intertidal groups of “good old Gammaridae^ of northwestern Europe (I H, Stock concept), the sexually dimorphic, powerfully subchclata form of the gnaihopods* utilized in sexual preeopolatory carrying behaviour in the male, was considered by many workers as l he basic or ancestral amphipod reproductive form (e g , Barnard. 1 969a). More recently, however, ex- tensive comparative morphological studies have been con- ducted on gnathopods and other phyJetically significant characters (e g. Bousfidd. 1979a, 1982a, 1983, 1986), and the scope of their function in reproductive behaviour (e g Borowsky . 19S4 l Conlan, 1991a). These studies have corre- lated gnathopod morphology and sex ual dimorph ism , across a rather broad spectrum of amphipod supcrfamilies, w ith a pre-amplexing and/or mate-guarding form of reproductive behaviour. As summarized partly by Sehram (1986). this form of reproductive behaviour is now considered by most workers as relatively highly evolved and specialized! w ithin the Ampbipoda as a peracaridan group. What then might be the probable ancestral f cmn of the guathopods, and concomitant ancestral reproductive life style within the Amphipttda? Wc might TiTSt look at gnathopod structure in members of various superfamilies that arc classified as primitive on the basis of other pies iom orphic character states (per Bou&fieJd 1979, 1983, etc.). The Ly sianassoidea is one such superfarnily group for which the distal portions of gnalhopods I &. 2 of species representative of (A more primitive component families ( Valeuiidae and Unstidae) are detailed in Fig. 12 . In the very primitive genus Valeniopsis Holmes (see Barnard and Ingram. 1990), thecarpusaudpmpodofboihgnathopodsfin IwiLh sexes) arc subsimilar moderately slender and elongate, each wilh subparalicl anterior (upper) and posterior flower! margins. The propod is weakly hut normally subchelata, the ciacty I short and close J y Fitting the si Ig htl y obi ique palm, in the slightly more specialized genus Hirotidelia . the carpus Of gnathopod 1 is relatively short and shallowly Inhale below, The propod is lightly narrowed distal I y, with an excavate palm. overlapped by the Lip of the dactyl. in gnathopod 2, the propod is relatively short, and the palm slightly oblique forwards (panache la le). In the genus Veniiettu , gnathopod 1 is little different, but in gnathopod 2. the propod hji-s become much shortened* and the palm and dactyl much reduced in size to form a miero-subcbela that is typical of die more advanced families and genera within I ysiaiutssoidea. Within QrchomeneUa (family UrisLidite), in addition to the micra-subchelate form of gnathopod 1 gnathopod l has also become structurally modi Red in having a much shortened carpus, with relatively narrow and deep posterior lobe, and ibe propod has become broadened, and the palm and dactyl enlarged and slightly parse hekte In summary, despite minor modifications within an in- creasingly sophisticated generic series, we may note that the plesiomorphic form of both gnathopods may be de- scribed as non sexually dimorphic and weakly subcbelate, with s lender carpus arid proprid. With i n the Ly.sianassoidea, characterized by gnathopod* of the above type, maiing iakes place freely and rapidly in the water column, and there is no pre-amp Eexus or mate -guarding phase, Giuithoporis within Natuntia. lf we examine a much broader range of superfamiiies in which reproductive or mating style i.s free within the water column, and the taxa are relegated to the subgroup Natan tia. a correspondingly broad range of gnathopod types cun be idoniificd (Figs, l£, 13), Within the primitive fossorial Phoxocephaloidea. gnathopod types range from the basi- cally plesiomorphic toon outlined in the Lysianassokfca (above), to a cusiroidean form with powerfully sub- or para- chelate propod and dactyl, and slender posteriorly lobate carpal wrist. In some specialized lys]anassids(hyperiopsids), eusiroideans t leptainphopids), siegocephaloldeans, pardaliscuideans, 5 yn op lot deans* dexammoideans { tepechinellldx), ampeliseoideans, and some melphidip- poideam (Mdpbidippidac), the carpus and propod (of both gnathopods) may be secondarily abnormally elongated and slender. In others* especially the highly modified and specialized members of the tossoriat, micro-carnivorous family Oedicermidae. the gnathopods are raptorial or fmsorial. but typically unlike in form, and the carpus is often much AMPHIPACIFICA VOL t NO J fX’TOBLtt 15, 1994 95 GNATHOPOD 1 GNATHOPOD £ Vatettiopsis VALETTHDAE Qrchomenelta URISTIDAE Hirondetfa Vent tel fa FIG, 12- FORM OF GJN AT HOPODS 1 & 2 IN LY SI AN ASSOIDE A [after Barnard & Ingram and oilier snortes] AMPHfi 1 ACIF1C. A. VOL I NO 3 OCTOBER 15. 1994 % EUSIROIDEA CALLIOP1IDAE GAMMARACANTHIDAE FALKLANDELLIDAE PONTOPOREIIDAE PONTOPOREIOIDEA HAUSTORHDAE OEDICEROTOIDEA OEDICEROTIOAE EXOEDICEROTI DAE FIG. 13. FORM OF GNATIIOPODS 1 & 2 IN SUPliRFAMlLlES OF AMPHIPQDA NATANTIA (from various sources) AMTHIPACIFICA VOL I NO. 3 OCTOBER IS. 1994 97 shorieued and strongly produced posteriorly. Hk- entire ap- pendage fralctionmg perhaps as a digging tool as well as a raptorial chela. In hyperiids, the gnathopods are usually short and simple, nearly alike in Idnru ami rnav serve mainly as accessory mnuthparts i maxillipeds, as in decapod crusta- ceans). In the examples above, and in nearly all compon- ent family members of those super! amtlies,, the gnathnpods are non- sexually dimorphic. However, exceptions to this general trend within the Naianiiaare noted here. Thus within the vegetal ively fossorial family FoMopnreiidae* although the reproductive style is pelagic and free within the water column, the gnaihopotb are also weakly but distinctly sexually dimorphic (see also Bousfield, 1987k Such a morphological anomaly may be vestigial, and represent a clue tophyietic relationships with other superfamitics such as the Gammaroidea. Thus, in such a scenario, we may presume a non-fossorial and pelagic common ancestor to both groups. However, in order to exploit food resources of physically harsh, lolic, intertidal, estuarine and fresh-water habitats, the ancestor may have become secondarily reproductive ly benthic, and developed weakly sexually dimorphic gnathopods and pie-amplexing mating behaviour. Today, ils dependents that developed even more strongly sexually dimorphic and pre-amplextng gnathopods (i,e. now within the Gammaruidea) are wide- spread and highly successful in those physically rigorous habitats. B y contrail those that became fossorial in bottom sediments li.e. now within the Pontoporeiidaei are today confined to tentie, lacustrine, or stibtidaf habitats within those environments that are still accessible to non- preamplexing reproductive life styles. In another evolution- ary direction within that same superfamily, members of family Haustoriidae are characterized by weakly suhchelatt. non-amplexing gnathopods yet almost certainly mate di- rectly on or within the bottom sediments, not in the water column. A parallel set of life stylus and morphologies mark the Cheidae and most genera of Urolhoidae within the austral fossorial counterpart superfamily Pboxocephaloidea, This phenomefloo of superficial similarity has been demonstrated as an example of convergent Dr homoplusious evolution in otherwise phy led call y very distant groups (see Bousfkld, l9g9) T ratherthan an indicator of dose natural relationships as proposed by Barnard and Drummond 1 19S2) and main- tained by Barnard & Karaman (1991). Weakly sexually dimorphic gnathopods are also typical of some Dcxamifioidea, and most of the Melphidtppoidea (including the foasorial Megaluropidac), On the basis of other character states, and of some earlier Held observations (e.g.ofEnequist, 1950), members of both superfamilies pre- sumably mate freely within the water column. However, many members w ithin the se groups are nestlers, commensals, or otherwise in the process of penetrating shallow -water, especially of anchialine brackish habitats of tropical and warm -water regions, where a pre-amp lexing reproductive life style is likely advantageous. In a similar scenario, In which phyletic reiat kinships are sought, we can reasonably look to a common ancestor for (he Dexanunoidca and for the fossorial Ampelicoidea in which I he gnathopods are non sexually dimorphic. However, the morphological specialize aiions and tube -building capabilities of the fossorral ampedseoideans have resulted in their enormously success- ful diversification arid dominance in marine sedimentary habitals. even becoming major food items for eschrictid baleen whales. Thepfesumedpara-smcestralDexaminoidea. however, are common in gondwanian regions (e.g. Austral- ian coastal waters) but arc now relatively rare and virtually relict in shelf habitats of the northern hemisphere [ Bousfleld & Kendall, 1994) Within the Melphidtppoidca, sexual dimorphism of the gnathopods is weak!/ lomodteratcly strongly expressed, but is distinctly present in all members. It is also characterized (til the male) by a consistent similarity in appearance of gnathopods 1 & 2, although Ihese differ markedly (between themselves) in size and Form (Fig.lt> bottom ). The.se gnathqwd characteristics arc found else where widely w ithin the Hadrioidea (especially in the Melitidae) that are now much more widespread in tropical and temperate coastal marine and brackish habitals, In combination with other character states (e g, of the antennae, uropods, and idson, etc.), these gnaihopod similarities may be extended, perhaps less strongly, lo !he Fftre^logammaridae of brackish and fresh waters of New Zealand le.g. in Sous field and Ruftb, unpublished), possibly even to Ihe hypogcan brackish- and fresh- water BogididJoidca. an d even to N oicn be yom ’ s ( 1 9£fi) remarkable, hypogcan (but calc CO l ate) Sensohd&ir, In this yei n , we are left with t he e xci ting possibili ty , req u iring m udi further investigation however, that present members of tive marine and semi-relict supcrfamily Melphidippoidea are dose to a postulated common ancestor to all of the above taxonomic groups (see phylogenetic tree, p, 1 26). Finally, we may note within the group of supertainUies of Nathntia, sexus 1 dimorphism weakly expressed in gnathopods of certain austral freshwater members within certai o Ires h water members of superfamil y Eu siroidca . but more Strongly expressed within fresh and brackish water members of Exoedkerolidae and Paracalliopiidae (see alao BousFidd. 1983). The freshwater cusiroklciin species of I'aikttirtdetl-a Schellenberg, 193 1 , and PraefaikkuufeUti Stock & PiatvneL. 1991 (as in counterpart AZAC specie of Fcna- leptamphopus) are characterized by a dominant gnathopod I that is weakly sexually dimorphic, and may have a pre- amplexing function. However, peracopod 3 of Falkland? Hu is also strongly sexually dimorphic, being carpochelate in the male (as in so me species of Fammetita (Crangonyctaklca) and i n many aq uatic asellid isopods). This latter appendage may function in pre-amplexus, as it does in the Lsopods, but pertinent behavioural studies have rot yet been made on these remote and presumably relict freshwater amphiped groups. In the antipodean oedieerotid families (above), the gnalhopds are typically strongly sexually dimorphic, with gnaihopod 2 doth i mint in males. A pro-ample xing carry itig AMPHTP ACDPIC A VOL I NO 1 OCTOBER 1994 9g STEGOCEPHALOIDEA 5TEGOCE PH ALI DAE HYPERIIDEA HYFEfilfDAE SYNOPIOIDEA SYNOP1IDAE PARDALISCIDAE PARDALISCOIDEA ATYLIDAE DEXAMINOIDEA LEPECHUMELLINAE OEXAMINIDAE AMPELISCOJDEA AMPEUSCIDAE MELPHIDJPPGMDEA MEGALUROPQDAE FIG. 14, FURTHER FORMS OF GNATHOPODS 1 & 2 IN SUPERFAMILIES OF AM PHI POD A NATANT1A (from various sources) AMTHIPACITICA VOL I NO 3 OCTOBER ] 5, 1994 99 of females by males is typical (Chapman & Lewis. 1976; personal observation l Members of these l wo fossoriai amphipod families am almost entirely micrtidals c.siu aiine and fresh-water in their ecological affinities, Their form of gnathopod morphology, and pre-amplexing benthic repro- ductive behaviour is typical of the Reptantia, Within a superfamiiy of Nalanlia, these characteristics have virtually certainly been independently derived and arc homoplasious with the condition in their gammaroidean -taxonomic and ecological counterparts of the northern hemisphere. We may conclude therefore that amphi pod superfam dies herewith grouped within the category Natan tia arc typified by pelagic reproductive (mating) behaviour, and by non- sexually dimorphic gnathopods that are primitively weakly subehc late and sub srimi Ear in form A fe w subgroups wa thi n certain natant superfamilies evince a more replant form of reproductive behaviour and gnat ho pod morphology, These exceptional instances can he explained, ai least tentatively, on the basis of (1 ) 3 secondary use of sedimentary benthic substrata as a ^fluid" mating medium wherein sexually dimorphic gnathopod* and pre-amplexing mating behaviour may not be required (e.g. in Haustoriidae; Cheidae. Urohaustortidac};C2) an independent or convergent evolu- tion within geographically isolated sub-taxa that have been exposed id similar, mainly ecological, evolutionary stresses le g. southern families of Oedtc&rotoidea); (3) a morph- ology vestige of presumed ancestr al types whose evolution- ary "ihrusT devolved mainly into other super-family groups that are, today, essentially Teptant" in reproductive life style (e.g. in Pontoporeiidae); or (4) a probable extant precursor of more successfu l {biogcographicaUyandecptogicaJly more widespread and diverse) descended modem taxonomic groups (t,g. in Dtxaminoidea^ Mdphidippotdea). Gnathopod structure and function in "Rcptantia" The types of grtathopods representative of component superfamiiies of the reprod actively benthic and/or pre- amplexing category Replan tia are illustrated in Figs. 1 5. 16. 