|ANflPHIPACmCA| * v Journal of Aquatic Systematic Biology^ P Volume 3 Number 1 Contents May 16, 2001 Norma E. Jarrett, 1931-2001: A tribute 1 The amphipod superfamily Leucothoidea on the Pacific coast of North America: Family Amphilochidae: systematics and distri- butional ecology. P. M. Hoover & E. L. Bousfield . . . : 3. The genus Anisogammarus (Gammaroidea: Anisogammaridae) on the Pacific coast of North America. E. L. Bousfield 29. An updated commentary on phyletic classification of the amphi- pod Crustacea and its application to the North American fauna. E. L. Bousfield 49. National Library of Canada ISSN No. 1 1 89-9905 AMPHIPACIFTCA is an international jour- nal of aquatic systematic biology aimed primarily at publication of monographic studies that are overly large (25 - 50+ printed pages) for traditional taxonomic journals. The journal has featured studies on crustaceans of the faunistically rich and palaeobiologically ancient North American Pa- cific coastal marine region. However, the geo- graphic scope extends to other marine regions, and to adjacent fresh waters. Faunistically, it encom- passes other free-living and parasitic invertebrate taxa, and aspects of aquatic vertebrate animals that may involve systematics, ecology and behaviour. AMPHIPACIFICA is scheduled to appear approximately quarterly, with a run of 250-300 copies per issue, each of about 125 pages, and a total volume pagination of approximately 500. The printed page size is 8.5 X 1 1 inches (22 X 27.5 cm). Paper quality accommodates line cuts and half tones at 300-600 d.p.i., and a limited number of colour plates at author cost. Manuscripts are to be submitted in “camera-ready” computerized for- mat (IBM- or MAC-compatible diskettes), and also in hard copy, that have previously been ref- ereed (name to be supplied) and text-edited at the author’s instigation. Suitability of manuscripts, based on content and adherence to submission regulations, 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 (CAN) or $40 (US) per volume, and by page charges to the authors of $25.00 per printed page (including line illustra- tions). For other instructions and a subscription form, please see the inside back cover. Further information may be supplied on request. AMPHIPACIFICA is published by Amphi- pacifica Research Publications under the Ontario Business Program License No. 100013820, Janu- ary, 2000. Editorial Board. Dr. E. L. Bousfleld, Managing Editor, Ottawa, Dr. D. G. Cook, Technical Editor, Greely, ON. Advisory Board. Dr. D. R. Calder, Royal Ontario Museum,Toronto, ON, Canada. Dr. F. R. Cook, North Augusta, ON, Canada. Dr. P. H. LeBlond, Galiano I„ BC, Canada. Dr. Z. Kabata, Nanaimo, BC, Canada. Dr. D. E. McAllister, Perth Mills, ON, Canada. Dr. G. G. E. Scudder, University of British Colum- bia, Vancouver, BC, Canada. Dr. C.-t. Shih, Taiwan Fisheries Research Instiute, Keelung, Taiwan; Canadian Museum of Nature, Ottawa, ON, Canada. Dr. C. P. Staude, Friday Harbor Laboratories, Friday Harbor, WA, USA. Translation Services. M, A. Bousfleld, MSc, Wolfe Island, ON, Canada. Sponsoring Agencies. Canadian Museum of Nature, Ottawa, ON, Canada; Friday Harbor Laboratories, University of Washington, Friday Harbor, WA. Royal British Columbia Museum, Victoria, BC, Royal Ontario Museum, Toronto, ON, Canada. Offices. (1) Subscriptions and Correspondence: Dr. E. L. Bousfleld, Managing Editor, Ottawa, ON, K2B 8E3. (2) Mailing: Canadian Museum of Nature, Research Division, Ottawa, ON, Canada, KIP 6P4. Registration. The journal AMPHIPACIFICA is registered at the National Library of Canada, Legal Deposit Office, 395 Wellington St.. Ottawa, Canada, as ISSN Number 1189-9905. Printing. Elm Printing Incorporated, 280 Albert St., Ottawa, ON, Canada. POSTMASTER: Please send changes of address to: Amphipacifica, att: Dr. E. L. Bousfleld, Managing Editor, 1710-1275 Richmond Rd., Ottawa, ON, Canada K2B 8E3. A MPH I PACIFICA JOURNAL OF AQUATIC SYSTEMATIC BIOLOGY Volume 3, Number 1 May 16, 2001 AMPHIPACIHCA, Journal of Aquatic Systematic Biology (ISSN No. 1189-9905) is published quarterly by Amphipacifica Research Publications, 1710-1275 Richmond Rd.,Ottawa, Ontario., Canada, K2B 8E3. Annual subscription rates are $40. US or $50. Canadian funds. DEDICATION The Joil^kMPffiPACIFICA is - ...-tV* r ied to me promotion of systematic biology and ■****fb the conservation of Earth’s natural resources. This issue is dedicated to the memory of Norma Eleanor Jarrett, 1931 - 2001. Cover design: Adapted from the title page of S. J. Holmes (1904). “Amphipod Crustaceans of the Expedition.” AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 1 Norma Eleanor Jarrett, 1931-2001: A Tribute With the death of Norma Eleanor Jarrett, on April 25, 2001, the scientific community lost a major con- tributor to aquatic systematic research in Canada. Norma began her scientific career relatively late in life. Following her marriage to John W. Jarrett in 1952 in Ottawa, and the raising of three children to adulthood, in 1975 she answered a call for volunteer research assistance at the National Museum of Natural Sciences. She first worked with staff scientist Dr. Rita O’ Clair on collections of marine polychaete worms and, during the following summer, Norma experienced her first marine field work on the Pacific coast of Canada. Following Dr. O'Clair's departure for the marine labo- ratory at Auke Bay, Alaska, Norma began systematic work on selected groups of marine amphipod crusta- ceans of the North American Pacific coast. With museum staff scientist. Dr. E. L. Bousfield, she first published an extensive list of NMNS west coast field stations, followed by a revision of the lysianassid amphipod genus Hippomedon. In the meantime, at nearby Carleton University, she attended day courses in biology essential to a BSc degree, but not com- pleted. Norma continued identification of Pacific amphipod collections, first at the museum's "Beamish building" laboratory on Carling Avenue and later at more commodious research quarters in the Holly Lane building in southeast Ottawa. During the late 1 980’s, the museum's research pub- lication series "Syllogeus" and "Bulletin" were termi- nated. These had offered important outlets for large copiously illustrated papers on the systematic s of ex- tensive Canadian Pacific aquatic inveretbrate faunas that traditional journals found difficult or impossible to accommodate. To fill this hiatus, creation of the privately funded research journal Amphipacifica in the early 1990's enabled Norma to continue meeting the challenge of publishing upon these taxonomically difficult,biologially diverse, but mainly unknown "mid- dlemen" of marine food energy cycles. She publ- ished, with help from line illustrator Susan Laurie- Bourque and in co-authorship with other scientists, 6 major lists and research papers. These treated more than 120 amphipod species, and included one new subfamily, 7 new genera, and 37 new species. Her final research paper, with Dr. Traudl Krapp-Schickel, Bonn, Germany, was published a few months before her death, and an additional three papers await publication. This lovely, warm, and capable person was cher- ished by her husband, her family, and her grandchil- dren, and will be greatly missed by her many friends and museum colleagues. The Editors Research Publications of Norma E. Jarrett 1 . Bousfield, E. L., & N. E. Jarrett 1981 . Station lists of marine biological expeditions of the National Museum of Natural Sciences in the North American Pacific coastal region, 1966-1980. Syllogeus No, 34: 66 pp., 13 figs. 2. Jarrett, N. E., & E. L. Bousfield 1982. Studies on amphipod crustaceans of the northeastern Pacific region. I. 4. Family Lysianassidae, genus Hippomedon. Nat’l. Mus. Nat. Sci., Publ. Biol. Oceanogr. 10 : 103-128, 9 figs. 3. Jarrett, N. E. ,& E.L. Bousfield 1994. The amphipod superfamily Phoxocephaloidea on the Pacific coast of North America. Family Phoxocephalidae. Parti. Metharpiniinae, new subfamily. Amphipacifica 1 (1): 58-140, 31 figs. 4. Jarrett, N.E.,& E. L. Bousfield 1994. The amphipod superfamily Phoxocephaloidea on the Pacific coast of North America. Family Phoxocephalidae. Part II. Subfamilies Pontharpiniinae, Parharpiniinae, Brolginae, Phoxoceph- alinae and Harpiniinae. Systematics and distributional ecology. Amphipacifica 1 (2): 71-150, 36 figs. 5. Jarrett, N. E., & E. L. Bousfield 1996. The amphipod superfamily Hadzioideaon the Pacific coast of North America. Part 1. The Melita group: systematics and distributional ecology. Amphipacifica 2(2): 3-74, 41 figs. 6. Krapp-Schickel, T., & N. E. Jarrett 2000. The amphipod family Melitidae on the Pacific coast of North America. Part II. The Maera-Ceradocus complex. Amphipacifica 2 (4): 23-61, 14 figs. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 2 Journal of the Marine Biological Association of the United Kingdom Published for the Marine Biological Association of the United Kingdom JMBA is an international journal with a worldwide distribution publishing original research on all aspects of marine biology. It includes current international research developments and features much of the pioneering work taking place today on major issues concerning marine organisms and their environment. Subjects covered include: ecological surveys and population studies of oceanic, coastal and shore communities; physiology and experimental biology; taxonomy, morphology and life history of marine animals and plants; and chemical and physical oceanographic work which relates closely to the biological environment. Papers are also published on the rapidly developing techniques employed at sea for the sampling, recording, capture and observation of marine organisms, and chemical analyses of sea water. JMBA is produced from a working marine laboratory by scientists actively involved in research. Recent Articles Chlorophyll-a and primary production during spring in the oceanic region of the Oyashio Water, the north-western Pacific, A, Shiomoto Sagittal otolith size and shape vaiability to identify geographical intraspecific differences in three species of the genus Merlussius, G.J. Torres , A. Lombarte&B. Morales-Nin The size range of suspended particles trapped and ingested by the filter-feeding lancelet Branchiostoma ^ri^{Cephalochordata: Acrania), E. £ Ruppert, T. R Nash &A.J. Smith Subscriptions Volume 81 in 2001: February, April, June, August, October and December institutions print plus electronic: £304/5498 Special arrangements exist for members of the Marine Biological Association of the United Kingdom ISSN 0025-3154 Take a closer look - free Please send me a free sample copy of : Journal of the Marine Biological Association of the United Kingdom Send this coupon to: Customer Services, Journals Division, Cambridge University Press, The Edinburgh Building, Cambridge, CB2 2RU, UK or 40 West 20th Street, New York, NY 10011-4211, USA name address to contact the Customer Services Journals Division in Cambridge: tel +44 (0)1223 326070 fax +44(0)1223 325150 email journals_subscriptions@cambridge.org in New York: tel (914)9379600 fax (914)9374712 email journals^subscriptions@cup.org Cambridge l MYl KSIIY 1’ K 1 SS I lie I'.dinluiriih Building, (..jinbridgt*. ob2 :ri , I K 40 WVm 20th .Street. New York, nv iooii - 4211, USA AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 3 THE AMPHIPOD SUPERFAMILY LEUCOTHOIDEA ON THE PACIFIC COAST OF NORTH AMERICA: FAMILY AMPHILOCHIDAE: SYSTEMATICS AND DISTRIBUTIONAL ECOLOGY. P. M. Hoover 1 , and E. L. Bousfield 2 ABSTRACT Previous definitions of component genera and their probable relationships within the gammaridean amphipod family Amphilochidae were found to be unsatisfactory, necessitating realignment of taxonomic groupings within the northern hemisphere. Apolochus n. g., encompassing the Mediterranean regional type species A. neopolitanus (Della Valle, 1893), and A. litoralis (Stout, 1912), A. barnardi, n. sp., andA. staudei, n. sp. from the North Pacific region, is separated from Amphilochus Bate, 1862, based on the North Atlantic type species A. manudens Bate, 1862. Hourstonius, n. g., based on the North Pacific type species H. vilordes (J. L. Barnard, 1962), is segregated from the Arctic and North Atlantic genus Gitanopsis Sars, 1895, based on G. bispinosa Boeck, 1871. Also redefined is Gitana Boeck, 1871, based on the Arctic type species G. sarsi Boeck, 1871, and including Gitana eltisi, n. sp., from the northeastern Pacific region. Numerical analysis of 20 generic-level characters and character states suggests that Hourstonius and Apolochus are closely related, boreal and warm-temperate, North Pacific and North Atlantic generic com- plexes. By contrast, the primitive Arctic and North Atlantic genera Gitana Boeck, Gitanopsis Sars ( sens , str.l and Amphilochus Bate (sens, 3fL) are closely related and possibly antecedent to the more advanced Mediter- ranean and "Pangean" genera Amphilocho ides Sars, Paramphilochoides Lincoln, and Amphilochella Ledoyer. The diversity of eastern North Pacific species of Amphilochidae is below that of equivalent latitudes of the western North Pacific and eastern North Atlantic regions, but reasons for these differences are speculative. INTRODUCTION Amphilochids are small, colourful, benthic leuco- thoidean amphipods commensal with sea fans, hydr- oids and other sessile marine invertebrates. During the past century and half of faunistic explorations of the Pacific coast of North America, only three species had previously been recorded, none prior to the turn of the century (Stimpson 1857; Stebbing 1906). The first regional species was described from south- ern California by Vimy Stout in 1912. Three additional new species were described from California by J. L. Barnard (1962, 1969b). He also listed species from deeper waters and submarine canyons (1966), and described the ecological occurrence of species in the rocky intertidal of south-central California (1969b; 1975). North Atlantic, Black Sea, and Arctic (Barents Sea) regions, and Shoemaker (1955) recorded Gitanopsis arctica from Pt. Barrow, Alaska. Barnard (1970) described several new taxa from the Hawaian archi- pelago. However, few amphilochid species had been recorded elsewhere in the North Pacific until the exten- sive work on Japanese coastal marine species by Hirayama ( 1 983) . These, and more recent records from Japan, were summarized in phyletic classification by Ishimaru (1994). The present study encompasses the previously un- treated amphilochid fauna of the Canadian Pacific and adjacent coastal marine regions and places it in the context of systematic concepts developed else- where. Among semi -popular accounts, Ricketts & Calvin (1968) included only "Amphilochus neopolitanus" among "common intertidal amphipod species" of the North American Pacific coast. Staude ( 1 987) included four amphilochid species in lists and keys from the northwestern Pacific region, and Austin (1985) listed the same species from this general region, but none actually from the coast of British Columbia. Gurjanova (1951) had listed amphilochids from the ACKNOWLEDGEMENTS Completion of field work, 1955-1980, was greatly assisted by several marine research agencies and their staffs, and were acknowledged in published stations lists (Bousfield 1958, 1963, 1968; Bousfield & Mc- Allister 1962; Bousfield & Jarrett 1981). The authors are indebted to the Pacific Biological Station, Nanai- mo, the Bamfield Marine Station, and the Friday Harbor Laboratories, WA, with special thanks to bi- 1. Assistant Curator, Marine Invertebrates, Natural History Museum of L. A. County, Los Angeles, CA, USA, 90007. 2. Research Associate, Royal Ontario Museum, Toronto, ON, CAN M5S 2C6. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 4 ologist Dr. Craig P. Staude. Dr. D. V. Ellis kindly provided laboratory facilities at the University of Vic- toria for much of the identification work. Dr. Pierre Brunei, University de Montreal, provided illustrations and helpful commentary on a proposed new amphil- ochid taxon from the St. Lawrence estuary. The authors are grateful for facilities at the Holly Lane laboratory provided by the Canadian Museum of Nature, Ottawa, during 1992-1995, and to the collec- tions management staff of the CMN, especially Judith Price, for more recent help in cataloguing and label- ling type material. The original line illustrations were prepared with the capable assistance of artist Floy E. Zittin, Cupertino, CA. SYSTEMATICS AMPHILOCHIDAE Boeck Amphilochidae Boeck, 1871: 107; — Stebbing 1906: 149;— Barnard 1962: 116;— Barnard 1969a: 132;— Lincolnl979: 146; — Krapp-Schickel 1982:70; — Bous- field 1982: 266.— Barnard & Karaman 1991: 92. Type Genus: Amphilochus Bate, 1862 (N. Atlantic). Genera: North Pacific: Git ana Boeck, 187 1 ; Hourstonius, n. g. (p. 11); Apolochus n. g. (p. 15); Paramphilochus Ishimaru & Ikehara, 1986; Afro gitanopsis Karaman, 1980 (Indian Ocean-Japan). Fxtralimital : Gitanopsis Sars, 1895 (Arctic-N. Atlan- tic); Amphilochoides Sars, 1895 (N. Atlantic-Mediter- ranean); Gitanogeiton Stebbing, 1910 (Southwestern Pacific); Amphilochopsis Stephensen, 1925 (Arctic); Amphilochella Schellenberg, 1926 (Antarcdc-Indian); Rostro gitanopsis Karaman, 1980 (S. African); Cyclotelson Potts, 1915 (Indo-Pacific). An unde- scribed genus is listed from the St. Lawrence estuary by Brunei el aL (1998). Diagnosis: Body small, smooth; abdominal segments separate,generally unomamented. Anterior head lobe acute or rounded; rostrum distinct; eyes rounded. An- tennae short; accessory flagellum minute or lacking. Flagellum of antenna 1 longer and more richly armed with aesthetascs in male. Mouthparts modified: upper lip apically notched, lobes often asymmetrical; lower lip tall, inner margins often "notched”, inner lobes essentially lacking. Man- dible: molar various, often much reduced; spine-row strong; palp slender. Maxilla 1 , inner plate small, outer plate strongly spined and/or toothed, palp 1- or 2- segmented. Maxilla 2 small, weakly setose. Maxilli- ped inner plate slender; outer plate broad; palp me- dium, dactyl not falciform. Coxae 2-4deep, increasing posteriorly; coxa 1 small, partly hidden by coxa 2. Gnathopods usually subchel- ate, often dissimilar in form and size, not sexually dimorphic. Gnathopod 2 usually the larger; posterior KEY TO NORTH PACIFIC GENERA OF AMPHILOCHIDAE (see also Fig. 1) 1 . Peraeopods 3-7 prehensile; gnathopod 2 sexually dimorphic Afrogitanopsis (Japan) Peraeopods 3-7 ordinary, ambulatory; gnathopod 2 similar in both sexes 2. 2. Tel son elongate, sharply acute, apex minutely dentate; coxae 2-4, lower margin serrate; accessory flagellum lacking or very minute 3. Tel son apex smoothly and sharply rounded, rarely acute; coxae 2-4, lower margin smooth; accessory flagellum small, 1 -segmented 4. 3. Gnathopod 2 distinctly parachelate, propod large; maxilla 1, palp 2-segmented; mandibular molar not triturative Paramphilochus (Japan) Gnathopods 1 & 2 , propods small, nearly simple, palm weakly (or not) distinct from hind margin; max- illa l,palp 1 -segmented; mandibular molar normally large, triturative (Fig. IB) .... Gitana (p. 6) 4. Mandibular molar large, triturative, with marginal fringe of slender spines (Fig. lc); gnathopod 1, propod broadening distally, sub-triangular; peraeopod 5, brood lamella longer than basis, with 8 + marginal setae; peraeopods 3 & 4, dactyls medium, slender Hourstonius (p. 1 1) Mandibular molar vestigial, triturating surface weak or lacking; gnathopod 1, propod slightly (or not) broadening distally, anterior margin convex; peraeopod 5, brood lamella small, with 5-6 marginal setae; peraeopods 3 & 4, dactyls short Apolochus (p. 15) AMPHIPACIHCA VOL. 3 NO. 1. MAY 16, 2001 5 Fig. 1. Characters and Characters states of Gitanopsis, Gitana, and Hourstonius. A. Gitanopsis inermis (modified from Lincoln (1979). B. Gitana sarsi (modified from Sars ( 1895); C. Hourstonius laguna (after McKinney 1978). [see p. 28 for figures legend] AMPHIPACIHCA VOL. 3 NO. 1. MAY 16,2001 6 lobe of carpus typically extended forward behind pro- pod. Coxae 5-7 deep, regularly postero-lobate. Per- aeopods slender. Peraeopods 5-7 usually subsimilar in size and form; segment 4, posterodistal process not strongly overhanging segment 5. Uropods slender, rami narrowly lanceolate. Uropod 2 small, rami unequal, rarely exceeding elongate pe- duncle of biramous uropod 3. Telson longer than wide, distally (sub)acute, minutely tridentate, or rounded, weakly (or not) keeled below. Brood plates broad, decreasing posteriorly, margins long-setose. Coxal gills simple, on pereopods 2-6. Remarks; Family Amphilochidae is a member of superfamily Leucothoidea, closely related to family Pleustidae (see Bousfield 1983; Bousfleld & Shih 1 994). Character states of coxal plates and uropods are alsosuperficiallysimilartothoseoffamilyStenothoidae near which it often closely positioned (e.g., Stebbing 1906; Barnard 1962). Barnard & Karaman (1991) include family Amphilochidae within an amphilocoid group that encompasses families Cyproideidae, Pseudamphilochidae, and Bolttsiidae. Family Amphilochidae presently encompasses 13 genera and about 70 species world-wide. A further yet undescribed genus, based on an undescribed Gitan- 1/2 propod; peraeopod 7 basis widest distally . . G . sarsi Boeck 3. Gnathopods 1 & 2, propodal palm short, distinct, oblique G. ellisi n. sp. (p. 6) Gnathopods 1 & 2, palm elongate, horizontal, continuous with posterior margin 4. 4. Gnathopds 1 & 2, propod distinctly longer than carpus, tel son slender, elongate, length >2X basal width, apex acute, simple G. calitemplado Barnard Gnathopod propod shorter than carpus; tel son shorter, apex tidentate G. liliuokilaniae Barnard almond-shaped, outer row of facets largest. Anten- marginally smooth, distinctly exceeding tel son. na 1 shorter than antenna 2, lacking accessory flagel- Telson narrowly subtriangular, apex tridentate lum; peduncular segments short, flagellum 6-seg- Brood plates 2-4 large; 5 small with 3-4 distal mented. Antenna 2, flagellum 9-10 segmented. marginal setae. Mandibular molar stout triturati ve; spine row with 7- 8 slender blades; left lacinia ~8-dentate; palp segment Etymology: The species name recognizes the extens- 3 slender, shorter than segment 2, apex narrowly trun- ive contributions of Dr. Derek V. Ellis, University of cate, with 2-3 Iongish simple setae. Maxilla 1, outer plate oblique apex with 8 stout spine-teeth and proxi- mal tuft of fine setae; palp stout, 1-segmented, with several apical setae. Maxilla2, inner distal margin with several slender setae; outer plate stout, with 4 apical setae. Maxi lliped, inner plate narrow, apex subtruncate ; outer plate medium broad, with inner marginal sub- apical incision; palp segment 2 exceeding outer plate; segment 3 shorter than 2. Coxa 1 narrow , rounded below ; coxae medium deep, increasing posteriorly, smooth below. Gnathopods 1 & 2, propods small, subrectangular; palms short, ob- lique, convex, denticulate, not sharply demarcated at posterior angle; dactyls stout, pectinate behind, unguis overlapping palm. Gnathopod 1, carpal lobe very short, little produced. Gnathopod 2 larger, carpus shorter than propod, posterior lobe short, stiff-setose, extending about half length of propod. Peraeopods 3 & 4 very slender, weakly armed; dactyls slender, medium long. Coxae 5-7 deeply posterolobate, margins unarmed. Peraeopods 5-7 regu- larly and subequally homopodous; bases regularly ex- panded, posteriorly slightly increasing in size; seg- ment 4 long, 5 short; dactyls slender, medium long . Pleon side plate 3, lower margin straight, hind comer slightly produced. Pleopod rami 9-10 segmented, slightly longer than thick peduncle. Uropod 1 slender, rami subequal, margins spinose. Uropod 2, rami marginally short-spinose; outer ramus >1/2 inner. Uropod 3, rami much shorter than slender peduncle. Victoria, in the development and teaching of coastal marine ecological and environmental concepts. Distribution: Only 7 specimens known; from shallow- sublittoral depths off southern Vancouver Island, B. C. Remarks: Gitana ellisi is closest to G. sarsii Boeck; however, its larger eyes, more distinctly subchelate gnathopods with less produced carpal lobes, smaller coxal plates, and much less markedly reduced man- dibular palp are more plesiomorphic character states than in G. sarsi . Gitanopsis Sars (restricted) Gitanopsis Sars, 1895: 223;— Stebbing 1906: 153;— Gurjanova 1951: 302. Gitanopsis Lincoln 1979: 164 (part);— Barnard 1962: 130 key (part);— Barnard & Karaman 1991: 97 (part). non: Gitanopsis McKinney 1978: 140.— Karaman 1980: 44;-Hirayama 1983: 124. Type species: Amphilochus bispinosa Boeck, 1871, original designation by G. O. Sars. Species: Gitanopsis inermis Sars, 1883; G. arctica Sars, 1895;— Shoemaker 1955; G. abyssicola Sars, 1895; Gitanopsis sp. A, Just ,1980; Gitanopsis sp. B, Just, 1980. AMPHIPACIHCA VOL. 3 NO. 1. MAY 16,2001 9 KEY TO NORTH PACIFIC SPECIE S OF HOURSTOMUS ( Gitanopsis iseebi Yamato transferred to Afrogitanopsis) 1 . Gnathopods 1 & 2, propods subsimilar in size and shape, palmar angle squared, defined by small tooth and spine; carpus, posterior lobe short, little produced H. pusilloides (Shoemaker) Gnathopod 2 distinctly larger than gnathopod 1, palmar angle obtuse, defined by spine(s) only; carpus of gnathopod 2 strongly produced posterodistally below propod 2. 2. Gnathopods 1 & 2, posterior angle of propod defined by single spine; maxilliped, outer plate mark- edly incised mediodistally; Hawaiian islands H. pete (Barnard) Gnathopod propods, posterior angle with paired spines; maxilliped, puter plate little or not incised mediodistally 3. 3. Telson apically acute, minutely bifid; maxilliped palp, segment 2 distinctly longer than segment 3; mandibular spine row short, ~6-bladed H. japonica (Hirayama) Telson, apex subacute or rounded; maxilliped palp segment 3 short, not longer than segment 3; mandibular spine row with 8-12 blades 4. 4. Gnathopod 2, propod with two prominent anterior submarginal spines; gnathopod 1, anterior mar- gin of basis setose throughout; maxilla 2, inner plate weak, little broader than outer 5. Gnathopod 2 lacking anterior submarginal spines; gnathopod 1, anterior margin of basis nearly bare; maxilla 2, inner plate normal, setose, broader than outer plate 6. 