3 7. & l£. Within Reptanlia, gnathopod morphology is basically different, and the range of motpbotypes is con ski' etably greater, than that already demonstrated in the Natan tia (above). Thus, in most superf amities of Reptantia the gnathopods are characteristically sexually dimorphic and strongly subebftlalc or chcliform. especially in males. However, many exceptions to these overall trends have been noted, and are hopefully plausibly accounted for. in the discourse below In phy led call y more primitive superfam ilies {so deter- mined from previous si u dies and from other character states above) such as the continental freshwater Crangonyctoidea and the holarctie fresh- and brackish - water Ciammaniidca (Fig. 10). the gnathopods are variously (usually markedly) sexually dimorphic, with gnathopod 2 usually “tfotiftinaAt**. In Crangonyctoidea (as in Natantia), the mature male stage (usually calttOlate) is terminal lor suhterminal. fide Con Lank as i n most Natantia . Preeopulaiory carrying of the female by the male is not documented, although ii is suspected to occur in epigean mem hers of the Paramei a i idae (e.g . in Parameliia , and Austragammantil. where males are distinct ly larger Cham females. In the holarvtic family Crangonyetidae. whose members (especially faypogean species) appear to be mainly raptors, gnathopods of both mates and females are often quite large and powerful, However, males are typically much (he smaller of the two sexes and presumably physically incapable of carrying females. In true am plexus, the male first gnaihopo&s are used to grasp the female laterally by the coxa! plates, and Ihc second jirtathopods remain free, pre- sumable to fend off other males (personal observation; Cortlan com mimic,). In most Gammuroidea, however, males are typically larger and more powerful than females, and pre-eopuiatory carrying i s the re product! ve norm In Cam ily Gamm&ri dae, the first gnaihopods typically have a very oblique palmar margin, enabling the pair to he employed in a "fore-and-aft" seizing of the firsl and fifth peraeonai (body) plates of the female. The larger second gnathopod are employed in agonistic behav iour io other males (and occasionally in earn ivory of newly moulted female of their own and other species!) (Borawsky. 1984; Costello, 1993, this sympo- si um ) . Wi ih i n fam L 1y Ani^ammaridae , (he pal m of gna [ho- ped I is vertical* studded with “‘peg-spines^ and presumably better suited to lateral grasping of the anterior margin of coxal plate 4 than peraeonai plates (Bousfield, 19£6 T pers, observation (in Eogammant.x). Within iheTaiitroidealFig. 16 ) pre-amplexus is typical of the intertidal and brackish- water family Hyalidae, the inter tidal fossorial Dogielinotidac, the coastal marine and fresh- water Hyatellkiae, and the more primitive members of the supraiidal family Talitddae. The gnathopodsare strongly sexually dimorphic, and in the usually larger male, gnathopod 2 is especially powerfully subehelate. probably for use in agonistic display, and in fending off other males. In carrying activity within most Hyalidae, Hyalellidae., and Dtigiel- inoddae, gnaihopod I is modified lo grasp the margin of a special pre-copulatory notch in the antero-ventral margin of peraeon 2 of the receptive female (see Borowsky, 19&4: Bousficld 1936, 1993). However, in the most terrestrial landhoppcr groups (Boris field 1984, 1988), in the most specialised aquatic inquiiinous families (e.g. Eophliamidae ), and in [he kelp- borers (Najnidae), the gnathopod* are weakly (or not) sexually dimorphic, and pre-amplexus is lacking, apparently lost seem daily. Within the Hadrioklea (especially family MchtidaeKFig. 17), gnathopods are typically strongly sexually dimorphic, and pre- ample xing reproductive style prominent in all bui the most hvpngeau subgroups. Using gnathopod I, the pro pod and dactyl of which may be specially modi Red LO clasp ihe female by an antero-ventral process of coxa 6 (in Abludometfia and rdativcs-XBorowsky. 1934* Bousrreld, pm. observation). The much larger male gnathopod 2 is held freely, and functions in agonistic behaviour toward other males, In the tropical and warm-temperate marine AMFHTPAOFICA VOL- I NO. ? OCTOBER 15, IW CRANGONYCTOIDEA AUSTROGAMM ARI DAE Austrogammsrus sp. CRANGONYCTIDAE GAMMA Rl DAE GAMMAROIDEA MESOGAMMARIDAE Mesogammarus americanus Cratigonyjt. sp. Uroctena sp. FIG. 15. CN ATHOPODS 1 & 2 IN PRIMITIVE SUPERFAMILIES OF AMPHIPODA REPTANTI A [after Bousfield (1958; 1979) and other sources! AMP HIP AC [Fir A VOL. I NO 3 OCTOBER 15, 1994 10 1 A MPH J PO D A RE PT ANTI A I A 1 1 males except w here indicated ] i from van o us sources ) AMPH1P ACIFIC A VOL I NO. 1 OCTOBER 15. 1994 }Q2 genus Ditlkhieibf cither (he right or left gnaihopod 2 of the male is enormously developed (Fig. 17). The dactyl is greatly enlarged, and its lip fits into a socket in the anlero vemrally produced palmar angle of the propod. Us overall form is grossly similar lo I he morph ology of the gnalhopod of the decapod “snapping shrimp" (genus Aiphat-uJt'}} sug- gesting that il functions in percussive sound production, either to attract receptive females or to warn away other males, However, in most of the hypogean hadzkndeans (e.g. weckeLds. metaniphargids. uietacrangoDyctid!;. etc.), whether the gnathopocls are strongly or weakly raptorial sexual dimorphism is weak or lacking ( Stock, 1985; Holsinger, 1992b). Within the mhc-building Corophiotdea occurs perhaps the greatest range of gnathopod sexual dimorphism of any replant amphipod supertainily (Fig, IB J. In the male. the gnacbopodsare typically strongly suhdielate orcarpocbelate, bu t very urd ike in form and size . The second gnai hopods are usually very much the larger, more complex, and dominant, except in die Aoridae and Cheluridac where gnachopod I is the larger. Since eorophkiidcans are sequestered in open- ended tubes of their own construction they have become, effectively, seini- sessile, and stray liule from a fixed loca- tion, Such a life style may have resulted in secondary loss of prccopulatory '■carrying*' of the female. Instead, the male "guards" the female in her tube and employs the enlarged gnat hoped 2 mainly in agnostic behaviour towards compet- ing males who might approach hi* reproductive terriiory rBomwsky, 19H4, Con Ian. 1988, 199 la). However, pro- am plexus is retained in the free-dinging family PodoceridaC and in the presumed desccndenl Caprdlidea, 1 including Cyamidac) in nearly all species of which the gnaihopods are var lously strongly sexually dimorphic (see Laubitz, 1970, 1979. 1993); TakeuChi, 1993). Across the replant elassifiealory board, however, some important exceptions to this general picture should be noted. Within the relatively plesiom orphic reptani , super-family Liljeborgioidea (Tig 17, topi, sexual dimorphism of the gnathopods is most strongly pronounced in the free-living families Liljeborgiidae, Sehidac. and the sponge-dwelling Colomastigidae. but L$ weak or virtually non-existent within the hypogean Salentinellidae and Paraerangonyeiidae. Waihiti other hypogean super family groups, especially those believed to be micro -predators (e.g. Bogidielloidca. Ingolfiellidea). the gnaihopods are powerfully subchelate ni carpochelaie and raptorial, but appear weakly (or not) sexual dimorphic. Finally, within the diverse and possibly potyphyletic assemblage of families currently assigned to the exclusively marine superfamily Leurothpidea, a corre- spondingly immense diversity of gnathupod types may he seen. Giuiihopodx I & 2 axe often much enlarged and of unusual or bizarre form, and often very different from each other in form and size. Taxa within families Leucothoidae, Amphilocbidae, and Pleustidae. etc., whose vegetative life styles ate commensal, inquilmoux* or parasitic, exhibit viriu- ally no sexual dimorphism of the gnathopods. However, in free - living groups such as the Slcnotboidae and some qf the Pleustidae, especially those of intertidal and brackish habi- tats (e.g. "Pvrcipt nixies'" den), the gnaihopods are variously (often strongly) sexually dimorphic. In summary, within component superfluities of Rep lamia, we may conclude that sexual dimorphism of the gnat hopods and benthic pre-amplexing reproductive styles are dominant and characteristic of member groups thai are vegeiaii vely free-living and epigean to physically rigorous habitats such as coastal shallows, estuaries, and fresh- wa- ters. Conversely, in members that haw become (presum- ably secondarily) symbiotic-ally associated with other ani- mals or plants of marine environments, or penetrated into hvpog i-an brackish- and fresh-water, or fully terrestrial habb tats, sexual dimorphism uf the gnathopods is expressed weak I y or not at all Asa group, the replant s include the most derived ainphipod morphotypes, that exploit unusual or restricted food resources unde rph y si call y rigorous or unusu- ally specialized environmental conditions. In the corre- sponding reproductive evolutionary sequence, a pre* ample* ing reproductive (mating) style is presumed to he an elTe ctlve means of ensuring species continuity. Thus.atthe precise time of ovulation during the female monk cycle, the newly taid eggs [ with in the female brood pouch) must be fertilized by the male. Without the ensured presence of the male at that time the species could not remain in place within the specialized h abi Lai nor remain viable as a species. How* ever, where such a mechanism is no longer needed lo ensure such close contact (as in lenftc hypogean habitats, or under confined symbiotic ctmdi lions), or the carrying mechanism become physically impossible io maintain (as in terrestrial habitats k the gnathopods lose (presumably secondarily) the .sexually dimorphic form, and neolenieally revert to a mor- phology suited to the vegetative life style of both sexually mature adults and immature stages. AMPHIFACIF1CA VOL. I NO. 3 OCTOBER 15, 1994 | Q3 Ntffltla nttids B0GID1ELL0IDEA BOGIDIELLIDA E LILJEBORGIOIDEA LILJEBORGIIDAE COLOMASTIGIDAE PARACRANGONYCTIDAE INGOLFIELL IDEA INGOLFIELLIDAE Ingoifietta sp. SALENTINELLIDAE Di/f+um ftp. MELTTIDAE Duitchieiia sp Bogldlefia btvtitnl Psmidingolftefla sp. tdumli a sp. Q&ioma^tix sp. HADZIOIDEA Satentineifa ep. HADZIIDAE FIG* 17. GNATHOPODS 1 & 2 IN ADVANCED SUPERFAMIUES OF AMP HI POD RE PT ANT I A [Males unless spec ified J ( from se ve r a J sou rces) AMPHJPACIHCA VO|,. E NO J OCTOBER 15. 1994 104 COROPHIOIDEA A OR I DAE Aoroides sp. Lamboa sp PODOCERIOAE CAPRELLIDEA FIG, IS. GNATHOPODS 1 & 2 IN COROPHIOIDEAN AND CAPRELLIDAN AMPHIPODA [males unless specified) (from several sources) AMFH1PACIFICA VOL. J NO. 3 OCTOBER 15. 1994 1()5 Muting Behaviour Within the Am phi pod a C onlan ( 1 99 3 ) has summarized recent advances in work on the Significance of precopulatory mating behaviour and sexual dimorphism in phyletic rclai ions hips of amphipod crustaceans. Amphipods employ two basic reproductive strategies to ensure proximity of males and females ai the lime of female ovulating eedysis; ( 1 ) mate-guarding, in which the males are either fa) carriers involving pre*am picking and concomitant modification of male gnathopodsfoi the purpose, crib) attendees, where they remain domiciled with the female and employ tiicgnathopods mainly in agonistic manner to ward off competing males. (2) non -mate-guarding in which the mature male simply seeks out females wherever they may be at the time of ovulation. These males are classified as (a) pelagic search- ers if the female is in the water column, or lb) benthic searchers if the female is on or in the bottom substrata. In either case the gnathopod* are tittle or not sexually dimor- phic, and no pre-am plexus lakes place. Both strategies wk determined hy the period of ovulation of the female, at which lime the male must he present if fertilization of the eggs is to take place. For a short period immediately following mouth mg, the cuticle ol Hie female is sufficiently flexible to allow for release of the eggs into the brood pouch or mnrsupium. Sperm is deposited there by the male during copulation, and fertilization of the eggs can then take place. Conlan f loe, cit .) has concluded that (he searching strategy is a primitive, and mate-guarding an advanced, form of reproductive behaviour in amphipo&s, This conclusion provides the principal basis for present semi-phyleiie classi- fication of amphipod superfamilies fftg. 30, p_l26j. In these mating strategies, she reproductive morphology of !he mature female is seldom si gnilicanlly different from thai of the vegetative or feeding stages, except in some species of M&ttia, some aquatic talitroideansantl a few others (see below). However, the breeding frequency and fecund* ily reflect overall differences in mating strategy. Thus, females of mate guarded tend to be itcroparous, with several broods in a life lime, whereas those of non-male -guarders tend to he semclparous, with only one brood in a lifetime. Examples of amplexus or copula within superfamilies of Amphipoda are illustrated m Fig. 19. Inset figures C and E are representative of superfarm lies of Natanfia; E, D, F, G, an: representative of Lbe Reptaolia, For comparative pur- poses, l he copulatory position of an outgroup mysid pair (j Ifetopodopm onemalisi is included (from Muir, 1939), The ventral "head- lo-ta i I " posi t ion of the male m y si d perm its direct access of the pents papillae to the posterior opening of the ipursupium, and presumably facilitates temporary clasp- ing of the female abdomen by the male peraeopods, The function of the modified and elongated pleopods 4 & 5 has nni been described; their position beneath the anterior end of the female would Suggest a tactile, rather than sperm-transfer role. The mating position in amphipod* contrasts with that in my aids except that, in bxith groups, the process is relatively rapid and takes place usually in darkness, inmost wrpcrfamiJy groups within Naianlia, contact between the mate-seeking male and ihc female lakes place only during actual copula- tion, and i Ls d urai ion i s brief (Con la n, 1 99 1 ) . in superfamil y Eusiroidea, family Pontogeneidae, the smaller male of Pa ramp fra Columbians lies across the thoracic region of the female, grasping her by the peraconal and coxal plates "fore and aft", using both pairs of gnathopods. Within the bent hie and less mobile tnetnbers of the Parampbithoidae, the male O fEpimcria cornigera holds the female crosswise under the specially curved lower margins of hi* coxal plate* 4 & 5 (Moore, 19$ 1 ), the gnathopods apparently playing little part in the action. Within the Reptantia, and lit the primitive superfamily Crangonyctoidea (c.e, SrnurefUa chambertaim). the smaller male grasps the female sidewise by the coxal plates, and inserts the dactyls of gnathopods 1 & 2 between the lower anterior margins of coxae 3 & 4 respectively, The paired antennae arc pressed closely to the body of the female, with the eatceoli nearly everywhere in contact with the female’s body surface, In family An Lsog ammariac (Gammanoidea) the dorsally positioned male grasps the female by the anterior margin of coxal plates 4 & 5, using gnaibopod 1 (Fig, I9D). In the semi- terrestrial Talitridae (Taiitroidca), (he male crouches across the female, lying on her side, and positions her by means of his gnaibopods and the enlarged peduncles Of antenna 2 working in concert (Fig, 19D). Pre-amplexing positions are illustrated in F\g, 20, Preamplexing is rare within (he superramilies of Natantia, and where it does occur, briefly, differs little from ample* us (Fig, 19 A). Within the Reptantia, however, prc-amplexus is nearly ihc rale. In the primitive Gamtnaroklea, males of Antsogammaridae (e.g. Eogammarus oefairi) carry the Smaller female by grasping die ba.se of coxa 4, usually by means of gnathqxxl 1 . In Gmmmts (family Gammaridae), ihc mule carries the female by means of a 'Tone-arid aft” duichingoflhcameriorcdgeofpcraeonplare I and posterior edge of peraeon 5, using gnathopod l, facilitated by its very oblique palms. Within the Hadzioidea, the male of Meiiia nttida grasps the female by the specially modified anterior lobe of her coxa 6, using the smaller gnathopod l for ihc purpose . The much enlarged male gmnhopod 2 may be used in fending off competing males. In many aquatic Tafitroidea, especially in HyakJla and AUorchestes (Hyalellidae) and in tiyale and Paraltottfmm (Hvalidae), the dorsal I y posi- tioned male inserts the dactyl d gnathopod 1 inaprecopulatory notch in the lower anterior margin of peraeon 2 of the smaller female. Again, the much enlarged gnathopod 2 apparently functions agOnislicaUy, In some species of Hyoie* however, gnalhopod 2 may be inserted into the female notch. These reproductive strategies are basically similar at super family level but differ in detail internally. They do demonstrate the widespread phenomenon of convergent evolution of similar mating strategies, with differing tactics and morphologies ai ihc family arid subfamily levels. AMPHIPACIHCA VOL I NO. 3 OCTOBER Its, 199J IQ^ A. MY SJ DA iMwopodop as omn laiti) C. EUSIROI DEA : PONTOG ENEHDAE (Parsmoara Columbian^ 8. CRANGONYCTOf DEA : CRANGONYCTIDAE (Symtrella chambarfami) D. GAMMA RQlOEA ANIS.OGAMMAfilDAE lEogammarus odairi) F. TALITROIDEA: TALITRIDAE {Tafftrus sattaior} E EUSIRO!DEA:PARAMPHlTHOIDAE (Epimetfa cornigeraj G. TALITROIDEA : TALITRIDAE (Orchedia gammaraltua} (author sources} FIG. 19. AMPLEXING POSITIONS IN REPRESENTATIVE SUPERFAMILIES OF AMPHIPOM* ANBMYSIDA A. (after Nair, 1939) F (after Moore, 1961} F (after Williamson, 1951} G. (after WWamsoh , 1951} B, D, C. (authors sources} AMPHlRACIFtCA VOL I NO, 3 OCTOBER 15. 1994 107 A. EUSIROIDEA: PONTOGENEWOAE ( Pammoenr co/uJTPfe/flna } B. TALITROIDEA: HYALIDAE. (Hyatt? scticornis) C. TAUTROIDEA: HYALELLIDAE (HyateHa ezt^co) D. HADZIOIDEA: ME LIT! DAE (Meitta nr tm) E. GAMMAROIDEAl ANISOGAMMARIDAE { Eogammants octafii J 9 F. GAMMAROIDEA: GAMMA RID AE (Gamma ms fascfatiiR ) FIG. 2D. PRECOPULA IN REPRESENTATIVE SUPERFAMILIES OF AM PH I POD A ' r REPTANTlA” (after Borowsky (1984) and authors sources! AMPHIPAC Dr 1C A VOL I NO J OCTOBER L 5. IW |QE Ph yle tic Significant of frropod 3. The significance of uropod ^ tn the general description and classification of amphipod crustaceans has always been primary {Stabbing, 1906; Gurjanova, 1951; Barnard, 1969a; Barnard and Karaman, 1991), Its character states have proven especially valuable in preparing taxonomic keys to regional and world faunas, at generic and family levels (e. g. Staude, 1987; Barnard^ Barnard, 1983 , Jfsroleinphylctic and semi-phyletic classification of umphipods (except in some Corophioidea, and the Caprdlidea where the abdome n js variously reduced and uropod 3 vestigial or lacking) has been considered previously (e.g, Lincoln, 1979; BousfieJd. 1979a, 1982a, 1983; Bowman and Greiner, 1973), However, more detailed study of its form and function in relation to broader aspects of phylctic classification would seem fruit- ful, and therefore fomns a principal pari of this overview study, in the Amphipoda, uropod 3 lorms the terminal set of paired body appendages, It is distinguished from uropods 1 & 2 by its form and function, Uropod 3 is primarily a swimming appendage, whether functioning in propulsion or steering. The rami are typically broadened or flattened, and (he margins lined with long plumose setae that provide a large surface area for effective paddling or steering action. U no pods I & 2 ait used mainly in strengthening the caudal portion of the body to permit jumping or flipping, by rapid ties ion of the urosome {Barnard & Karaman, 1991); they are secondarily modified for dopulatory or tactile function in specialized habitats but are seldom modified for swimming. The most plcsiomorphie and general ised form of uropod 3 is typical of the Natan t La and more primitive Reptantia (Figs .21, 22 upper), The paired rami are large, lanceolate, and typically subequal in length {aequiramous condition), and die inner and outer margins variously lined with plumose setae and/or short spines (Figs. 21A-D), The terminal seg- ment of the outer ramus is present (ptesiom Orphic condition? in the more primitive superfamilies such, as the Pboxo- cephaloidea, Lysiaimsioidea, and P&rdahscoidea (Figs.21 A- D), but trends to loss or fusion with the: proximal segment in advanced cal ly nophora tes ( e ,g .Stcgocephai oidea t Fig, 2IN, O) or in vegeta lively benthic forms such as Poutopcreioidea (Fig, 21 G,H>. In many pelagic groups ( within Eusiroidea. Oedicerotoidea. Synpioidea, Hy peri idea, Melpbidipptidca and pelagic mates of Decaminoidea and Ampehscoideak the terminal segment is totally lacking (Fig, 2iE, F, L, M, p, Q, Tv V, X). In more advanced, especially .abyssal-benthic forms (e.g, Lepcchinellinae). both rami may be reduced in size and Swimming setae lost, or nearly so (Fig, 21 U). Within the Nalantia, especially the Pardaliscoidea, and Hyped idea havi Agape lag ie life sty le, sexua ! dj morphism of uropod 3 i$ generally slight, the rami being scarcely mere ^ongly setose m the male than in the female. However, in vegetative! y bembitand repnoductively pelagic faxa such as Phoxocqpbaioidca and Pontoporeioidm se xtiai di morph ism of uropod 3 is often pronounced. In female and imina tines (he appendage is much smaller, ihe inner ramus is often reduced in size (parviramus condition. Fig, 2 1 B 1 J), and rarnal margins usually lack swimming setae. Exceptionally, in some of the POfltoporeioidea (e.g. Haustoriidae) and many of the Oediccrotoidea (Osdiccrotidae). mature males may be secondarily infaunal and/or mate within the substratum, and show little or no recent ion of the natatorial form of oropod 3 (Fig. 21 K. L), Apomorpbie conditions of unoptxi 3 characterize the more advanced super family groups within the Replantia (Fig, 22, lower, Fig, 23), Only within primitive crang- onycloideans, gamma] oidea ns. hadriodeans and liljeborgi- oideans is the folly biramous and/or marginally setose con- dition encountered (Fig.22A-F; Fig, 22 L), In the hypogean Bogididloideiu the rami remain essentially aequi ramus and not sexually dimorphic, despite overall reduction in size, general lack of marginal setae, and loss of the terminal segment of the outer ramus (Fig, 220, P), This feature suggests close natural relationship of the Bogidiclloidea to Ihe epigean Melphidtppoidea, In more advanced hypogean forms, the terminal segment of the outer ram us may be much enlarged and especially distinctive in males (as in Er topis a Fig. 22H, Ginmiphargus (Williams and Barnard, 1938), in several Species of Affocmttgottyx an d PseudoHlphdrgus and in many Niphargidae (Barnard & Barnard, 1983). In these loans, the primary function may be tactile, as in the elongate antennae and elongate, setose peraeopods. More often, how- ever, one or both rami arc reduced, often markedly so. wilh loud loss of marginal setae, and/or spines, as in infaunal or hypog can crangonycioideans, hadrioideans , gammaroideam and Li Ijeborgio ideate (Fig. 2 2£, G, J, K. M, N), Within Amphipoda Reptantia, sexual dimorphism of uropod 3 is variously expressed, often strongly so, depend- ing to large degree on reproductive life style. In primarily benthic taxa, with pre-amplexing or benthic reproductive style, uropod 3 is moderately sexually dimorphic in freely ambulatory groups, both epigean and hypogean in primitive Crangouyctoidea and Gammaroidea, less so in primitive Hadzioidea and marine Li tjeborgioidea). In groups that have apparently become secondarily aquatic (non terres- trial Talrtroidea), the rami have are very short vestigial or lacking (Fig, 23D, E). Sexual dimorphism of uropods is entirely lost (or nearly so) in tube-building, inqudinou*, commensal, advanced hypogean, and saltatory groups (e.g, most Corophi oidea, Leueothoklea, Liljeborgioidea. and TaJitroidea). Here the appendage is often highly modified or specialized, in both form and function, in both sexes (Fig,23 A, B, C). Within the domicoEous Corophiokiea, uropod 3 is much reduced, with rami typically short and slender, but remains biramoui (even with terminal segment of outer ramus in some primitive Tseaeidae) in all but the most advanced Aoridae and Coraphiidae (Fig T 23H t J), In the Ampithoidaeand I^fcyroceridae, the outer ramus is equipped dis tally with hooks and spines for the purpose of retaining hold of its tube while foraging from ihe entrance or repelling i n vadcr s (Figs . 2 3F, G, J ) . 1 n the ad vanced Podoceridae , uro- pdd 3 is vestigial (Fig. 23L), Within suborder Caprellidea, AMPfflPACIPlCA VOL I NO. 3 OCTOBER J 5, JQ9 FlG.2L FORM OF UROPOD 3 IN SUPER FA Ml LIES OF AMPfflPODA "NATANTIA " A, B - PHOXOCEPH A LOl DE A; C, D - LYSIANASSQ1DEA; E s F ■ EUSIROIDEA; C, H, J - PONTOPOREIOlDEAj K - HA USTORI OIDE A : L. M - OEIXCEROrOIBEA N, 0 - STEG OCEPH A LQI DEA; P - HYPERHDEA; Q - S YNOPIOl DE A ; R, S - PAR DAL! SCO IDEA T, D - DEX A MINOT DEA ; V - AMPELISCOIDEA; W, X * M E L PH i Dl PPOi DEA * [after Barnard , 1969, and other sources] AMPKIPACIFICA VOL. [ NO..? (OCTOBER 15, 1994 \\Q BOGIDIELLOIDEA HADZIOIOEA CRANGONVCTOIDEA GAMMAROIDEA FIG. 22. FORM OF UROPOD 3 IN PRIMITIVE AND INTERMEDIATI AMP HI P OD A " REPTANTI A " (from severs I sources ) \ t B - Auttrogautmnru*, Crangitnx C, D, E- Gammants, Gammnroporeia F, C, H , J r K ■ Hadzjn, Fimmopui. Eriopixa, M elite, Metacnmgonyx L, M n N - Lis trie Ha, Salentinella, PseudineolfieUa O, P- Kersutrlenwia AMPHI PACIFIC A VOI, I NO 3 OCTOBER 1 5 . 1994 1 | J LEUCGTHOIDEA TALITRQIDEA FIG. 23. FORM OF UROPOD 3 IN ADVANCED AMPHIPODA 'REPTANTIA' A ■ PLEUSTIDAE: B - LAFYSTltDAE: C - STENOTHOIDAEr D - HYALtDAE: E - TAUTRIDAE; F - AMPITHOIDAE G * ISCHYROCERIDAE H - CHELURIDAE J - COROPHJIDAE; K - ICILIIDAE; L- PODOCERIDAE; M - CERCOPIDAE (from several sources! I he abdomen is vestigial in all but (he mOSl primitive species, and uropod 3 is entirely lost (Fig. 23M). In summary, we may note that with lew exceptions, in ad amphipod superfamilies in which the reproductive (mat- ing) style is pelagic, uropod 3 (in ihe male) is of Lhe large natatory, usually aequiramous type, even where the vegeta- tive life style is he nib ic and/or i n faunal. This plesiomorph ic form of uropod 3 is diagnostic of the phylctically primitive, gammaridean and hyperiidean superfamily groups, within the Natantia, By con irast „ in all superfamily groups thai have become secondarily benthic or infaunal, and r productively benthic orprc-amplexing, the form of uropod 3 is typically of the non-swimming, tactile form. Here the rami are second- arily, and thus apomarphically, reduced or modified in form and function. Only in vegeta lively benthic or infaunal forms wi thin the Natantia and within free-living, pre-amplexing super! ami lies of Replant sa is uropod 3 found to be strongly sexually dimorphic. Extreme reduction and/or modi fi ca- tion of uropod 3 is associated with domieolous, commensal, fossoriai, hypogeau or nearly sessile aquatic life styles, or with colonization of supraiidai and terrestrial environments, lb these forms, Ihe orignal natatory function of the append- age has been lost and/or modified for secondary functions that have presumably enabled the species to penetrate new new environments, new niches and utilize new food re- sources. Thus, the form of uropod 3 may be utilized as a valuable and useful indicator of phyletic classificatory rela- tionships within die Amphipoda. AMPHJPACIFICA VOL I MO. 3 OCTOBER 15. 1994 } |2 PHYLETIC SIGNIFICANCE OF THE FORM OF THE TEL SON, As analyzed previously {BotisfieJd 1979. 1983, 1986). the deeply bil abate form of the telson is deemed the plesiomorphic condition within amphipodan, peracaritlau, and i ndeed, ail malacoslracan crustaceans, Con verse ly, the entire, pi ate I ike. or "fleshy" form of the tel son is concluded to be the typical apomorphie state. as in Lcticoihoidean and Coropbioidean subfami J ies, a nd represents { typical ly ) a distal fusion of Uie- Iwo primary lobes. A very advanced condition is seen in (he Thaumaidsonidae, and many Hypcriidea, where the plate-like telson is fused with the uroscrme, A less frequent, presumably apom orphic, condition occurs where the lobes become separated I hrou^ hoot their entire length fas i i l most Garnmaro idea and ceriai n H adz i 01 dea ) and attai n s an extreme separation dorsall y on tiresome 3 (aboininal seg- inent 61 in the advanced IbssoriaJ genus Eohausiorius (Ponioporeiokkak A panoramic view of telson types across the spectrum of higher amphipod taxa is provided in Figure 24, The prototype am phi pod is depicted with a bdobale telson, the apes of each lobe having a "notch and spine” configuration. This stale may be derived from a pelagic peracaridan foe primitive malacostraoijii) ancestral outgrtap in which the tips of the telson lobes may actually represent vestiges or primordial caudal fureae, as in the phyletic relict Lophoga&trida and Etiphaiisiacea, Following evolutionary lines oul wards from this base, through each superf amity group, we find that member species and genera having the ereat-est number of plesiomorphie character states (those nearest the base) also tctid to have fully or partially bi-lohate telsons. Conversely, member species and genera with the most apomorphie or derived character states, in balance, u suail y sho w the most strong I y fused or plate - 1 ike form o f the telson. The totally bilub-ate apomorphie form may be noted in advanced members of ihc Gammaroictea and in some members of the Pontoporeioidea f family Haustoriidae), Derivation ofa phyletic ally "critical" sigftificaiicc to the overall form of the telson is not siraighifor-wanl. however, because of the obvious independently hooptasimis develop- ment or the plate- like telson within nearly every superfamily group. Thus, to derive a superfamily group based solely on a plaic-iike telson would embrace members of at least ten different major groups, and be solidly artificial, However, if we look more closely at these evoui ionary trends, we may note that within "naiant” pelagitially mating sup-erf ami lies, e.g. Lysianassoideiu Eusiroidea, Pardaliscoidea, Synopioidea. etc., the dearly dominant (typical) form of the telson t& deeply bilobate. Conversely, within the more advanced ■replant" superf am il ies such as the Le ucothoidea. T id itroi dea, Bogidielloadea. and Corophioidca. the dominant state is dimlly noi died or p! ate- 1 ike. Perhaps in confirm ing these general phyletic