5. Gnathopod 2, carpal lobe elongate, extending almost to palmar angle; telson elongate, length >2X maximum width; N. American Pacific H. vilordes (Barnard) (p. 11) Gnathopod 2, carpal lobe short, ~ 3/4 length of posterior margin of propod; telson short, length 1.5 X maximum width; Gulf of Mexico H. laguna (McKinney) 6. Telson narrow, elongate, length > 2X maximum width; gnathopod 2, carpal lobe extending be- yond posterior palmar angle 7. Telson short, broader, length not > 2X width ;gnathopod 2, carpal lobe not quite reaching poster- ior palmar angle H. brevteulus (Hirayama). 7. Abdominal (epimeral) side plate 3, hind comer slightly produced; uropod 2, outer ramus normal , >1/2 length of inner ramus H. robastodentes (Hirayama) Epimeral plate 3, hind comer squared or rounded; uropod 2 outer ramus short, not greater than 1/2 length of outer ramus //. longus (Hirayama) Diagnosis: Rostrum medium; anterior head margin usually acute; eye round. Antennae usually subequal in length; accessory flagellum lacking or very minute. Upper lip shallowly notched, lobes subs ymmetri cal . Lower lip, inner margins strongly notched. Mandible: molar process large, triturating surface without raised marginal spines; spine row with few blades; palp seg- ment 3 elongate, setulose near apex. Maxilla 1 normal, palp 2-segmented. Maxilla 2 normal, inner plate nar- row. Maxilliped palp slender, segment 2 often short. Coxae 2-4 large, lower margin weakly serrate. Gna- thopods \ & 2 medium to weakly subchelate; prododal palm not demarcated from posterior margin; carpus of medium length, lobe produced. Peraepods 3 & 4, dactyls slender, medium. Coxae 5-7 regularly post- erolobate. Peraeopods 5-7, dactyls slender, medium long. Epimeral plate 3, hind comer subquadrate . Uropod 3 relatively short, little exceeding uropod 1. Telson elongate, narrowing, apex acute, minutely tridentate. Distribution: Holarctic and northern North Atlantic, 0 -875 m. in depth. Remarks: Gitanopsis s ens, lat. (e.g., Barnard & Kar- aman, io^sil.) here consists of three groups: Gitanopsis Boeck sens, str. (Arctic and northern North Atlantic in distribution); and Hourstonius new genus, mainly in AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 10 Fig. 3. Hourstonius vilordes ( Barnard). 9 ov. (3..9 mm). Hinks I., N. end Aristazabei I., B .C. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 11 the North Pacific region, with some species in the Gulf of Mexico, and possibly coastal waters of the African South Atlantic region. A third generic group, infor- mally recognizable and yet unnamed within Gitanopsis (sens* iaL), may be represented by G. marionis (Steb- bing, 1888), as figured by Bellan-Santini & Ledoyer (1974), G. tai Myers, 1985, and several other largely Indo-Pacific and southern hemispheric species (ex- cept G. squamosa (Thompson, 1880) as listed by Bar- nard & Karaman (1991). Principal characters and character states utilized in keys, diagnoses, and numerical analyses pertaining to the genus Gitanopsis ( sens, lat .) are represented in Fig. 1 (p* 55). Hourstonius , new genus Gitanopsis Sars, 1895: 224 (part);— Barnard 1962: 130 (key, part);— Hirayama 1983: 124 (key);— Bar- nard & Karaman 1991: 97 (part);— Ishimaru, 1994: 52 (species list). mmGitanopsis Gurjanova. 1951:302;— Lincoln 1979: 164. Type species: Gitanopsis v Hordes J. L. Barnard, 1962 Species: Hourstonius breviculus (Hirayama, 1983); H. japonica (Hirayama, 1983);//. longus (Hirayama, 1 983) ; H. robastodentes (Hirayama, 1 983 ) ( J apan) ; H. laguna (McKinney, 1978); H. tortugae (Shoemaker 1942)(Rorida); H. pusilloides (Shoemaker, 1933; H. pusilla (K. H. Barnard, 1916)(S. Africa); H.pele (J. L. Barnard, 1970); H. baciroa (J. L. Barnard, 1979) (Galapagos);//. magdai( Reid, 195 l)(Trop. Atlantic) non: Afrogitanopsis paguri (Myers, 1974) (W. Indian); A. iseebi (Yamato, 1993)( Japan). Diagnosis: Anterior head lobe generally rounded. Ac- cessory flagellum 1 -segmented or minute. Upper lip notched, lobes asymmetrical; lower lip , inner marginal “notch” weak or lacking; mandible, molar distinct, outer triturating ridge with raised spines; maxilla 1, palp slightly modified; maxilliped, palp segment 3 short; coxa 2, lower margin smooth (not serrate); gnathopods 1 & 2 strongly subchelate, propod with paired spines at posterodistal angle demarcating palm; gnathopod 2, carpus narrow, posterior lobe elon- gate; Epimeral plate 3, hind comer squared or rounded; telson linguiform, medium to long, apex broadly or sharply rounded (acute in H. japonica). Distribution: Mainly North Pacific, with some spe- cies in the Gulf of Mexico, and possibly coastal waters of the African South Atlantic region. Etymology: The genus is named in honour of the late Alan S. Hourston, fisheries scientist with the Pacific Biological Station, Nanaimo, British Columbia. Dr. Hourston, his wife Barbara, and their family provided much help and encouragement to the senior author and his family during the 25 -year period of field work for the Pacific amphipod program, 1955-1980. Remarks: Members of this genus are readily distin- guished from species of Gitanopsis Sars ( sens, str. ) in possessing an accessory flagellum, more strongly sub- chelate gnathopods with shorter carpus; weakly (or not) serrated lower margin of coxa 2; and linguiform telson with rounded apex. Moreover, species of Gitan- opsis (sens* sIl) are mainly Arctic and North Atlantic in distribution. Members of the third generic group (G. marionis , G. squamosa , G. tai , and several others (see Gitanop- sis above). Members appear superficially more closely similar in some character states (e.g., of accessory flagellum, upper 1 ip) to Hourstonius than to Gitanopsis sens, sir., but detailed analysis is beyond the scope of the present study. Hourstonius vilordes (J. L. Barnard, 1962) (Fig. 3) Gitanopsisvilordes Barnard, 1962: 131,fig.6;— 1969b: 82;- 1975: 344, fig. 190;- Austin, 1985: 593;- Staude, 1987: 379. Material Examined: ALASKA SE Alaska, ELB Stns., 1961: A3, Little Daykoo, Dali I. (54°42'N, 132° 42' W); MW-LW, May 3 1 - 2 99, NMCC 1992-02 15. BRITISH COLUMBIA. Queen Charlotte Islands, ELB Stns., 1957: W4b, Small bay, north shore Hippa Passage (53°27’N, 132° 58'W), 6-10 m dredge, gravel, stones, shells, Aug. 10. - 1 9, NMCC 1992-0204. North Central coast, ELB Stns., 1964: H3, Cockle Bay, Lady Douglas I. (52°21'N, 128°23'W), sand, Zostera, kelp, LW, July 9. - 4 99, NMCC 1992-02 19. H5, Hinks I., north end Aristazabal I. (52°38'N, 129°05' W), stones, PhylIosp[adix, kelp LW, July 10.- 9 (ov.) (Flg’d specimen), 2 99; 3 Cfc? , 57 imm. ; slide mounts: 99 ( 1 .8mm, AMPHIPACIHCA VOL. 3 NO. 1. MAY 16,2001 12 2.2 mm, 2.3 mm, 2.4 mm. 2.4 mm, 2.6 mm, 2.7 mm,2. 9 mm), NMCC 1992-0220. H26, Cox Pt., inlet at mouth (53°08'N, 129°45'W), shelly sand, kelp, LW, July 19. - 2 99, NMCC 1992-0225. H65, Christie Pass, cove on south side Hurst I. (50°50'N, 127°35’W), shelly gravel, kelp, MW-LW, Aug. 1 1. - 8 99, NMCC 1992-0232. Northern Vancouver I., ELB Stns., 1959: 04, Browning Inlet (50°30'N, 128°06’W), shelly sand, eel grass, LW, July 19. - 1 9, NMCC1992- 0213? V5, Lemon Pt., Nigei I. (50°51'N, 127°46'W), Phyllospadix, Corallina, fucoids, LW, Aug. 7. - 1 9, NMCC 1992-02 10. V7, Lady Ellen Pt. .Broughton Str. (50°36’N, 127°07W) - 1 9, NMCC 1992-021 1. V 10,AlertBay,BroughtonStr. (SO^S'N, 126°56'W) muddy sand, kelp, LW, Aug. 1 - 1 9, NMCC 1992- 0212. V17, Boat Bay, Cracroft I. (50°31'N, 126°34W), coarse sand, eelgrass, LW, Aug. 5.-3 99, NMCC1992- 0217. V20, Brown Bay (50°10’N, 125°22'W), coarse sand, kelp, LW, June 22. - 1 CT, 2 99, NMCC 1 992-02 14. N 1 1 , Port Progress, Queen Charlotte Str. (SO^'N,^ 0 ^’ W), sandy mud, Tostera , LW, Aug. 4 - 399, NMCC 1992-0205. Southern Vancouver Island, ELB Stns., 1955: F6, Telegraph Bay (48°27'N, 123°17W) - 1 9 NMCC 1992- 0202. Victoria region (48°N, 123°W), KEC Stn.?, 1981. - 1 9, NMCC 1992- 0249; Ibid., - 3 d*Cf, 1 9 ov. (2.5 mm), slide mount, 10 females, NMCC 1992-0250; Ibid., - 19, NMCC 1992-0251. Diagnosis: Female ov. (3.9 mm) (fig’d specimen). Rostrum medium. Eye large, subreniform. Anten- nae short, subequal. Antenna 1, flagellum 5-6 seg- mented; accessory flagellum minute. Antenna 2, flagellum 6-segmented. Upper lip, apex distinctly asymmetrical. Lower lip with weak inner “shoulders”. Left mandible, lacinia 8-9 cuspate; spine row with 10-12 slender blades; palp segment 3 distinctly longer than 2, inner distal margin finely pectinate, apex sharply acute.. Maxilla 1, inner plate with 7 apical spine teeth and 4-5 inner apical seta; palp and segments stout. Maxilla 2, plate medium, weakly armed. Maxilliped, plates narrow; palp seg- ment 1 large, length nearly equal to 2 & 3 combined. Coxa 1 narrow elongate; coxae 2 -4 large deep, smooth below. Gnathopods 1 & 2 propods large, palms smoothly convex, nearly vertical, sharply demarcated at posterior angle by paired spines; dactyls slender, finely pectinate behind except on unguis. Gnatho- pod 1, carpal lobe spinose, produced 2/3 length of posterior margin of propod. Gnathopod 2, carpus narrow, lobe extending almost to posterior angle of propod. Peraeopods 3 & 4 slender, weakly armed; dactyls slender. Coxae 5-7 normally posterolobate, margins unarmed. Peraeopods 5-7 regularly and subequally homopodous; bases regularly expanded, posteriorly slightly increasing in size; dactyls slender, medium long. Pleon side plates 2 &3, hind comers weakly acumi- nate. Pleopod rami slightly longer than peduncle, 7-8 segmented. Uropod 1, rami markedly unequal, mar- gins weakly spinose. Uropod 2, ramus marginally short-spinose; outer ramus ~= 1/2 inner. Uropod 3, rami short, broad, marginally smooth, slightly exceed- ing telson. Telson elongate, smooth, apex narrowly rounded. Brood plates medium broad; plate 5 with 10-12 distal marginal setae. Distribution: Shallow littoral depths from SE Alaska to southern Vancouver Island, south to central and southern California. Remarks: In character state similarity, Hourstonius vilordes appears somewhat closer to species of Hourst- onius from the Gulf of Mexico than to species of "Gitanopsis" ( sens. lat .J described by Hirayama ( 1983) from Japanese coastal marine waters (see key to spe- cies, p. 9). H. vilordes differs from the Hawaiian spec- ies, H. pele (Barnard, 1970), in its larger coxal plate 2, larger mandibular palp, and narrower telson, as well as character states of the gnathopods given in the key. Amphilochus Bate (restricted) Amphilochus Bate, 1862: 107.— Stebbing 1906: 149 (part);— Lincoln 1979: 148 (part);— Krapp-Schickel 1982: 74 (part);— Barnard & Karaman 1991: 96 (part). Type species. Amphilochus manudens Bate, 1862, monotypy. Species: Amphilochus tenuimanus Boeck, 1871; A. planierensis Ledoyer, 1977. Diagnosis: Rostrum medium; anterior head lobe acute; eyes rounded. Antennae subequal in length; accessory flagellum lacking or minute. Upper lip, apical incision shallow, lobes sub-sym- metrical. Lower lip, inner margins smooth. Mand- ible: molar very reduced, knob-like, lacking triturating ridges; spine-row medium; palp segment 3 elongate. Maxilla 1 normal, palp 2-segmented. Maxilla 2 regu- lar, outer plate slender. Maxilliped palp slender. Coxal plates 2-4 large, lower margins serrate. Gnath- opods 1 & 2 medium to strongly subchelate, palmar AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 13 KEY TO NORTHERN PACIFIC AND ATLANTIC SPECIES OF APOLOCHUS (see character states of subgroups 1 & 2 of Fig. 4, p. 64 ) 1. Mandibular molar small, knob-like, lacking triturating ridges; maxilla 2, plates reduced, weakly armed; peraeopods 3 & 4, dactyls short, thick (North American Pacific subgroup) 2. Mandibular molar distinct, apex acute with few triturating ridges, or flat, with several ridges; max- illa 2, inner plate broad; peraeopods 3 & 4, dactyls usually slender, medium (Atlantic-Mediterr- anean regional subgroup) 4. 2. Gnathopod 2, carpal lobe short, <1/2 propod margin; mandibular spine-row 10-12 bladed; uropod 2, rami nearly bare; telson elongate, 2 1/2X width A. Utoralis (Stout) (p. 16) Gnathopod 2, carpal lobe long, reaching posterior palmar angle; mandibular spine row long, 14- 16 bladed; uropod 2, rami marginally spinose; telson shorter, length 2X width 3. 3. Antenna 1 shorter than 2; gnathopod 2, palm shallowly convex, nearly perpendicular; maxilliped outer plate wide, little longer than broad A. barnardi , n. sp. (p. 18 ) Antennae 1 & 2 short, subequal in length; gnathopod 2, palm strongly convex, oblique; maxilliped, outer plate medium, longer than wide A. staudei , n. sp. (p. 19 ) 4.. Mandibular molar elongate, apex subacute, with few (or no) triturating ridges; gnathopod 2, carpus short, < 1/3 anterior margin of propod 5. Mandibular molar short, apex sub truncate, flat, with several triturating ridges; gnathopod 2, carpus medium, length >1/3 anterior margin of propod 7. 5. Gnathopod 2 , propod large, carpal lobe extending almost to palmar angle; accessory flagellum very minute or lacking A. picadurus (J. L. Barnard ) Gnathopod 2, propod medium to small, carpal lobe extending little more than half way to palmar angle; accessory flagellum distinct, 1 -segmented 6. 6. Uropod 3, rami subequal; maxilliped, outer plate broader than long A. borealis (Enequist) Uropod 3, inner ramus distinctly longer than outer ramus; maxilliped outer plate longer than broad. A. pillaii (Barnard & Thomas) 7. Antennae subequal in length; coxa 1 serrate below; mandibular spine-row with 8-10 blades 8. Antenna 1 short flagellum little beyond peduncle of A2; coxa 2 smooth below; mandibular spine row with ~14 blades A. neopolitanus complex, N-E. Atlantic 9. 8. Gnathopod 2, propod with 4 stout antero-marginal spines A. casahoya (McKinney) Gnathopod 2, propod with 2 stout antero-marginal spines A. delacaya (McKinney) 9. Uropod 2, ramal margins spinose; coxa 1 subrectangular, lower margin smooth; telson short, length 1.5 X maximum width A. neopolitanus (Della Valle) (fide Krapp-Schickel)* Uropod 2, margins of rami nearly unarmed; coxa 1 subtriangular, lower margin with distinct notch; telson regular, length about twice width . . Apolochus sp. (= A. picadurus Krapp-Schickel, 1982). margin not demarcated from posterior margin by paired Pleon segment 3, hind comer with small cusp, spines; carpus medium, lobe variously produced; dact- Pleopods regular. Uropod 3, inner ramus slightly yls finely denticulate. Peraeopods 3-4, dactyls slender, broadened. Telson elongate, narrowing distally, apex medium long. Coxae 5-7 deep, shallowly posterolob- acute, minutely dentate, ate. Peraeopods 5-7, dactyls slender, medium long. * " Amphilochus " neopolitanus of Lincoln (1979) differs from that of Krapp-Schickel (1982) AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 14 Fig. 4. Selected Characters and Character States within Amphilochus and Apolochus. A.Amphilochus manudens Bate [modifed from Lincoln (1979), Sars (1895)]; B. Apolochus borealis (Enequist, 1950); Q. Apolochus sp. [= A. picadurus (Krapp-Schickel ( 1982)1 ; D. Apolochus delacay a (McKinney, 1978). E. Apolochus neopolitanus Della Valle (after Krapp-Schickel (1982)]; F. Apolochus picadurus (Barnard, 1962). [see p. 28 for figure legend] AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 15 Distribution: Northern North Atlantic, N. American [Brunei el aL. 1998) and boreal European-Mediterra- nean region. Habitat: On scleractinian and homy corals, 0-600 m. Remarks: Amphilochus Bate (sens, str.) is defi ned by: anterior head lobe acute; accessory flagellum lacking; upper lip apical lobes subsymmetrical; gnathopod propods lacking posterodistal palmar spines; coxae 2- 4 with serrate lower margins; peraeopod dactyls elon- gate; and tel son apically acute, minutely tridentate. The form of the gnathopods, mouthparts, and telson suggest that Amphilochus sens. sir* may have been para-ancestral to regionally more advanced genera such as Amphilochoides Sars, 1895, and Paramphil - ochoides Lincoln, 1979. However, these latter two genera may themselves require redefinition. Thus, the type species of Amphilochoides [A. boecki Sars (= A. odontonyx Sars, 1895)] possesses a unique combina- tion of generic-level character states: upper lip, apical lobes subsymmetrical; maxilla 2 plates small, partly fused; dactyls of both gnathopod 1 & gnathopod 2 with proximal nodiform process; and epimeral plate 3, hind comer toothed. At least three species with normally reduced mandibular molar, presently included in the genus Amphilochoides by Barnard & Karaman (1991), differ in the above character states. Thus, " Amphil- ochoides ” longimanus Chevreux, 1888, A. serratipes Norman, 1869, and a distinctive Mediterranean species figured as A. serratipes by Krapp-Schickel (1982) might justify separate generic recognition. Despite aberrencies in the form of the gnathopods, in overall character-state similarity the genus Paramphilochus Ishimaru & Ikehara, 1986, appears more closely similar to Pacific members of Apolo - chus and Hourstonius than to Amphilochus ( sens. sIl). Apolochus, new genus Amphilochus Sars 1895: 215 (part); Stebbing 1906: 1 49(part) ; — Barnard 1969a: 1 36 1/2 inner. Uropod 3 elongate, rami unequal, marginally spinose, greatly exceeding telson. Telson smooth, length about twice width, apex narrowly rounded. Brood plates medium broad; 5 with 2-3 distal mar- ginal setae. Distribution: Intertidal and LW habitats, from south- ern California, north through Oregon,Washington, and B. C. to southern S.E. Alaska, Taxonomic Commentary: As indicated in the key, Amphilochus litoralis and other American Pacific coast species are distinct from North Atlantic and Gulf spe- cies . Apolochus barnardi new species (Fig. 6) Amphilochus ?neopolitanus cf. Della Valle fide B ar- nard, 1962: 126;-Bamard 1964: 105(?)-Bamard 1969b: 82. Material Examined: None. Diagnosis (partly after Barnard 1962): Femaleov.(2.5 mm): Rostrum medium. Eye medium, narrowly subovate. Antennae medium short, subequal. Antenna 1, peduncular segments 1 & 2 short and deep, 2 with posterodistal tuftof setae;3 small, narrow; flagellum 8- 9 segmented; accessory flagellum minute. Antenna 2, flagellum 9-10 segmented. Mandibular molar small, broadly triangular. Left mandible, lacinia 8-9 cuspate; spine row with 15-17 slender blades, distinctly largest distally; incisor multidentate; palp segment slightly longer than 2, inner margin finely pectinate, apex with single short seta. Maxilla 1, apical margin of inner plate nearly vertical, with 7 spine-teeth and 6-8 setae at inner angle; palp segments stout, segment 1 large. Maxilla 2, plates medium, inner plate with 6 marginal setae, proximaily plumulose. Maxilliped, inner plates tall, narrow, apex subtruncate; outer plate short, broad, distal margin with several setae and stout spine, inner margin smooth; palp segment 2 distinctly exceeding outer plate; seg- ment 3, inner distal margin with narrow denticles. Coxa 1 short, little occluded by 2; coxae 2 -4 succes- sively deepening, rounded below. Gnathopods 1 & 2 distinctly subchelate. Gnathopod 1, propod slightly expanding distally, palm convex, oblique; dactyl slen- der body denticulate behind, unguis elongate, slightly exceeding palmar angle; carpus narrow, lobe medium, extending about 1/2 posteriormarginof propod. Gnath- opod 2, propod large, broadening distally, palm con- vex, finely crenulate, nearly vertical, sharply demar- cated at posterior angle by paired spines; dactyl slen- der, body finely pectinate behind, unguis elongate slightly exceeding palm; carpus short, posterior lobe elongate extending length of posterior margin of pro- pod. Peraeopods 3 & 4 medium, segment 6 with 4 poste- rior marginal spines; dactyls short, unguis short. Coxae 5-7 normally posterolobate, hind lobes rounded below, posteroventral margin of 7 weakly spinose. Peraeo- AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 19 Fig. 6. Apolochus bamardi , new species (=A. neopolitanus cf. Della ValleJMe J. L. Barnard ,1962) (modified from Barnard, 1962). pods 5-7 regularly homopodous; bases regularly ex- panded, posteriorly slightly increasing in size; segment 4not elongate; distal segments marginally spinose, esp. anteriorly; dactyls short. Pleon side plates 2 & 3 convex below, hind comers not acuminate. Pleopod rami distinctly longer than peduncle, 9-10 segmented. Uropod 1 slender, rami unequal, margins spinose. Uropod 2, rami marginally short-spinose; outer ramus <1/2 inner. Uropod 3 elon- gate, greatly exceeding telson; rami medium, unequal, marginally spinose. Telson subtriangular, length slightly less than twice width, apex narrowly rounded. Brood plates medium broad, 5 narrow. Distributional Ecology: Apolochus barnardi occurs in the Phyllospadix and Egregia sublittoral zone to depths of ~20 m, mainly on bottoms of coralline algae, stones, and sessile invertebrates, from Central to South- ern California. Etymology: The name is a tribute to the early recogni- tion of this distinctive form by the late J. L. Barnard, and to his major contributions to knowledge of the North American Pacific amphipod fauna. Remarks: Apolochus barnardi is distinct from A. neopolitanus DellaValle 1893, as figured by Krapp- Schickel (1982) in characters of the key (p. 13 ), It is also distinguished from A. staudei by characters of the key and as noted elsewhere (below). Apolochus staudei new species (Fig. 7) Amphilochus neopolitanus Della Valle, fide Staude, 1987: 379, fig. 18. 40?; -Austin 1985: 593. non : Amphilochus neopolitanus cf. Delle Valle fide Barnard 1962 (California). Material Examined: 15 specimens at 7 stations: BRITISH COLUMBIA North central coast, ELB Stns., 1964: H12, Stephens I., NW end (54°U'N, 130°48'W), Phyllo- spadix, kelp, LW, July 13, - 1 $; NMCC 1992-0222. H20, McCauley I., NH end (53°43'N, 130°22'W), fine sand, LW, July 17 - 2 99, N MCC 1 992-022 1 . H21, N. end Banks I. (53°25'N, 130°10'W), 40-60 m. - 1 d* ; 2 99: NMCC 1992- 0223. H22, 1/2 mile off Larsen Hd., Banks I. (53°34'N, 130°34'W), kelp on sand and shell, 20 m dredge. July 17. - AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 20 Fig. 7. Apolochus staudei, n. sp. Female ov. (2.4 mm) Holotype. N end of Rennison I., B. C. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 21 1 9, 2 imm, NMCC-0224. H30, N. end of Rennison I. (52°51 'N, 129°2 1 'W), kelp and sand, 8-25 m dredge, July 20. - 9 Holotype (slide mount); 34 99 Paratypes, 3 d*C f Allotypes, 9 im, Paratypes, NMCC 1992-0226. H53, Townsend Pt., St. John Harbour (52°12'N, 128°28'W), Phyllospadix, kelp, Corallina, bedrock, LW, Aug. 7. - 1 9, NMCC 1992-0230. San Juan's Brown I., E. side. Queen Charlotte Str. ( 51°19'N, 127°46'W), 50-60 m dredge, FRB?, 1983. - 1 9, NMCC- 1992-0253. Diagnosis: Female ov. (2.4 mm) (holotype). Rostrum medium. Eye large, subovate. Antennae short, subequal. Antenna 1, peduncular segment 1 & 2 short and deep, 3 small; flagellum 6-7 segmented; accessory flagellum minute. Antenna 2, flagellum 6-7 segmented. Mandibular molar small, broadly triangular, with apical setules. Left mandible, lacinia 8-9 cuspate; spine row with 15-17 slender blades, distinctly largest distally; incisor multidentate; palp segment slightly longer than 2, inner margin finely pectinate, apex with single short seta. Maxilla 1 , apical margin of inner plate nearly vertical, with 7 spine-teeth and 6-8 setae at inner angle; palp segments stout, segment 1 large. Maxilla 2, plates medium, inner plate with 6 marginal setae, proximally plumulose. Maxilliped, inner plate tall, narrow, apex subtruncate; outer plate medium broad, tall, inner margin smooth; palp segment 2 only slightly exceeding outer plate. Coxa 1 short, little occluded by 2; coxae 2 -4 succes- sively deepening, rounded below. Gnathopods 1 & 2 distinctly subchelate. Gnathopod 1, propod slightly expanding distally, palm convex, oblique; dactyl slen- der body denticulate behind, unguis elongate, slightly exceeding palmar angle; carpus narrow, lobe medium, extending about 1/2 posterior margin of propod. Gnath- opod 2, propod large, broadening distally, palm con- vex, finely crenulate, nearly vertical, sharply demar- cated at posterior angle by paired spines; dactyl slen- der, body finely pectinate behind, unguis elongate slightly exceeding palm; carpus short, posterior lobe elongate extending length of posterior margin of pro- pod. Peraeopods 3 & 4 medium, segment 6 with 4 poste- rior marginal spines ; dactyl s short, unguis short. Coxae 5-7 normally posterolobate, hind lobes acute below, margins unarmed. Peraeopods 5-7 regularly homopod- ous; bases regularly expanded, posteriorly slightly in- creasing in size; segment 4 not elongate; dactyls short. Pleon side plates 2 &3, gently convex below, hind comers not acuminate. Pleopod rami distinctly longer than peduncle, 9-10 segmented. Uropod 1 slender, rami slightly unequal, margins weakly spinose. Uropod 2, rami marginally short-spinose; outer ramus >1/2 inner. Uropod 3 elongate, greatly exceeding telson; rami medium, unequal, marginally spinose. Telson narrowly triangular, length about twice width, apex sharply rounded. Brood plates medium broad; plate 5 narrow, with 5-6 longish distal marginal setae. Etymology: The species name recognizes Dr. Craig P. Staude, Friday Harbor Laboratories, for his outstand- ing contribution to knowledge of the systematics and ecology of amphipods of the northeastern Pacific mar- ine region. Distributional Ecology: Known only from the Queen Charlotte Sound coast of north central British Colum- bia south to northern Queen Charlotte Strait, LW and shallow sublittoral to 60 m in depth. Remarks: Although most closely related to A. barn - ardi, Amphilochus staude i differs mainly in the form of the gnathopods, especially gnathopod 1, the distinctly posteriorly pectinate dactyls, theform of the maxilliped plates and palp, and the slightly longer and more acutely pointed telson. DISCUSSION Phyletic Reclassification In phyletic revisions of the Gammaridea, family Amphilochidae has been placed within superfamily Leucothoidea(Bousfield 1979, 1982, 1983,2000,2001 ; Bousfield & Shih 1994). Lowry & Myers (2000) have recently proposed superfamily Iphimedioidea which combines former leucothoidean families Iphimedidae, Lafystiidae,andLaphystiopsidaewitheusiroideanfam- ilies Epimeridae and Amphithopsidae. Remaining within Leucothoidea are families Leuco- thoidae, Anamixidae, Amphilochidae, Pleustidae, Stenothoidae,Thaumatelsonidae,andCressidae. How- ever, the Stenothoidae, Thaumatelsonidae, and Cress- idae differ markedly in lacking a conspicuous rostrum, but exhibit sexually dimorphic gnathopods, strongly reduced maxilliped plates, frequent fusion of urosome segments and/or telson, and 2-segmented outer ramus of uropod 3. Furthermore, the gnathopods are fre- quently sexually dimorphic, Bousfield (2001b) for- mally utilizes the superfamily name Stenothoidea to encompass these three families. Based on the principal AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 22 Fig, 8. Phenogram of morphological similarities abnd possible phyletic relationships within Gitana , Gitanopsis , and Hourstonius . character state differences noted above, and pending further detailed analysis, the superfamily name Stenothoidea is herewith formally recognized. Family Amphilochidae therefore remains within superfamily Leucothoidea. Reorganization of genera Taxonomic placement of species within northern hemispheric genera of amphilochid amphipods has long been unsatisfactory. Difficulties encountered, and partial generic revisions attempted by Lincoln (1979), Karaman (1980), and to some extent Krapp- Schickel (1982) and Hirayama ( 1983), are reflected in the number of character state "variables" specified by Barnard & Karaman ( 1991), especially within the gen- era Gitana, Gitanopsis, Amphilochus, and Amphiloch - opsis. Likewise, the present study encountered a per- sistence of unsatisfactory taxonomic categorizations within North Pacific species, and absence of previous numerical analysis. The authors have here attempted a realignment of species on more comprehensive and more natural generic conceptualizations. The present treatment of species and genera utilizes a semi-phyletic modification of the UPGMA system of Sneath & Sokal (1973), as in previous analysis ofother NorthPacificamphipodgroups(e.g.,Jarrett&Bousfield 1 994; Bousfieid & Chevrier 1996) . Character states are ordered plesio-apomorphically and relative phyletic placement of a taxon is represented by a numerical sum of plesiomorphic, intermediate, and apomorphic char- acter states values (0, l, and 2, resp.) in a Plesio- Apomorphic (P.-A.) Index. Tabular data on which the resulting phenograms are based are considered overly bulky and repetitive for publication here, but may be supplied on request. Fig. 8 graphically portrays morphological similari- ties within species of Gitana , Gitanopsis sens, sir* (effectively encompassing the arctic monotypic genus Amphilochopsis ), and boreal-warm temperate. North Pacific and North Atlantic species of Gitanopsis (= AMPHIPACIF1CA VOL. 3 NO. 1. MAY 16,2001 23 > H tr < O) Fig. 9. Phenogram of morphological similarities and possible phyletic relationships within the genera Amphilochus and Apolochus. Hourstonius). Species groupings are distinct above the 75% similarity level within Gitana, Gitanopsis sens. str. T and Hourstonius. The former two genera cluster at about the 75% similarity level, with the genus Amphilochopsis somewhat intermediate between the two. However, the genus Hourstonius remains dis- tinct below the 60% similarity level. The eastern Pac- ific species, H. v Hordes (J.L. Barnard) is similar to species of both the western Pacific and Caribbean (Gulf of Mexico) regions, described mainly by Hiray- ama and McKinney, respectively. Not unexpectedly it is relatively remote from species of the South Atlantic (e.g., H. magdai, H. squamosa ) described elsewhere. This limited analysis tends to validate generic rea- lignment of species within Gitanopsis Sars, 1895, and formal recognition of the generic concept Hourstonius. However, further study of relationships of species of the southern hemisphere, and of groups commensal with crustaceans, is clearly needed. Fig. 9 portrays character state similarities within the North Atlantic genus Amphilochus Bate (sens. str. L based on the type species A. manudens, and species of the boreal -warm-temperate North Pacific, North At- lantic and Mediterranean-Indo-Pacific regions (= Apolochus, n. g.). Species groupings are distinct above the 75% similarity level within both groups, but the two genera remain distinct below the 50% similarity level. Apolochus here encompasses three subgroups: (1) an eastern Pacific complex of A. litoralis (Stout, 1912 ) andtwocloselysimilar species ne wl y de sc ri bed herei n ; (2) a more speciose, essentially Atlantic (Caribbean- Mediterranean) subgroup encompassing A. neopol- itanus Della Valle, A. picadurus (J. L. Barnard) and several superficially similar species [e.g., figured but unnamed by Lincoln (1979), Krapp-Schickel (1982)]; and (3) a Mediterranean Indo-Pacific subgroup encom- passing A. brunneus, A. spencebatei and species attrib- uted to "A. neopolitanus" by Ledoyer (1977). AMPHIPACIFICA VOL. 3 NO. 1. MAY 16,2001 24 Fig. 10. Phenogram of morphological similarities and possible phyletic relationships within genera of Amphilochidae. Species realignment within AmphilochusSars, 1895, andformal recognitionof the generic concept Apolochus seem clearly in order. Species attributable to genus Apolochus exhibit significantly higher A.-P. indices than the three more primitive species here attributed to Amphilochus Bate sens, str. However, further study, especiallyof Indo-Pacificregional taxa,is muchneeded. Fig. 10 is a phenogram of morphological similari- ties within genera of family Amphilochidae, mainly of the northern hemisphere. The genera form three main subgroups at, or slightly below, the 75% similarity level: (1) an Arctic-North Atlantic complex of the primitive genus Gitana and the closely related Gitanopsis, Amphilochopsis and Amphilochus, having low P.-A. values ranging from 12 to 19; (2) a Mediter- ranean-Indian oceanic complex of advanced genera Amphilochoides, Paramphilochoides and Amphill- ochella, with intermediate P.-A. values of 19-21; and (3) a moderately advanced group of the closely related genera Hourstonius and Apolochus "satellite genera" Paramphilochus and Afrogitanopsis respectively, all with intermediate to high P.-A. values of 20-23. The two principal genera encompass about half the known species within the entire family. The analysis further tends to confirm generic rea- lignment of species within the new generic concepts of Hourstonius and Apolochus. The overall morphologi- cal closeness of these two genera underscores a need for use of multiple-character anal ysi s, and avoidance of single- or few-character diagnoses, in defining taxo- nomic concepts at generic level and higher. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 25 Phyletic and Biogeographic Conclusions Members of family Amphilochidae are ectocom- mensals on gorgonians, hydroids and other sessile marine invertebrates. Their overall body and limb morphology is relatively primitive, in many ways simi- lar to that of leucothoidean family Pleustidae. Such is expressed in the distinct rostrum, deep coxal plates, homopodous peraeopods, posterolobate coxae, and lanceolate uropod rami. However, morphological specializations including near-total loss of an acces- sory flagellum, modification of mouthparts for cam- ivory, and development of strongly subchelate gnatho- pods, are considered apomorphic. Members of the primitive genus Gitana are mainly arctic-boreal and deep water, those of the more ad- vanced genera Gitanopsis ( sens. str .L Amphilochopsis, and Amphilochus ( sens, str,) are actic-boreal and tem- perate, whereas those of the most advanced genera Hourstonius, Apolochus and Paramphilochus exhibit temperate, warm-temperate distributions in the north- ern hemisphere. These trends somewhat reflect the significance of higher phyletic classification in biogeo- graphical relationships of North American marine amphipod taxa (Bousfield 2001). In this scenario, the most primitive higher taxa tend to occur mainly in Arctic waters, secondarily along the Pacific coast, and the most advanced taxa in the North Atlantic and Gulf regions. This phenomenon may reflect two major long-term evolutionary factors. Firstly, morphologi- cal evolution proceeds at a higher rate at higher ambient temperatures, and conversely at a lower rate at lower temperatures. 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Gitana domin— ica, a new species from the Caribbean Sea (Am- phipoda: Amphilochidae). Proc. Biol. Soc. Wash. 103(3): 617-623. Thomson, W. 1880. A new species of Crustacea from New Zealand. Ann. Mag. Nat. Hist, ser .5, (6): 1-6, pi. 1. Yamato, S. 1993. A new amphilochid species (Crust- acea: Amphipoda: Amphilochidae) from a spiny lobster. Publ. Seto Mar. Biol Lab., 36 (l/2):99- 106, 3 figs. AMPHIPACIHCA VOL. 3 NO. 1. MAY 16,2001 28 Legend for Figures A1 antenna 1 A2 antenna 2 ACC FL accessory flagellum ABD abdomen BrPL brood lamella CX coxal plate EP abd. side plate GN1 gnathopod 1 GN2 gnathopod 2 HD head LFT left LL lower lip (labium) MD - mandible MX1 - maxilla 1 MX2 - maxilla 2 MXPD - maxilliped P5-7 - peraeopods 5, 6, 7 PLP - pleopod RT - right SP - spine T - tel son U - uropod cf - male 9 - female AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 29 The Amphipod genus Anisogam mams (Gammaroidea: Anisogammaridae) on the Pacific coast of North America. E. L. Bousfield, Research Associate Royal Ontario Museum, Toronto, ON, Canada MSS 2C6. ABSTRACT The genus Anisogammarus Derzhavin (sens, lat.) (Amphipoda: Gammaroidea: Anisogammaridae) is represented in the eastern North Pacific coastal marine region by A. pugettensis pugettensis (Dana), A. slatteryi , n. sp., A. epistomus, n. sp., and A. amchitkana , n. sp. Anisogammarus tvetkovae , new species, occurs in the northwestern Sea of Japan and Okhotsk Sea, along with the western Pacific subspecies A. pugettensis dybovskyi Derzhavin. Anisogammarids are free-living omnivores, occurring in the shallow sublittoral of cold-water, high salinity coastlines. The large coxal gills , each bearing accessory gills, are presumably advantageous for survival in partly anoxic habitats where they commonly occur. Introduction The first anisogammarid species was described un- der the name Gammarus pugettensis by J. D. Dana (1853) from material collected in Puget Sound by the U. S. Exploring Expedition. Common regional amphi- pods described by Stimpson (1857) contained the sole anisogammarid, Gammarus confervicolus . Stebbing (1906) summarized records of four Pacific aniso- gammarid species under various names within Gammarus (sens, lat.), mostlywi thin family Gammar- idae (sens. lat.). However, Stebbing (loc. cit.) assigned Dana's "Gammarus pugettensis" ,a\so\hted by Holmes (1904), to genus Liljeborgia. During the first half of the 20th century, few aniso- gammarids were recorded from the North American Pacific coast. Barnard ( 1954) more fully illustrated Dana’s Anisogammarus pugettensis based on extensive collections from Oregon, and Shoemaker (1955) de- scribed A. macginitiei from Pt. Barrow , Alaska. Exten- sive amphipod material from British Columbia and adjacent regions was collected by National Museum of Canada marine biological expeditions during 1955- 1980 (see below). Mainly from this material, the author (1979, 1981) described and illustrated a num- ber of new anisogammarid genera and species. Aniso- gammarus macginitiei was also transferred to the new genus Barrowgammarus where is used as an outgroup in later analysis (p. 45). Some of these, and earlier records, are embodied in the general faunistic guides and catalogues of Ricketts & Calvin (1968), Barnard (1975), Austin (1985), and Staude (1987). Gurjanova ( 1951 ) summarized early work on west- ern Pacific anisogammarids, updated by the compre- hensive study of Tzvetkova ( 1975) and the world-wide compilation of Barnard & Barnard (1983). Ishimaru (1994) summarized earlier records from Japan. The present study attempts to provide a more com- plete analysis of the systematics and distributional ecology of North American Pacific species of Aniso- gammarus. Acknowledgments For assistance with field work the author is greatly indebted to regional marine laboratories, notably he Pacific Biological Station, Nanaimo, B. C., the Pacific Environmental Institute, West Vancouver, B. C., the Bamfield Marine Station, B. C., and the Friday Harbor Laboratories, WA. Much of the detail has been acknowedged in previous station lists (Bousfield 1958, 1963, 1968; Bousfield & Jarrett 1981; Bousfield & McAllister 1962). Dr. Craig P. Staude, Dr. Colin B. Levings, the late Dr. Josephine F. L. Hart, and the late Dr. Dan B. Quayle were especially helpful with the field collecting. Drs. Peter Slattery and Charles E. O'Clair provided valuable study material from the Bering Sea and Aleutian Islands regions. Dr. Nina Tzvetkova, Zoological Museum, St. Petersburg, pro- vided helpful commentary and loan of valuable west- ern North Pacific material. Dr. Chang Bae Kim, Seoul National University, kindly provided commentary and figures on new material from South Korea, The original taxonomic work was performed as a staff member of the National Museum of Natural Sci- ences at the Hoi ly Lane Laboratory, Ottawa, ON, 1 979- 1983. Original line illustrations were prepared with the capable assistance of artist Roy E. Zittin, Cupertino, CA. The author is grateful to staff of the Canadian Mu- seum of Nature, especially to Ed Hendrycks for assist- ance i n preparation of si ide mounts and commentary on Anisogammarus epistomus, and to collections mana- ger Judith Price for help in cataloguing and labeling. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 30 SYSTEMATICS Anisogammar idae Bousfield Anisogammaridae Bousfield 1977: 295;— Bousfield 1979: 307 (+key to genera);— 1981:72-76,figs. 1-4;— : 259;— 1983:267;— Barnard &Karaman 1991: 1 14;— Ishimaru 1994: 46;— Bousfield & Shih 1994: 129. Ani sogammarids: Barnard & Barnard 1983: 582 (+ key to genera). Gammaridae (part): Stebbing 1906: 364;— Gurjanova 1951: 760;— Barnard, 1969 (part): 242;— Tzvetkova 1975 (part): 30. Type Genus: Anisogammarus Derzhavin, 1927: 8. Genera: Barrowgammarus Bousfield, 1979: 321; Eogammarus Birstein, 1933: 149; Spinulogammarus Tzvetkova, 1972: 954; Spasskogammarus Bousfield, 1979: 332; Locustogammarus Bousfield, 1979: 322; Jesogammarus Bousfield, 1979: 335; Ramello- gammarus Bousfield, 1979: 337; Carineogammarus Bousfield, 1979: 343. Diagnosis: Head, rostrum very short; inferior antennal sinus large, occasionally with narrow posterior notch. Antennae medium, subequal, accessory flagellum prominent, Antenna 2, peduncle large, flagellum occa- sionally calceolate. Mouthparts regular, little modified. Lower lip, inner lobes variously developed. Mandible: left lacinia 5-dentate; spine-row strong. Peraeon dorsally smooth. Coxal plates 1-4 medium deep, regular; plates 5 & 6 shallowly anterolobate. Gnathopods powerfully subchelate (male); gnatho- pod 1 larger than 2; palmar margins bearing peg spines (male), simple or pectinate (female); carpus short, lobe small. Peraeopods 5-7, bases weakly heterpodous; dactyls short. Peraeopods 2-7 with large coxal gills, 2- 5 with 2, P6 with 3, and P7 with 1-2 accessory gills. Pleosome and urosome variously dorsally carinate, toothed, spinose, or smooth. Uropods 1 & 2, rami usu- ally short, linear, spinose. Uropod 3 large, sub- aequiramous, terminal segment small. Tel son bilobate, with marginal and apical spines. Female brood plates large, unequal, with numerous long marginal setae. Remarks: During amplexus, males position them- selves dorsally and grasp the anterior margin of coxal plate 4, typically by means of the dactyl and propod of gnathopod 1 (Bousfield & Shih 1994). Ailometric growth changes are often noticeable. Compared with the adult stage, juveniles tend to be armed with fewer but relatively large spines and dorsal tooth of urosome 2, and the inner ramus of uropod 3 is relatively short. Anisogammarus Derzhavin AnwogammarwsDerzhavin, 1927:8;— Gurjanova 1951 : 776;— Tzvetkova 1975: 94 (part, + key to species);— Barnard & Barnard 1983: 584; — Ishimaru 1994: 46. Type species: Gammarus pugettensis Dana, 1853 Species: Anisogammarus pugettensis dybovskyi Der- zhavin, 1927; A. slattery i n. sp. (p. 34); A. epistomus n. sp. (p. 36); A. amchitkana , n. sp. (p. 39); A. tzvetkovae , n. sp. (p. 41). Diagnosis: Anterior head lobe acute above, rounded below, with shallow lateral notch. Eye medium, reniform. Antenna 1 shorter than 2; accessory flagel- lum well developed. Antenna 2, peduncle stout, seg- ment 5 shorter than 4, often with clusters of fine spines with extended tips (male); flagellum lacking calceoli (male). Mouthparts basic, with few modifications. Lower lip, inner lobes incompletely developed. Mandible: left lacinia 5-dentate; palp slender, terminal segment with well developed " D" spines and "E" setae, but only one cluster of "A" setae (of Cole 1 980). Maxilla 1 , palp 2-segmented. Maxilla 2, inner plate with full row of oblique facial setae. Maxilliped, inner plate with 3 apical spine teeth; palp setose, not raptorial. Coxae 1-4 medium deep, smooth, rounded below; coxae 5 & 6 weakly anterolobate. Gnathopod 1 (male) larger than 2 but usually similar in form; in the female it is much larger and of different form than gnathopod 2; carpus short; propodal palm (male) with peg-spines variously developed. Peraeopods 3 and 4, segment 5 short, dactyl s short. Peraeopods 5-7, bases broadened proximally, weakly heteropodous; segment 5 longer than 4; peraeopod 7 not longer than 6. Coxal gills on peraeopods 2-5 and 7 each with 2 and on peraeopod 6 with 3, linear accessory gills. Female brood lamellae broad, with numerous long simple marginal setae. Pleosome smooth above, lacking spines or setae; Epimeral plate 3, hind comer quadrate or slightly produced. Pleopods regular, outer ramus basally with split-tipped “clothespin” spines. Urosome 1 with mid- dorsal hump and 3 groups of spines; uropod 2 with AMPHI PACIFICA VOL. 3 NO. 1. MAY 16, 2001 31 KEY TO SPECIES OF ANISOGAMMARUS AND BARROWGAMMARUS 1 . Antenna 1 and 2 subequal in length; peraeopods 5-7, bases sublinear, not broadened posteriorly; urosome segments 1 & 2 lacking dorso-lateral spines or teeth; uropods 1 & 2, rami lanceolate, unarmed Barrowgammarus macginitiei (p. 44) Antenna 1 distinctly shorter than antenna 2; peraeopods 5-7, bases broadened proximo-poster- iorly; urosome segment 1 with dorsolateral spines; uropods 1 & 2, rami linear, with marginal and apical spines 2. 2. Gnathopod 2 (c? ) small, weakly subchelate, as in 9; uropod 3 (cP ,9), rami with marginal spines, lacking setae; telson short, length not greater than width A. amchitkana (p. 39) Gnathopod 2 (d 1 ) large, subsimilar to gnathopod 1; uropod 3, margins of rami with spines and setae; telson normal, length distinctly longer than basal width 3. 3. Gnathopods 1 & 2 (cf ), propodal palmar margins with heavy blunt peg spines; mandibular palp elongate; peraeopod 4, segment 4 elongate ~2X segment 5 A. tzvetkovae (p. 41) Gnathopods (<$ ), spines of palmar margins regularpeg spines, tips not broadened ; mandibular palp normal; peraeopod 4, length of segment 4 ~ 1.5 X segment 5 4. 4. Antenna 1, peduncular segment 2 short, length <1/2 segment 1; epimeral plate 3, hind comer squared or slightly acuminate; urosome 1 with 1-2 weak dorsolateral spines; mandibular palp segment 3 short, “D” spines enlarging distally 5. Antenna 1, peduncular segment 2 normal, length >1/2 segment 1; epimeral plate 3, hind comer acute, produced; urosome segment 1 with 3-4 medium strong dorsolateral spines; mandibular palp segment 3 regular, “D” spines of uniform size throughout 6. 5. Coxae 1-3, lower margin richly armed with longish setae; uropod 3 inner ramus markedly shorter than (2/3 length of) outer ramus; telson. distal marginal cluster with one very large,elongate spine (>2X length of other spine) A. slatteryi (p. 34) Coxae 1-3, lower margin weakly armed with short to medium setae; uropod 3, inner ramus large, length >3/4 outer ramus; telson, distal marginal cluster of spines not markedly unequal in size, (longest <2X other spines) A. e pis tom us (p. 36) 6. Antenna 1 (c? ), peduncular segment 5 with scattered clusters of slender, tip-extended spines; uro- pod 3, outer ramus broad nearly straight A. pugettensis pugettensis (p. 31) Antenna 1 (cf ), peduncular segment 5 with clusters of slender spines in distinct rows; uropod 3, outer ramus slender, medio-distally curved A. pugettensis dybovskyi (p. 34) acute mid-dorsal tooth, and weak postero-lateral cusp on each side. Uropods 1 & 2 short, stout. Uropod 3 subequally biramous, margins spi nose and /or plumose- setose; outer ramus with short terminal segment. Lobes of telson each with two groups of lateral spines. Distribution: Panboreal North Pacific, in algae, mainly on sedimentary bottoms, low intertidal to ~30 m. Anisogammarus pugettensis pugettensis (Dana) (Fig. 1 ) Gammarus pugettensis Dana, 1853: 957, fig. 1;— Holmes 1904; 239. Anisogammaru$?ugettensisGur}anova. 1 95 1 (part) : 777, fig. 541;— Barnard 1954: 13, pis. 12- 14; — Tzvetkova 1 975 (part) : 98, fig. 35; — Bousfield 1979:310 (key);- Bousfield 1982: 72, fig. 1;— Barnard & Barnard 1983 (part): 584, fig. 38; — Austin 1 985: 607; — Staude 1 987: 383. Material Examined: More than 600 specimens in 99 100 lots. ALASKA. SE Alaska, ELB Stns., 1961 (see Bousfield & McAllister, 1962): A5, Tongass Narrows, near Ketchikan - Cf ( 17 mm) ; lot #2 - 9 ov ( 1 1 .5 mm), fig’d specimens, CMNC 1980-0053; A7, AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 32 BostwickBay, June 2/61- 9 (br. II) (14.5 mm), slide mounts, CMNC 1980-0084; Lot#2 - 2 $$); Lot#3 (18cfcT);lot #4-599 ov,CMNC 1980-0087; CMNC 1980-0092; CMNC ^980-0094; All (100 &<3 & 99 ov, small im); Lot #2 ( 9, lim);Lot#3 (<$, im);A12(22c?d\33 9 9 ov,5im),Lot#2 - l9ov);A18(lim);A16(lim);A20 (Id*, l9ov, 2im); A25 ( 9, 2 im) ; All ( 100 large <$& (to 15.5 mm) ; All (Lot #2- 12 99 ov); A30 (4 99 ov, 4 im); A33 (5 99 ov, 3 juv); A34(5d'C?,499 ov, 5im);A37(ld*);A43(l9ov), Lot #2- 12d*cf, 1599, 20 im; #3 - 14d’d\3399ov, 15im);A48 (1 99 ov); A54(l juv); A55(l cf, 19, Lot #2 - 12 im); A65 (Id 1 ); A67 ( 1 9 , 14 juv) ; A71 ( 12 C?C?, 20 99, 14 im; lot #2 -30im); A73(12d’d’,999ov, 5im);Lot#2(8d'd\ 1499, 2 im) ; A 8 1 ( 10 C?C?, 7 99 ov, 30 im) ; A83 (3 tfc?, 10 99 ov, 17 juv); A 84 (60 tfc?, 99 ov large); A86 (Icf); A88 (3cfd\ 12 99, 2 im); A93 (24 im); A 136 (1 d\ 13 im); A 139 (3 CfC? 5 99, 5im);A140(9d’d', 15 99, 30im&juv); lot #2(1 9,3 im); A 141 (lcf); A153 (1 im); A 171 (2 juv); A174(l d\ small); Lot # 2 (25 adult, 17 juv). ELB Stns., 1980 (see Bousfield & Jarrett 1981): S14L1 (3 imm). BRITISH COLUMBIA: Queen Charlotte Islands, ELB Stns., 1957 (see Bousfield 1963): H5 (6 cfd\ 8 99, 12 im); N4 (2 juv). North Central Coast, ELB Stns., 1964 (see Bousfield 1968): H5 (1 juv); H13 (1 9 ov, 35 im); H16 (2c?d\ 3 99, 12 im); H17 ( 1 9, 12 im); HI 8 ( 1 juv); H39 ( 1 juv); H50 (2 im); H56 (4 cfd\ 10 99, 13 juv); Lot #2 (8 im). Pearl Harbour, nr. Prince Rupert, silty sand, eel grass, LW, D. E McAllister, June 23/65 - ld\ Northern Vancouver Island, ELB Stns., 1959 (see Bous- field 1963): V 17 (7d > cf, 10 99 ov, 1 1 im); V 18 ( 1 d 1 , 3 im); V22 (75 spms., mostly large 99 ov); lot #2(1 d\ gnathopods dissected); N 18 (lc?, 4 99, 17 juv). Southern Vancouver island, ELB Stns. 1955 (see Bousfield 1958): F3 ( 1 im) ; F4 (9 ov, 13 im) ; F6 ( 1 d\ 43 99, 5 im, dried); M2 (1 im); M2 (1 Cf, 2 99); M5 (29 im & juv); G4 (3 juv); G10 (5 im & juv); G1 1 (8 im & juv); G15 (1 d\ 3 99, 2 im). 1970 (for Stns. of 1970-80, see Bousfield & Jarrett 1981): P717 (sev. im). 1975: Friiday Harbor, May /75 - 2 CfC? (20 mm). 1976: Pacific Environmental Institute, West Vancouver, in halibut tank, June 8 - Id 1 (18 mm). 1977: B2 7 99, 5 im); B7a .Willis Beach, Oak Pt„ Viet- oria. May 19 - 18 cfd\ 30 99 subad (16 mm), slide mount, CMNC 1980-0029; CMNC 1980-0038; CMNC 1980-0039; E2 (2 99 ov). Misc. CMN collections: Ladysmith Hbr., Vancouver I., B. C., D.B.Quayle coll., June 8/38 - 27 d’d' , 99 ov (broken specimens); Satuma I., Bruce Bight, B. C., J.F.L. Carl coll., Aug. 26/55 - 2 9 9 ov; Ibid. , night light over kelp, Aug 24/55 - 1 d\ 1 9, 2 juv; Porpoise Hbr, B. C., 20 m, M. Waldichuk coll.. Sept. 24/64 - 3 99; Ibid., Sept. 18/62 - 3 cfcf, 7 99 ov, 1 im, NMNS Cat. No. 6-90; Nass Hbr., Iceberg Bay, on dead fish in trap, S. Gorham coll., June 20/65 - 7 c?d\ 4 99 ov., 2 im Off Cordova, Orca Inlet, Prince William Sound, SE Alaska, 13 m dredge, K. E. Conlan, Feb. 18/89 - d\ 9 mating pair. WASH.-ORE, USA. ELB Stns., 1955 (see Bousfield, 1958): F8, Garrison Bay, San Juan I., 9 6 99, 4 im. ELB Stns., 1966 (see Bousfield & Jarrett, 1981): W3 (1 d\ 1 9, 10im);W4(l im); W5(l juv); W7, Meadow Point, Puget Sound, July 17. - d* (14.0mm); 9ov.(ll mm), slide mounts, CMNC 1980-0065; Ibid.. Lot #2-20 spms; W10 (4d*cr,7 99ov, 5im); Wll (1 juv); W 18(1 juv); W22 ( 1 im); W33 (d\ 10 im); W39 (6 juv); IhisL (Lot #2 - 3 d’d 1 , 7 99, 35 im); W44 (3 im); W69 (9 ov). Diagnosis Male (16 mm): Anterior head lobe slightly incised. Eye medium, sub-reniform. Antenna 1, peduncular segment 2 medium, length 1/2 peduncle 1 ; flagellum ~20- segmented, little exceeding peduncle of antenna 2. Antenna 2, peduncular segment 5=4, with few clusters of tip-extended slender spines; flagellum 17-segmented, shorter than peduncle. Mandibular spine row with 8-9 blades; palp rela- tively short; segment 3 > 2/3 segment 2, "D" spines uniform, extending 2/3 of inner margin; segment 2, beta and gamma setae very short. Maxilla 1 , palp little broadening distally. Maxilliped, inner plate apically truncate, outer plate little broadened; palp segment 3 regular, length > 1/2 segment 2. Gnathopods 1 & 2 stout, dactyls with short unguis; Gnathopod 1, palmar angle with 8-10 inner and outer rows of simple spines. Gnathopod 2, propodal postero- distal angle with inner submarginal row of 6 short simple spines. Peraeopods 3 & 4, segment 6 relatively AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 33 Fig. 1. Anisogammarus pugettensis pugettensis (Dana), c? (17 mm); Q ov. (11.5 mm). Tongass Narrows, SE alaska. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 34 short, arched, little longer than segment 5. Peraeopod 7, basis with slight postero-distal marginal excavation. Coxal gill on peraeopod 7 relatively large, broad. Epimeral plates 2 and 3, hind comer acute, distinctly produced. Urosome 1, mid-dorsal hump medium, with cluster of 8-10 medium spines; lateral clusters with 3- 4 spines. Urosome 2 with strong median tooth and single postero-dorsal cusp on each side. Urosome 3 with mid-dorsal and dorsolateral clusters of 2-3 me- dium spines. Uropods 1 & 2, rami shorter than pedun- cles, margins moderately spinose. Uropod 3, outer ramus medium broad, inner margin plumose-setose, slightly but distinctly longer than slender inner ramus; terminal segment short. Telson lobes medium, each side with proximolateral group of three spines, and distolateral longish marginal spine. Female ov. (14 mm). Gnathopod 1, propod relatively large, subquadrate, posterodistal angle with groups of 3 inner, and 5-6 outer submarginal simple spines. Gnathopod 2, propod subrectangular, postero-distal angle with submarginal row of 1 simple and 4 pectinate spines; brood plate large, broad, with numerous mar- ginal setae. Uropod 3, rami shorter than in male, inner margin plumose-setose. Distributional Ecology: Aleutian Islands and S. E. Alaska, through B. C. and Washington state south to Coos Bay, Oregon, and Northern California, low inter- tidal to subtidal, in Ulva and Enteromorpha, and in partially anoxic bottom deposits of wood chips (Waldichuk & Bousfield 1962). Remarks: A very similar form has been recorded under this name from the northern Sea of Japan and Sea of Okhotsk by Gurjanova (1951) andTzvetkova ( 1 975) . Anisogammarus pugettensis (Dana), subsp. dybovskyi Derzhavin (fig. 2) Anisogammarus dybovsky Derzhavin, 1927: 8 ;— Stephensen 1944: 47, figs. 10. 