trendy we may note that the form of the te Ison i n some of the most advanced super families of Malar da Ic.g. the Stegocephaloidea, Oediceretoidca, and the Hyperiidae) is predominantly tor entirely) plate- like. Hvperiids, how ever, are basically parasitoid, at least for part of their life cycle, and employ salpac medusae, and other pelagic invertebrates host sub- strata ; in this sense they are "re plan C i n life style, [ n the more prim itive families wiiltin selected superfumilks of Rcptantia (e.g. Crangonyetoidea, Gaminaroidea, and LiJjcborgioideal. retention of the deeply or partly bilobate condition is common. Undoubtedly, the function of the telson has an important bearing on both i|s overall and de-Uiited 1 Firm. In pelagic, free -swim in ing groups, the flexible, bilobate telson may function in balance and in aileron -I ike siabililizatioru taking over this function (partly from the antennal square e that is Lacking in amphtpods (see Walling 1983). In "thruster- swimmers” such as the oediccroiidas and hyperiids, the plate- like telson is part of the entire forward -thrusting tail- fan in which the urostmial segments may be fused and strength -ened. Here, the role of the telson may be subordinate lo Chut of the larger and presumably more cflec- live component uropods, the rami of which are effectively bilobaie and flexible. On die other hand* wilhin the ;“rcsptanf\ primarily benthic, infaunal, lube- building, commensal, andVor hypogean amphipod superfamilies, hydrodynamic functions of the telson are presumably gradually lost. Giber functions such as grooming tree Bowman, 197]), tubed wetting (sec Barnaul 1969; Myers ( 1988); Conlait (1990), or saltation las in Taljtnttac), appear better served by a short plate-like form, with various spcciali/cd spinosc marginal and apical modifications. A certain degree of sexual dimorphism is retained in the form of the telson, especially within hypogean groups such as ihc neoniphargid and stygobromid crangonyctoideans. flllocrangonyctids, riphorgoidears,ete. Here, ihc let son of the mature male is Often relatively elongate and more deeply eleft or notched distal! y I ban in the female (vestige of its primordial natatory function?). Unfor- tunately . demited and well-documented infonnauod on the precise rO leaf the lel.snn is lacking for many of the “replant”, as well asmore-difficult-to study "natanF groups. Lu summary, the prescru view [Revolutionary and phyletic trends m the form of the telson contrasts directly with the views of some others, in which the "fleshy” entire telson was considered pie siom orphic, and led io postulating the Corephiidae as a probable ancestral amphipod type (see Bare aid 1969. 1973, Barnard and Koraman, 1980) (Fig. 5) However, the brooder more comprehensive studies on the malacoslracoii telson by Bowman { 1974), Schminkc ( 1977), ami $chram ( 1986), while controversial and conflicting, tend little support to the Bamardian view. At this piti'ii we may safely conclude, from an over- w helirti ng area y of e vide rice, E hat tbe plesiomorph ic or primi - livecondilion of I be amphipod telson is bilobate, and that the apomorphie or advanced condition is typically plate- 1 ike or apical ly entire , Howe ver, the form of the telson is so fmugh i with life- sty le modifications at lower taxonomic levels (hat. per _sc . it may he phyletically significant only at faintly, subfamily, or even generic levels, or not at all. AM PHI PACIFIC A VOL 1 NO 3 OCTOBER 15. 19W 113 AMFHIPACIFfCA VOL ] SO 3 OCTOBER 15, IW4 [ [4 FIG. 24. INVOLUTION OF THIi TELSO.N WITHIN AMPIIIPOD SUPER FA Ml LIES SPECIAL TAXONOMIC AND PHYLETIC PROB LEMS WITHIN THE AMPHIPODA The phyletie classification ol ampbipod tTusUiceans is rendered especially difficult by the widespread occurrence tui' character convergence in unrelated tax a of sirnHbtr life styles, and by the difficulty of selecting suitable outgroup taxa. with or without the use of numerical taxonomic meth- odology. Assuming natural monophyly of the Amphipoda as an ordinal group within the Malaeostraea, an attempt is made here to establish closest pftylciiu relationships of: ( 1 ) Suborder Hy pen idea (2) Suborder Ingolfiellidea (3) Selected hypogean genera of uncertain classification having character states of potential ancestral significance. 0 1 Systematic* and phytogeny of the HyperUdea The Suborder Hyperi idea is divided into two infraorders. 6 superfamilies, and 2 1 families ( Bowman & Grtaner, 1 973). mfraorder Physosomaia is generally regarded to be more primitive (showing more pleskunorphic character states) and is thus closer to the presumed ancestral hypenid than is infiaordeFCephalosoma ta (Bowman & Garner, (973). In many respects some members of the Pbysosomata resemble some nou-calceolate cal I ynophorate members of Gainmaridea -Natanlia, including the broad pcraeonal body region, short- ened head that often telescopes into peraeon l, small peraeopod 7, and usual presence of a mandibular palp , The fused urosome segments 1 , 2 & 3. the fused inner ramus and peduncles of uropods 1-3. the 1 -segmented router ramus of hropod 3, and plate- like telson are advanced character stales that are only occasionally met with (and never totally in combination) in only a few gamin arideans (e.g. cyphocarid lysianassids) that tend to have pelagic and neiitic life styles that rtre similar to the hyperii deans Sveshnikov & Vinogradov (1987) considered the sub- order Hyperiidea toconsist of a heterogenous and apparently polymorphic group of pelagic carnivorous crustaceans. All are hyperi ids are pelagobioms; none are benthic. Member species can he grouped into two life form classes of which about 35% are fra-swiiiming predators, and the other 65% exclusively parasites and commensals of gelatinous animals, The former are all members of the advanced Physoccpbalata whereas the paranoids encompass all of the Physosomala and several groups within the CephaJosomata. Of the former, the primitive sciniform family members are commensals and (Uriel ectoparasites. These animals have a well de veloped plcon and urosome, but the grasping adapta- tions of the appendages arc poorly developed or absent. Since the scinid physosomatids arc among the most primi- tive forms of hvperiids. we might reasonably look for ances- tiaJ types among the gairnnaridean amphipods that are simi larly free-swimming and weakly parasitoid. Table II presents a character-state matrix pertinent to physoHumatid Jiyperiids, and to non-caJceoLa te tally nophorate superfamilies of Gaimnaridea-Natantia, The closest (or least different) match (score of 28740) with the scinid hyperiids is thai of superfamily Stegocephaloidea. Similarities with other gamm andean superfamilies (Lysiamssoidea andPard- aliseoito are smaller, in the 40-50% range. These levels are higher, however, than with advanced members of the benthic Reptaiuia, including the CoropMoidea, considered by some to be directly ancestral to the Hyperiidea (see p. 85). Some similarities with stcgocephaloideans are conspicuous. Synapomoiphics include a telescoped head, an asymmetri- cally notched upper lip, slightly dissimilar but mainly sim ple gnathopods, a weakened or shortened iiUisillipedal palp, shortened peraeopod 7, and nearly plate-like (apieally notched) teison.. However, stegocephaloideans are much less advanced in retaining an accessory flagellum, deep coxal plates, unfilled urosome segment 2 & 3, sometimes 2* segmented outer ramus of uropod 3, and the invariable presence of coxal gill on peraeopod 7. among other ples- iomorphie features. Figure £5 Ls a resulting phenogram of character state similarities between physosomatid hyperi ids and non- ealeeoiate gam mail dean Natanlia. This phenogram, derived through simple cluster analysis, shows an overall average similarity of hyperi ids to callynophorate gammari deans of about 55 % r Character state differences th at contribute to the relatively low morphological similarity include, in the hypemds. lack of antennal ealceoli and accessory flagellum, absence of a max imperial pa)p,and total fusion of urosome segments 2 & 3, and telson lobes. Conclusions. These observations suggest that hyperiids may have evolved from a gammaridean ancestral type that was nearest to the present sicgoeephaloidean body form. Bousfield ( 1 982b) has hypothesized a probable mid-Mesozoic most recent time of origin for callynophoratc gammarijeao groups, a diesis which, if reasonably correct, would suggesi an earlier common ancestry for hyperiid amphipods. The fact that hyperi ids exhibit several major differences from closest gammaridean relatives would also suggest that hyperi ids have diverged from a common ancestor over a con- siderable period of geological time, However, despite the remarkable diversity of form . function, and life style shown by members of the Hyperiidea, their derivation from a common ancestor within the much mom primitive Gam- maridea might justify consideration of their clas&ifieatory status as infraordinal within the Gaotmaridea Natantia, By analogy w ithin the world of vertebrate animals, might the hypenids be to the gammarideans what the birds are to the dinosaurian euryapsid reptiles? By similar analysis, members of suborder Capr el lirtea can be derived from a corophloidean ancestral type (Podoceridae, Laubilz, 1979, 1982) and thus justify reduc- tion of its current subordinal status to inffaordinal level By con trast , howe ver, t he Tngol fie!! idea (see also p. 1 20) possesses unique character stales Hint are more plesiomorphic ihan anything occurring within the Gammaridea ( sens, lari . These include I he short unpigmented eye lobes, elongate peduncular segment 3 of antenna 2. partially divided (in- AMPH1PACIFICA VOL 1 NO $ OCTOBER [ 5 . 1994 ] [5 TABLE L CHARACTERS AND CHARACTER STATES OF SUBORDER HYPE RUDE A. 1. CaJlynophore (A, mate) 2. Calceolii {A, male) 3. Accessory flagellum (male* female) 4. Maxilliped palp 5. Gnathopods I & 2 (sexual dimorphism) 6. Brood pi arc* slender (female) 7. Coxal plates 1-4 large 8. Pereopod 7 > Pereopod 6 9. CoxaJ gill of perenpod 7 10. PLeopods (male) re] to pleopods of female 1 1. Sexual dimorphism in pereopods 12. Sexual dimorphism of plcopod rami 13. Sexual dimorphism of uropods 14. Lower lip, inner lobes 15. Mandible, left lac ima dentition 16. Urosome segments 17. Telson Ig. Upper Lip 19 Maxilla I. inner plate, 20. Uropod 3, outer ramus , segment 4. Usually absent, rarely ^segmented rudiment 5. If present, not significant 6. No, all broad* bowed margins, smooth 7. If larger, usually significant 8. Usually smaller to subeiiual (Mimonectes excepted) 9. Always lacking 10. Always mone powerful 11. Slight if any 1 2. Never 13. Usually slight (strong in Lycae&psis) 14. No, never seen 15. fi- lo 15-den Late 16. Urosomites 2 and 3 fused 17. Entire I S. Notched 19. Usually not present 20. Always I -segmented CHARACTER CHARACTER STATE 1. Present 2. Absent 3. Absent TABLE II. CHARACTER STATE MATRIX I IV PERT IDE A ANDCAMMARIDEAN SIPERFAMILIES MAJOR TAXON CHARACTER NUMBER PJA IND 1 a 3 4 5 e 7 8 9 10 11 12 13 14 IS 16 17 IS 10- 20 A. Hyper- id« 0 2 2 ; 2 0 0 7+ 1 + 0 0 2 0 0 0 f 2 2 2 2 2 21 + B. Stegoeephaloidea 0 2 0 0 0 1 0 t 2 0 1- 1 0 0 0+ 0 0 1 i T 2 12 0. LysianaBBoidea (hyp*riop*idaa) 0 1 0 0+ 0 2 0 1 a 0 2 0 0 0 1 0 0 0 + 0 0 7+ D. Lyeianassoidea [Tr ischi; ostomalidae) 0 0 + 0 C 1 i T- 2 j 0 1 0 0 0 0 1 0 0 2 0 1 10 E. Pardallseoidea 0 2 04 0 0 0 T Of j 0 1- 1 0 1 1+ 1 0 0 2 0 1 fit F. Synopioidea a 2 0 0 0 T T 0 o 0 0 0 0 1 + 0 0 0+ 0 0 1 0 1 G, Dexamlncidea i 2 n 1 1 u Ot 0 1- 0 1 0 T 1 2 0 0 0 0 : 1 u* H. Stenotholdea 2 2 2' 0 2 0 0 r 2 1 2 2 0 2 It 0 2 0 1 2 24 J. Corophioiden 1 £ 2 1 0 2 0 1 0 2 2 2 2 2 2 2 1 2 0+ 0 2 21 LEGEND FOR CHARACTER STATES: 0 - PLE5IOMOHPHIC; 1 - INTERMEDIATE, ?- AFOMORPHIC. AMPHIP ACIFIC A VOI. I NO. 3 OCTOBER 15, 1994 \\fi 14 12 10 NATANTIA A. Hyperiidea B. Stegocophaloidea C. Lysianassoidea (Hyperiopeidae) 0, Lvsmnaeaoidea (Trischizostomatidae) EL Pardaifecoidea F. Synopioidea G. Dexaminoidea REPTANTIA H. Leucothoidea (Stenothoidae) J. Corophiaidea 25 FIG. 25, PHENOGMAM: HYPERIIDEA AND GAM MAR IDE AN CALLYNO- PHORATE AND NON-CALLYNOPHORATE SUPERFAMILIES completely fused) segment f of the maxi Hi peck subsimilar csrpOChelatC gnaihoptxis, and large urupod 2. All of these unique features strongly support continued full siibordmal recognition of the [ngolfiellidea. The distributional -ecological occurrence of th^ Ingalfiellidea, worldwide in marine and hypogean in conti- nental freshwater supports not only their clas&ificatory distinctiveness but their pmbabie great antiquity (late Paleozoic, per Bouslield & Con Earn 1990> Ilistrihutjuriul-Kcnlrtgy of hyperiid amphipuris Both hyperiids and slegoeepbaloi deans are exclusively marine, in fully saline f> 30# «■) waters, well away from the immediate influence of land run-off. Both groups are presen l over the shelf and slope, and in the abyss, or exhibit vertical diurnal migrations from below the euphotic zone. Stegocephaloi deans are mainly epibenthic, hut Paramionia. boeck is mesopelagie (Moo« & Rainbow, 1989), and lives in association with medusae (e.g. Atolte parmX Also stegocephaloideans arc found mainly in cold-water regions, as are the more primitive members of the Hyperiidea, the Physosomata. and some of the more primitive members of the Cephalnsomaia { of family Hypcriidae), As noted above, at some stage in their life history* most hyperiids are parasitotd, usually in relationship with the Coelenterata. Tunicata, and other jelly-like pelagic animals. Stegocephaloid-caris arc symbiotic with sponges, tunicate*, sessile coelenterates, and other cnidarians (Moore and Rain- bow. 1984. 1989). Such associations indicate lengthy evolutionary development, and classifies lory stability, fur- ther underscoring the suitability of stegocephalcidtans as a phylelic outgroup taxon for the Hy peri idea. AMPHIPACIHCA VOL. J NO, 3 OCTOBER 15, 1994 | 1 7 The HjjuMwNiidea Problem The phyletic classification of fn^sonaJ, free-burrowing amphipods having a the so-called rfiaustoriifF facies has tong posed a parboil ary difftcul! problem for sy$iart3tlSl&. The "hausioriicT superfamily concept variously encom- passes families al'HauaioriitS'Yikc animals and pontoporeids of northern coastal wafers, and ureiLhoids, nrabaiisloriids, phono cephalitis. pfooxocepbalopsids, platyischnopids, zobracboULs, cheids, condukiids, plus a Tew other enigmatic genera of mainly austral marine regions. Differing views on the taxonomic boundaries of family and superfamily diag- noses, and on the phyletic importance of certain "Tnssoriaf" character states, have resul led i n two pri n ci pal rceen i phylct ic classifications. In essence . ihe concept of the Hausinrioidea proposed by the tale J. L, Barnard broadly encompasses all of tltosc groups (Barnaul k Drummond, 1982; Barn and & Karama n. 199 1 >. A fa rthcr concept, proposed by one of us, restricts the HauslurLoidea to the norlhern families Haustomdae, Pontoporeiidae. and Bathyporeiidae, and rel- egates the austral families to the superfamily Phoxo- ccphaloidea (Bousficld, 1979a, 1982. 