11; — Ishimuru 1994 (part): 46. Gammarus pribiloffensis Pearse, 1913: 571, fig. 1. Anisogammarus pugettensis Gurjanova 1951 (part): 777, fig. 541;— Tzvetkova 1975 (part): 99, fig. 35;- Ishimaru 1994 (part): 46. Material Examined: 3 lots from East Kamchatka, USSR, K. Vinogradov coll., 1933 -cf (13 mm), slide mount; 2cfcf (13 mm); 9°v. (11 mm), slide mount (identified as A. pribilofensis by E.F. Gurjanova, 1933), Zoological Museum cotins., St. Peters- burg, Russia. Alaska-Bering Sea P. Slattery coll: Mukmuk Bay, St. Lawrence I., 40 ft. scoop, July 1/83 - 3 99 ov, 3 juv, IZ 1989-002. NE St. Lawrence I., July/83 - 1 Cf (18.5 mm). Unimak I, P. Slattery, June-Oct/82 - 2 cf Cf , 5 99 - Diagnosis: Male (16 mm). Very similar to Anisogam- marus p. pugettensis (Dana, 1853) but differing in the following features: Eye large, reniform. Antenna 2 (male), peduncle 5 subequal to 4, with numerous groups of tip-extended slender spines. Mandibular palp, segment 3 with few- er "A" and "E" setae. Coxa 1 more strongly setose below. Gnathop;od 1, propodal palmar spines shorter ?ind thicker, apex more blunt ; carpus, posterior lobe narrow, subacute. Peraeopod 7, posterodistal marginal excavation lined with fine setae; segment 6 with a few clusters of longish setae, rather than clusters of short spines. Epimeral plates 2 & 3, hind corner less strongly produced. Coxal gill 7 small and short relative to coxal gill of peraeopod 6. Uropod 3, outer ramus relatively narrow, length 4X width, curved distomedially. Tel- son lobes each with pair of distolateral short spines. Distributional Ecology: Western Pacific coastal ma- rine waters, northern Japan Sea and Sea of Okhotsk to werstemBering Sea , mostly along open coasts, onsandy and silty substrata, from lower intertidal to depths of 280 m (Tzvetova, 1975); waters around Japan (Ishi- maru, 1994); animals scavenge drowned dead human bodies (Kosek et al 1962). Remarks: This species has been synonymized with A. pugettensis , originally described from the eastern Pa- cific by Dana, 1853. However, sufficient differences exist (above, and key) as to distinguish the two forms at subspecies level. Anisogammarus s lottery i, n. sp. (fig. 3) Anisogammarus sp. 1, Austin, 1985: 607. Material Examined: ALASKA: St. Lawrence I., Bering Sea, 6 m sand, P. Slattery coll., June 6/87 - 7 juv (2-4 mm) ; Ibid- , lot #2- 7 juv (2-4 mm). Ibid. , Lot #3. - 40 juveniles, CMN collections. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 35 Fig. 2. Anisogammarus pugettensis dybovskyi Derzhavin. 2/3 segment 2; segment 3, "D" spines in short row, increasing in length distally. Maxilla 1, palp slightly narrowing distally. Maxilliped, inner plate apically oblique, outer plate little broad- ened, with long apical pectinate setae; palp segment 3 short, length ~ 1/2 segment 2. Coxae 2-4, lower margins with 8-10 mainly longish setae. Gnathopod 1 very stout, distinctly larger than gnathopod 2; dactyls with short unguis. Gnathopod 1, palmar angle with inner and outer submarginai rows of 6-7 and 3-4 simple spines respectively. Gnathopod 2, palmar angle with inner submarginal rows of 4 and 2 short simple spines respectively. Peraeopods 3 & 4, segment 6 straight, longer than segment 5. Coxae 3- 4 distinctly anterolobate. Peraeopods 5-7, bases dis- tinctly heteropodous; peraeopod 6 slightly the longest. Peraeopods 6 & 7, basis with slight postero-distal marginal excavation. Coxal gill on peraeopod 7 large, about equal in size to that of peraeopod 6. Epimeral plate 3 , hind comer squared. Urosome 1, mid-dorsal hump very low, with 1-2 small spines and weak lateral clusters of 2 spines. Urosome 2 with small median tooth, posterodorsal cusps lacking. Urosome 3 with single mid-dorsal and dorsolateral medium spines. Uropods 1 & 2, rami shorter than peduncles, outer ramus of uropod 2 lacking marginal spines. Uropod 3, outer ramus short, medium broad, margins spinose; terminal segmentdistinct;innerramus short, ~ 1/2 outer ramus, inner margin withfew plumose setae. Telson lobes short, each side with proximolateral group of three spines, and distolateral pair of unequal spines. Female ov. (6.5 mm). Antenna 2, flagellum 10-seg- mented. Gnathopod 1 medium large, subquadrate, spination of posterodistal angle similar to that of male. Gnathopod 2, propod short, subrectangular, postero- distal angle with inner submarginal row of 3 simple, outer row of 4-5 pectinate spines. Brood plate large, broad, but with relatively few (<30) marginal setae. Uropod 3, rami shorter than in male, margins spinose , with a few simple setae. Etymology; The name recognizes marine biologist Dr. Peter F. Slattery, who has contributed broadly to knowledge of marine benthic communities on the Pa- cific coast of North America. Distribution: Bering Sea south through Vancouver I. to Washington State, LW and subtidally, to depths of ~13 m, on sand and in feeding pits of the gray whale, Eschrichtius robustus. Remarks: The species is very similar to A. epistomus but differs mainly in its smaller size, normally un- produced epistome, and other character states of the key (p. 31). The small subadult female from Lulu I.,has mark- edly unequal rami of uropod 3, and long coxal setae. Although the small specimen from Leadbetter Pt. was not dissected, it exhibits some characteristics of A. slattery /, including a small mid-dorsal tooth on urosome 2, and relatively large and powerful gnathopods. The inner ramus of uropod 3 is relatively short and thin. Anisogammarus epistomus , n. sp. (Figs. 4, 4A) Anisogammarus sp. 2, Austin, 1985: 607. Material Examined. BRITISH COLUMBIA: Southern Vancouver I., ELB Stns. 1955: P6a, Long Beach, SE end Wickaninnish Bay, under algal debris on sand, LW, Aug. 2 -C? (13.0 mm), Allotype, (slide mount), CMNC 2001-0010; 1 1/2 segment 2. Gnathopods 1 & 2 large, strong; dactyls with short unguis. Gnathopod 1, palmar angle with 8- 10 inner and outer rows of simple spines. Gnathopod 2, propodal postero-distal angle with inner submarginal row of 6 short simple spines. Peraeopods 3 & 4, segment 6 relatively short, arched, little longer than segment 5. Peraeopod 7, basis with slight post-erodistal marginal excavation. Coxal gill on peraeo-pod 7 relatively large, broad. Epimeral plates 2 and 3 , hind comer acute, slightly produced. Urosome 1, mid-dorsal humpmedium,with cluster of 8-10 medium spines; lateral clusters with 3- 4 spines. Urosome 2, with strong median tooth and single postero-dorsal cusps on each side. Urosome 3, with mid-dorsal and dorsolateral clusters of 2-3 me- dium spines. Uropods 1 & 2, rami shorter than pedun- cles, margins moderately spinose. Uropod 3, outer ramus medium broad, inner margin plumose-setose, slightly but distinctly longer than slender inner ramus; terminal segment short. Tel son lobes medium, each side with proximolat- eral group of three spines, and distolateral longish marginal spine. Male (13.0 mm). Allotype: Antenna 1 elongate, flag- ellum of 22 segments; accessory flagellum 6-segmen- ted. Antenna 2, peduncular segment 5 with few sur- fac-ial clusters of slender spines. Upper lip, epistome prominently bulging anteriorly. Mandibular palp with 5 "A" setae. Coxae 2-4, lower margins nearly bare, armed spars- ely with short setae. Gnathopod 1, propod and dactyl powerful, propodal palmar spines regular, tips little or not broadened. Gnathopod 2, propod much less pow- erful, similar in form and armature to that of female but slightly more powerful. Peraeopod5,basisvery broad, hindmarginrounded. Uropod 3, rami subequal in length; outer ramus with 9- 1 1 groups of spines, inner margin distally plumose- setose; inner ramus, inner margin with spines and setae; terminal segment very short. Telson, lobes normal, longer than basal width. Etymology: From "epi" + " stomum ", alluding to the large epistome protruding over the upper lip. AMPHIPACIHCA VOL. 3 NO. 1. MAY 16. 2001 40 MXPD*' \ Fig. 5. Anisogammarus amchitkana n. sp. C? (15 mm), Holotype; $ov (14.0 mm), Allotype. Cyril Cove, Amchitka, Alaska. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 41 Distributional Ecology: Known only from the outer coast of Vancouver Island, from Pachena Bay to Wickaninnish Beach; under algal debris, on open surf- exposed sand, LW level, probably shallow subtidal. Remarks: Character states of Anisogammarus epi- stomus, especially of the male, suggest that the species is a member of the pugettensis group, with weakly developed palmar peg spines, but closer to A. amchit- kana , having weakly developed gnathopod 2 and large aequiramus uropod 3. Anisogammarus amchitkana, n. sp. (Fig. 5) Material Examined. Bering Sea- Alaska region: Lot #1 , Square Bay, Cyril Cove, Amchitka, Aleutian Islands, C. E. O’ Clair coll.. Mar. 24, 1969 - C? (15 mm), Holotype, slide mount, CMNC2001-0015; 1 9 ov. Allotype (14 mm), slide mount, CMNC2001-0016; 5 99 ov,Paratypes (19 ov, dissected), CMN collns. Constantine Harbor, Amchitka 1., among algae on dock pilings, P. Slattery coll., Sept. 7. 1969 - 1 C? (12 mm), CMN Acc. No. 1982-79. St. Lawrence I., SE Cape, in kelp & mysid swarms, P. Slattery coll., June 6/86 - 54 subadult specimens (3-5 mm), CMN collns. Kialegak camp, SW St. Lawrence Bay, Aug. 25, 1985 - 1 9 br. II (11 mm) + 499 itti (8-10 mm), CMN collns - Diagnosis. Male (15 mm): Anterior head lobe dis- tinctly incised. Eye medium large, subreniform. An- tenna 1, peduncular segment 2 medium, length >1/2 peduncle 1; flagellum ~20-24-segmented, little ex- ceeding peduncle of antenna 2. Antenna 2, pedunc- ular segment 5 = 4, with few clusters of tip-extended slender spines; flagellum ~20-segmented, nearly as long as peduncle., Mandibuar spine row with 7 blades; palp short, segment 3 > 2/3 segment 2. Maxilla 1, palp slightly broadening distally. Maxilliped, inner plate apically truncate, outer plate slightly broadened; palp segment 3 regular, length > 1/2 segment 2. Coxae 1-4 medium deep, lower margins rounded, weakly setulose. Gnathopods 1 & 2 very unequal in size; gnathopod 1 large, powerfully subchelate, gnath- opod 2 weakly subchelate, as in female; dactyls with short unguis. Gnathopod 1, palmar angle with inner and outer rows of 8-12 mostly peg spines, inner row respectively. Peraeopods 3 & 4, segment 6 relatively short, arched, litle longer than segment 5. Coxae 5 & 6 shallowly anterolobate. Peraeopods 5-7, bases weakly heteropodous. Peraeopod 7, basis relatively narrow, posterodistal margin straight. Coxal gill on peraeopod 7 small, narrow relative to gill on peraeo- pod 6. Epimeral plates 2 and 3, hind comers weakly acute. Urosome 1, mid-dorsal hump low, with cluster of 8-10 medium spines; lateral clusters each with 3-4 spines. Urosome 2 with ordinary median tooth and single posterodorsal cusps oneach side. Urosome 3 with mid- dorsal and dorsolateral clusters of 2-3 medium spines. Uropods 1 & 2, rami subequal in length to peduncles, margins moderately spinose. Uropod 3, outer ramus medium broad, margins with 6-7 clusters of short spines, distinctly longer than slender spinose inner ramus; terminal segment short. Telson lobes short, basally broad,each side with proximolateral group of three spines, and distolateral single short marginal spine. Female ov (14 mm). Gnathopod 1, propod relatively small, subovate, posterodistal angle with groups of 5 inner, and 4 outer submarginal simple spines. Gnath- opod 2, propod subrectangular, posterodistal angle with submarginal row of 1 simple and 4 pectinate spines; brood plate large, broad, with numerous mar- ginal setae. Uropod 3, rami shorter and broader than m male, margins spinose. Etymology: The species name acknowledges the type locality on the Aleutian Island of Amchitka. Distributional Ecology: Amchitka and Aleutian Is- lands, from LW intertidal to depths of ~10 m. Remarks: The small body size, relatively large size of both gnathopods 1 & 2 (male), and presence of more strongly developed propodal palmar peg spines re- move amchitkana from the A. pugettenis - slatteryi complex (see Fig. 8). Anisogammarus tzvetkovae, n. sp. (Fig. 6) Anisogammarus possjeticus Tzvetkova , 1975 (part?). extending well up palm. Gnathopod 2, propodal sub- rectangular, posterodistal angle with inner and outer submarginal rows of 5 and 6 short simple spines. Material Examined: Peter-the-Great Bay, Russia, LW in- tertidal, Nina L. Tzvetkova coll. - C? (24. 5 mm), Holotype; 9 ov. (18.0 mm), Allotype; slide mounts, loan material. Zoological Institute, St. Petersburg, Russia. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 42 Fig. 6. Amsogammarus tzvetkovae, n. sp. cf (24.5 mm), Holotype; 9 ov (18 mm) Allotype Peter-the-Great Bay, Sea of Japan. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 43 P6 Fig. 7. Barrowgammarus macgimtiei (Shoemaker, 1955). 1/2 segment 2, "D" spines short, extending proximad of facial cluster of "A” setae; segment 2 with 8-10 long alpha setae. Maxilla 1, palp distinctly broadening distally, apex with 7-8 short spines. Maxilliped, inner plate apically truncate, outer plate tall, broadened; palp segment 3 regular, length ~ 2/3 segment 2. Coxae 1-4 relatively deep, 1-3 narrow, lower margin with afew short setae. Gnathopods 1 & 2 stout; dactyls with short unguis. Gnathopod 1, pal mar margin rugose, file-like, posterior angle with 6-8 inner, medial, and outer rows of stout peg-spines; carpus with short nar- row posterior lobe. Gnathopod 2, propodal postero- distal angle with inner and outer submarginal row of 4- 6 short peg spines. Peraeopods 3 & 4, segment 4 elongate, segments 5 & 6 relatively short, subequal; dactyls very short. Coxae5&6shalIowIyanterolobate. Peraeopods 5-7, bases relatively narrow, little broad- ened; basis of peraeopod 7 with slight postero-distal marginal excavation. Coxal gill on peraeopod 7 large, broad, deep, nearly as large as gill of peraeopod 6. Epimeral plates 2 and 3, hind comer minutely acute, lower margins spinose, Urosome 1 , mid-dorsal hump large, with mid-dorsal “V” of -20 stout spines; dorso- lateral clusters each with 3-4 spines. Urosome 2 with strong median tooth and single posterodorsal short spines on each side. Urosome 3 with mid-dorsal and dorsolateral clusters of 2-4 medium spines. Uropods 1 & 2, rami shorter than peduncles, margins spinose. Uropod3, rami large, slender, inner slightly the shorter, margins with 8-10 clusters of short spines and setae; terminal segment small. Telson lobes medium long, narrowing distally, proximolateral and distolateral spines short. Female ov (18 mm). Gnathopod 1, propod medium targe, subrectangular, posterodistal angle with groups of 5 inner, and 4 outer submarginal simple spines. Gnathopod 2, propod slender, elongate-rectangular, posterodistal angle with inner submarginal row of 2 simple and 6-7 pectinate spines, and outer submarginal row of 1 simple spine and 5-6 pectinate spines. Brood plate on peraeopod 2 relatively small and slender, with ~30 longish marginal setae. Uropod 3, rami shorter than in male, and margins less setose. Etymology: In recogniton of Dr. Nina L. Tzvetkova, Zoological Institute, St. Petersburg, Russia, who has contributed in an outstanding manner to knowledge of gammaroi deans and littoral marine amphipoda of the northwestern Pacific Ocean. Distributional Ecology: Known only from the coasts of North and South Korea, the northwestern coast of Japan, and Peter-the-Great Bay, Russia. Remarks: Material and illustrations from the east coast of South Korea, kindly supplied by Dr. Chang Bae Kim in 1992 (pers. commun.) is virtually identical with that of Dr. Nina Tzvetkova from Peter-the-Great Bay (above). Barrowgammarus Bousfield Barrowgammarus Bousfield, 1979: 321;— Barnard & Barnard 1983: 586. Diagnosis: Body veiy large. Eyes small, oval. Anten- nae subequal in length, sparesly setose, notcalceolate. Mouthparts poorly described. Mandibular palp, segment 3 slender, "D" setae uniformly short, extend- ing proximally to distal group of "A" setae. Gnathopods powerfully subchelate, propodal palmar margins with peg spines (male). Peraeopods 5-7, bases little expanded, each with distinct posterodistal lobes; dactyls short. Coxal gills 2-5 with paired acces- sory gills; coxal gills 6 & 7 with single accessory gills. Pleon smooth above. Urosome segments 1 and 2 each with prominent mid-dorsal tooth. Uropods 1 & 2, rami lanceolate, lacking marginal spines. Uropod 3, inner ramus short, <1/2 outer ramus, inner margins of both are plumose-setose; terminal segment distinct. Telson lobes narrowing distally, fused basally. Barrowgammarus macginitiei (Shoemaker) (Fig. 7) Anisogammarus macginitiei Shoemaker, 1 955: 54, fig. 16;— Tzvetkova 1975: 103, fig. 37. Barrowgammarus macginitiei Bousfield 1979: 321; Barnard & Barnard 1983: 586. AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 45 s e 3 3 go GO 3 1 3d 5 •i»* si. '1- § 3 so s BQ CL < in 24 P. -A. INDEX T Fig. 8. Morphological smilarities and possible phyletic relationships among species of Anisogammarus and Barrowgammarus. Diagnosis: With the characters of the genus. Distribution: Beaufort Sea, Okhotsk Sea, sublittoral. Remarks: This monotypic taxon is undoubtedly a member of family Anisogammaridae, most closely related to the genus Anisogammarus , and is included here as an outgroup . The female i s about the same size as the male but has not been described in detail. The presence of dorsal protruberances on urosome seg- ments 1 & 2,and of dorsal armature on the pleosome, the form of the gnathopods (male) and the inaequi- ramous form of uropod 3 suggest a common ancestry with Anisogammarus pugettensis. Calceolation of antenna 2 (male) has not been confirmed. Discussion. The present treatment of anisogammarid species utilizes a semi-phyletic modification of the UPGMA system of Sneath and Sokal (1973), as in previous analyses of other North Pacific amphi pd taxa. Charac- ter states are ordered plesio-apomorphically and the relative phyletic placement of a given taxon is repre- sented by a numerical sum of plesiomorphic, interme- diate, and apomorphic character state values (0, 1, and 2, respectively) in a PI esio- Apomorphic (P.-A.) Index. Tabular data on which the resulting phenograms are based are considered overly bulky and repetitive for publication here, but can be supplied on request. Fig. 8 portrays character state similarities within the North Pacific genus Anisogammarus and the selected AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 46 outgroup species Barrowgammarus mac ginitiei (Shoe- maker). The outgroup species, formerly Anisogam- marus macginitiei, is now recognized at the generic level, Barrow gammar us, a decision in agreement with the "less than 50% similarity" that it here demonstrates with the six other species and subspecies of Anisogammarus. The two subspecies of A. pugetten- sis, and the species A. amchitkana and A. epistomus cluster above the 75% similarity level and these two fuse at the 68% level. The specialized sand-dwelling species A. slatteryi (P.-A. Index = 17) and the primit- ive western Pacific species A. tzvetkovae (P.-A. index = 12) join these at levels of 62% and 58% similarity respectively. Positive consideration of elevating some taxa to subgeneric rank might be justified. However, since other regional species may await formal recogni- tion, elevation of taxa at this time seems premature. The known species of Anisogammarus are cold- temperate (boreal) North Pacific in biogeographic af- finity. They are included in an updated list of N. American amphipod species on which comparative biogeogrographical studies were also based (Bousfield, 200 1 ). Four species are apparently excl usi vely North American, ranging from the Bering Sea region south- ward to northern California. Two species range from the western Bering Sea and Kamchatka peninsula, southward along the Asiatic coast to the northern Sea of Japan, consistent with the penetration of cold-water elements into that region (Derzhavin 1930). The distri- bution of only one full species, Barrow gammar us macginitiei , apparently overlaps the central Bering Sea divisional region. The biogeographic separation into eastern and western species groups appears to match the east-west distributional separation of species with- in other anisogammarid genera, notably the species- rich and more southerly ranging genus Eogammarus (Tzvetkova 1975; Bousfield 1979). However, cogni- zant of the current lack of a fossil record and other evidence of past distributions, reasons for these bio- geographical consistencies "across the anisogammarid taxonomic board" remain speculative. References: Austin, W.C. 1985. Amphipoda. In: An annotated checklist of marine invertebrates in the cold temp- erate Northeast Pacific. Khoyatan Marine Lab 3: 588-623. Barnard, J. L. 1954. Marine Amphipoda of Oregon. Oregon State Monogr., Studies Zool. 8: 9-36,9pls. Barnard, J. L. 1969. The families and genera of mar- ine gammaridean Amphipoda. Bull. U. S. Nat’I. Mus. 271: 1-535, 173 figs. Barnard, J. L. 1975. Amphipoda: Gammaridea. pp. 313-366, pis. 70-85. In R. 1. Smith & J. T. Carlton (eds). Light’s Manual: Intertidal Invertebrates of the Central California Coast, 3rd ed. Univ. California Press: 716 pp.. Barnard, J. L., &C.M. Barnard, 1883. FWamphipods of the World. Vols. 1 & 2: 830 pp., 50figs. 7 graphs, 98 maps, 12 tables. Mt. Vernon, VA. Hayfield Associates. Barnard, J. L.,&G. S . Karaman 199 1. The families and genera of marine gammaridean Amphipoda (except marine gammaroids). Part 2. Rec. Australian Mus. Suppl. 13 (Parts 1 & 2): 866 pp., 133 fig. Birstein, J. A. 1933. MaJacostraca der Kutais-Hohlen am Rion (Transkaukasien, Georgien). Zool. Anz., Bd 104: 143-156, 24 figs. Bousfield, E. L., 1958. Ecological Investigations on shore invertebrates of the Pacific coast of Canada. Nat’I Mus. Can. Bull. 147 : 104-115. Bousfield, E. L., 1963. Investigations on sea-shore invertebrates of the Pacific coast of Canada, 1957 and 1959. 1. Station List. Nat’I Mus. Can. Bull. 185 : 72-89. Bousfield, E. L. 1968. Studies on littoral marine inver tebrates of the Pacific coast of Canada, 1964. 1. Station List. Nat’ 1. Mus. Can. Bull. 223: 49-57. Bousfield, E. L. 1977. A new look at the sytematics of gammaroideanamphipodsof the world. Crustaceana Suppl. 4:282-316. Bousfield, E. L. 1979. The amphipod superfamily Gammaroidea in the northeastern Pacific region: systematics and distributional ecology. Bull. Biol. Soc. Washington 3: 297-359, 12 figs. Bousfield, E. L. 1981. Evolution in North Pacific Marine Amphipod Crustaceans . in G.G.E. Scudder & J. L. Reveal (eds.), Evolution Today. Proc. 2nd Intemat. Congr. Syst. Evol. Biol.: 69-89. 18 figs. Bousfield, E. L. 1982. Amphipoda: Gammaridea. pp. 254-285. in Synopsis and Classification of Living Organisms. S.B. Parker (ed.). McGraw-Hill, New York, Vol. 2.: 254-285; 293-294. Bousfield, E. L. 1983. An updated phyletic classifica- tion and palaeohi story of the Amphipoda. Crustac- ean Issues. A. A. Balkema, Rotterdam. 1: 257-278. Bousfield, E. L. 2001. Phyletic classification as applied to amphipod crustaceans of North America (north of Mexico). Amphipacifica 3(1): 49-1 19. Bousfield, E. L. & N. E. Jarrett 1981. Station lists of marine biological expeditions of the National Mus- eum of Natural Sciences in the North American Pacific coastal region, 1966 to 1980. Syllogeus 34 1 - 66 . AMPHIPACIFTCA VOL. 3 NO. 1. MAY 16, 2001 47 Bousfield, E. L, & Shih, C.-t. 1994. The phyletic classification of amphipod crustaceans: problems in resolution. Amphipacifica 1 (3): 76-134. Dana, J. D. 1853. Crustacea. Part II. United States Exploring Expedition 14: 689-1618, atlas of 96 pis. Cole, G. A., 1980. The mandibular palps of North American freshwater species of Gammarus. Crustaceana, Suppl. 6: 67-83, 4 figs. Derzhavin, A. N. 1927. Gammaridae. Kamchatka Ex- pedition, 1908-1909. Hydrobiol. Jour. SSSR, 6(1- 2): 1-15. [in Russian]. Derzhavin, A. N. 1930. Arctic elements in the fauna of the peracarids of the Sea of Japan. Hydrobiol. Jour. SSSR, 8 (10-12): 326-329 [in Russian]. Gurjanova, E. F. 1951. Bokoplavy moreii SSSR i sopredel'nykh vod (Amphipoda-Gammaridea). Akad. Nauk SSSR, Opred. po Faune SSSR 41: 1029 pp, 705 figs. Holmes, S. J. 1904. Amphipod crustaceans of the ex- pedition. Harriman Alaska Expedition: 233—246, figs. 118-128. Ishimaru, S. 1994. A catalogue of gammaridean and ingolfiellidean Amphipoda recorded from the vic- inity of Japan. Rept, Sado Mar. Biol. Sta. 24: 1-86. Koseki, T., S. Yamanouchi, and K. Nagata, 1962. The post-mortem injury in the drowned dead body at tacked by amphipods, Med. Biol 64 (3): 74-76 (in Japanese). Pearse, A. S. 1913. Note on a small collection of Amphipoda from the Pribilof Islands with descript- ions of new species, Proc. U. S. Nat’l. Mus. 45: 571- 573. Ricketts, E., & J. Calvin 1968. Between Pacific Tides (4th ed.). Stanford University Press: 614 pp. Shoemaker, C. R. 1955. Amphipoda collected at the Arctic Laboratory, Office of Naval Research, Point Barrow, Alaska, by G. E. MacGinitie. Smiths. Misc. Coll. 128(1): 1-78, 20 figs. Sneath, P. H. A., & R. R. Sokal 1973. Numerical Taxonomy. W. H. Freeman & Co., San Francisco. 573 pp. Staude, C. P. 1987. Amphipoda Gammaridea. pp. 346- 391. In Kozloff, A.(ed.). Marine invertebrates of the Pacific Northwest. Univ. Wash. Press: 51 1 pp. Stebbing, T. R. R. 1906. Amphipoda I. Gammaridea. Das Tierreich: 1-806, figs. 1-127. Stephensen, K., 1944. Some Japanese Amphipods. Vidensk. Medd. Dansk Naturh. Foren. Bd 108: 25- 99. Stimpson,W. 1857. The Crustacea and Echinodermata of the Pacific shores of North America. J. Boston Soc. Nat. Hist. 6: 1-92, pis. 18-23. Tzvetkova, N. L. 1972. K sistematike rode Gammarus Fabr. i novye vidy bokoplavov (Amphipoda, Gam maridea) iz se vero-zapadnoi chasti Tikhogo Okeana. Akad. Nauk SSSR, Trud. Zool. Inst., 52: 201-222, 7 figs. Tzvetkova, N. L. 1975. Littoral gammarid amphipods of the northern and far-eastern seas of the USSR and surrounding waters (in Russian). Izdat. Nauka, Akad. Nauk SSSR: 256 pp., 89 figs. Waldichuk, M., & E.L. Bousfield 1962. Amphipods in low -oxygen waters adjacent to a sulphite pulp mill J. Fish. Res. Board Can. 19 (6): 1 163-1165. Legend for Figures A1 antenna 1 A2 - antenna 2 ACC FL - accessory flagellum ABD - abdomen BR - brood lamella CX - coxal plate EP - abdominal side plate EPIST - epistome GN1 - gnathopod 1 GN2 - gnathopod 2 HD - head LFT - left LL - lower lip (labium) MD - mandible MX1 - maxilla 1 MX2 maxilla 2 MXPD - maxilliped P5-7 - peraeopods 5, 6, 7 PL - pieopod PLP - palp RT - right SP - spine T telson U - uropod UL - upper lip (labrum) UROS - urosome X - enlarged (S - male 9 - female im - immature juv - juvenile OV - ovigerous subad. - subadult AMPHIPACIFICA VOL. 3 NO. 1. MAY 16, 2001 48 The Canadian Field-Naturalist: 120 years of northern biodiversity publication. The Canadian Field-Naturalist publishes articles and notes on original research and observations on natural history relevant to Canada (therefore on northern portions of both Nearctic and Palaearctic regions) including distribution, faunal analyses, taxonomy, ecology, and behaviour, and items of news, comment, tributes, review papers, book reviews and new titles. The official publication of The Ottawa Field-Naturalists’ Club, it prints minutes ofthe annual meeting and awards presented by the Club. Since 1984, it has featured edited Status Reports for many individual species designated by the Convention on Species of Endangered Wildlife in Canada (COSEWIC), particularly those on fish and marine mammals. As well, recent special issues have featured the history of botanical investigation of Canada, St. Pierre et Miquelon, and Greenland; a biography of the Canadian ornithologist Percy A. Taverner, a history of the Canadian Wildlife Service, and an analysis of the Orchids of the Ottawa district. The Ottawa Field-Naturalists’ Club was formed in 1879 by scientists from embryonic federal departments, including the Geological Survey and the Dominion Experimental Farm, together with leading amateurs of the time; a similar mix remains as its strength to this day. The Club quickly emphasized publication, and for seven years beginning in 1880, it annually issued the Transactions of the Ottawa Field-Naturalists’ Club. With volume 3 in 1887, the Transactions became a subtitle for Volume 1 of The Ottawa Naturalist, a new monthly publication. With Volume 3 of The Ottawa Naturalist in 1889 emphasis shiftedfrom largely 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 ofthe Transactions, although this subtitle was soon omitted) . The issues per year were gradually reduced from 12 to 9 to 6 and, eventually, to 4; the latter beginning with Volume 67 in 1953, but the individual issue size increased. The annual pages published reached a record of 794 in 1988 (volume 102) and 1994 (volume 1 12). The largest single issue 254 pages was, however, published in 1996 as 1 10( 1). Since 1967, the Club has separately published a local (Ottawa area) natural history journal. Trail & Landscape, now also issued 4 times a year. Submissions to The Canadian Field-Naturalist and its predecessors have been peer reviewed since its inception, first through a “Publishing Committee”, later “Sub-editors”, and then “Assistant Editors” until the present designation Associate Editors was adopted in 1885. Currently, most submissions also go to at least one (often more) additional reviewers). Associate Editors are listed in every issue and, since 1982, additional reviewers been acknowledged individually in the Editor’s Report annually. A formal publication policy was published in The Canadian Field-Naturalist 97(2): 231-234. “Advice to Contributors” is publ-ished in one or more issues annually giving format and charges. The journal is entirely supported through a portion of club membership (40%), subscriptions (100%), annual interest on Club investment funds (80%) and publication charges for pages and reprints. The Ottawa Field- Naturalists’ Club at the beginning of 2000 had 957 members and The Canadian Field-Naturalist an additional 253 individual and 497 institutional subscribers in 22 countries, for a distribution of 1707 copies. The current annual subscription rate is $28 for individuals and $45 for institutions. Membership in The Ottawa Field-Naturalists’ Club (which includes receipt of The Canadian Field-Naturalist) is $28 for individ-uals and $30 for families. Postage outside Canada is $5.00 additional. Subscriptions should be sent toThe Canadian Field-Naturalist, Box 35069 Westgate P.0., Ottawa, Ontario, Canada K 1Z 1 A2. Manuscripts for consideration should be addressed to Dr. Francis R. Cook, Editor, Canadian Field- Naturalist, RR 3, North Augusta, Ontario K0G 1R0, Canada. Francis R. Cook Editor, Canadian Field-Naturalist fcook@achilles.net AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 49 An updated commentary on phyletic classification of the amphipod Crustacea and its applicability to the North American fauna. Edward L. Bousfield, PhD, FRSC, Research Associate Royal Ontario Museum, 100 Queen's Park, Toronto, ON K2B 8E3. ABSTRACT Bousfield & Shih (1994, Amphipacifica 1(3):76-134) provided a phyletic classification of the Amphipoda consistent with superf amity-level standards of classification in use for the Hyperiidea, Caprellidea, Ingolfiellidea, and Gammaridea. For gammaridean amphipods, the basis for phyletic classification is reproductive form and behaviour. Detailed character-state analyses support the view that the ancestral amphipod was a “swimmer-clinger”, rather than a benthic “crawler-burrower”. This study comments on difficulties posed to moiphological classification by near- universal occurrence of homoplasy within major character states. Thepresent phyletic classification is here applied to a list of ~ 1650 scientific names of amphipod crustaceans from marine, freshwater and terrestrial habitats of North America (north ofMexico), updated to the end of the 20th century. Character state variation of antennal callynophore, brush setae, calceoli, uropods, and telson, and sexual dimorphism of gnathopods are further analysed. Suborders and gammaridean superfamilies are phyletically classified and annotated in tabular form. Although phyletic classifica- tion is presently controversial, alternative or more suitable phyletic groupings proposed by cladistic and/or rDNA analyses are yet lacking or unproven. Broad acceptance and/or usage of gammaridean superfamilies (or equivalents) outlined here provide demonstrably greater meaning and functionality to taxonomic interrelationships, and therefore greater research credibility than simple alphabetical listings of families and genera. INTRODUCTION Classification is the naming of essentially discreet groups of living organisms in a manner that reflects their probably correct phylogenetic hi story. Develop- ment of a classification requires input by scientists who are knowledgeable in animal systematics, and experi- enced in recognition of the significance of morphologi- cal characters and the probably correct ordering of the character states within the group concerned. Ideally, classification discriminates true phyletic relationships from homoplasious (artifical, convergent) similarities. Phyletic classification is thus distinct from, and far more useful than, an alphabetical listing of previously described taxa. If the Darwinian theory of evolution is essentially correct for multi -cellular organisms, it follows that amphipod crustaceans evolved in only one manner, and left only one biohistorical "track record". As a coroll- ary to that thesis, all species were atone time or another linked by so-called "intermediate" forms which, espe- cially if extant, tend to mask the "clean" separation of lineages into pragmatically distinct clades or higher taxonomic groupings. For several reasons, however, phylogenists are unlikely to discover that record pre- cisely. These factors include: (1) lack of a significant (long-term) amphipod fossil record (not earlier than Cenozoic); (2) incomplete description of extant taxa, especially of species from hypogean waters and the deep sea; and (3) a relatively undeveloped state of broadly applicable phyletic analysis. Clues to natural relationships are provided mainly by analysis of exter- nal and internal morphology, behaviour, physiology, and distributional ecology of extant species. Methods of phyletic analysis, whether intrinsic, phenetic, cladistic, genetic, or in combination, require careful research input. Particlarly in treatment of speciose higher-level taxa, methodologies to date have proven neither "infallible", nor "guaranteed" to provide a realistic, credible result. Thus, in cladistic analysis, prior choice of ingroup/outgroup taxa, selection of numbers and kinds of morphological characters, and ordering of character states, all constitute subjective (and fallible) decisions that directly effect the quality of the results. Thus, sheer numbers of characters and character states, if inappropriately selected and/or wrongly ordered, may produce results that are actually misleading, internally conflicting, or otherwise of low credibility, particularly when compared with results employing other methodologies. Nor can a correct result be assumed because of the “sophistication” of methodology or computerized format. The main text of this paper was first presented at the 10th International Colloquium on Amphipoda held at Heraklion, Crete, April 16-21, 2000. The purpose of the work is to review the status of phyletic classifica- tion of the Amphipoda, and demonstrate its applicabil- ity to a recently compiled list of amphipod families, genera, and species recorded to date from the North American continent north of Mexico. AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 50 ACKNOWLEDGMENTS The author is grateful to many colleagues for their stimulating taxonomic and phyletic input, especially Dr. Michael J. Thurston (deep-water taxa), Dr. Jorgen Berge (stegocephalid reclassification), Dr. J. Emmett Duffy (genetic methodologies). Dr. C.-t. Shih (hyper- iid classification), and Dr. J. K. Lowry (lysianassoid and podocerid phyogeny). Compilation of the list of North American amphi- pods (Appendix I) involved the services of many col- leagues, co-ordinated through a Committee on Scien- tific and Common Names of Invertebrate Animals (CNIA), chaired by Dr. Donna D. Turgeon, NOAA, Washington, DC. A full list of contributors is to be included in a final CNIA report and publication. Espe- cially helpful to the author, who served as amphipod subcommittee chairperson and project co-ordinator, have the been the following amphipod systematists: Dr. Pierre Brunei, Universite de Montreal, (Atlantic gammarideans); Dr. Donald B. Cadien, Marine Biol- ogy Lab, Carson, CA (SW Pacific gammarideans); Dr. Kathleen E. Conlan and E. A. Hendrycks, Canadian Museum of Nature, Ottawa, ON (Arctic and Pacific gammarideans); Dr. John Foster, Panama City, FL (Hyperiidea, Gulf of Mexico); Stephen Grabe, Envi- ronmental Protection Commission, Tampa, FL (Gam- maridea, Ingolfiellidea: Gulf of Mexico); Dr. John R. Hoi singer. Old Dominion University, Norfolk, VA (freshwater amphipods); Diana R. Laubitz, Canadian Museum of Nature, Ottawa, ON (Caprellidea); Sara E. LeCroy, Gulf Coast Research Laboratory, Ocean Spri ngs,MI(Gammaridea: Gulf ofMexico) ;Dr. Chiang- tai Shih, Fisheries Research Institue, Taiwan (Hyperiidea); and Dr. Craig P. Staude, Friday Harbor Laboratories, WA (Pacific Gammaridea). CLASSIFICATORY SYSTEMATICS. The malacostracan order Amphipoda has long been considered an especially difficult problem of phyletic classification (Riley 1983; Schram 1986). The prob- lem of internal classification of this ordinal crustacean group is complicated by extreme diversity of body form ranging from thick-bodied spiny-legged burrow- ing haustoriids; big-eyed fast-swimming oceanic hy- periids; slender-bodied skeleton shrimps, to eyeless, vermiform infaunal ingolfiellids. How might we find commonality of relationships among widely diverse external pigmentation, from the pure white of burrow- ing phoxocephaloideans, through beautifully cryptic maculation of "swash-zone" pontogeneiids and calliopiids, to the vertical striping of odiids and multivariate pigmentation of "thick nosed" pleustids and minute commensal stenothoids? What natural ordering, if any, might exist between such diverse feed- ing types as free-swimming predaceous eusiroideans and pardaliscodeans, longicomiculate trypton-feeding podocerids, and vertically tube-building ampeliscids? Phyletic classification seeks to provide answers to these questions and bring a semblance of natural order out of almost chaotic diversity of form and behaviour. The history of development of amphipod class- ificatory systems has been outlined by Bousfield and Shih (1994) and is briefly summarized here. In es- sence, during a period of taxonomic discovery lasting approximately two centuries since the time of Linneus (1758), phyletic (superfamily-level) classifications fi- nally came into standard use for the Hyperiidea through theworkofBowman&Gruner( 1973), for Ingolfiellidea by Stock ( 1 977) , and for Caprellidea notabl y by Vassil- enko (1974) and D. R. Laubitz (1993). Within the diverse and taxonomically more difficult suborder Gammaridea, however, the story is more complex. For nearly two centuries (to the mid- 1950’s) gammaridean classification had been essentially phyletic, stabilized by the semi-phyletic, non-alpha- betical arrangements of families proposed by Sars (1895) and Stebbing(1906). This system was broadly accepted and utilized by amphipod systematists at least until the early fifties (e.g.. Shoemaker 1930; Gurjanova 1951 ; Dunbar 1954). However, two major weaknesses in these classifications remained: (1) several large families such as "Gammaridae" and "Lysianassidae" were weakly defined, effectively polyphyletic, or oth- erwise "unwieldy", and (2) other, mostly smaller fami- lies "begged" for inclusion within higher "umbrella" categories that would recognize their close phyletic similarities. In the second instance, Bulycheva (1957) proposed the super-family name Talitroidea to encom- passthenaturaIlyrelatedfamiliesHya!idae,HyalelIidae, and Talitridae. J. L. Barnard (1973) combined a num- ber of domicolous families within superfamily Corophioidea. In the first instance, the formal task if unravellingfamily-level units within polyphyletic fam- ily "Gammaridae" was initiated mainly by Bousfield (1973, 1977). Recombination within superfamily categories, of several older family names and those newly proposed, soon culminated in a fully phyletic classification of suborder Gammaridea (Bousfield 1979, 1982a, 1983). This classification was adopted to various degrees by Riley (1983), Schram (1986), and Ishimaru (1994). Some superfamily concepts were also revised and expanded by others [e.g., Crang- AMPHIPACIFICA VOL. 3 NO. 1 MAY 16, 2001. 51 onyctoidea by Holsinger 1992a; Lysianassoidea by Lowry and Stoddart 1997). As updated by Bousfield and Shih (1994), the "new" phyletic classification proved basically not unlike the semi-phyletic family "arrangement" of Sars (1895) and Stebbing (1906), since both recent and older systems were presumably based on si milar conceptual ordering of character states of reproductive morphology and behaviour. In the interim, however, J. L. Barnard had become dissatisfied with perceived anomalies of the Sars-Steb- bing classification and the apparent intractability of their ready solution. Although he informall y diagram- med suggested relationships between known amphi- pod families, based on a "Gammarus- like" prototype, he commenced listing gammaridean families and gen- era in alphabetical sequence ( 1958, 1969) . The prag- matics of a simple alphabetical treatment of higher gammaridean taxa, thenapproaching lOOfamily names, was soon widely adopted. In further updatings and expansions of these original compendia (Barnard & Barnard 1983; Barnard &Karaman 1991), a number of anglicized concepts of some higher groups were pro- posed. These included the names "gammaridans", "hadzioids", etc., and later (Williams & Barnard 1988) "crangonyctoids", as well as a broadening of some original formal family-level concepts (e.g., Eusiridae, Corophiidae). Notably perhaps, these names corre- sponded, with about 75% similarity, to superfamily concepts formally proposed earlier in the phyletic literature. However, with Gordan Karaman ( 1 99 1 , p. 7), Barnard steered away from formal phyletic classi- fication and concluded this final major work with an alphabetical listing of all families and component gen- era. During the past two decades, some major regional faunistic studies have utilized mainly alphabetical list- ings and retained older treatments of higher taxa such as "Gammaridae" (e.g., Ruffo el &L 1982,1988, 1993, 1998; Camp( 1998). However, with increasing sophis- tication of cladistic analytical methodology (e.g., Lowry& Myers, in prep.), earlier supeifamily concepts are now being re-analysed [e.g., Serejo 2000 in press (Talitroidea); Berge and Vader 2000, in press (Stegocephaloidea)], and new superfamily taxa pro- posed (e.g., Iphimedioidea Lowry & Myers, 2000). In the light of recently proposed phyletic studies utilizing genetic methodology (e.g.. Shram, 2000; Macdonald 1999), a resumption of development of phyletic classi- fication of the gammaridean Amphipoda now seems promising. Character State Analyses As noted above, the present analysis of phyletic classification within the order Amphipoda is based mainly on reproductive morphology and behaviour, updated from earlier work (Bousfield & Shih, 1994). To some degree, modified repetition of material here compensates for the limited original circulation of that source paper, now out of print. The present analysis, however, utilizes only seven mostly reproductively significant, characters and character states. These include sensory organelles of the antennae (callyn- ophore, brush setae, and calceoli); form of the tel son, and degree of sexual dimorphism and use of the gnath- opods during amplexus. To these has been newly added the form of the rami of uropods 1 & 2. The character states vary widely and homoplasiously from group to group, as do those of the mouthparts, coxal plates, peraeopods, and uropod 3 of the earlier study. Nonetheless, collectively and judiciously , they provide a consistent and verifiable morphological basis for phyletic grouping of higher amphipod taxa. In general, the ordering of character states is based on an assumed plesiomorphic condition in more primi- tive "outgroup" members of the superorder Peracarida, such as the Mysidacea and Cumacea, and more primi- tive members (shrimp-like groups) within the Deca- poda. Thus, in members of phyletically primitive amphipod groups ( "swimmers "), the sensory organelles of the antennae are well developed, the telson is typi- cally bilobate, and sexual dimorphism of the gnatho- pods is rare or lacking. Since the mating process usually takes place in the open water column, precop- ulatory "holding" of the female by the male gnathopods is apparently not developed. Conversely, in members of phyletically more advanced gammaridean superfamilies ("crawlers”), the antennal sensory fea- tures are much reduced or lacking and the telson lobes are oftenfusedapically. Since mating usually occurs on (or in) the bottom substrata, often in strongly lotic waters, the male gnathopods are typically strongly modified for pre-amplectic grasping and holding of the female and/or agonistic behaviour with other males. The Antennal Callynophore The callynophore consists of a bundle of close-set aesthetases on the postero-medial margin of the fused (or conjoint) basal segments of the flagellum. This organelle occurs typically withinpelagic ordinal groups of the higher Malacostraca and, within the Amphipoda, characterizes superfamily groups of the “Natantia”, especially the Hyperiidea (Fig. Id). Its primary func- AMPHIPACIFICA VOL. Ill NO. 1. MAY 16, 2001. 52 Pacifoculodes bruneli A. LYSIANASSOIDEA (Hyperiopsidae) B. STEGOCEPHALOIDEA C. PARDALISCOIDEA D. HYPERIIDEA E. OEDICEROTOIDEA Brush setae F. AMPELISCOIDEA AmpeUsca sp. Fig. L Types of antennal callynophores [after Barnard (1969), Bowman & Gruner(1973), Bousfield & Chevrier (1996), and unattributed sources]. don is almost certainly chemosensory. Its presence mainly in the final adult male instar would seem to be of direct reproductive significance in the detection of females within the water column. However, in some lysianassoidean and synopioidean subgroups, callyn- ophore-like structures may also be present in mature females and subadult stages, perhaps indicating a pos- sible secondary role in detection of food resources. Representative forms of callynophores within the Amphipoda are illustrated in Fig. 1 . Lowry (1986) has described a one-field arrangement of the callynophore within families Platyischnopidae, Urothoidae and Phoxocephalidae (Phoxocephaloidea), a condition he considers primitive, and in some hyperiids (e.g.. Archaeoscinidae), perhaps convergently. In all other taxa the arrangment is two-field. The possible significance of the callynophore in phyletic classification was first introduced by Lincoln and Lowry (1984) and amplified formally by Lowry (1986). Although strongly developed in pelagic carni- vores and necrophages, especially where calceoli are weak or lacking (e.g., Synopioidea, Pardaliscoidea, Stegocephaloidea, and Hyperiidea), the organelle is generally weak or lacking in reproducti vely pelagic but vegetatively benthic groups such as the nestling Dex- aminoidea and tube-building Ampeliscoidea, and in the fossorial Phoxocephaloidea and Pontoporeioidea. It is virtually lacking in several “natant” subgroups AMPHIPACIHCA VOL. Ill NO. 1. MAY 16, 2001. 53 where the entire life cycle is essentially infaunal (e.g., Haustoriidae), or commensal or parasitic (e.g., some Lysianassoidea) and/or where preamplexing reproduc- ti ve behaviour has secondarily and convergently devel- oped (e.g.JParacalliopiidae and Exoedicerotidae within Oedicerotoidea). Curiously, the callynophore is sur- prisingly weakly developed in the mainly marine but mainly acalceolate family Oedicerotidae and even within the Eusiroidea (e.g., in the pelagic, primitive family Eusiridae, but not found in Pontogeneiidae, nor Calliopiidae). The callynophore is essentially lacking in reproduct- ively benthic Reptantia, including the Caprellidea and Ingolfiellidea, and notfound in freshwater taxa, even in those that have apparently become secondarily pelagic such as Macrohectopus within the Gammaroidea. However, callynophore-like structures have been re- ported from a few Amphilochidae (e.g. Austro- pheonoides , Peltocoxa ) and Cressidae ( Cressa crist- ata ) within primitive subgroups of superfamily Leucothoidea (Lowry 1986). The presence or absence of a callynophore may therefore offer a useful criterion of reproductive life style. Although its occurrence appears subject to homoplasious tendencies, such aberrancies may be correlated with non-reproductive features of life style and are thus predictable. In broader perspective, the presence of a callynophore is a plesiomorphic, or basic feature of malacostracan reproductive morphology. As concluded previously (Bousfield & Shih 1994), the callynophore provides a primary basisfor development of a phyletic classification within the Amphipoda. Antennal Brush setae The term "brush setae" applies to dense tufts or clusters of short brush-like setae that variously line the anterior margins of peduncular segments 3, 4, and 5 of antenna 2. Brush setae may occur also on the posterior (lower) margins of peduncular segments 1-3 of antenna 1 (e.g., in Dexaminoidea). Similar types of setae occur in other peracaridan taxa, including the Cumacea and Mysidacea. Within the Amphipoda these organelles have been found only in the terminal male stage of pelagicaliy reproductive amphipod superfamilies, and not in sub- adult males, females, and/or immature stages. Their function is yet unknown and conjectural. Although brush setae may not have been studied in ultrastructural detail, their gross morphology is similar to modified setae rather than thin-walled aesthetascs. Their role may be tactile when, during the process of copulation, the male is briefly in close contact with the female. The potential usefulness of brush setae in phyletic classification was previously suggested by Bousfield ( 1 979) ; Bousfield & Shih ( 1 994). These organelles are most strongly developed in non-calceolate primitive superfamilies of Natantia (e.g., Pardaliscoidea, Synopioidea), and moderately developed in some calceolate "natant" taxa (e.g., Lysianassoidea, Phoxocephaloidea, Eusiroidea, Oedicerotoidea), and acalceolate "transitional" super-families (e.g., Dexam- inoidea, Ampeliscoidea, and Mel-phidippoidea). They are less well developed or rare within the Stegocephaloidea and Hyperiidea (Fig. 1). The presence of brush setae in males only indicates that their function is reproductively significant. Their limited distribution within the Natantia and total ab- sence from the Reptantia indicates a potentially pri- mary value in phyletic classification. The Antennal Calceolus The calceolus is a slipper-shaped membranous microstructure attached variously to the anteromedial segmental margins of the flagella and peduncles of both antenna 1 (antennule) and antenna 2 of some gammaridean Amphipoda. Principal features of these micro- structures have been described, across a broad range of higher taxa, by Lincoln and Hurley (1981) and, with special reference to genera within the primitive “replant” superfamilies Crangonyctoidea and Gam- maroidea, by Godfrey etal( 1988). The calceolus is not to be confused with the aesthetasc, a sublinear thin-walled microstructure of mainly chemosensory function, found only on flagellar segments of antenna I in most species of Amphipoda. The aesthetasc also occurs widely across malacostracan ordinal subgroups, including the Decapoda. The calceolus is also structur- ally readily distinguishable from brush setae and other seta-like structures co-occurring on antennal pedunc- ular and flagellar segments. Representative types of amphipod calceoli are illus- trated in figs. 2 & 3. Calceol us-like structures are found on the proximal flagellar segments of antenna 1 (male) of a few other malacostracans, notably within the Syncarida (e.g., Koonunga cursor ) and the Mysidacea (e.g., Xenacanthomysis pseudomacropsis). Such structures are not considered calceoli by Lincoln (pers. commun.) since they may be convergent in form and/or of differentfunction. However, these organelles are included here as of possible phyletic significance within the Malacostraca and, in my view, merit further comparative micro-anatomical and behavioural study. AMPHIPACIHCA VOL. Ill NO. 1. MAY 16, 2001. 