1983 >, Since compo- nent groups encompass most of the littoral marine sand- burn) wing amphrpotls of the world, form an important cle- ment of marine food energy cycles, and are proving lobe use- ful indicators of sedimentary environmental quality, prob- lems concerning their natural classification merit our further systematic attention. An assessment of phyletic relationships of hawslorrid amphipods was undertaken and presented relatively recently by one of us, but the results remain formally unpublished (Bous field, 1989), Characters found to be of important phyletic significance included general body form, size and shape of the rostrum, presence of antennal sensory organ- elles, structure and “dactylaiiotT of thepostcriorperacopcsds and nasiliipetlal palp, form of the pk0puds> type of moulhparts, and differences in character states of the tel son. uropuds, and other appendages. The major difficulty in sorting out the phytogeny of f assorts] animals is the "look- alike" problem, Le. the high incidence of convergent evolu- tion wi thin nearly every character and character state, of all the fami ty groups investigated. Howe vet, close and careful examination of these c huracter states, i n relation shi p to those that tend to be relatively independent of fossorlal life style (eg, significant in reproduction, feeding, and swimming), in combination, provides a more reliable basis for sorting out homopbsious siimlarii ies from true phy let ic si mi larities. Or this methsxiological premise, evidence from the evolution- ary direction, or trending, within pertinent character states suggested a basic phyletic difference between the two major groups. Thus, the northern hanstoriids appealed to be more ctosedly related to gamnwoidean amphipods, and of rela- tively recent origin, perhaps associated with the opening of the Atlantic Ocean during the Mesozoic Era. The sou them group was found phyletically more primitive and isolated from other major taxa, and of greater antiquity, originating probably prior to the Gondwanian continental breakup. In this brief recapitulations of the 1 989 analysis, we here consider in detail one main character state, (he form of the rostrum {Fig. 26). The upper row shows a dorsal outline of the head rostrum and proximal peduncular segments or representative species of fossorial ponlogammarids within (he Gauunaroidea {A), and of a bathyporeiid and two haustoriids within the Pofitoporeiokfca (B, C D). The middle row gives similar views of representative species within urothoidlEi, urohausloriid(F), phoxocephalopsid(G). and zobrachoidtH) family complexes, within urothoid type phoxocephaloidears The bottom row gives similar views of species within subfamilies of Phoxocephaiidae (H, J. K). Chcidae CL), and Plalyisehnopidae (M). Trends and key differences in the form of the nostrum sire pronounced. Thus in the "hooded heads' " f Phoxocephaiidae land "shark-snouts" rcheids and plaiyischnupiods'Kbuiiuni row) ihe rostrum is variously elongate and ex ends much beyond the lateral head lobes. In Lhe urothoid type animals (middle row), the rostrum is. short but extends distinctly beyond the lateral head lobes. In the gamin arids, pontoporeiids, and hauxtoi iidx {top row), however, the rostrum is vestigial or very short, and extends little or not beyond the lateral head lobes. In these latter groups, the substrate- penetrating function performed by the prow-like rostrum of the phoxocephalids and urothoids is apparently performed by the distally narrowing and closely approximated peduncular segments of the first antennal pair. Other major character stales have been correlated with differences in form ol the rostrum {Bousfteld. 1989), Thus, family members of the upper row all possess strongly deflexed urosomes ("bent backs"), weakly dactyl ale (or adactylate) peraeopods and muxilliped palp, variously dissimilar and weakly sexually dimorphic gnatbopods 1 & 2, unreduced tgammaroidean) mouthparb, pleopods reproductive!)' non sexually dimorphic, broad to medium broad brood plates, and advanced, gammaroidean -type antennal calceoJi (when present), among ocher differences. Family members of the middle and lower rows, all possess weakly deflexed urosonies ("straight backs"), strongly daetylaie peraeopods and maxilliped palp, subsimilar and non sexually dimorphic palbopods I & 2, wrongly reduced or modified {carnivo- rous) moulhpart&> high incidences of rcprftducl9ively sexu- ally dimorphic pteopods, linear or sublinear brood plates, and primi tive crangonyclcadean -type caiceoli (when present). In all these instances, these differences arc here consid- ered of major phy le tic significance rather than of convergent similar it y . According l y , members of die family H austoriidae arc included herewith the phy letieaJJy related Pontoporeiidae and Balhyporciidac. within superfamily PonlOpOreiriideh, and allied with superfamily Gammaroidea Of the northern hemisphere {Fig. 30, Table III). Members of the southern fossorlal family groups are here maintained within superfamily Phoxuccphaloidea,that is phyletically isolated from other marine superfamily groups, but exhibits character slates that perhaps indicate dislini relationships to the Crangonycloidea, now restricted to continental fresh waiters of the world. AMPHJPACD-ICA VOL. I NO, 3 OCTOBER 15, m 4 MS FIG* 26. SIGNIFICAN C E OF ROSTRUM IN SUPERFAMILY RELATIONSHIPS TOP ROW MIDDLE ROW BOTTOM ROW A Ponfugumtiuiridae B Ba thy pore! Idae C Haustoriitlae (Protohanstarius) D T la ustmiidae ( H a astorim ) E Urothoinae F L rohausloriinae G Pho\ocepha(ops1nae H Zobrachoinae Phtfvocephalidae J Tipimeglnae K Brolginue L PhoKoceph&Ilnaje M Cheidac H Platyischnopidae ANlPHlP ACIFIC A VOL I NO J OCTOBER 15. 1994 \\q The Classification and Phyletic I Visit ion of the Lngolf- tellidea. A s noted by Sohram ( 1 9S6), ttie classification of the Ingolfidlidea has been the subject of modest controversy. Following discovery of the fust species of Ingoifielia by Hansen (1903), the animals were first classified as a new family within the Gammaridea (c,g, Stubbing, 1906). Fol- lowing Reibisch ( 1927), and discovery of further new fresh- water and. marine species, die group was elevated to separate suhordinal status within the Arnphipoda, a classification now accepted by most workers, However* Bowman and Abele (19S2) and Bowman (pers. comm uni c.. 1992) would relegate the group to family level status within the Gammaridea. Here, we briefly reexamine its major charac- ter states and re-evaluate their significance in phyletic clas- sification (see also pp, 125-26), Suborder Ingolfidfida encompasses a small group of blind, vermiform, hypogean and foxsorialamphipods occur- ring nearly worid-wide in both marine and fresh waicr habits (Bousfidi 1982a; Stock. 1977). They occur over a remark- able range of hypogean and infaunal habitats, and are die only freshwater am phi pods presently known from fresh waters of south-central Africa, north of Zimbabwe. About 4fj species have been described to date. They are classified in several genera and subgenera belonging to two families, the IngolfieHtdae Hansen. 1903 and the McLaingoli'illidae Ruffb, 1969, The Lalier family is tnonotypic and in some features more primitive than members of the speciose family Ingolftellkke. The former is here considered likely lo reveal ancestral character states that might link the suborder with other ampbipod types and with other pern can dan taxa. Some of the principal morphological features of M et a mgo IfleUa mirahifi.'i Ruffe, 6969, arc shown in Fig# 7. Descriptive details can be found in Ruffb's original work Hoc dt.) and in fam ily - level compe ntlia by Bousfi eld ( 1 982a) and others. This large species exhibits the following mor- phological features mostly previously considered to be of major taxonomic and phyletic significance: 1. Antenna shorter than antenna 2, with accessory flagellum 2. Antenna 2, peduncular segment 3 elongate , >1/2 length of segment 4 3. Antenna 2, segment 3 free, not concealed by lateral head margin 4. Unpigmented ocular lobes present, at the lateral anterior head process, 5. Paired maxilllpcds with distall y separated Umfuscd) basal segments 6. Gnatbopods large, dissimilar, raptorial, strongly carpocheliform (carpus witli palm. against which closes the combined propod and dactyl ) : rot sexually dimorphic, 7. Peraeopods 3-7, dactyls very short, 8. Pleopods biramous. rami annulate, pleopod 1 com- plexly sexually dimorphic. 9. Uropod 2 much larger and longer than uropod 1 . almost pleopod-like 10, Telson lobes fused to a narrow plate, with paired dislal penicillate setae Character states 2, 3, 4 , 5, 6, 8, and 9 are all considered plesioniorphic an dfn und nowhere else within the Ampliipodu, let alone in hypogean families and superfamilies within suborder Gamsnaridea, This taxon is therefore morphologi- cally unique w'ithin the Amphipoda. cannot be classified within suborder Gammaridea, as presently conceptualized* and therefore merits full subordinal status of it own, It is difficult to extrapolate character states of a highly modified vermiform ampbipod to a form in which these characters might have existed in the presumed epigean ancestors of the Ingolfidlidea. Horn opinions reduction of locomotory appendages and mouLhparts, and loss of pro* nounced sexual dimorphism, is almost the rule in fully hypo- gean amphipods. As noted in the hypothetical phyletic free of the Arnphipoda (Ftg, 30, p. 126), the ancestral epigean ingolficllid was almost certainly callynophorate, with primi- tively cal cfco Late antenna, much as in modern crangonyct- oidearts, and with a terminal male stage. The eye lobes may have borne pigmented stalked eyes, and peduncular segment 3 of antenna 2 a vestigial fcquame. The gnathopods were almost certainly non sexually dimorphic and non preamplexing. However, as noted previously, character nos. 2. 3.4. 5 and 8 occur, in more conspicuous form, within some extant petal opht halm id My sidacea but, to date, nowhere else within potential ancestral outgroup perdcaridans, As noted above, the Ingolfidlidea occur widely in both fresh and salt water, from the shore line to the abyss, nearly w orld wide On the other hand, both the H yperiidea and the Caprellidea ate strictly marine and of restricted ecology and life style, Ingolfiellidsoverlapdi-StributiiHiaiJy and ecologi- cally with many other hyopogean ampliipod groups, espe- cially with bogidieloideaos and niptiargids but are readily distinguishable. Whereas the IqgOlffeHids possess several symplesiomorphies but no synapomorpfiies vis-a-vis the Gumma ride a, the reverse is true of the Mypeilidea and Caprellidea, We therefore cor elude [hat the case for contin- ued recognition of the Ingoltielliea at subordinal level is strong whereas that for the H yperiidea and Caprellidea merits further consideration. Phyletic Relationships of Large Hypogean Amphipods A,x in the Fosfcorial amphipods, the phyletic placement of hy|w>gean amphipods is subject to problems of convergent evolution because of the specialized but relatively uniform nature of the phreatic environment, However, such prob- lems tend to be evidenced in rather different and mainly nor reproductive! y related aspects of their systematic^, Holsinger (1993) has comprehensively reviewed the distribution of the world fauna of 740 hypogean amphipod species that are di stributed amor g 36 fain ilics and S 2 supe rfamil ies or equ i va- Lcnt groups. Most of these occur in die northern hemisphere, bui diversify irrelatively high among groundwater amphipods AMPH1PACIHCA VOL, T NO 3 OCTOBER 15, 3994 120 FIG. 27 MORPHOLOGICAL FEATURES OF INGOLFIFLLIt) E A {mainly after Ruffe, 1069) AMPHiPACIHCA VOL ! NO. 3 OCTOBER 15. 1994 \2 \ of l he southern conti nen ts. Most species are read i \y assign ■ able to family md superftuniJy categories, but some taxa reanctin enigmatic and difficult of satisfactory ptiyletic place- ment. In 1986, Notenboom described a relatively large, carn- ivorous amphipod species from wells and a cave lake near Valencia in eastern Spain. The animal appears basically gEiammaroidean in general features, but is remarkable in possessing calceoli on both antenna of both sexes. As noted in fig, 1.1> ' this is a strongly plesiDmorptnc feature that is found only amon g the relatively prim i live su pertain i 1 y groups and within very few other epigean gammaroidean subgroups (c.g. Paranu'.wgamitmrus}. The species was fully figured and described by Notenboom and is re figured here for comparison with possible closely related ingroups (Fig Sensonaior vatentien&is appears more gammaroidean than any other superfamily group, especially in character states of the antennae, some moulhpartx (eg- simple lower lip), anterolobate coxae, dorsal abdominal spi nation, urepod 3, telson* and surface ultrasiructure. However, males are smaller than females, (he gnaihopods are non sexually di- morphic, and some mouthpans. especially the mandible, are rather strongly modified for an apparently specialized feed- ing rote. After comparing the species with member of the LiljebOfgudac : , Pardaliscidae, Niphargidae*Crangonycddae, Bogidiell idae, Paeudoniphargus , and other hypogean groups, Noieuboom was unable to place the animal phylelically. However, he refrained for formally proposing a new 1 family or higher level taxon for its reception, and hence has left the matter open for further consideration As seen in our Fig . 