54 Fig. 2. Types of antennal calceoli in gammaridean Amphipoda, and positionally similar organelles in other malacostracan Crustacea (modified from Bousfield & Shih 1994). AMPHIPACIFICA VOL III NO. 1. MAY 16, 2001. 55 The presumed "advanced" form of the calceolus is grossly similar to that of a parabolic radar "dish" (Fig. 2C,D). Combined with its anterior antennal location, this morphology suggests that the organelle functions primarily as a mechanoreceptor for aquatic acoustical vibrations. However, its innervation and connection to the brain has not yet been ascertained, nor have micro- acoustical studies yet confirmed its true function (Lin- coln & Hurley 1981). The distribution of calceoli between the sexes sug- gests that calceoli developed initially in males only, presumably as a device for detection of vibrations from swimming females of its own species. In free-swim- ming raptors (e.g., Gammarellidae within the Eusir- oidea), special types of calceoli have apparently devel- oped in females and immatures, and occur alongside the reproductive ly functional form of calceolus in terminal stage males. As described by Steele & Steele (1993), these organelles appear to have became more complex structurally, presumably, and possibly sec- ondarily adapted, for detection of escape vibrations of free-swimming prey. However, the primary reproduc- tive function of calceoli apparently diminished or dis- appeared in concert with changes in life style from pelagic to benthic, neritic to abyssal, lotic to lentic, marine to freshwater, epigean to hypogean, and corre- sponding development of pre-amplexing gnathopods (see p. 61). As indicated in Fig. 3, reduction and disappearance of calceoli occurred initially in antenna 1 and subsequently in antenna 2. Within the latter, the sequence of loss was initially from the peduncle and distal flagellar segments, and finally from the proximal flagellar segments. However, as noted above, calceoli persisted in both males and females of some epigean freshwater groups (e.g., some Gammaridae, Anisogammaridae) and/or cave pool amphipods where life styles presumably remained free-swimming and raptorial (e.g., Crangonyx packardi and Sternophysinx calceola (Crangonyctoidea); Sensonator valentiensis (Melphidippoidea?), and several eusiroideans of south- ern continental land masses (Bousfield 1980). The possible significance of antennal calceoli in phyletic classification of the Amphipoda has been alluded to variously by Bousfield (1979, 1983), Lin- coln and Hurley (1981), Lincoln & Lowry (1984), and more recently by Godfrey gjtaL (1988), Stapleton el al ( 1988), Hoi singer (1992a), and Steele & Steele (1993). These views were analysed and expanded upon by Bousfield & Shih (1994) and are here summarized and updated, with special application to the North Ameri- can amphipod fauna (Appendix I). The external morphology of the calceolus within theprimitivereptantsuperfamilyCrangonyctoidea(cat- egory 9, Lincoln and Hurley 1981) appears to be the most simplified, and thus probably the most plesio- morphic extant form (Figs. 2 A & 3). It consists only of a basal stalk and elongate body that bears numerous (20+ ) elements of similar simple structure. Holsinger ( 1992a) has distinguished two subcategories of calceoli within the Crangonyctoidea. In members of holarctic fami 1 y Crangonyctidae (Crangonyx, Synurella, pp. 101- 104) the form is slender and elongate, with a simple branched internal “tree trunk” configuration. Some separation of basal elements in Crangonyx rich- mondensis , illustrated by Godfrey el aL (1988), are suggestive of "protoreceptacles". By contrast, the calceolus within austral families Sternophysingidae and Paramelitidae is typically broad, paddle-shaped, and its internal tree-trunk configuration has more nu- merous indistinct branches, a seemingly more plesiomorphic condition. In slightly more advanced types of calceoli (Fig. 3,upper: Phoxocephaloidea), the elements are fewer ( 10-15 in Platyischnopidae; 4-6 in Phoxocephalidae) and the body may be short and spatulate, or barrel -shaped. With respect to the sexes, the more plesiomorphic types of calceoli occur (with very few exceptions) in the 'males only' category of presumed most primitive superfamily taxa such as the Crangonyctoidea, Phoxocephaloidea, Pontoporeioidea, and most of the Lysianassoidea (Fig. 2, i, ii; Fig. 3, upper two rows). In more advanced types of calceoli (Fig. 2, iii), the basal element is broadened and forms a receptacle that is weakly developed in Pontoporeioidea and Gammar- oidea but strongly so in Eusiroidea (Fig. 2, iv). The basal stal k i s di stall y expanded i nto a bulla or resonator, weakly and more strongly in those same groups respec- tively. In some Pontoporeioidea (Bathyporeiidae), finger-like processes protrude over the proximal ele- ments. In the most advanced types of calceoli (viz., in some Eusiroidea: Gammarellidae, Eusiridae), and in some pelagic Lysianassoidea (e.g., Ichnopus spp., Lowry and Stoddart 1992), the distal elements are few andwidelyseparatedfromoneormorelarge,cup-shaped receptacles, and the bulla may be prominent. The evolutionary morphological sequence within calceoli portrayed here is believed to match more closely the phylogeny of corresponding superfamily groups, based on other character states (see below), than does the somewhat pragmatic sequence originally provided by Lincoln and Hurley ( 1981 ). A graphical plot of the types of calceoli and their neponnf>n o « - « h o « ». w o * n »» <= n o o t* > c * «« AMPHIPACIFICA VOL. Ill NO. 1. MAY 16, 2001. 56 MYSIDA A1 only OUT-GROUP • — I- \ PROTO- AMPHIPOD - d \ • PROTO-CRAMGONYCTID DISTRIBUTION OF CALCEOU A1 + A2 'N Extinct epigean ancestors LACKING - ingolfiellidea Austrogammarus^ PARAMELITIDAE _ STERNOPHYSiNGIDAE x , . CRANGONYCTOIDEA Extinct marine ancestors \ "V *— NEONIPHARplDAE , CRANGONYCTIDAE Other extinct marine amphipods PLATY1SC HNOPIDAE PHOXOCEPHALOIDEA — P Hny OCE PHV ID AETt- u " OTHOiDAE ^ i o 1°"^ HYPERIOPSIOAE VALETTIIDAE PONTOPOREIIDAE PONTOPOREIOIDEA - PriscUlina? I URIST1DAE ' [C YPHOCARIDIDAE y I d 'Qy, Psammonyx j C f |{Q loss 1 secondary) I lysianassoidea BATHYPOREIIDAE MESOG AMMARID AE I I ^ gammaroidea l v EUSIROIDEA • EUSiRIDAE/ OED|CEROTO | DEA J i ..N fpARALEPTAMPHOPIDAE Exoecttcerolcjes /\ / I \ ' PONTOGENEIIDAE PARACALLIOPIIDAE EXOEDiCEROTIDAE Paramoera I Lysianassa EOCRANGONYCTIDAE TALITROIDEA CHEIDAE CONDUKIIDAE ZOBRACHOIDAE Z UROHAUSTORilDAE s phoxocephalopsidaeI HAUSTORIIDAE ACANTHOG AMMARIDAE TYPHLOGAMMARIDAE MACROHECTOPIDAE GAMMAROPOREIIDAE GAMMARIDAE ANISOGAMMARIDAE BATEiDAE Bathyporeiapus Metoediceros \ OEDiCEROTIDAE Notoediceros\ Patuki X NIPHARGIDAE PHREATOG AMMARIDAE | MELPHID1PPOIDEA bogidielloidea HADZIOIDEA COROPHIOIDEA CAPRELLIDEA LEUCOTHOIDEA STENOTHOIDEA(?) IPHIMEDIOIDEA "" DEXAMINOIDEA -SYNOPIOIDEA# \ ^ # HYPERIIDEA \ AMPELISCOIDEA PARDALISCOIDEA • N JEBORG 1 O.DEA • STEGOCEPHALOIDEA _________ CALLYNOPHORE • Strongly developed / present in most species « Moderately well developed / present frequently • Weakly developed / present in few species Fig. 3. Suggested phylogenetic relationships within the Amphipoda based on distribution of the 8 calceoli on the antenna and between the sexes (modified from Bousfield & Shih 1994). AMPHIPACIFICA VOL III NO. 1. MAY 16, 2001. 57 distribution by antennal site, sex, and higher taxon, can be linked by means of a branching arrangement with relationships that, in part, are remarkably similar to phyletic arrangements derived elsewhere from analy- sis of other character states (Fig. 3). In the first two categories, this arrangement goes somewhat beyond the relationships proposed by Lincoln & Lowry ( 1 984) on the basis of the taxonomic (classificatory) distribu- tion of calceoli. In the present chart, the positions of the major taxa in the various “boxes” are correlated prima- rily with the distribution (or lack) of calceoli on one or other (or both) antennae, along the horizontal axis and withthe morphological typeandits sexual occurrences the vertical axis. The vertical and horizontal axes also simulate, fanwise, an approximate evolutionary time scale for the probable frst appearance of the ancestral type of each major taxonomic group. The arrangement of calceoli is here rooted in a presumed mysid-like out-group in which calceol us-like structures are known, at least on antenna 1 of the male. Such structures may have occurred in presumed former epigean and pelagic marine ancestors of the now hypogean relict suborder Ingolfiellidea, and of the con- tinental freshwater-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 from earlier considerations (e.g., Bousfield 1982b). In this two-dimensional scheme, all members of the seven calceolate superfamilies, and the enigmatic hypogean calceolate Sensonator valentien - sis Notenboom, 1986 (Melphidippoidea?), cannot be confined cleanly within any given graphical box. Such variance is attributable to parallel development, diver- sification, and subsequent loss of calceoli from the an- tenna of both sexes, presumably in response to chang- ing life styles within the various taxonomic subgroups (above). Notably, the more strongly calceolate super- family groups (calceoli on both A1 and A2, left col- umn) are those in which members are primarily pelagic and/or reproduce freely in the water column. These include most Phoxocephaloidea, Pontoporeioidea, Lysianassoidea, Eusiroidea, and Oedicerotoidea. The less strongly calceolate superfamilies (with rare excep- tions, calceoli on A2 only, right column) are found in the most primitive members of benthic superfamilies of the Reptantia (Crangonyctoidea, Gammaroidea). The position of acalceolate superfamilies is tentative, but is suggested partly by the presence or absence of other presumably plesiomorphic, often vestigial characters such as antennal callynophore and brush setae (above). The presence or absence and type of antennal calceolus are character states of undoubted phyletic significance. However, their restricted distribution among extant gammaridean superfamilies limits their use to cases of phyletic classification where other parameters of broader classificatory applicability (e.g, form of uropods, coxal plates, gnathopods) are known. Uropods 1 & 2. The uropods of amphipods are biramous append- ages of the three posterior abdominal segments. They function mainly in forward propulsion during swim- ming or crawling activities. The uropods are well developed and conspicuous in most gammarideans, hyperiideans, and ingolfiellideans, but minute or lack- ing in caprellideans. The rami are seldon equal in size, the outer usually being noticeably the shorter. Only within the Ingolfiellidea is uropod 2 typically larger than uropod 1. Morphological variation in the rami of uropod 3 and its utilization in phyletic classification have been analyzed previously (Bousfield and Shih 1994). In this study, the form and armature of the rami of uropods 1 & 2 are similarly investigated. In nektonic forms, the rami are often lamellate or lanceolate, whereas in benthonic crawling or burrowing forms the rami are typically styliform (Schram 1986). The rami may also be modified for specialized functions in domicolous and/or commensal species, and for presumed copula- tion (in males) widely across the taxonomic spectrum (e. g., in some Lysianassoidea, Crangonyctoidea, Tal- itroidea, and Gammaroidea). At higher taxonomic levels, armature of the peduncle may also prove phyl- etically significant, particularly the development of baso-facial spine(s) in gammaroidean superfamilies, and distolateral spines in gammarioideans and some fossorial superfamilies (e.g., Phoxocephaloidea and Pontoporeioidea) . Figure 4 illustrates three main types of rami of uropods 1 & 2 and their occurrence in representative gammaridean superfamilies. Lanceolate rami (A) are generally slender and taper distally to an acute apex that lacks distinct apical spine(s) or spine clusters; marginal spines (when present) are typically arranged in oppos- ing, evenly spaced series. Lanceolate rami typify the most primitive superfamilies of reproductive "swim- mers" (Natantia), including the Lysianassoidea, Phoxocephaloidea, Pardaliscoidea and most Eusir- oidea. Linear rami (C) are generally thick and robust (styliform), with subparallel margins that tapering only slightly distally; the apex is rounded or blunt, and usually bears a distinct cluster of spines of unequal length. These rami typify mostly benthonic crawling or burrowing superfamilies, with reproductive! y pre- AMPHIPACIFICA VOL. Ill N0.1. MAY 16, 2001. 58 Fig. 4. Form of rami of uropods 1 & 2. A. Lanceolate; B. Transitional; C. Linear. (After Bousfield (1973) and unattributed sources) amplexing gnathopods (Reptantia), such the Crang- 2 apparently transcends these categories within a few onyctoidea, Talitroidea, Gammaroidea, and Coroph- gammaridean superfamilies (e.g., Pontoporeioidea). ioidea. Transitional rami (B) taper variously to a sub- Also, within family Podoceridae, the dulichiid sub- acute apex that may bear a single spine or a few very group possesses lanceolate uropod rami that are atypi- short spines; marginal spines are usually present and cal of superfamily Corophioidea, to the other character serially arranged (e. g., Dexaminoidea and Mel phi dip* states of which the dulichiids conform reasonably well . P°idea). The vestigial uropod rami of cercopid caprellidean The form and armature of the rami of uropods 1 & amphipods are also lanceolate. Such a character state AMPHIPACIFICA VOL. 3 NO. 1 MAY 16, 2001. 59 Fig. 5. Suggested evolutionary relationships of the telson within the Amphipoda, (modified from Bousfield & Shih 1994). AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 60 "anomaly" apparently supports direct phyletic ancest- ry of dulichiids to the cercopid line of Caprellidea, as proposed by Laubitz ( 1993). It may suggest a possible diphyletic origin of the Corophioidea, and considera- tion of a possible leucothoidean ancestry for the dulich- iid podocerids (see fig. 7; Bousfield & Shih 1994). However, broadly across families of most gammar- idean superfamilies, the uropod ramal condition is remarkably stable and correlates well with the phyletic status of other character states in those same taxa. Thus, the lanceolate condition is typical of super- families that exhibit plesiomorphic states of antennal sensory organelles and sexually similar gnathopods. Conversely, the linear ramal condition is associated most frequently with apomorphic reduction of anten- nal sensory organelles, presence of sexually dimorphic gnathopods, and reproductive pre-amplexing behav- iour. Not surprisingly, the transitional ramal form occurs mainly in higher taxa with a phyletically "inter- mediate" status of other character states. Thus, the form and armature of uropods 1 & 2 appear to be character states of high-level classificatory signifi- cance. The Telson. The form of the telson has long been considered a character of prime taxonomic significance (Stebbing 1906; Barnard & Karaman 1991). Its probable function in both free-swimming and benthonic life styles, and its overall significance in supeifamily level classification has been reviewed by Bousfield & Shih (1994). The deeply bilobate form is generally deemed the plesiomorphic condition within amphipodan, peracar- idan,andindeed,allmalacostracancrustaceans(Schram 1986). Conversely, the entire, platelike, or "fleshy" form of the telson, presumably represents a distal fus- ion of the two primary lobes (e.g., as in superfamilies Leucothoidea and Corophioidea respectively) and thus the typical apomorphic state. A very advanced condi- tion is seen in the Thaumatelsonidae and many Hyperiidea, where the plate -like telson is fused with the urosome. A less frequent but presumably apomorphic condition occurs where the lobes become separated throughout their entire length (as in most Gammaroidea and certain Hadzioidea) and attains an extreme separa- tion dorsally on urosome 3 (abominal segment 6) in the advanced fossorial genus Eohaustorius (Pontoporei- oidea). A panoramic view of telson types across the spec- trum of higher amphipod taxa is provided in Figure 5. The prototype amphipod is depicted with a bilobate telson, the apex of each lobe having a "notch and spine" configuration derived from a presumed pelagic pera- caridan (primitive malacostracan) ancestral outgroup. Following evolutionary lines outwards from this base through each superfamily group, we find that member species and genera having the greatest number of plesiomorphic character states (those nearest the base) also tend to have a fully or partially bilobate telson. Conversely, member species and genera with the most apomorphic or derived character states, in balance, usually show the most strongly fused or plate-like form of the telson. The totally bilobate apomorphic form may be noted in advanced members of the Gammar- oidea and in some members of the Pontoporeioidea (family Haustoriidae). However, the overall form of the telson proves not directly significant in development of a phyletic classi- fication. As noted in fig. 5, development of a plate-like telson takes place independently and homoplasiously within nearly every superfamily group. Derivation of a superfamily group based solely on a plate-like telson would encompass members of at least ten different major groups, and thus be totally artificial. However, within component families of the most primitive superfamilies of "Natantia" (e.g., Lysianassoidea, Phoxocephaloidea, Eusiroidea, Pardaliscoidea) the deeply bilobate form of the telson is dominant. Con- versely, within the more advanced “natant” super- families such as the Oedicerotoidea and Leucothoidea, the Hyperiidea, and among most reptant superfamilies (e.g., Crangonyctoidea, Talitroidea, Bogidielloidea, Corophioidea), the distally notched or plate-like form is dominant. Despite contrary views of some (e.g., Barnard & Karaman 1981 , 1 99 1), the balance of evidence strongly supports the overall conclusion that a deeply bilobate telson is the plesiomorphic or ancestral condition with- in the Amphipoda. Conversely, a plate-like or apically entire telson is the apomorphic or advanced condition. However, structure of the telson appears to be more dependent uponfactors of life-style at lower taxonomic levels rather on the more broadly “stable” features of reproductive biology. Character states of the telson may therefore be phyletically significant only at fam- ily, subfamily, or even generic classificatory levels. Phyletic Significance of Gnathopod Structure The external morphological features of the gnath- opods (peraeopods 1 & 2 of formal malacostracan terminology) have previously been considered one of the most significant and fundamental indicators of high AMPHIPACIFICA V0L.3N0.1 MAY 16, 2001. 61 level phyletic relationships within suborders Gammar- idea (Stebbing 1906); Barnard & Karaman 1991) and Caprellidea (Laubitz 1993;Takeuchi 1993). A cross- secti on of amphipod gnathopod types was al so anal yzed by Bousfield & Shih 1994. Early taxonomic studies had long detailed the sexu- ally dimorphic, powerfully subchelate form of the gnathopods of intertidal and freshwater species of Gammarus of northwestern Europe. In females and immature stages, these anterior appendages were used mainly as implements of food-gathering but, in sexu- ally mature males, are utilized in precopulatory carry- ing of the female, thus ensuring close proximity of the sexes at the time of her "mating" moult (ecdysis). Justified or not, Gammarus was considered by many workers to be the basic or ancestral amphipod repro- ductive form (e.g., Barnard 1969). More recent studies (e.g., Borowksy 1984; Conlan 1991a) have investigated gnathopod morphology and sexual dimorphism across a broad spectrum of amphi- pod superfamilies. The results have been compared with a pre-amplexing and/or mate-guarding form of reproductive behaviour in species of Gammarus of northwestern Europe. As indicated by Schram (1986), this form of reproductive behaviour is now considered by most workers as relatively highly evolved and specialized within the Amphipoda. The search for a probable ancestral form of the gnathopods first centred on members of supeifamilies that were classified as primitive on the basis of other plesiomorphic character states. Over a range of family and generic morphotypes within the primitive super- family Lysianassoidea (e.g., Barnard and Ingram 1990; Lowry & Stoddart 1997), the distal portions (carpus, propod and dactyl) of both gnathopods in both sexes, are found to be consistently subsimilar. Despite minor modifications within an increasingly sophisticated ge- neric series, the plesiomorphic form of both gnatho- pods may be described as weakly subchelate, with slender carpus and propod, and clearly not sexually dimorphic. Within the Lysianassoidea, mating takes place freely and rapidly in the water column, there i s no pre-amplexus or mate-guarding phase, and by corol- lary represents the plesiomorphic reproductive (mat- ing) behaviour. Amphipod supeifamilies grouped within the cat- egory Natantia (Table I, p.67) are typified by pelagic reproductive (mating) behaviour, and by nonsexually dimorphic gnathopods that are primitively weakly sub- chelate and subsimilar in form. Exceptions can be explained, at least tentatively, on the basis of (1) independent or convergent evol ution wi thin geographi - cally isolated sub-taxa that have been exposed to simi- lar, mainly ecological , evol utionary stresses(e.g. , south- ern families of Oedicerotoidea); (2) a morphological vestige of presumed ancestral types whose evolution- ary “thrust” devolved mainly into other superfamily groups that are, today, essentially “reptant” in repro- ductive life style (e.g., in Pontoporeiidae); or (3) a probable extant precursor of more widespread and diverse modem taxonomic groups (e.g., in Dexaminoi- dea, Melphidippoidea). Within subcategory Reptantia, gnathopod morphol- ogy is basically different, and the range of morphotypes is considerably greater than seen in the Natantia (Bousfield & Shih 1994). Thus, in most component superfamil ies the gnathopods are characteri sticall y sex- ually dimorphic and strongly subchelate or cheliform, especially in males. However, many exceptions to these overall trends have been noted. These plausibly include a secondary use of sedimentary benthic sub- strata as a “fluid” mating medium wherein sexually dimorphic gnathopods and pre-amplexing mating be- haviour may not be required (e.g., in Haustoriidae). In summary, within component superfamilies of Reptantia, sexual dimorphism of the gnathopods, and benthic pre-amplexing reproductive styles are typical. These types are mainly vegetatively free-living and epigean in physically rigorous habitats such as coastal shallows, estuaries, and fresh-waters. Conversely, in members that have become secondarily commensal with other marine animals or plants, penetrated into hypogean brackish- and fresh-water or the deep sea, or have become fully terrestrial, sexual dimorphism of the gnathopods is markedly reduced or lacking. Second- arily and neotenically, the sexually dimorphic form may revert to a morphology suited to the vegetative life style of both sexually mature adults and immature stages. Mating Behaviour Within the Amphipoda The significance of precopulatory mating behav- iour and sexual dimorphism in phyletic relationships of amphipod crustaceans has been broadly investigated by Conlan (1991) and summarized by Bousfield & Shih (1994). To ensure proximity of males and females at the time of female ovulating ecdysis, amphipods employ two basic reproductive strategies: ( 1 ) mate-guarding, in which the males are either (a) carriers involving pre-amplexing, with concomitant modification of male gnathopods for the purpose, or (b) attenders, where they remain domiciled with the fe- AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 62 A. EUSIROIDEA: PONTOGENEIIDAE (Paramoara cotumbfana) (Hyafe aettcomi*) D. HAOZIOIDEA: MEUTIOAE (UHItanm t) E. GAMMAROIDEA: ANISOGAMMARIDAE (Eogamman isocftlrl) C. TALITROIDEA: HYALELLIDAE (Hyalai lamtteca) F. GAMMAROIDEA: GAMMA RIDAE (Gammarua taacfatoa ) Fig. 6. Precopula in representative superfamilies of gammaridean Amphipoda. (after Bousfield & Shih 1994) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 63 male and employ the gnathopods mainly in agonistic manner to ward off competing males. (2) non-mate-guarding, in which the mature male sim- ply seeks out females wherever they may be at the time of ovulation. These males are classified as (a) pelagic searchers if the female is in the water column, or (b) benthic searchers if the female is on or in the bottom substrata. In either case the gnathopods are little or not sexually dimorphic, and no pre-amplexus takes place. Both strategies are determined by the period of ovulation of the female, at which time the male must be present if fertilization of the eggs is to take place. For a short period immediately following moulting, the cuticle of the female is sufficiently flexible to allow for release of the eggs into the brood pouch or marsupium. Sperm is deposited there by the male during copulation, and fertilization of the eggs can then take place. Coni an (1991) concluded that the searching strat- egy is a primitive, and mate-guarding an advanced, form of reproductive behaviour in amphipods. This conclusion provides the principal basis for the present updated semi-phyletic classification of amphipod superfamilies (Table I, p. 67). In these mating strategies, the reproductive mor- phology of the mature female is seldom significantly different from that of the vegetative or feeding stages, except in some species of Melita and some aquatic talitroi deans where the coxae are modified to accept the dactyl of the precopulatory gnathopod of the male. However, the breeding frequency and fecundity reflect overall differences in mating strategy. Thus, females of mate guarders tend to be iteroparous, with several broods in a lifetime, whereas non-mate-guarders tend to be semelparous, with only one brood in a lifetime. In the most primitive superfamily groups within Natantia, contact between the mate-seeking male and the female takes place only during actual copulation, and its duration is brief (Conlan 1991). These positions have been illustrated for a number of representative families and superfamilies of both Natantia and Reptantia. (Bousfield & Shih 1994). The positions vary according to the relative size of males and females, and on environmental conditions. All ensure rapid sperm transfer at the time of the female's ovulation moult. Some pre-amplexing positions are illustrated in Fig. 6. Preamplexing is rare within superfamilies of Nat- antia, and where it does occur briefly, differs little from that of amplexus. Within Reptantia, however, pre- amplexus is nearly the rule. In the primitive Gammar- oidea, males of Anisogammaridae (e.g., Eogammarus oclairi. Fig. 6E) grasp the smaller female by the base of coxa 4, usually by means ofgnathopod 1. In family Gammaridae (e.g., Gammarus, Fig. 6F), the male car- ries the female by means of a ‘Yore-and aft” clutching of the anterior edge of peraeon plate 1 and posterior edge of peraeon 5, facilitiated by the very oblique palm of gnathopod 1. Within the Hadzioidea, the male of Melita nitida (Fig. 6D) employs his small gnathopod 1 to grasp the female by the specially modified anterior lobe of coxa 6. His much enlarged gnathopod 2 may be used in fending off competing males. In many aquatic Talitroidea, especially in Hyalella and Allorchestes (Hyalellidae, Fig. 6C) and in Hyale and Parallorchestes (Hyalidae, Fig. 6B), the dorsally positioned male in- serts the dactyl of gnathopod 1 in a precopulatory notch in the lower anterior margin of peraeon 2 of the smaller female. Again, the much enlarged gnathopod 2 appar- ently functions agonistically towards other males. In some species of Hyale , however, the dactyl ofgnatho- pod 2 may be used in grasping and/or rotating the fe- male. The widespread phenomenon of convergent evolu- tion of high-level characters states is well illustrated by these superficially similar mating strategies, that differ morphologically and tactically at family and subfamily levels. The Phylogenetic Tree. Possible natural relationships among subordinal and superfamily groups may be represented in the form of a phylogenetic tree (Fig. 7). This dendrogram is modified from Bousfield & Shih (1994) to include superfamilies Iphimedioidea Lowry and Myers, 2000 and Stenothoidea Bousfield, 2000, and reflect the in- fluence of additional characters and character states. The plesiomorphic character states, especially of the antennal sensory organelles, are most strongly evinced in taxa, extant or extinct, that are closest to the trunk and main branches. The apomorphic or advanced and specialized features are best developed in taxa placed nearthe branching extremi ties. The phylogenetic“tree” may be viewed, in effect, as a form of cladogram in which the character states are ordered and arranged "parsimoniously", but without numerical basis. The present version is little changed from the earlier tree (1994). During the past 10 years the number of species in each group has increased, variously, by only about 5-10%, few major new taxa have been discov- ered, and the ordering of character states has remained basically unchanged. However, as noted above (Fig. 4) the form of the rami of uropods 1 & 2 have here been AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 64 \V; V -A HYPERIIDEA \ + +.+ V / CAPRELL1DEA BOGIDIELL- OIDEA LEGEND CALLYNOPHORATES - inside CALCEOLATES inslda oooooooooo TAXA WITH BRUSH SETAE - Inside + + + Fig. 7. Phylogenetic tree of suggested natural relationships within suborders and superfamilies of the Amphipoda (modified from Bousfield & Shih 1994). AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 65 suggested as significant indicators of phyletic relation- ships. The degree of anterolobation of coxae 5&6, and of heteropody of pereopods 5 -7, deserve further study as indicators of phyletic significance. Emphasis on such parameters has here altered the position of the main trunk which now centrally subtends superfamilies of Natantia leading to the Hyperiidea, the most highly modified (advanced) of fully natatory taxa. The phyletic position of the Liljeborgioidea, a group not yet rigorously defined, remains uncertain. North Ameri canfamily inclusions (p. 100) are mainly benthic, commensal, deep-demersal, or hypogean in fresh wa- ter. Antennal reproductive sensory organelles are lacking in all subgroups, and most have developed sexually dimorphic gnathopods and pre-amplexing mating behaviour. Paradoxically it seems, component families retain lanceolate or transitional type uropod rami, posterolobate or weakly anterolobate coxae 5 & 6, and peropods 5-7 are basically homopodous. Other enigmatic family- or perhaps superfamily-Ievei groups elsewhere include the Niphargidae, Phreatogam- maridae, and the monotypic Sensonator valentiensis Notenboom, 1986. In phylogentic analysis of the 10 suborders of the Isopoda, Brusca & Wilson (1991) have employed cladistic methodology leading to major classificatory recommendations. However, the universal applicabil- ity and adequacy of cladistic analyses for this purpose has been questioned by some (e.g., Gosliner & Ghiseln 1984). Relative to the taxonomically "difficult" order Amphipoda, the superficially similar peracari dan order Isopoda is more uniformly benthic in life style, with much greater development of both external and internal parasitic forms. It is palaeohistorically more ancient (Bousfield & Conlan 1990), and thus with perhaps fewer "intermediate" stages that frustrate creation of neatly defined phyletic units based on one or two character states only, A full cladistic 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 (e.g., in Berge £l al 2001). However, a phyletic tree based on "first principles" here provides a useful visual basis for eventual numerical establishment of a true phyletic classification of the Amphipoda. PHYLETIC ARRANGEMENT OF HIGHER TAXA The present phyletic classification of higher amphipod taxa (Table I) is based on relatively few characters and character states, most of which exhibit high classificatory value. The North American species list (Appendix I, p. 75) follows this arrangement of higher taxonomic names. The present analysis recognizes the Ingolfiellidea and Gammaridea as distinct and valid subordinal divi- sions of Order Amphipoda. However, the Hyperiidea and Caprellidea are of lesser significance, here sub- merged within subcategories of Natantia and Rept- antia, respectively. This conclusion agrees in part with the results of a limited cladistic anlysis of the Amphi- poda by Berge el al (2001). That study likewise combines hyperiids and caprellids variously within the Gammaridea, but is less demonstrative of the subord- inal distinctness of the Ingolfiellidea. Within suborder Gammaridea, the pragmatic terms "Natantia" and Reptantia" continue to encompass al- most the same superfamily groups as earlier proposed (Bousfield & Shih 1994). Introduction of the form of the rami of uropods 1 & 2 as primary phyletic indicators reinforces the applicability of those subcategories, at least on a semi-phyletic basis. Thus, the newly pro- posed uropod-descriptive terms "Lanceolata"and "Lin- eata", are essentially interchangeable with the original terms "Natantia" and "Reptantia", since they encom- pass virtually the same respective superfamily goups. Two major subgroups may be recognized within the Natantia: the primary Lanceolata, and the transi- tional Lanceolata. Member of the former are typically fully marine, have a mainly free-swimming life style, their antennal sensory organelles are well-developed, but the gnathopods are not sexually dimorphic. The "Transitionals" are not strictly marine, exhibit a wider variety of benthonic (commensal) life styles, and ex- hibit varying loss of antennal organelles, but corre- sponding development of sexually dimorphic, pream- plexing gnathopods. The Reptantia may be subdivided into: (1) primit- ive superfamilies having posterolobate coxae 5 & 6, and (2) advanced superfamilies in which these coxae are mainly anterolobate. The more primitive "antero- lobates" encompass the pontoporeioidean and gam- maroidean superfamilies ("gammarida" of Barnard & Barnard 1983). The advanced "anterolobates" contain the most highly evolved groups of gammaridean amphipods, marked by very specialized morphologies and life styles. As noted above, the position of the Liljeborgioidea remains enigmatic. In conspicuous morphological character states and life style, component members seem clearly assignable to the "Reptantia". However, the condition of the posterior peraeopods and uropod AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 66 rami is plesiomorphic and characteristic of most "Nat- antia". A tentative, but not entirely satisfactory sol - ution is to place the Liljeborgioidea among the "Ad- vanced Transitionals” within Natantia (Table I). Inearlyphyleticstudies(e.g.,Bousfield 1979, 1983), the author utilized several other external morphologi- cal features, some of which tend to support the present categories. Thus, members of the Natantia usually possess a distinct rostrum, coxal gill on per-aeopod 7, well-developed natatory uropod 3, but relatively short antenna 1 ; in the Reptantia, however, the rostrum, and coxal gill of peraeopod 7 are usually lacking, uropod 3 is often reduced and non-natatory, and antenna 1 is usually elongate. Sternal gills, of various form and presumed osmo- regulatory function, occur only in freshwater taxa, but may have phyletic significance nonetheless. Thus, withinNatantia, ail superfamilies that encompass fresh- water families and genera contain some species bear- ing sternal gills (e. g., in Gammar acanthus, Pseud- amoera, and Falklandella within Eusiroidea; Para- calliope within Oedicerotoidea; Phreatogammarus within Melphidippoidea; and Paracrangonyx within Liljeborgioidea). In the Reptantia, however, sternal gills are characteristic of the more primitive super- families Crangonyctoidea (all families), Talitroidea (Hyalellidae), and Pontoporeioidea ( Monoporeia , Diporeia ). Sternal gills are lacking in all freshwater gammaroideans and hadzioideans (e.g., weckeliids, pseudoniphargids), to which may be added the Euro- pean-Mediterranean regional species of Niphargidae, Sensonator, and all members of superfamily Bogidiell- oidea. Attempts at utilizing other seemingly phyletically promising characters and states have proven frustrat- ing and ineffective, largely because of homoplasious character state similarities at superfamily level . Mouth- part morphology tends to reflect feeding style and is thus useful mainly in family level classification [e. g., in Stegocephalidae (Berge 2000)]. Seemingly "in defi- ance of" other phyletic trends across both Natantia and Reptantia, the morphology of female brood lamellae varies between the broad, marginally setose, presumed plesiomorphic condition, and the narrow, strap-like, apomorphic form. A very few characters have been little utilized to date, and may merit further investigation. Pleopod morphology is seldom figured or described in detail, especially in the early literature. What little is known of theircharacterstates(e.g.,typeof retinacula, "clothes- pin spines") tends to be conservative "across the taxo- nomic board". Small morphological differences may therefore be significant at high classificatory level. Presence or absence and size of the accessory flagellum seems not phyletically accountable; its length appears secondarily increased in some deepwater gammarids of Lake Baikal. However, its position of origin (ant- erior in some Phoxocephalidae and Liljeborgiidae, mediolateral in nearly all other taxa) merits further study. Character states of surface ultrastructure are little known but may be especially promising as phyl- etic indicators when the difficulties of terminology and function have been resolved ( Halcrow & Bous- field 1987). CONCLUSIONS Analysis of plesio-apomorphic conditions of se- lected external morphological characters and repro- ductive behaviour has resulted in a revised classifica- tion of the amphipod Crustacea. Introduction of new characters has lent support to recognition of only two suborders, the primitive Ingolfiellidea, and the more advanced and much more diverse Gammaridea. The analysis also lends further support to the phyletic significance of previous gammaridean subcategories "Natantia and "Reptantia", interchangeable with newly proposed terms "Lanceolata" and "Lineata" respec- tively. These basic gammaridean morphotypes are re- presented in Fig. 8 as an assist to vizualizing or concep- tualizing morphological relationships among the spe- cies of North American amphipods (Appendix I). Because of homoplasious occurrence of some char- acter states "across the taxonomic board", these sub- category names combine elements of phyletic signifi- cance with pragmatic usefulness. Cognizance of such variation within all component species requires that superfamilies be realistically diagnosed by a "best-fit" consensus of character states, rather than by rigorous conformity to one or two morphological criteria. The classification outlined in Table I may be used as a form of "key" to subordinal and superfamily groups listed in Appendix 1. This extensive list of marine, brackish, freshwater and terrestrial species contains all known suborders and superfamilies, and many of the families allocated to each superfamily (see Martin & Davis 2001). Phyletic classification has many advantages, not the least of which is conformity with phyletic classifica- tions elsewhere within Class Crustacea, and major ord- inal groups within the Animal Kingdom. Superfamily grouping of the North American fauna (Appendix I) has also facilitated comparative biogeogeographical AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 67 TABLE I. Phyletic Classification of the Amphipoda suggested by character states of the uropods superimposed on those of reproductive morphology and belaviour, and other characters. I. AMPHIPODA INGOLFIELLIDEA (uropod 2 >1; eyes stalked; maxillipeds partly separated basally; peduncle 3 of antenna 2 elongate, body vermiform; 2 families * hypogean, marine and freshwater). II. AMPHIPODA GAMMARIDEA (uropod 1 > 2; eyes sessile; maxillipeds fused basally; peduncle 3 of antenna 2 not elongate; -150 families - epigean and hypogean, marine, freshwater, and terrestrial). A. LANCEOLATA (=NATANTIA) (rami of both uropods 1 & 2 lanceolate, often with serially arranged marginal spines and lacking apical spines; antennae strongly sexually dimorphic, male with sensory antennalorganelles; gnathopods not (or weakly) sexually dimorphic; uropod 3 usually large, biramous). I. Basic Lanceolates (uropods 1 & 2 rami lanceolate; gnathopods not sexually dimorphic). 1 . Callynophorates (with antennal callynophore and brush setae in male) Lysianassoidea (antennae calceolate, head not rostrate) Pardaliscoidea; Stegocephaloidea: Hyperiidea (non-calceolate; head rostrate) Hyperiidea (maxilliped lacking palp; coxae 1-4 small; A2 short in female) Synopioidea (callynophore weak or non-existant, but brush setae present); 2. Phoxocephaloideans (callynophore seldom and brush setae infrequent; calceoli plesiomorphic, receptacle and bulla lacking, body with few distal elements; head strongly rostrate; peraeopod 5 dactylate); 5 families iossorial, marine, mainly anti boreal). II. Transitionals (uropods 1 & 2 transitional inform; callynophore & brush setae reduced or lacking,; gnathopods weakly sexually dimorphic, or not). 3. Primitive Transitionals (antennae often calceolate, coxae 5 & 6 posterolobate) Eusiroidea (mostly pelagic; pereopods 5-7 homopodous, segment 4 produced behind) Oedicerotoidea (fossorial; peraeopods 5 & 6 homopodous, P7 elongate; gnathopods sexually dimorphic in 2 families). Leucothoidea (benthonic) (uropod 3, outer ramus I -segmented; gnathopod rarely sex. di morph.) Iphimedioidea (benthonic): uropod 3, outer ramus 1 -segmented, gnathopods weak not dimorph). Stenothoidea (benthonic), uropod 3, outer ramus 2-segmented; gnathopod often sex. dimorphic) 4 Advanced Transitionals (male antennae non-calceolate,with brush setae, callynophore rare; coxae 5 & 6 anterolobate, uroppod 3 biramous, often natatory) Dexaminoidea and Ampeliscoidea (urosome 2 & 3 fused; U3 rami large, natatory) Melphidippoidea: (urosome 2 & 3 separate; U3 lanceolate, weakly sexually dimorphic) Liljeborgioidea (gnathopods sexually dimorphic; life style commensal or freshwater hypogean. B. LINEATA (=REPTANTIA) (uropod rami linear, with apical spines, lateral marginal spines irregular; gnathopods sexually dimorphic, usually strongly; usally benthic reproductive behaviour) I. Posterolobate reptants (Coxae 5 & 6 posterolobate; uropod 3 short, rami reduced) Crangonyctoidea (Antenna 1 elongate, wth accessory flagellum; A2 calceolate in male); Talitroidea (Antenna 1 the shorter, lacking accessory flagellum; A2 non-calceolate) II. Anterolobate Reptants (Coxae 5 & 6 anterolobate; uropod 3, one or both rami large) 1. Primitive Anterolobates (telson bilobate; free-swimming. free-burrowing,or commensal) Pontoporeioideans (appendages fossorial, P5 adactylate; gnathopods weakly or not sexually dimorphic; may retain pelagic reproduction, with primitively calceolate antenna 2 (male) Gammaroideans (appendages seldom fossorial; gnathopods subsimilar in size and sexually dimorphic; antennae weakly or not calceolate, coxal gill on peraeopod 7; mainly freshwater) Hadzioideans (appendages rarely fossorial; antennae not calceolate; gnathopods unlike and strongly sexually dimorphic; antennae not calceolate; P7 lacking coxal gill; marine and brackish) 2. Advanced Anterolobates (telson plate-like or entire; domicolous or excl. hypogean life style) Bogidi ell oi deans (vermiform; uropod 3 subequally biramous; telson plate-like; f.w. hypogean). Corophioideans (body depressed; peraeopods 3 & 4 glandular; uropod 3 reduced, telson fleshy. animals marine, domicolous (tube-buidling); male gnathopods mate guarding). Caprellideans (body slender, cylindrical; coxae lacking; abdomen vestigial; marine, epigean, semi-sessile; 2 infrorders: Caprellida (skeleton shrimps) and Cyamida (whale lice). AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 68 Hyperiopsidae (near Lysianassoidea ) 3 4 5 \ \ t Peraaopodo ^ w 7123 Uropod© Plaopod* le] Elm mop us (MeUtldae) (Uadzioidea) Fig. 8. Representative morphotypes of basic categories of phyletic classification of the Amphipoda A. Lanceolata (=Natantia). B. Lineata (=Reptantia) (after Bousfield & Shih 1994). AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 69 analysis of subregional faunas (Bousfleld 2001). In summary, Arctic and Pacific coastal marine amphipod faunas are relatively primitive, possibly reflecting the long-term (biohistorical) stability of those regions. The east coast faunas are more advanced, compare phyletically with those of the Mederranean region, and presumably reflect the relatively recent origin of the North Atlantic Ocean (since late Jurassic). Gulf coast amphipods encompass the highest percentage of ad- vanced, and lowest percentage of primitive super- families, consistent with its relatively high year-round temperature regime. Thus, within the Amphipoda, ev- olution of apomorphic features (e.g., sexually dimor- phic gnathopods) "classically” proceeds most rapidly in tropical regions; conversely, plesiomorphic features (e.g., antennal sensory organelles) are most frequently retained in cold-water regions and in the deep sea where evolutionary rates are presumably much slower. This biogeographic-phyletic analytical methodology has been extrapolated from North American superfam- ily groups to other well-studied regional faunas to conclude that the world's most primitive marine assem- blages presently occur in the Antarctic. The North American freshwater amphipod fauna is much more diverse than was believed during the mid 1900's, thanks mainly to the extensive recent work of Dr. John R. Holsinger and colleagues, with much new material yet to be published (per. commun.). Itcontains a high percentage of ancient relict types with sternal gills, dominated in hypogean habitats by members of the Crangonyctoidea, and in epigean habitats by the exclusively neotropical Hyalellidae (Bousfleld 1996) and the arctic-boreal pontoporeioidean genus Diporeia (Bousfleld 1987). The more modem gammaroi deans and hadzioideans, lacking sternal gills, are widely diverse throughout Eurasia. In North America, how- ever, these advanced groups are represented only pe- ripherally, and by small numbers of species and few families, of which some are recently introduced (e.g., Witt elal, 1997). A few relict species within Gammar- acanthidae, Sebidae, and Bogidiellidae complete the North American freshwater complex. The need for full return to phyletic classication of the Amphipoda, inevitable though it may be, remains urgent. Present analysis indicates that a fully satisfac- tory phyletic classification still eludes us. Cladistic methodology (e.g., Berge el al, 2001) has not yet solved the problem of sui table outgroups and/or homoplasious occurrence of character states widely accross the taxo- nomic board. The problem may yet be solved through pooling of results from all analytical methodology, and employment of some of the characters and character states here developed. Especially promising is genetic methodology, both DNA hybridization and rDNA sequencing (Schram, Duffy, pers. commun.). Al- though these methodologies have special limitations of their own, they seem minimally affected by homoplasy of external character states, thereby providing a more reliable basis for phyletic classification. The present arrangement of superfamilies is not fully phyletic and is farfrom afinal answer. 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A review of the genus Iphimedia (Crustacea: Amphipoda) with de scriptions of three new species from Australia, Papua, New Guinea, and Florida. Invert. Taxon- omy 5: 469-485. Thomas, J. D., and J. L. Barnard 1991b. Phot is traph- erus, a new elephantine species from the Caribbean Sea (Crustacea: Amphipoda). Proc. Biol. Soc. Wash. 104(1): 96-100. Thomas, J. D., & J. L. Barnard, 1991c. Two new species of Netamelita from the Caribbean Sea (Crustacea: Amphipoda: Gammaridea). Proc. Biol. Soc. Wash. 104 (3): 583-592. Thomas, J. D., & J. L. Barnard 1992a. Podocerus cheloniphilus , a testudinous amphipod newly re- corded from the western Atlantic Ocean. Bull. Mar. Sci. 50(1): 108-116. Thomas, J. D.. & J. L. Barnard 1992b. Podocerus kleidus , new species from the Florida Keys. (Crust- acea: Amphipoda, Dulichiidae). Bull. Mar. Sci 51 (3): 309-314. AMPHIPACIFICA VOL 3 NO.l MAY 16, 2001. 75 Williams, W. D., & J. L. Barnard 1988. The taxonomy of crangonyctoidAmphipoda(Crustacea)from Aus- tralian freshwaters. Rec. Austral. Mus., Suppl. 10 : 1-180. Witt, J. D. S., P. D. N. Hebert,& W.B. Morton, 1997. Echinogammarus ischnus : another crustacean in- vader in the Laurentian Great Lakes Basin. Can. J Fish. Aquat. Sci. 54: 264-268. APPENDIX I. PHYLETIC LIST OF AMPHIPOD CRUSTACEA OF NORTH AMERICA, NORTH OF MEXICO. The study of phyletic classification presented in the main text was developed mainly through analysis of North American species listed here. The North Ameri- can amphipod fauna contains about 1650 species, rep- resentative of all known suborders and superfamilies, about 2/3 of families, and perhaps 1/4 of the total number of species world-wide. As noted in the ac- knowledgements (p. 50), the list was developed, over a 15-year period, by a subcommittee of the Committee on Scientific and Common Names of Aquatic Inverte- brates, chaired by Dr. Donna D. Turgeon, NOAA, Washington, D. C. The list encompasses marine, brackish, freshwater and terrestrial faunal components. An additonal ~200 species have been recognized from continental North America north of Mexico, including Canada and Alaska, but not Greenland, Bermuda, or the Bahamas. Others are known from the U. S. mid- Pacific state of Hawaii. These undescribed taxa are in the process of being treated by systematic specialists. Their work will be added to an updated final list, including common names where possible, to be pub- lished in a special volume on the Crustacea of North America jointly sponsored by NOAA and the Ameri- can Fisheries Society. The system of higher classification of amphipods of this list is essentially phyletic, including superfamily level taxa, following standards proposed for Ingolf- iellidea by Stock (1977), Hyperiidea by Bowman & Griiner (1973); Caprellidea by Laubitz (1993) and Gammaridea by Schram (1986), updated by Bousfield and Shih (1994) and Bousfield (2000). Although the arrangement of superfamilies follows that of Table I of the main text (p. 67), the component families and genera are listed alphabetically. Newly proposed subordinal categories of classification are omitted for the present, but if reasonably widely ac- cepted by colleagues, may be introduced in the final CNAI crustacean volume. The former subordinal- level names Hyperiidea and Caprellidea are retained in situ within the list, mainly for pragmatic reasons, even though they have been merged within suborder Gam- maridea. The merged older names have yet to be re- assessed at suitable classificatory levels. As noted above, the phyletically arranged list of North american amphipods provides a basi s for biogeo- graphical analysis of it subregional marine and fresh- water faunas. This study, currently in press (Bousfield 2001b), also contains a detailed numerical analysis of numbers of species by subregion, supeifamily, and family level categories. Ocurrence Legend A Arctic AC - Acadian At - Atlantic AL - Alaska ALEUT - Aleutians BAR - Pt Barrow BC - British Columbia BER - Bering Sea C - Carolinian CAL - California CHES - Chesapeake Bay CUBA - Cuba E - Eastern FL - Florida G - Gulf of Mexico HAT - Cape Hatteras HAW . Hawaii LA Louisiana LABR Labrador MI Mississippi N Northern NC North Carolina NFLD Newfoundland P Pacific ORE Oregon SE Southeastern STL St Lawrence Gulf V Virginian WA Washington State FW Freshwater ST Semiterrestrial T Terrestrial TEX Texas W Western YUC Yucatan AMPHIPACIFICA VOL. 3 NO. 1 MAY 16, 2001. 