28 hie differences between Sensema ror and other major regional groups of hypogean amphipods such as niphargids, lyphiogammarids, and bogidiellids are fairly obvious and need noi be detailed here. However, if general features of ihe species arc compared with regional littoral marine species within the Mclphidippoidea, some strikingly similar character states may be noted- Thus, North Atlantic species of Cheit&cratus and Case® have similarly sharply incised inferior head sinuses, antenna 1 much shorter than 2, anterior coxae diminishing in size posteriorly, gnathopods unequal in size (2 the larger), pcracopods 5-1 long and nearly homopodous, with short dactyls and ten- dency to strong distal relation, strongly aequiramous uropod 3, and tel son short and bilobate, These species also have narrow brood plates and lack a coxal gill on peraeopod 7, We concur with Notenboom’s evolutionary scenario in which a frees-swimming marine ancestor probably invaded macroporous biotopes in the littoral karst. We would suggest that as far back as the Cretaceous ancestral md phiddipoideuns may have been cafceolate and much more numerous than their present relict status might indicate. Such ancestral type* uiay once have occupied littoral biotopes now taken over by the more advanced badzioideans (melitids). In our view* modern melphidippoideans merit further study as an extant relict group (hat may well have sprung from the .same common ancestor as Senwmaittn A somewhat similar problem of phyfetic classification has concerned Phreawgammarus fragilis described by Chilton more than 100 years ago from stream beds in the South Island of New Zealand. He assigned the species lo family Gatiiinarfdae were it remained following its (^description Eind the addition of further species by Hurley (1954), The species is refigured here, for comparison with other regional epigean species and with other hypogean world genera of possible pbyletic relationships (Fig, 29), The animals superficially resemble some g&mmar- oideans of the northern hemisphere, including specie* of TypMogam ma rus (Fig £ ^Characters of stiongesi sim i lari ty are found in (he elongate antennae, with strong accessory flagellum, large, sexually dimorphic gnatbopods (2 the larger), elongate peraeopods with antero-lobate coxae, dorsal I y spinose urosome, and large brood plate*. However, differ- ences may be noted in the moutbparts, peraeopod dactyls, urupods, teLson, and a form of sternal gill is present, all of which precludes direci assignment within any known mod- em group of gammaroi deans. Although Phreatogammarus fragilis in continental in New Zealand, it bears a superficial resemblance to medium and Huge hypogean specie* such as grandis Stock from wells in the Caribbean continental island of Haiti, and to Caniarimetita jansiocki Bousficld from anchialinc cave pools in the volcanic Ha wai- inn Islands (Fig.a$. Although Phreatogamrnarus is readily distinguishable from these two form*, especially in the mewthpart*, coxal gills and uropods, these two forms appear at least remotely phyletically related and meril further inves- tigation in this regard. By fortunate chance, one of us (ELB) was able to collect material of additional epigean estuarine and freshwater spe- cies. here designated tePkreaiogammarus sp, I and Phrecitogatnrrmeus sp, 2 respectively (Fig, 29), A prelim - inary report on this material was presented at the Interna- tional Crustacean Symposium in Sydney, in 1980, but the new laxa have not yci been formally described. These species arc similar to the phreatic species, except for their smaller size, pigmented eye*, and more strongly sexually dimorphic gnathopods. They form a taxonomic and ecologi- cal series, from marine and estuarine, through fluvial cpigean to fluvial hypogean biotope*. We might /^seasonably con- jecture. therefore, that (hi* series reveals a direct pathway of egress by which littoral marine organisms have penetrated hypogean fresh waters in the past, not only in austral regions, but world- wide. Except for the relatively short tel son lobes, the estuarine specie* also demonstrates a remarkable overall similarity to species of Homrilki. a tropical and Indo-Pacific genus within superfamily Mdphidippoidea, The genu* Phreato- gatnmttms may well have shared a common ancestor wilh present-day littoral marine melphidippoidean*. Thus, pend- ing more detailed comparison over a broader spectrum of material, the two groups are placed tentatively on the same major evolutionary' branch of the revised and updated amphipod phyletk tree (Fig. 30). AMPHIP ACIFI C A VOl . t NO. 3 OCTOBER 15, IW 122 A, Typhogammarldae: Typhfogaromanjs sp D. Cheirqcralklae: Chefrocratus s undevatfi calceoll Ci CheJfocratJdae Casco bigelowt B Family incerta Sensofiator s * c,i: vaientiensis Nipfiargidae: Niphargus gp, F Bogidiellidae Bog id fella bredini FLG, 2S CONVERGENT MORPHOLOGIES IN LARGE HYPOCEAN AMPH1PODS | from. Notenboom (19S6) and various sources] AMFHIPAC1FICA VOL. I NO =5 fXTTOBER !5 H *994 J23 FIG. 29 PBREATOGAMMARUS SPP. AND SELECTED M ELPHIDI P POT D F A AND HADZIOIDEA A. Homeltia sp B. Phraatogammarus sp. f C. Phreatogammarus sp. 2 D. Pfrreafo - gammarus fragilis Chilton E r Pintaweckeiia grandis Stock F. Carnari meiita stock i Bous field AMFHIEAQMC A VOL. 1 NO. 3 OCTOBER 15 . 1994 124 Phylogenetic tree* Long before the advent of numerical taxonomic analy- sis- hypothetical phylogenetic relationships between higher categories of clarification of organisms had classically been presented in a branching tree like arrangement, In a mor- phological treatment, ihe plesiomopphtc character states are most strongly evinced in taxa. extam or extinct, that am closest to Ihe trunk and main branches, and the apom orphic or advanced and specialized features are best developed in taxa placed near ihe branching extremities, In effect, the phylogenetic "tree" may be viewed as a form of dadogram in which the character states are ordered and arranged “parsimoniously”, but without numerical basis, Brusca & Wilson (1991) have employed dadistie methodology in phylogentic analysis of the Isopoda, resulting in major ciasrificatory recommendations for the 10 suborders within this very diverse* primarily benthic, and relatively ancient group of peracaricJans, However, the universality applicabil- ity and adequacy of dadistie analyses for this purpose has been questioned by some (e,g. Gos liner ^GtriSeln, 1984). A full dadistic analysis of the Amphipoda is beyond the scope of this paper. Serious problems concerning character state homoplasy, and the status of so-called "Intermediate” taxa have yet to be resolved. However, a phyleiic tree based on “first principles” is bene provided as a useful visual basis for eventual numerical establishment of a true phyletic classifi- cation or the Amphipoda. In this respect, Bous field {1979a) has proposed a tree* tike arrangement for amphipod suborders and super families thai is here refined and updated on die basis of new informa- tion and expanded analysis of major characters and character slates (see Fig. 30). The thickness of the branches was roughly proportional to ihe number of extant species In each subtended major category, hi Lhc early version, the “stem' lay within a boundary or envelope of {hose possessing a pelagic reproductive and terminal mate stage, Envelopes of .selected ptesiomorphic character states such as the pre^ tacc of postero-lobaie coxae of peraeopods 5-7,aiidcalceoJate antennae also encompassed superfamities, closer to the tips of the branches, in which mature nudes were benthic, preamptexirtg, and of indeterminate life stage. Hie present version of the tree (Fig. 30} is essentially similar. During die past 15 years the number of species in each group has increased, variously, by only about 5-10%, few major new taxa have been discovered, and the ordering of character sta te.s has re mained basically unchanged. H o w- ever, the callynophore (Lowry, I9S6), calceoli (Lincoln and Hurley i 1931), brush setae, and other sensory and swimming structures of reproductive males (p, 831 have since been developed as significant indicators of phyletic relationships. Emphasis on such parameters has here altered Ihe position of the main trunk which now centrally subtends supermini lies ol Nalantia leading to the most highly advanced and modi- fied Hyperitdea, These laxa are marked by the pies iomorplte of Table I (p, ) that include, in the mate, a more slender and flexible urosome, powerfully natatory plcopods, and well- developed, usually plumose-setose uropod 3 and tail fan. Calceolalc antennae are restricted to die more primitive members of Natanlia and to more advanced subfamilies that have variously penetrated lode-water environments of coastal estuaries and fresh waters (e,g. somepontogenei ids and callio* piids within Eusinridea; paracaltiopuds and exoedicerotids within Oediccroto ideal The Lysianassoidea is the only group within Natan lia to retain callynophore, calceoli, and brush setae, thus remaining closest to the presumed gammaridean ancestral lype and confirming the classical ancestral position set forth by Sara {1895) and Stebhing (1906), The pelagic mates of nestling and tube-building Ltexami noidea and Ampdiscoidea have virtually lost the callynophore, but retain antennal brush setae, powerful tail fan, and bile bate tel son. In this respect, the Melphidippoidea are similar, but in their development of weakly sexually dimorphic gnathopods, appeal transitional to members of the Reptamia. The monotypic gen tis Sensonaror (of Nolenbooin, 1 906) is here proposed (p, ) as u primitive early offshootlhai still retains antennal calceoli of the presumed marine ances- tral melphidippokiean. Primitive members of the fossorial Pontoporeioidea (excluding. Hausloriidae) and the Pboxocephaloitteu (most) have totally lost the antennal eallynophore but have retained brush setae and calceoli, As noted previously, naiam superfamilies wilh calceoli are primarily cold-temperate and arctic in distribution, those without are primarily tropica) and warm -temperate. The coxal gill of peraeopod 7 is retained widely within the Natanlia* and is ptesiomorphically pleated or dendritic in pelagic males of Lysianassoidea, Eusinyidea, Dexami noidea, and Ampeliscoidca. The superfamilies ofReptamia are placed nearer to the branch tips. Those on the right side of the tree tend to possess more plcsiomorphjr character states such as homo-podous pcraeopotls 5-7, with posterodobitte coxae, and generally lack an eedesial (Imo-facial) spine on uiopod I, Among fresh -water members, the occurrence of various types of sternal gills is widespread (e g. most crsifligdiiyc lids, hyatellin talitroidecms, pontogeneiid eusirids and Falkland?! la, and paraerangonyctid liljeborgijds). Categories of Reptantiaon the l eft side of the tree are advanced in those same character states and, in fresh water members (e.g. of Ganunaroidea, Hadzioidea, Bogidiellnidea), sternal gills are lacking or very- rare. The coxal gill of peraeopod 7 is retained only in the most pn mi Live mem burs of Reptanda (eg. most Gammaroidea andCrangonyctoidca) and that of peraeopod 1 is lost in many corophioideans and till caprellidans, On [he left side of (he tree, the primitive hypogean and fossorial lu^olfiellidea (p. 126) diverged early from the many evolutionary trunk. Its presumed epigean free-living ancestors were almost certainly call ynophora te and caleeolate bui little except some mystd-Jike character states can be deduced from comparative morphology (p, 80) and no trace remains in the very limited amphipod fossil record. The hadzioidean Etndcorophioton supeifamilies underwent pro- gressive reduction of antennal sensory structures, dimimr- AMFHIPACIFICA VtJI. ] NO. 3 OCTOfSKR 15, 1994 [25 AMPHIPACIHCA VOL. I NO. 3 OCTOBER 3 5, 1994 ]26 tioa of plcopods, uropod 3 and tail (au, and fusion of telson lobes, but enormous development of pre-amplexing and mate-guarding gnathopds in the male ip, 105), apparently associated with benthic and domicoloui 3 tie styles, The relatively primitive, mainly fresh water gammaroideans have lost callynophore and brush setae, hut many have retain amennal calceoli. and fairly strong development of uropod 3, tail fan, and trilobate tel son. Gnathppods of the male func- tion in pie-amplems , rather than in agonistic mate -guarding. Presumably related to the natantian melphidippoideans are the allocrangonyctids and pseudoniphargids, on the one hand, and thephreaiogaimnarids and bbgidielloideaJison the other (p, 126). On the right side of the tree, the crangonyetoideans are clearly replant. Having lost the callynophoie and brush setae, Undergone strong reduction of pleopods. uropod 3 , and fusion of telson lobes, and are predominantly tiypogean in life style. The Crangonyctoidea ranks as the most primitive of reptant superf ami lies in which males are typically smaller than females, with non- preamp] ex ipg gnathopods, atul rer* minal in life stage.. Their widespread retention of calceoli. of a very plesiomorphic form, provides a plausdble link with the marine Phoxoceptialoidea. Indeed, Perthia (the most primitive crangonyctoidean) possesses a natatory- uropod 3,. strongly hilobate telson, primitively calceolale antenna 1 (male only), specialized carnivorous mouth parts, squarish coxal plates, sexually non -dimorphic rapiorial gnathopods, and elongate peraeopod (ri features that are reminiscent of many Australian PhoxocephaJidae (see alk> Williams & Barnard, 1988), In association with their freshwater and terrestrial evolutionary thrust, and ability to salts te in air, the talitrotdeans have undergone very marked reduction of the antennae and sensory structures, of pleopcd and uropod 3, and fusion of tel son, and powerful development of agonistic and/or pre-amplcxing gnafhopods, but have otherwise re- mained generalized and primitive in general body form. Miirine members of the Liljeborgioidea (e.g, ofLiljeborgudae, Sebidae, Colomastigidac) are variously specialized for commensal life style, with strongly sexually dimorphic gnathopods. The freshwater members (of Sebidae, Salcniinelfidae, and Paracrangonyclidite) are hypogean and gnathopods may have become secondarily weakly or not sexually dimorphic. Within the marine leucotboideans. however, members, that are morphological Ey modified in commensal association with lunicates. Sponges., and other sessile marine in vertebrates (eg. most Leucothoidae, Vicmuaiidae, some Plcustidae, etc), show little or no sexual dimorphism of the gnathopods, except in the tmcrocamivorous S lertoLhoidae, and (he Anaitiixidae where modification is extreme (Thomas & Barnard, 1983). Within the Pleustidae. the neopieustiosd branch, may ha ve given rise to the Podoceridae (and perhaps the Tciliidae), currently classified within the Carophioidea. These in turn, having strong sexual dimorphism of gnathopods and substrate- clinging life style, have probably given rise directly to the "mainstream” form of Lhe Capre Hide a ( Laubitz, 1 979. 1 993 ). However, the possibility of a polyphyieiic origin of the CaprelJidea remains investigative (e.g. Laubite, 1993; Takeuchi, 1993), as is the origin of the Corepbioidea (as presently defined). The leucothoid-podocerid-capfellid clade lias remained strictly marine, with strong reduction of abdom- inal and locomotory appendages, and a strong tendency to semi-sessile, commensal, or eetoparasitk- life styles. Conlan (1991) has utilized the earlier version of the phyletic tree to illustrate the taxonooitc distribution of male - guarding and non -mate -guarding behaviour in die Am- phipoda. Mate-guarding behaviour had ihen beer formally described within the GtUnmaroidea.TaliteoideaBadzioidea, Coropbioidea, and Captellidca, here categorized within the Rep tan! j a. Non mute-guarding behaviour bad been observed within the Lysianassotdea, Eusiroidea, Pboxouephaloidea, PoncopOtreioidea, and Ampeliseoidea. all categorized here wi thin | heNaia n tia. She also fo und dial species of Crangon vx (Crangon yctoidea) and Hausroriui and Amphiporeia (Ponioporeoidea) did not leave the bottom in mate search* ing, yet also did not mate-carry or mate-guard. Such behav- iour, overlapping beteen Natanlia arid Reptantia. is not unexpected, and may reveal how similar mating strategies evolved eon ver gently in phyletically unrelated groups ex- posed to similar environments and edaphk conditions. Revised StTni-ptaytetk c lass ifj cation of the A m phi pod u Pbyletically oriented classifications of the Amphipoda proposed by Bousfield (1979a, 1 982a, 1983) and embodied in Sehrarn [ 1986) are revised and: updated here (Table HI), A closely similar version was published recently by Bousfield anti Staude ( 1 994), Although fiiesiiNyrdina! and superfamily concepts remain essentially the same, their semi-phyletic arrangement has been altered significantly to conform with the concept of'’Natarttia-Reptaii|ia" relationships developed in previous sections, and graphically presented in Fig. 3D. As we may note below, the families encompassed within several .superfamilies have been expanded or modified in the light of recent discoveries and taxonomic advances. Within the “Natantia" superfamily Lysianassidae is restored to the basic, ancestral position of earlier authors (Sars. 1895; Guijia nova 1951). The list of member families is expanded to include; (I) the Hyperiopsidae and Cyphoearidae, all rnenbersof which are neritic, pelagic, and bathy pelagic, and the primitive Vatettiidue of deep coastal and offshore waters; (2) the fish -parasitic Trischizo- stomatidae; and (3) the benthic commensal, and modified Cpflicostomatidae. All of these possess, variously, in combination, the typical lysianassid character stales of short swollen peduncular segments and strongly callynophoraie flagellum of antenna I ; short rostrum; mouthpam variously minified for carnivory or necrophagy; weakly subchelate, long wristed gnalhopod 2 (often with elongate ischium); pleated or convoluted coxal gills; slender or linear brood plates; and (variously) calceolale antennae. Although the Phoxoecpkaloidea possess more primitive cakeoli, and are strong] y rostrate , t hey are ranked ph y k u ea I ly higher because AMPHIPACIF1CA VOJ,. L no. 3 OCTOBER 3 5, IW4 1 27 TABLE 11L SEMI -PHYLETIC CLASSIFICATION OF THE AMPHIPOD CRUSTACEANS- [Gammaridea and Itgolfiellidea ate SougficLd 1982a* 1983: Wyperitdea after Bowman & Gruner, 1973: CapmEIito after McCain, 1970) L AM PH I POD A «N AT ANTI A” Superfamily LYSIANASS1DAE (Gammaridca) 1. Valettiidae 2. Hyperiopsidae 3. Cypbocaridae 4. Uristidae 5. Lysianassidae 6. Conicosiomatidae 7 r Trischi/jQStomatidae fh lficerta sedis Superfamily PHOXOCEPHALOJDEA L Utotfioidae* 2. Plaiyisehnopidae 3, Cheidae 4. Phoxooephalidae 5, Condufclite Superfamily SYNOPfOIDEA I . Synopiidae 2 r Argissidae Superfamily PA R DALI SCO IDE A 1, StUipedidae (inch Asiyridac) 2, Pardaliscidac 3, Vitja/ianidac Superfamily STEGOCEPH ALO I DE A L Stegocephalidac SUBORDER HYPER TIDE A INFRAORDER PHY SOSOMAT A Superfamily SONOIDEA I. At'chaeoscinidae ,2, MimouecUdac 3. Proscinidae 4,. Soinidae S uperfamily LANCEOLlDEA 1, Microphasmidae 2, Chuacolidae 3, Lanoeolidae INFRAORDER PHYSOCEPHALATA Superfamily VTBILIOIDEA 1, Vibiliidae 2, Cystosomalldac 3, Paraphronimidae of their loss of callynophore, and (heir more highly modified moutbpam (lower lip with inner lobes), and unpleated gills. The Synopioidea, PardaJiscoideaajidStegoceplialoidcafonn a nomcaiceolate core group within Natantia leading to the Superfamily PHRONIMOlDEA L Hyperiidae 2, Dairellidae 3. PhronLmidae 4 r Phrositiidae Superfamily LYCAEOPSOIDEA 1 . Lycaeopsidae Superfamily PLATYSCELOIDEA L Prouoidae; 2, Anftpronoidae 3, Lycaeidae; 4. Oxycephatidae 5. Platyscelidiie; 6 . ParasceEidae Superfamily DEXAM I NOIDEA {Gammarideaj L Atylidae (+ Lepechinellinae) 2 . Dcxamirtidat: (+ Piophliantidae) Superfamily AMPELISCOfDEA !. Ampelisridae Superfamily PONTOPOREIOIDEA 1. Pontoporeiidae find. Barhyponeiidae) 2, Haustonidae Superfamily EUSIROIDEA 1 „ Ponlogeiieiidae 2. Eusiridae 3 r Bateidae 4, Calliopitdae 5, Parale ptamphopi d a e (Inch FalkiandeMidae) G, Gammarellidae 7. Amphithopsidae £. Gammaraeanthidac 9 r Paramphithoidae Superfamily OEDICEROTOTDEA h Paracalliopiidae 2, Exoediceratldae 3. Oedtcerotidae Superfamily ME LPH1DIPPOIDE A 1. Sensomtor group (monotypic) 2 r Cheirocrdtidae f-Hortielitidae} 3, Melphidippidae 4, Megaluropidae 5, Niphargidae? (incert. sed, ) 6 , Phieatogammaridae? (insert . s*d,) advanced, parasitoid Hyperildea. the internal classification of which has been established by Bowman & Grurer ( 1973). The weakly or non -rostrate de xaminids and ampelsscids are yet mane highly advanced in near loss of call ynophore, AMPHtPAClFlCA VOL. 1 NO 3 OCTOBER 15, 1994 128 TABLE 111, fcnnt’ri). 11, AMPH1PODA "REFT ANTI A” SUBORDER INGOLFTELL1DEA 1 . Engoifiellidae 2 Metaingolfiellidae Superfamily CRANGONYCTOlDEA 1, Neoitipliamdae (+ Penhiidae) 2, Paramelitidae 3, Stemophysingidac 4, Eocrangonyctidae 5, Granger yctidae Superl’amiiy LIU E BORG 101DE A 1- Liljeborgndue 2. Paracrangonyctidae 3. Sebidae 4. Coiomastigidae 5. SalcnttadJidae Super family TALITROIDEA L Hyalidae (inch Hyaielbdae?) 2 . Dogiteliootidae 3, Najnidae 4, Ceinidac 5. Eophliantidae 6, Phliamidac 7. Kuriidae E. TitliEridae (4 subgroups) 9. TemnophJianlidae Superfamily LEUCOTHQIDEA 1. ViemuSiidae 2. Pleustidae ( 12 subf) 3. Acantoioto^atnatidat (inch 3 subf ) 4. Lafystiidae 5. Laphystiopsidue 6 . Ochlejiidae 7. Amphilochidae (2 subf„) 8. Stenolhoidae 9. Cressidae 10. Thaumatelsonidae 1 1 . Maxillipiidae 12 . Nihotungidae 13 . Pagetinidae J4. Leucoflioidae find Anamixidae} 3. Mesogammaridac 4. Typhlogatnmaridae 5. Gammaridae 6, Pontogammaridae 7, Acaniiicgammaridae 8, Macrohectopidac 9. CaspicoJidae? 10, Inccrt, sed. Superfamily BOGID1ELLOIDEA, 1, Artesiidae 2, Bogidiellidac 3, Kerguelen itiUdae 11 Superfamily HADZIQIDEA 1. Hadnidae (+ sev. subL) 2, Metaeraogonyctidaje 4, Nuuanidac 5, Melilidae 6, Carangoliopsidae 7, Aetiopedidae (transfer from CoropMidae) 8, Allocrangonyctiidae (= Pseudoniphargidae) Superfamily COROPHIOIDEA L Ampitbt>idae + 2 . Biancolinidae* 3. Aoridae 4. Chduridac 5 - Isaeidae 6 . Isehvroeendai; 7. Neomegamphnpidae 8. Corophiidae 9 r Podoceridae* SUBORDER CAPRELLIDEA INFRAORDER CAPRELLIDA Superfamily PHTTSICOIDEA 1, Phtisieidae 2 „ Dodecadidae Superfamily CAPRELLOJDEA 1 . CaprtJgammaridae 2 . Paracercopidae 3. Caprellidae 4. Aegindlidae Superfamily GAMMA ROIDE A) [NFRAORDER C YAM ID A 1. Anisogammaridae 2. Gammaroporeudac * Possibly convergent within Corophloidea Supcrlamity CYAMOIDEA 1. Cyamidae arid weak devilopmem of sexually dimorphic griathbpods. dimorphic gnat hopc>ds and a rostrate head. Freshwater mem- The Pontoporeioidea have retained elongate calceolatc bm passes sternal gills. Members ofbe true Haustoriidae antennae (male), but lack coxal gill of peraeop^nJ 7, and are exhibit many character states that are homoplasious with allied to the replant Gammaroidea in possessing sexually phosocephaloidcan genera (Bousfield, 1989). Despite the AMPHIPACIFICA VOL I NO. 3 OCTOBER IS. 1994 [29 lack of calc eoli and (with rare exceptions ) loss of gnathopod sexual dimctfpbisiru family Hausioriidac continues here to he classified within the Pontoporcioidca on the basis of head form, mouth pan morphology, EKJaetylateprateCpods* strongly deflexed urosome and dose similarity of itsmosi primitive members to the sympatrie ponloporeiid genus Amphiporeia Within the Eusiroidea may be recognized two main groups: (1) an essentially pelagic, small to medium sized animals that mostly relain brush selae, calceoli, and strongly natatory pleopods and tail fan, and {2) and essentially benthic group of medium to large-sized animals, including the Paramphiihoidac, Aitiptaithopsidae* and Gammaraeambidae Bous field, 1989. have become benthic but lack sexually dimorphic gnathopod* and rein ai n es.se n ti al I y h wine. Within subgroup (l) the calceotale pomogeneilds and calltopiids have apparently give rise to various groups of Paramoera and allied genera, and to the paraleptamphopid and falklandclJid family groups of austral fresh waters, These all tend to have a much reduced u/opod 3 and tail fan. but males (in many genera) have developed a pre-amplexing gnathopod l . Withi n the closely related but distinctive supeifamily Oed iceroloidea, some marine members have retained both caJceoM and callynophore. but within estuarine and fresh water exoedieerotids and paracalliopiids (southern hemisphere), males have become large r than females and have con verged 1 1 y (to gammaroideans of the northern hemisphere) developed strongly preamplcxing gnalhopods and the reproductive "carrying” habit, features convergent with those of gammaroideans of the northern hemisphere, As outlined above J t he superfam ily Meiphidippoidca now encom ■ passes the frec-swunmi ng marine Chelrocratidae, thefossorial Megaluropidae, and the primitive para-ancestral freshwater hypogean genus Sci? sonator. The phyletie status of the fresh- water hypogean family Nipbargidae, endemic to the Europ- ean ■ Mediterranean region, is considered peripherally melphi- dtppoidean, but remains essentially enigmatic. The order of listing of .superfainilies and suborders of Repiamia is essentially that previously arranged in the fam- ily tree (p. 12&). The primitive Ingolfiellidea axe here considered fully subordinally distinct from the Garmnaiidea ( see also p, l2fy\Yithin the Crangonvetoidea, the rationale of flolsingej (1992a) in scp:irating the Stemophysingidae from the Paramelilidae is recognized here, but family Penhiidae, proposed by Williams and Barnard { 1988), is readily encom- passed within family Neomphargjdac.Thc sponge 'dwell ing Colomastigidae is here formally to transferred from the Leucothoidea to the Liljcborgioidea. Family composition within the Talitroidea remains unchanged, although the freshwater Hyalellidae ha ve proven to be closely allied with Aihtrdmtes and other marine genera and may soon be relegated to subfamily staLus within the Hyalidae. The concept of superfamily Leucothoidea has been broadened to encompass the Lafystiidae, Acantho- nctozomalidae, and Qchlesi&ac tali transt erred from Stegrxephaloidea), and the unique pkustiddike Vicmusiidae Just. 1990, recorded from Bass Strait Canyon, Australia. Despiie considerable recent taxonomic work on both superfamilies, the family composition of the Gammamdea and Bogidielloidea remains lillle changed. The taxonomic refinements within the Hadzioidea have resulted it] several new family proposaiserf which AJIocrangonyltdaeHolsingcr, 1989: Nuuanidae McKinney & Barnard, 1977; and Metacrangonyctidae Boutin & Missouli. 1988, are provi- sionally listed here. The family Aetiopedidae Moore and Myers, 1988, based on an enigmatic new form froan the Bass Strait region of Austral ia h was originally placed within the Coroph roidea, but is here transferred to the Hadzioidea. The type species, Aetiopedes gracilis, possesses a number of strongly rnelphidippoideumhadzioideaii and: non -cor- ophioidean character states. These include a short antenna I, posteriorly decreasing size of coxae M, elongate carpus of gnathopod:*, non-glandular bases and unguifatffi dactyls of peraeopods 3 &4, ful ly biramou* andpluniosc-sctoseumpod 3, and linear brood plates. The inouthparts appear hadzioidem and non-corophioidear . especially in the form of maxilla 2 and maxi lliped, and in the notched and slightly asymmetrical form of the upper lip. The concept of iupcrfamily Corophioidea has remained stable following numerous studies by A, A, Myers flw,cjt. ) but. under closer numerical taxonomic scrutiny, the concept may prove to be pol yphy tenc . Th us T ampi thoid-biancolinids may form one group, aorid-chelurid-corophiidx a second, i saeid 'lschyrotx-nds a thinl. and the podoeerids a fourth, ad related more closely to outgroup 1'ansi ties within other superfatn ilies than to ca eh other. Superfamily and fam 1 1 y concepts within the Caprellidtea accepted here are basically those of McCain C 1 970) that also take account the high prob- ability ofpolypbylelic ancestries proposed by Laubitz( 1993) and Takeuchi (1993). In this presentation, we have delved into the pertinency and usefulness of some morphological features for phyletie clawi ti cation of amphipod crustaceans. A more comprehen- sive study might have included the class ifientory signifi- cance of sexual dimorphism of the pleopods. of the form of the rostrum, of segmenlion of peraeoptxfe, and of several other major characters. We look, to eventual establishment of a data base of non-homoplasiou^ diameter states suffi- ciently large to employ eladlstical analytical methododology with confidence. We urge further study on the significance of surface ultras luxe in amphipod pjbylogeny, currently being advanced hy Kevin Hatcrow (Halorow Si Bousfiefe 1987: Halcrow anil Ptiwell ! 992; Halcrow, 1993), The pro- tein electrophoretic approach that is now providing answers to species level relationships (Bulnheim & Scholl, 1981; Stew ait. 1993) might prove applicable at much higher taxo- nomic levels. Finally, the Fundamental work of Sibley and Ahlquist i 1983. gj ^eq i j.i in which DN A-DNA hybridization techniques were utilized in major phyletie reorganization of avian classification, may eventually be adapted to providing genetic data of exceptional value forthe phyletie classifica- tion of amphipod crustaceans. AMPHIPACIFtCA VOL, 1 NO 3 OCTOBER 15, 1994 | 3Q SELECTED REFERENCES BiinianJ, J, L-., 1958, Index to families, genera, and species of the gammaridean Amphipoda (Crustacea), Occa*. Pap. Allan Hancock Foundation. 19: 3-1 45, i 1969. The Families and Genera of Marine Gammaridean Amphipoda, Bull. U.S. Natl, Mus, 271: 534 pp., 173 figs. 1973. Revision of Corophiidae and Related Families (Amphipoda). Smiths. Contr. Zool. 151: I 27, i rig. + 1974, Gammaridean Amphipoda of Austra- lia, Part l. Smiths. Contr, Zool 139: M48, 83 Figs, , & C, M. Barnard, 1983. Freshwater Amp hi- poda of the world. Vols. I & 11, Hayfield Associates. Mt, Vernon, VA, 830 pp., 50 figs. , & M M. Drummond, 1978. Gammaridean Amphipoda of Australia. Pan III. The Phoxoeephal- idae. Smiths. Cantr. Zool. 245:3-551. , 1982. Gammaridean Amphi- poda of Australia. Pari V. Haustonoidea, Smiths, Contr. Zool. 360: M 48, 58 .figs. ? & C, Ingram, 1990. Smiths, Contr. Zool, 499: 1-80, 43 figs. . & G. S, Karainan. 1980. Classification ofgammyrid Amphipoda, Crustaceans, Supp(,6: 5-16 , 1983. Australia as a major evolutionary centre for Amphipoda (Crustacea} Pfoc. Int. Conf. Biol, acid Evol, Crustacea, Austral, Mus., Sydney: 45-61. . , „ 199], The Families and Genera of Marine Gammaridcan Amphipoda (Ex- cept Marine GammaroitKl. Pis, 1 & 2. Rec„ Austral. Mus,, Suppl, 13, Pis. 1 & 2, f 866 pp. Bate, C. S„ 1862. Catalogue of the specimens ofamphipod Crustacea in the collections of the British Museum : 399 pp,. 58 figs. London, Borow&ky, B., 1984. The use of lire males 1 gnathopods during prccopula in some gammaridean Amphipoda. Crustaceans 47(3): 245-250. Bousfiuld, H. L„ 1958, Fresh- water amphipod. crustaceans of glaciated N. America, Can Field Nat. 72(2}:55-l 13. , 1973. Shallow -water gammaridean Am- phipoda of New England, Cornell Univ. Press,, Ithaca N.Y.* 312 pp, 7 1977, A new look at the sv sterna tics of gammuroidean am pfo ipods of the world- Crustaceana, Supplement 4: 282-316* f fig. i 1979. A revised classification and phyto- geny of the amphipod Crustacea. Trans, Roy, Soc, Can. 4(14): 343-390. L980, Studies on the freshwater ampin- pod crustaceans of New Zealand and Tasmania, P roe Int. Conf. BioL and Evol. Crustacea, Austral Mus., Sydney, May, 1980 Oral Pres & Abstract, . 1981. Evolution in North Pacific coastal marine amphipod crustaceans, in: Evolution Today. G. Scudderand J, Reveal teds.). Proc. 2nd Ini. Congr, Syst. & Evol, Biol.: 69-89, 18 figs, * 1982a. Amphipoda. Gam markka. In: S. P. Parker led. I- Synopsis and Classification of Living Organisms, McGrow Hill, New York,, vol 2; 254-285, and 293-294. _+ 1982b- Amphipoda: Palaeohistory. jji McGrow -Hill Yearbook of Science and Technology for 1982 4983: 96-100 ■ 1983, An updated phyletic classification and pulaeuhi story ol the Amphipoda, Crustacean Is- sues. A, A, Balkcma, Rotterdam. 1: 257-278, ■ 1984. Recent Advances in the systemat- ic* and hiogcography of land hoppers (Amphipoda: Taiilridae) olThc Indo-Pacific region, Bernice P. Bis- hop Mus,. Spec-, Fubl, 72: 171-210+ - 1987, Revised morphological relation- ships within the amphipod genera Pontoporem and Gammaracamhus and the “glacial relict" significance of their postglacial distributions. Jour. Fish Aqu. Sci, 46(10): 1714-1725. * 1988, Ordered Character States as a ba- sis forphytetic classification within the .Amphipoda, Abstract, Crustaceana. Suppl, 13: 279-280. j 1989, Convergent morphologies in sand- burrowing members of phyleiicaJly unrelated gammar- i clean superfamilies. Proc, VHth Inter, Collnqu. Amphi- poda- Walpole, Maine, Sept. 14^ 16+ 1989- Oral pres- entation and Abstract, - 1990. A new r genus and species of hadzi- oidean amphipod crustacean from gncbiaUne pools in Hawaii. Beaufortia 4 1 (4): 25-30. - 1992. The phylelic classification of Am- phiptid crustaceans - Problems in resolution. First Euro pean Crustacean Conference, Paris, Aug. 31, 1992, Abstract. , &- K E. Coolan, 1990, Malacostraca, in Green et al: CrifcLaoea, Encyclopaedia Britannica, Macropaedia, 1990: 854-859, . & E, A. Hendry cks> 1994, A Revision of the Family Pleustidae (Crustacea: Amphipoda Leuco- thoidea), Part f, Systematics and biogcograpohy of component subfamilies. Amph ipacifica 1(1): 17-57, ■, & N. E. Jarre it, 1981. Station lists of mar- ine biological expeditions of the National Museum of Natural Sciences in the North American Pacific coastal region, 1966 to I9S0, Syllnge us 34: 1-66. _, & J. Kendall, l994 r The amphipod super family Dexaminoidea on the North American Pacific coast: Families Atylidae and Dexaminidae: systemat- ic* and distributional ecology, .Amph ipacifica 1(3): 3-66.31 figs. . &. G, O. Poinar, Jj. P 1994. A new terrestrial amphipod from tertiary amber deposits of Chiapas pros AMPHIPACJFICA VOL. I NO. 3 OCTOBER 15, 1994 ] 3 | incc. southern Mexico. Hist. Biol. 7: 105-1 14 r &. C. R Stiiude, 1994. The anpaci oi J. L Biimiuxl on th North American Pacific amphipod search, Amphipacifica HI): 3-16. Boufim C„ Sc M. Messouli. 1988. Mttacrangonyx giwtL h. spM d'nne source du haul- Atlas Marocain et 9a famille an an ten - nal receptor of gainmaridean Amphipoda (Custacca). Jour. Nat, Hist 19; 921-927* , Sl D. £. Hurley; 198 1. The ealeeolus* a sensory structure of gam m andean amph ipods. Bull. Brit, Mus, (Nat, Hist.). 40: 1 03- 1 16- Low ryj . K„ 1986- The cal lyuophore, aeuearidan/peraeuridan, a sensory organ pro valent among the Am phipoda (Crust- acea). Zool.Scr. 15(4): 333-349. _ , & R. J. Lincoln. 1984. Suggestion fora proposed phyletic classification based on the molar flak c. callynophore, and caJoeoCL Amphipod Phyletic Classification Workshop, Natl. Mus, Nat, Sci. Ottawa, August* 1984, Absd. & Oral PresentaUoo, _ ,1992 A revision of the genu* IcfmopUS (Crustacea: Amphipoda: Lysianassoidea; Lhisfidae) , Rec. Austral Mus, 44: 185-245. Mwne. P. G. 1981. A functional interpretation of coxal morphology in Epimeria comigero t Crustacea: Ain phi - poda: ParampHthoidae) Jour, Mai. Biol. Assoc, U-K, 61: 749-757. & A. A. Myers, 1988- An enigma from Aus- tralia; a new variation on the corophioid theme. (Crust- acea: Amphipoda t. Jour, Nat. Hist. 22: 1665-1675 1989, Feeding biology of the iiicsispeiagic gammuridean amphipod Pamndania Twdfci (Stebbirg. 1888) (Crustacea: AmpWpoda: Stegoceph- alidae): from the Atlantic Ocean. Ophelia 30: J-19. Myers, A. A,. 1988. A dadistic and biogeographic analysis of the Aorinac, subfamily nov, Crustaceans. Suppl, 13:167-192. Nair, K. B„ 1939. The reproduction, oogenesis, and dcvcl- opmeid of Mtxxf&psispritfttfaUs Tau_ Proc. Ind. Acad. Sci, 12, Sect, B (4); 175-222. Notenhoom J., 1986 Senmnaior valemimds, n. g„ n. sp. (Amphipoda). from different biotopes in southern Val- encia. Bidj. Dierk, 56(1): 60-74, Read, A. T. t and D. D. Williams, ]990, The role of the eakeoli in precopul atory behaviour and mate recogni- tion of psrudofimruieus Bousfield (Crusta- cea, Amphipoda), Jour, Nat- Hist. 24: 351-359. Reibisch.l.. 1927, Amphipoda, ifl.Kuken(hyl, W. Handbueh dcr Zoologie> Bd. 111. 1:767-808, Ridley. 1983. The explanation of organic diversity. Clarendon Press, Oxford. 262 pp Ruffo, S., 1969. Desc rizionic di Metiiingoiffelta mirabitis n. gen. n. sp. (Crustacea Amphipoda, Meiaingolftfillidae lain nova) del 1c aeque sotterance del salento nell Italia meridionals Mem. Mus. Civ. Stor. Natur XVI: 239- 260. , |989, Amphipoda of the Mediterranean. Mem, lust. Oeeanngr, Part 2. Gammaridea; 365-576. Sars. G O . 1895. Amphipoda, An aecoutu of the Crusta- cea of Norway with short descriptions and figures of all the species, Christiana and Copenhagen, vol. 1. Amphi- poda. pp, i-viii, 1-71 L pis. 1240, 8 suppl, pis. Sclidtenberg. A., 1931. Gammaridcn und Caprelliden des Megallangebieis. Sudgcorgiens, und der West-antarktis. Further zoological results of the Swedish Antarctic Expedition 1 90 M TO, 2(6): 1-29, figs I -136, 1 pi. Schminke, H. K., 1977. The ubiquitous telson and the de- ceptive furca, Crustaccana 3(X3): 292-300^ 13 figs. Schram. F, R„ 1984 Relationships within Eumalacostracan Crustacea, Trans. San Diego Soc . NaL. H isE . 20( 16) : 30 1 - 312, T 1986. Crustacea. Oxford Univ, Press,, New York, 606 pp, p 1994. Re view of; "Aniphipod£,anobleobscs- sion: essays in Tneiiu>ry of J, Laurens B arnard (1928- 199 1 ) h L. Watling and P. G. Moore ( eds.). Jour, Crust. Biol. 14(3): 612-613, Shaw, P., 1989. New amph ipods from geothennal vent AMPHIPACIFICA VO[ i NO 3 OCTOBER 15, 1994 133 sites off the west coast of Vancouver Island, British Columbia, wiih a reappraisal of the amphipod family Sebidae. Can, Jour. Zoo!. 67; ISS2-189G. Shoemaker, C R, a 1930. The Amphipoda of the Chetieamp Expedition of 191 7. Contr. Can. Biol. & Fish.n. h, 5(10): 1-14 1, figs. 1-54. » (955. Amphipoda collected ay the Arctic Laboratory. Office of Naval Research, Point Barrow, Alaska, by G, E. MacGinitic. Smithson. Mist, Coif 128(1): 1-78, 20 figs. Sibley, C. G., and J. E. Ahlquisi, 1983. Phytogeny and classification of binds based on the data of DNA-DNA hybridization. ioJCunent Ornithology. T. R. F. Johnston (cd.) Plenum Publ., New Haven, Cb. 9; 245-292, Slewing. R. T 1963, Studies in maJacostraeati morphology: results and problems. Phytogeny and Evolution of Crustacea. Mu*. Comp. Zook Spec Pub!., 1963; 85- 103. Slattery, P, N., &. J. Oliver, 1985. Life histories of infaunal amphipods from subiidal sands of Monterrey Bay, Cali fomia. Jour. Crust. Biol. 5(4); 635-649. Sncath, P T H, A.* and R. R, Sokal, 1973, Numerical Tax- onomy. Freeman and Cou, San Francisco 573 pp. Stapleton, J. L., W. D. Williams, & J. L. Barnard, 1988. The morphology of the caJccolus of an Australian crangonyetoid freshwater ampbipod. CmsJaceana 55: 157-162, 1988. Maude, C. P., 1987. .Amphipoda Gainmaridea, pp. 346-391. to E. N, Knztoff fed.}. Marine Invertebrates or the Pac- ific Northwest. Univ. Wash. Press, Seattle. 511 pp, S rebbrng, T, R. R. s 1888. Repon on the Amphipoda collected by H, MS. Challenger during the years 1873-1876. Challenger Rep. 29: 1-1737, 210 pL * (906. Amphipoda 1; Gammaridea, Das TierrtJCh, 21: 1-806, figs, 1-127. Steele, D. H., 1988. Some evolutionary trends in the Am- phipoda. Proc. VI rh fm_ ColJoqu. Amphipod Crusta- cea. Cmstaeeana, Suppl. 1 3, Studies on Amphipoda ; 107-121. — * 1 988. What is the amphipod life style? Crust- aceana, SuppL 13. Studies on Amphipoda: 134-142. Steele, V, J r , & D, H. Steele, 1993, Pre^ncc of two types of catoeoJi in Gammtirtiius atigulo^us ( Amphipoda: Gammarideal Jour, Crust. Biol* 1 3(3)1 538-543, Sfigs. Stephensen, K, 1923, Crustacea Malacostraea, V. (Am- phipoda I J. Danish Ingolf Expedition. 3(8); 1- UK). Slew-art, B. A., 1 993. The Use of protein electrophoresis for determining species boundaries in atnphipods. Cntsiac- eana 65(2); 255-277. Stock, J. H., 1977, The Zoogeography oflhe Crustacean suborder Ingolflelbdca. Stud. Fauna Curacao LV; 13 L 146. : I9S 3 . The taxonomy and zoogeography of the family Bogid ie I lidae (Crustacea, Amphipody), with em- phasis on the West Indian laxa. Bijdr, Dierk. 5 1 (2): 345-374. .i 1985. Slygobtont amphipod crustaceans of the hadzioid group from Haiti. Bijdr. Dierk, 55(22); 13 1- 426 p, 67 + 8 figs. — . , & D. Ftatvoet. 1991, The freshwater Ain - phipoda of Lhc Falkland Islands. Jour Nat, Hist. 25: 1469-1491. Sveshntkov. V. A., & G. M. Vinogradov, 1987. Life forms of hyperiid amphipods. Dokl. Akad Nauk SSSR. Vol. 293(4): 101 1 1015. Takeuchi, I.* 1 993. Are the CapreJIidea a monophylelic group? Proc. J, L. Barnard Mem, Svmp, 1992. Jour. Nat. Hist, 27: 977. Thomas* J. D., &. J. L. Barnard. 1983, Transformation of the Leucothoidts morph to the Attamixis morph (Ainphi- poda). Jour Crust, Biol. 3(1): 154-157, Vassilenko, S, V„ 1974. Capcctlideaof the seas of the USSR and Etdjacent waters. Opred. Faunc SSSR. Akad. Nauk SSSR 107: 1-288 (in Russian) . Warling, L„ 1981, An alternative phytogeny of pcracarid crustaceans. Jour. Crust Biol, 1:201-210. 3 1983. Feracaridan disunity and its bearing to eumalacostracan phytogeny with a redefinition of cumalacoslracan supenmlers. R. Schram fed.) Crustacean Ph> logcny. Crustacean issues No. 1 (Rot- terdam; A, A. Balkema), pp. 213-228. Williamson. D. L 1951. On the mating and breeding of some scmi-icrrestrial amphipods, Dove Mar. Lab. Rep., Third Ser. No, 12: 49-65, 3 figs. Williams, W. D., & J. L. Barnard, 1988. The taxonomy of crangonyetoid Amphipoda (Crustacea) from Australian fresh waters: foundation studies. Ret. Austral, Mus.. Suppl, 10: I- 1 80, 104 figs. Yamaguchi, T. 199G. The most primitive Balanomorpha. S eal pel tomorphu, & Verrueomorpha, ail from hydro- thermal fields of the western Pacific, tot. Crust. Conf.. Brisbane, July 2, 1990. Oral presentation and Abstract. AMPHIPACIFICA VUL I NO 3 OCTOBER 15, 1994 134 AMPHIPACIFICA INSTRUCTIONS TO AUTHORS Manuscript submission. Manuscripts submitted for pub- Jication should represent original contributions that have not been published elsewhere. Under special circumstances, some reviews, advertisements, and pertinent short articles may be considered for publication. The text should be writ- ten in English or French, with the Abstract preferably pre- pared in the other language. Abstracts should be suitable for separate publication in an abstract journal. Manuscripts must be submitted primarily on 3 1/2 inch high density diskettes, utilizing either IBM- or M AC-compatible computerized pub- lishing systems (e.g. Aldus PageMaker, Quark Express), pref- erably in 2 -column form. Diskettes must be accompanied by one 8.5 X 1 1-inch (22 X 29 cm) hard copy (printed manu- script), with I -in. text margins. Papers must have been previously refereed and text-edited. Authors should submit the name, address, and telephone number of the referee. Acceptance of the pa- per is also contingent on prior payment of printing charges of $ 15. per page (CAN- ADIAN FUNDS). Applications for page subsidies, from authors lacking supporting institutions, will be considered. Diskettes must be submitted in publishable form since virtu- ally no editorial (mechanical) services are performed on them here or by the Printing House, and page proof is not custom- arily sent. Authors each receive 5 free copies of the complete issue but, regretably, no reprints. Text, The publication style of most standard international taxonomic nals is accept- able, with the fol- lowing provisos. Genus and species names m ust be i n I tali cs , and in Boldface italics when utilized as headings. Titles of sect- ions and subsections are to be in Boldface Roman. The text, tables, and figures combined should total not less than 40, but not more than about 1 00, primed pages, but shorter or larger papers will be considered. Prime recommended type font is "Times'’ 10-point, as in this copy, but acceptable non- serif fonts are "Geneva 11 and "Helvetic a". Figures and Tables, All figures and tables must be clearly n umbered an d referred to i n the text. Line i 1 1 ustr ations and or half-tone photographs must meet acceptable standards of clarity and quality; the former should be accompanied by PMT’s of publishable size (max. width of 6.5 inches (16.5 cm), and the latter by glossy prints and/or negatives. Tax- onomic papers are expected to include illustrations. References. Works should be quoted and listed in a font} utilized by standard (refereed) international taxonomic jour- nals. Standard abbreviations for the journals’ names should be used. Distribution. Mailings of this journal reach more than 200 libraries of museums, biological laboratories, and other re- search institutions world -wide, in addition to individual sub- scribers. Please enter my subscription to AMPHIPACIFICA; Name: . . Agency . Address 0 VOLUME I . (4 issues, 1993-94) renewed on an annual basis, at $50. CAN. FUNDS (= $40. U.S. FUNDS) . . 0 Cheque (Money Order) enclosed Date 0 Please invoice me Please address all correspondence to: Dr. E. L. Bousfield, Managing Editor, AMPHIPACIFICA, c/o Royal British Columbia Museum, 675 Belleville St., Victoria, B. C. CANADA. V8V 1X4. Phone: (604) 380-3787. FAX (604) 387-5360.