76 SCIENTIFIC NAME OCCURRENCE SUBORDER INGOLFIELLIDEA HANSEN, 1903 Family Ingolfiellidae Hansen, 1903 Jngolfiella fuscina Dojiri & Sieg, 1987 At-G (SC-W FL) SUBORDER GAMMARIDEA LATREILLE, 1803 Superfamily Lysianassoidea (Bousfield, 1979; Lowry & Stoddart, 1997) Family Lysianassidae Dana, 1849 Subfamily Lysianassinae Dana, 1849 Acidostoma laticorne G. O. Sars, 1879 Aruga oculata Holmes, 1908 A. holmesi (Barnard, 1955) Bonassa bonairensis (Stephensen, 1933) Concarnes concavus (Shoemaker, 1933) Dissiminassa homosassa Lowry & Stoddart, 1997 D. dissimilis (Stout, 1913) Eclecticus eclecticus Lowry & Stoddart, 1997 Lysianopsis alba Holmes, 1903 L. cubensis Shoemaker, 1933 L. hummelincki (Stephensen, 1933) L. ozona Lowry & Stoddart, 1997 L. subantarctica (Schellenberg, 1931) Macronassa macromera (Shoemaker, 1916) M. par iter (J. L. Barnard, 1969) Menigrates obtusifrons (Boeck, 1861) Shoemakerella cubensis (Stebbing, 1897) S. nasuta (Dana, 1853) At (N, slope) P (CAL P (WA-CAL), G (W FL) G (FL) G(FL) G(FL) P(S CAL) G(FL) At (V-C), G (FL) G (FL.) G (FL) G( WFL) G (FI -tropic?) P(S CAL) P (CAL) At (G, N, slope) G (W FL) G (FL) (see Shoemaker, 1948) Subfamily Tryphosinae Lowry & Stoddart, 1997 Allogaussia recondita Stasek, 1958 P(BC-CAL) Hippomedon coecus (Holmes, 1908) P(S CAL) H. columbianus Jarrett & Bousfield, 1982 P(BC-ORE) H . denticulatus (Bate 1857) At (N, slope) H. granulosus Bulycheva, 1955 P(BER-BC) H. holbolli (Kroyer, 1946) At (ST L) H. pensacola Lowry & Stoddart, 1997 G (W FL) H. propinquus Sars, 1890 At (ST. L-HAT) H. serratus Holmes, 1905 At (AC-CHES) H. subrobustus Hurley, 1963 P (CAL?) H. tenax Barnard, 1966 P(S CAL?) H. tricatrix Barnard, 1971 P (ORE, deep) H. zetismus Hurley, 1963 P (CAL, deep) Koroga megalops Holmes, 1908 P (BC-WA, offshore) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 77 Lepidepecrella charno Barnard, 1966 Lepidepecreoides nubifer Barnard, 1971 Lepidepecreum eoum Gurjanova, 1951 L. nubifer Barnard, 1971 L. garthi Hurley, 1963 L. gurjanovae Hurley, 1963 L. serraculum Dalkey, 1998 L. serratum Stephensen, 1925 Orchomenella decipiens Hurley, 1963 O. holmesi (Hurley, 1963) Orchomenella minuta (Kroyer, 1846) O. pacifica (Gurjanova, 1951) O. perdido Lowry & Stoddart, 1997 O. thomasi Lowry & Stoddart, 1997 Orchomene depressa Shoemaker, 1930 O. holmesi (Hurley, 1963) O. limodes Meador & Present, 1985 O. macroserrata Shoemaker, 1930 O. magdalensis (Shoemaker, 1942)' O. nugax (Holmes, 1904) O. obtusa (Sars, 1891) O. pectinata Sars, 1882 O. serrata (Boeck, 1861) Paralibrotus setosus Stephensen, 1923 Paratrypkosites abyssi (Goes, 1866) Psammonyx longimerus Jarrett & Bousfield, 1982 P- nobilis (Stimpson, 1853) P- terranovae Steele, 1979 Rimakoroga floridiana Lowry & Stoddard, 1997 & rima (J. L. Barnard, 1964) Schisturella pulchra (Hansen, 1887) Tmetonyx cicada (Fabricius,1780) T- gulosus (Kroyer, 1845) Tryphosella apalachicola Lowry & Stoddart, 1997 T. compressa (Sars, 1891) ^ groenlandica (Hansen, 1887) T. gulosus (Kroyer, 1845) T. index (Barnard, 1966) T. metacaecula Barnard, 1967 T. nanoides (G. O. Sars, 1895) T- orchomenoides Stephensen, 1925 T- rotundata (Stephensen, 1923) T- spitzbergensis (Chevreux, 1926) T- triangula (Stephensen, 1925) Wecomedon wecomus (Barnard, 1971) W. similis Jarrett & Bousfield, 1982 W. wirketis (Gurjanova, 1962) P (CAL-deep?) P (ORE-deep) P (BER-CAL) P(ORE, deep) P(BC-CAL) P (BC-CAL) P ((CAL) At (G, N, slope) P (CAL) P (BC-CAL) P (AL-ORE)-A-At (STL) P (BER-CAL) G(WF) G(WFL) At (AC, shelf) P (CAL) P (CAL) At (AC, shelf) P(S CAL?) P(BER-WA) P(SEAL-CAL), At (STL) At (ST L) At (AC-V) At (ST L, slope) A-At (G-BCN) P(BC-ORE) At (AC-DEL) At (AC-NFLD) G (W FL) P(S CAL) At ST L slope) At (ST L -AC) At (STL) G (W FL) At (ST Lslope) At (ST L slope) At (N, slope) P (CAL) P (CAL) At (St.L) A-At At (ST L, slope A-At (ST L, slope) At (STL) P (SEAL-ORE) P(BER-SEAL) P(BER-AL) Family Uristidae Hurley, 1963 Anonyx adoxus Hurley, 1963 P(ORE-CAL) AMPHIPACIF1CA VOL. 3 NO.l MAY 16, 2001. 78 A. barrowensis Steele, 1 882 P-A (BAR) A. beringi Steele, 1982 P(BER) A. comecrudus J. L. Barnard, 1971 P(ORE) A. compactus Gurjanova, 1962 P (BER?), At (STL) A. dalli Steele, 1983 P (BER?) A. debruyni Hoek, 1882 At (STL) A. epistomaticus Kudrjaschov, 1965 P (BER?) A. filiger Stimpson, 1864 P(WA?) A. hurleyi Steele, 1986 P(AL?) A. laticoxae Gurjanova, 1962 P (BER?) A. lilljeborgi Boeck, 1871 P (AL-CAL)-A-At (AC) A. makarovi Gurjanova, 1 962 P (BER)A-At (STL) A. m. rcwgar (HYBRID, Brunei MS)) At (STL) A nwgax (Phipps, 1774) P(BER-CAL?)-A-At A. ochoticus Gurjanova, 1962 P (BER?)-A-At A. pacificus Gurjanova, 1962 P(AL-WA) A. petersoni Steele, 1986 P (BER?) A. schefferi Steele, 1986 P (AL?) A. sculptifer Gurjanova, 1982 P (BER?) A. shoemakeri Steele, 1983 P (BER?) Centromedon pavor Barnard, 1966 P(ORE-CAL) C. pumilus (Liljeborg, 1865) At (ST L) Euonyx laquaeus Barnard, 1967 P (deep) Gronella groenlandica (Hansen, 1887) At (ST L) Hirondellea fidenter Barnard, 1966 P (CAL)? Kyska dalli Shoemaker, 1964 P (ALEUT) Onisimus (Onisimus) litoralis (Kroyer, 1845) A-At (N) Onisimus ( Boekosimus ) edwardsi (Kroyer, 1846) At (G, N) 0. (B.) glacialis (G. 0. Sars, 1900) At (G) 0. (B.) normani (Sars, 1891) At (ST L slope) 0. (B.) plautus (Kroyer, 1845) At (ST L slope) Paronesimus bare nisi (Stebbing, 1894) A-At (N, slope) Paratryphosites abyssi (Goes, 1866) At (deep) Schisturella cedrosiana Barnard, 1967 P (CAL) S cocula Barnard, 1966 P (BC) S. dorotheae (Hurley, 1963) P (CAL) S. grabenis Barnard, 1967 P (deep) S. totorami Barnard, 1967 P (CAL) S. tracalero (Barnard, 1966) P(S CAL) S. zopa Barnard, 1966 P(S CAL) Sop hro syne robertsoni Stebbing & Robertson, 1891 P (CAL) Stephonyx biscayensis (Chevreux, 1908) G(FL) U.ristes californicus Hurley, 1963 P (CAL) U. dawsoni Hurley, 1963 P (CAL) U. entalladurus Barnard, 1963 P (CAL?) U. perspinus Barnard, 1967 P (ORE, deep) U. umbonatus (Sars, 1882) At ST L slope) A MPHI PACIFICA VOL. 3 NO.l MAY 16, 2001. 79 Family Scopelocheiridae Lowry & Stoddart, 1997 Paracallisoma coecum (Holmes, 1908) P (AL-BC, offshore) Family T rise hizostomatidae Lilljeborg 1865 Trischizostoma sp. (Bousfield, 1987 proposed) P-At (AL-CAL, deep) Family Opisidae Lowry & Stoddart, 1995 Opisa eschrichti Kroyer 1842 P (BC-CAL), At (AC) O. tridentata Hurley, 1963 P (BC- CAL) O. odontochela Bousfield, 1987 P(SE AL-BC) Subfamily concept Conicostomatinae Lowry & Stoddart proposal; Barnard & Karaman 1991? Acidostoma hancocki Hurley, 1963 P (BC-CAL) A. obesum subsp. ortum J. L. Barnard, 1967 P (CAL, deep) Ocosingo borlus Barnard, 1964 (= Fresnillo Barnard) P (BC-CAL) Pachynus barnardi Hurley, 1963 P(WA-CAL) Prachynella lodo Barnard, 1964 P(WA-CAL) Socarnes hartmanae Hurley, 1963 P (CAL) S. vahli (Kroyer, 1838) At (ST L, slope) Socarnoides illudens Hurley, 1963 P (ORE-CAL) Family Cyphocarididae Lowry & Stoddart, 1997 Cyclocaris guile Imi Chevreux, 1899 P(SCAL?) Cyphocaris challenger i Stebbing, 1880 P (AL-CAL) C. faurei K. H. Barnard, 1918 P(S CAL?) C. guile Imi Chevreux, 1899 P (AL-CAL) C. richardi Chevreux, 1905 P (BC-CAL) C. anonyx Boeck, 1871 P (BC, offshore) C. tunicola Lowry & Stoddart, 1997 G (W FL) Metacyphocaris helgae Tattersall, 1906 P (AL-CAL) Family Aristiidae Lowry & Stoddart, 1997 Aristias captiva Lowry & Stoddart, 1997 A. expers Barnard, 1967 A. pacific us Schellenberg, 1936 A . topsenti Chevreux, 1900 A. tumidus (Kroyer, 1846) A. veleronis Hurley, 1963 Boca campi Lowry & Stoddart, 1997 B. elvae Lowry & Stoddart, 1997 B. megachela Lowry & Stoddart, 1997 G(WFL) P (CAL?) P(BC-WA) At ST L, slope) P(WA), At (STL, slope) P (BC-CAL) G(WFL) G (E & W FL) G(WFL) Family Endevouridae Lowry & Stoddart, 1997 Ensayara entrichoma Gable & Lazo-Wasem, 1990 G (W FL) E. ramonella Barnard, 1964 P (S CAL?) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 80 Family Hyperiopsidae Bovallius, 1886 (near Cyphocarididae Lowry & Stoddart?) Parargissa americana Barnard, 1961 P (CAL, BC, deep) P. galatheae Barnard, 1961 P (CAL?) Family Valettiidae Stebbing, 1888 Valettiopsis dentata Holmes, 1908 P (BC-CAL, deep) Cedrosella fames (Barnard, 1967) P (CAL)? 1. Incerta sedis Eurystheus grillus Lichtenstein, 1882 P (abyssal) Superfamily Stegocephaloidea Bousfield, 1979 Family Stegocephalidae Dana, 1855 Subfamily Adanieniexinae Berge, 2000 Andaniexis abyssi Boeck, 1871 P (deep), A- At (AC) A. elinae Berge & Vader, 1997 A A. gracilis Berge & Vader, 1997 A A. lupus Berge & Vader, 1997 A Parandania boecki (Stebbing, 1888) P (BC) Parandaniexis mirabilis Schellenberg, 1929 P (BC?) Subfamily Andaniopsinae Berge, 2000 Andaniopsis nordlandica (Boeck, 1871) At (BF) Andanieopsis pectinata (Sars, 1882) A- At (NFLD) Subfamily Stegocephalinae Berge, 2000 Bousfieldia mammilidacta (Moore, 1992) Gordania camoti (Barnard, 1967) Phippsia romeri Schellenberg, 1925 Pseudo viscaina (Barnard, 1967) Stegocephalexia penelope Moore, 1992 S hancocki (Hurley, 1956) S. minima (Stephensen, 1925) S. pajarella (Barnard, 1967) Stegocephalus ampulla (Phipps, 1774) S. abyssicola (Oldevig, 1959) S. inflatus Kroyer, 1842 S. cascadiensis (Moore, 1992) S. similis (Sars, 1895) P(BC) P (CAL) A P (CAL) P(BC) P (S CAL, deep) A-At (NFLD) P (CAL) A A PA (BER)-A-At (ST L) P (ORE, deep) A AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 81 Superfamily Pardaliscoidea Bousfleld, 1979 Family Pardaliscidae Boeck, 1871 Caleidoscopsis tikal (J. L. Barnard, 1967) Halice abyssi Boeck, 1871 H. malygini (Gurjanova, 1936) H. ulcisor Barnard, 1971 Halicoides lolo (Barnard, 1971) H. synopiae (Barnard, 1962) H. tambella (Barnard, 1961) Pardaliscella symmetrica Barnard, 1959 P. yaquina Barnard, 1971 Pardaliscoides fictotelson J. L. Barnard, 1966 Parahalice mirabilis Birstein & Vinogradov, 1962 Rhynohalicella halona (Barnard, 1971) Tosilus arroyo Barnard, 1966 P (CAL) At (ST L) A P (ORE) P(ORE) P(ORE) P (CAL) P (CAL) P (ORE) P (CAL, deep) P (abyssal) P(BC-CAL) P (S CAL, deep) Family Stilipedidae Holmes, 1908 Subfamily Stilipedinae Holmes, 1908 (revised Holman & Walling, 1983) Stilipes distincta Holmes, 1908 P (AL-CAL) Subfamily Astyrinae Pirlot, 1934 (revised Holman & Walling, 1983) Astyra abyssi Boeck, 1871 At (STL) Family Vitjazianidae Birstein & Vinogradov, 1955 Vitjaziana gurjanovae Birstein & Vinogradov, 1955 P (BER, deep) Family Vemanidae Bousfleld 1979 (see Thurston, 1989) Vemana lemur esa Barnard, 1967 P (B CAL, deep) Superfamily Synopioidea Bousfleld, 1979 Family Synopiidae Dana, 1855 Bruzelia tuberculata Sars, 1866 B. inlex Barnard, 1967 B. guayacura Barnard, 1972 B. ascua Barnard 1966 Bruzeliopsis cuspidata Barnard, 1962 B. turba Barnard, 1964 Priscosyrrhoe priscis (Barnard, 1967) Garosyrrhoe bigarra (Barnard, 1962) G. cf. bigarra (Barnard, 1962) G. laquei Ortiz, 1985 Pseudotiron pervicax Barnard, 1967 P. golens Barnard, 1962 P (AL-CAL), A-At (ST L) P (CAL) P (CAL?) P (CL, deep) P (CAL) P (CAL) P(S CAL) P(S CAL) G(FL) G (FL - CUBA) P (CAL) P (CAL) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 82 P . coas Barnard, 1967 Synopia ultramarina Dana, 1853 S. scheeleana Bovallius, 1886 Syrrhoe crenulata Goes, 1866 S. longifrons Shoemaker, 1964 S. oluta Barnard, 1972 Syrrhoites columbiae Barnard, 1972 S. cokasseta Barnard, 1967 S. dulcis Barnard, 1967 S. lorida Barnard, 1962 S. silex Barnard, 1967 S. terceris Barnard, 1964 S. trux Barnard, 1967 Tiron biocellata Barnard, 1962 T. spiniferus (Stimpson, 1854) Metatiron cf. be Hair si (Just, 1981) M. triocellatus (Goeke, 1985) M. tropakis (Barnard, 1972) P(CAL) G(FL) G(SEFL) P (AL-CAL), A-At (ST L - AC) P(BC-CAL) P(CAL) P (ORE, deep) P (CAL) P (CAL) P (CAL) P (CAL) P (CAL) P (CAL, deep?) P(BC-CAL) A-At (AC) G(FL) G(FL) P (CAL?), At (V-C) G (FL?) Family Argissidae Walker, 1904 Argissa hamatipes (Norman, 1869) P (BER-CAL), A-At (ST) G (NW FL) SUBORDER HYPERIIDEA MILNE EDWARDS, 1830 Infraorder Physosomata Pirlot, 1929 Superfamily Scinoidea Bowman & Gruner, 1973 Family Scinidae Stebbing, 1888 Scina borealis (G. O. Sars, 1882) S. crassicornis (Fabricius, 1775) S. wanaWagler, 1926 S. rattrayi Stebbing, 1895 S. tullbergi (Bovallius, 1885) Proscina vinogradovi Shih & Hendrycks, 1996 Cheloscina antennula Shih & Hendrycks, 1996 P (BER-CAL)-At P (ORE-CAL) P (CAL) P (BC-WA, slope)-At P (CAL)-At (G) P ( AL) (54 40’N 155 10’W) P (AL) (53 20’N 155 16’W) Family Mimonectidae Bovallius, 1885 Mimonectes sphaericus Bovallius, 1885 P (BER)-A-At M. gaussi Woltereck,1904? P (BC-WA) Superfamily Lanceoloidea Bowman & Gruner, 1973 Family Lanceolidae Bovallius, 1887 Scypholaneola aestiva Stebbing, 1888 S. vankoeffeni Woltereck, 1909 P (WA-CAL, deep)-At P (BC-WA) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 83 Lanceola loveni Bovallius, 1885 L. serrata Bovallius, 1885 L. paciflca Bowman 1973 L. sayana Bovallius, 1885 P (ORE, deep)-At P (CAL, deep) P (BC-WA) P(BER) Family Chuneolidae Woltereck, 1909 Chuneola parasitica Vinogradov, 1956 P (BER-W ALEUT) Infraorder Physocephalata Bowman & Gruner, 1973 Superfamily Vibilioidea Bowman & Gruner, 1973 Family Vibiliidae Dana, 1852 Vibilia armata Bovallius, 1887 V. australis Stebbing, 1888 V. viatrix Bovallius, 1887 K gibbosal Bovallius 1887 P (ORE-CAL) P (BC-WA)-At - G P(CAL) P (CAL) Family Cystosomatidae Willemoes-Suhm, 1875 Cyctosomafabricii Stebbing, 1888 C. pellucidus (Willemoes-Suhm, 1873) P (BC-CAL)-At, deep P (SE AL-CAL)-At Family Paraphronimidae Bovallius, 1887 Paraphronima crassipes Claus, 1879 P- gracilis Claus, 1879 P (BER-CAL, slope)-At (G) P (BC-WA, deep)-At (Gulf) Superfamily Phronimoidea Bowman & Gruner, 1973 Family Phronimidae Dana, 1853 Phronima atlantica Guerin, 1836 P. bowmani Shih, 1991 P. dunbari Shih, 1991 P. pacifica Streets, 1877 P. sedentaria (Forskal, 1775) P. solitaria Guerin, 1836 P. stebbingi Vosseler, 1900 Phronimella elongata (Claus, 1862) P (BER-CAL)-At-G P (CAL) P (CAL) At-G P (BC-CAL)-At At-G At-G P (ORE)-At-G Family DaireUidae Bovallius, 1887 Dairella californica (Bovallius, 1885) p (ORE-CAL, oceanic) Family Phrosinidae Dana, 1853 (=Anchylomeridae) Anchylomera blossevillei Milne-Edwards, 1830 P (WA-CAL)-At-G AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 84 Phrosina semilunata Risso, 1822 P (CAL)-At-G Primno abyssalis (Bowman, 1968) P (BC-CAL) P. brevidens Bowman, 1978 G P. johnsoni Bowman, 1978 At-G P. lateillei Stebbing, 1888 P(CAL) Family Hyperiidae Dana, 1852 Hyperia antarctica Spandl, 1927 H. bengalensis (Giles, 1887?) H. galba (Montagu, 1813) H. leptura Bowman, 1973 H. medusarum (O.F.Mueller, 1776) H. spinigera Bovallius, 1889 Hyperietta stephenseni Bowman 1973 H. vosseleri (Stebbing, 1904) H. luzoni (Stebbing, 1888) H. stebbingi Bowman, 1973 Hyperoche medusarum (Kroyer, 1842) Hyperioides longipes Chevreux, 1900 Hyperionyx macrodactylus (Stephensen, 1924) Iulopsis loveni Bovallius, 1887 Lestrigonus bengalensis Giles, 1887 L. schizogeneios (Stebbing, 1888) L. crucipes (Bovallius, 1889) L. macrophthalmus (Vosseler, 1901) L. latissimus (Bovallius, 1889) L. shoemaker i Bowman, 1973 Parathemisto abyssorum Boeck, 1870 Phronimopsis spinifera Claus, 1879 Themistella fusca (Dana, 1853) Themisto pacifica (Stebbing, 1888) T. libellula Lichtenstein, 1822 T. guadichaudii Guerin 1842 P(AL-CAL) P (CAL) P (BER)-A-At P (CAL) P (BER-CAL)-A-At P (BC-CAL)-At P (BC-CAL)-At(G) P (CAL)- At (G) P (CAL)-At-G P(CAL)-At-G) P-A-At P (CAL)-At-G At-G At At -G P (CAL)-At-G At-G At-G At-G P (S CAL) P (BER)-A-At, deep At-G At-G P (BER-CAL) P (BER)-A-At A-At Superfamily Lycaeopsoidea Bowman & Gruner, 1973 Family Lycaeopsidae Chevreux, 1913 Lycaeopsis themistoides Claus, 1879 L. zamboangae (Stebbing, 1888) Superfamily Platysceloidea Bowman & Gruner, 1973 At-G P (CAL)-At Family Pronoidae Claus, 1879 Eupronoe armata Claus, 1879 E. minuta Claus, 1879 Paralycaea gracilis Claus, 1879 Sympronoe parva (Claus, 1879) At-G P(CAL)-At-GULF P (CAL)-At-G P(S CAL)-At-G A MPHI PA Cl FI C A VOL. 3 NO.l MAY 16, 2001. 85 Family Anapronoidae Bowman & Griiner, 1973 Anapronoe reinhardti Stephensen, 1925 P (CAL) Family Lycaeidae Claus, 1879 Lycaea pulex Marion, 1874 p (CAL) L. vincenti Stebbing, 1888 At-G L. bovallioides Stephensen, 1925 G L bovallii Chevreux, 1900 G Brachyscelus crusculum Bate, 1961 p (BC-CAL)-At? B. globiceps (Claus, 1871) At (CUBA)? B. rapax Claus, 1871 Q Family Oxycephalidae Bate, 1861 Oxycephalus clausi Bovallius, 1887 O. piscator Milne Edwards, 1830 Cranoecephal us scleroticus (Streets, 1878) Leptocotis tenuirostris (Claus, 1871) Rhabdosoma whitei Bate, 1862 Simorhynchotis antennarius Claus, 1871 Streetsia challengeri Stebbing, 1888 S. mindanaonis (Stebbing, 1888) S. pronoides (Bovallius, 1887) P (BC-CAL, deep)-At-G At-G At-G At-G At-G G P (BC-CAL, slope)-At-G G P (CAL) Family Platyscelidae Bate, 1862 Amphithyrus bispinosus Claus, 1879 G A. sculpturatus Claus, 1879 At-G Hemityphus rapax (Milne-Edwards, 1830) At-G Par aty phis maculatus Claus, 1879 At-G Platyscelus serratulus Stebbing 1888? p (S CAL) P- ovoides (Claus, 1879) At-G Tetrathyrus forcipatus Claus, 1879 At-G Family Parascelidae Bovallius, 1887 Thyropus edwardsi (Claus, 1879) At-G T. sphaeroma (Claus, 1879) At-G T. typhoides (Claus, 1979) P(CAL)-G Schizoscelus ornatus Claus, 1879 At-G AMPHIPACIFICA VOL. 3 NO. 1 MAY 16, 2001. 86 Superfamily Phoxocephaloidea Bousfield, 1979 [=Haustorioidea Barnard & Drummon, 1982 (part)] Family Platyischnopidae Thomas & Barnard, 1983 Eudevenopus honduranus Thomas & Barnard, 1983 E. metagracilis (Barnard, 1964) Skaptopus brychius Thomas & Barnard, 1983 Tiburonella viscana (Barnard, 1969) At (FL-SC), G P (S CAL) At (V-C, slope), G P (S CAL) Family Urothoidae Bousfield, 1979 Urothoe denticulata Gurjanova, 1951 U. rotundifrons Barnard, 1962 U. varvarini Gurjanova, 1953 P (BER?) P (CAL) P(BC-CAL) Family Phoxocephalidae G. O. Sars, 1895 Subfamily Metharpiniinae Jarrett & Bousfield, 1994a Grandifoxus aciculatus Coyle, 1982 P(AL-BC) G. acanthinus Coyle, 1982 P(AL) G. constantinus Jarrett & Bousfield, 1994a P (BER) G. dixonensis Jarrett & Bousfield, 1994a P (BC) G. grandis (Stimpson, 1856) P(BC-CAL) G. lindbergi (Gurjanova, 1953) P(BER-BC) G. longirostris (Gurjanova, 1938) P(BER-BC) G. nasutus (Gurjanova, 1936) P(AL) G. pseudonasutus Jarrett & Bousfield, 1994a P (ALEUT) G vulpinus Coyle, 1982 P(AL-BC) Beringiaphoxus beringianus Jarrett & Bousfield, 1994a P (BER) Majoxiphalus major (Barnard, 1960) P(SEAL-CAL) M. maximus Jarrett & Bousfield, 1994a P(AL-BC) Foxiphalus aleuti (Barnard & Barnard, 1982) P (AL) F. apache Barnard & Barnard, 1982 P (S CAL) F. cognatus (Barnard, 1960) P (S CAL) F. falciformis Jarrett & Bousfield, 1994a P(BC-ORE) F. fucaximeus Jarrett & Bousfield, 1994a P(WA) F. golfensis Barnard & Barnard, 1982 P (S CAL) F. obtusidens (Alderman 1936) P (ORE-CAL) F. secasius Barnard & Barnard, 1982 P(S CAL) F. similis (Barnard, 1960) P(BC-CAL) F. slatteryi Jarrett & Bousfield, 1994a P (BER) F. xiximeus Barnard & Barnard, 1982 P(BC-CAL) Metharpinia coronado i Barnard 1980 P (S CAL) M. floridana (Shoemaker, 1933) P (CAL?), G (FL) M. jonesi (Barnard, 1963) P(S CAL) Rhepoxynius abronius (J. L. Barnard, 1960) P(BC-CAL) R. barnardi Jarrett & Bousfield, 1994a P(BC-CAL) R. bicuspidatus (Barnard, 1960) P(BC-CAL) R. boreovariatus Jarrett & Bousfield, 1994a P(BC) R. daboius (Barnard, 1960) P(BC-CAL) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 87 R . fatigans (Barnard, 1960) P (BC-CAL) R. gemmatus (Barnard, 1969) P (S CAL) R. heterocuspidatus (Barnard, 1960) P (S CAL) R. homocuspidatus (Barnard & Barnard, 1982) P (S CAL) R . lucubrans (Barnard, 1960) P (S CAL) R. menziesi (Barnard & Barnard, 1982) P(S CAL) R. pallidus (Barnard, 1960) P (BC-CAL) R. stenodes (Barnard, 1960) P(S CAL) R. tridentatus (Barnard, 1954) P (ORE-CAL) R. variatus (Barnard, 1960) P (BC-CAL) R. vigitegus (Barnard, 1971) P(BC-ORE) R. epistomus (Shoemaker, 1938) At (V-C?) G (FL?) R. hudsoni Barnard & Barnard, 1982 At (V-C) G (FL?) Subfamily Pontharpiniinae Barnard & Drummond, 1978 Mandibulophoxus alaskensis Jarrett & Bousfield, 1994b P (AL-BC) A/. gilesi J. L. Barnard, 1957 P (BC-CAL) M. mayi Jarrett & Bousfield, 1994b P (SE AL-BC) Subfamily Parharpiniinae Barnard & Drummond, 1978 Eyakia robusta (Holmes, 1908) P(SEAL-CAL) Eyakia sp. 1 (= E. robusta Barnard & Barnard, 1981) P(CAL) Eyakia calcarata (Gurjanova, 1938] P (CAL) Subfamily Brolginae Barnard & Drummond, 1978 Eobrolgus chumashi Barnard & Barnard, 1981 P(AL-CAL) £. pontarpioides Gurjanova, 1953 P(BER) E. spinosus (Holmes, 1905) P?-At(V), G (E FL?) Paraphoxus beringiensis Jarrett & Bousfield, 1994b P(BER) P. communis Jarrett & Bousfield, 1994b P(BC) P. gracilis Jarrett & Bousfield, 1994b P (BC-CAL) P. oculatus Sars, 1879 At (ST L) P. pacificus Jarrett & Bousfield, 1994b P(BER-BC) P. rugosus Jarrett & Bousfield, 1994b P(BER) P. similis Jarrett & Bousfield, 1994b P (BC) P. simplex Jarrett & Bousfield, 1994b P (BER?) Subfamily Phoxocephalinae Barnard & Drummond, 1978 Cephalophoxoides homilis (Barnard, 1960) Leptophoxus icelus Barnard, 1960 Metaphoxus frequens Barnard, 1960 Parametaphoxus fultoni (in Barnard, 1960 in part) Parametophoxus quay lei Jarrett & Bousfield, 1994b Phoxocephalus holbolli (Kroyer, 1842) P (BC-CAL) P (CAL) P(SEAL-CAL) P(AL-CAL) P(BC-ORE) A- At (AC-CHES) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 88 Subfamily Harpiniinae Barnard & Drummond, 1978 Coxophoxus hidalgo J. L. Barnard, 1966 Harpinia antennaria Meinert, 1893 H. clivicola Watling, 1981 H. cabotensis Shoemaker, 1930 H. pectinata G.O. Sars,1891 H. plumosa (Kroyer, 1842) H. propinqua Sars, 1891 H. serrata Sars, 1879 H. truncata Sars, 1894 Harpiniopsis fulgens J. L. Barnard, 1960 H. emeryi Barnard, 1960 H. epistomata Barnard, 1960 H. fulgens Barnard, 1960 H. galera Barnard, 1960 H. gurjanovae Bulycheva, 1936 H. naiadis Barnard, 1960 H. percellaris Barnard, 1971 H. petulans Barnard, 1966 H. profundis Barnard 1960 H. triplex Barnard, 1971 Heterophoxus affinis (Holmes, 1908) H. oculatus (Holmes. 1908) H. conlanae Jarrett & Bousfield, 1994b H. ellisi Jarrett & Bousfield, 1994b H. ellisi variant Jarrett & Bousfield ,1994b H. nitellus Barnard, 1990 Pseudharpinia excavata Chevreux, 1887 P. inexpectata Jarrett & Bousfield, 1994b P. sanpedroensis (Barnard, 1960) P(CAL) AP, At (V, deep slope) At (off DEL) At (AC) AP, At (S to Hatteras) (see Watling) At (St.L) At (AC) C Hat. Watling, 1981 At, G deep? At (to Mid At) (see Watling) P(BC-CAL) P (CAL?) P (S CAL) P (CAL) P (CAL) P(BER) P (S CAL) P(ORE, deep) P (CAL) P (CAL?) P (ORE, deep) P(SE AL-CAL) P (S CAL) P (SEAL-ORE) P(BC-ORE) P (BC) P (S. CAL) P (CAL) P(BC) P(S CAL) Superfamily Eusiroidea Bousfield, 1979 Family Amathillopsidae Pirlot, 1934 (transferred to Iphimedioidea by Lowry & Myers, 2000) Amathillopsis spinigera Heller, 1875 P -At (pelagic) Family Bateidae Stebbing, 1906 Batea catharinensis Muller, 1865 G(FL) B. bousfleldi Ortiz, 1991 G(WFL) B . lobata Shoemaker, 1926 P(S CAL) B . transversa Shoemaker, 1926 P (S CAL) Carinobatea cuspidata Shoemaker, 1926 G(WFL) C. carinata Shoemaker, 1926 G(FL) Family Eusiridae Stebbing, 1888 Cleonardo moirae Bousfield & Hendrycks, 1995a Eusirella elegans Chevreux, 1908 P (BC, pelagic) At (ST L) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 89 E . multicalceola (Thorsteinson, 1941) P (BC-WA, pelagic) Eusirogenes deflexifrons Shoemaker, 1930 At (ST L) Eusiroides monoculoides (Haswell, 1879) P (CAL) Eusirus columbianus Bousfield & Hendrycks, 1995a P(BC) Eusirus cuspidatus Kroyer, 1845 P, At (STL) Eusirus longipes Boeck, 1871 At (ST L, slope) Eusirus propinquus G. 0. Sars, 1893 At (ST L, slope) Rhachotropis aculeata (Lepechin, 1780) A-P R. americana Bousfield & Hendrycks, 1995a P (BC) R. barnardi Bousfield & Hendrycks, 1995a P (CAL) R. boreopacifica Bousfield & Hendrycks, 1995a P(SEAL-BC) R. cervus Barnard, 1957 P(S CAL) R. clemens Barnard, 1967 P(CAL) R . conlanae Bousfield & Hendrycks, 1995a P(BC) R. distincta (Holmes, 1908) P (pelagic). At (STL) R. inflata (Sars, 1883) P (AL-CAL)At (ST L, slope) R. ludificor Barnard, 1967 P (CAL) R. luculenta Barnard, 1969 P(S CAL?) R. oculata (Hansen, 1888) At-A-P R. minuta Bousfield & Hendrycks, 1995a P(BC) R. natator (Holmes, 1908) P (pelagic) Rozinante jragilis (Goes, 1866) A-At (STL) Family Gammaracanthidae Bousfield, 1977 Gammaracanthus loricatus Sabine, 1824 A-At (AC) Pseudacanthus aestuariorum (Lomakinia, 1952) P (AL)-A-At (AC)(Dadswel 1,1974) Family Gammarellidae Bousfield, 1977 Gammarellus homari (L., 1768) A-At (AC) G. angulosus (Rathke, 1843) At (AC) Family Pontogeneiidae Stebbing, 1906 Accedomoera vagor J. L. Barnard, 1969 P(SEAL-CAL) A. melanopthalma (Gurjanova, 1938) P (SE AL-CAL) Nasageneia quinsana (Barnard, 1964) P(S CAL) N. yucatenensis Ledoyer, 1986 G(FL) Paramoera ( Paramoera ) columbiana Bousfield, 1958 P (SEAL-ORE) P. ( Paramoera ) mohri Barnard, 1958 P (CAL-WA) P. (Paramoera) bousfieldi Staude, 1995 P (SEAL-ORE) P . (Paramoera) serrata Staude, 1995 P(WA-CAL) P. (Paramoera) suchaneki Staude, 1995 P (SE AL-S CAL P. (Rhithromoera) bucki Staude, 1995 P(SEAL-WA) P. (Rhithromoera) carlottensis Bousfield, 1958 P(SE AL-BC) P. (Humilomoera) leucophthalma Staude, 1995 P(SEAL-WA) P. (Humilomoera) crassicauda Staude, 1995 P(AL) Pontogeneia inermis (Kroyer, 1838) P (BER-CAL)-A-At P ivanovi Gurjanova 1951 P (BER-WA)-A P. rostrata Gurjanova, 1938 P (BER-CAL)-A AMPHIPACIFICA V0L.3N0.1 MAY 16, 2001. 90 p. intermedia Gurjanova, 1938 P (BER-CAL)- p. (Tethygeneia) opaia Barnard, 1979 P(CAL) p. (T.) longleyi Shoemaker, 1933 G(F1) p. (T.) bartschi Shoemaker, 1948 G (FL-CUBA) Family Calliopiidae G. O. Sars, 1895 Apherusa bispinosa (Bate, 1857) A. cirrhus (Bate, 1862) A. fragilis (Goes, 1966) A. glacialis Hansen, 1888 A. megalops (Buchholz, 1874) A. retovskii Gurjanova, 1934 A. sarsi Shoemaker, 1930 A. tridentata (Bruzelius, 1859) Bouvierella carcinophila Chevreux, 1889 Calliopius behringi Gurjanova, 1951 C. columbianus Bousfield & Hendrycks, 1997 C. carinatus Bousfield & Hendrycks, 1997 C. laeviusculus (Kroyer, 1838) C. pacificus Bousfield & Hendrycks, 1997 C. sablensis Bousfield & Hendrycks, 1997 Cleippides bicuspis Stephenson, 1931 C. quadricuspis Heller, 1875 Dolobrotus mardeni Bowman 1974 Halirages bispinosus Stephensen 1916 H. fulvocincta (M. Sars, 1858) H. elegans Norman, 1882 H. mixta Stephenson, 1931 H. nilssoni Ohlin, 1895 H. quadridentata Sars, 1876 Haliragoides inermis (Sars, 1882) Laothoes meinerti Boeck, 1871 L. pacificus Gurjanova, 1938 L. polylovi Gurjanova., 1946 Leptamphopus paripes Stephensen, 1931 Oligochinus lighti J. L. Barnard, 1969 Oradarea longimana (Boeck, 1871) Paracalliopiella pratti Barnard, 1954 P. beringiensis Bousfield & Hendrycks, 1997 P haliragoides Bousfield & Hendrycks, 1997 P. kudrjaschovi Bousfield & Hendrycks, 1997 P. slatteryi Bousfield & Hendrycks, 1997 Weyprechtia pinguis (Kroyer, 1838) W. heuglini (Buchholz, 1874) At (GST L) A A-At (STL) A-P A-P(BER) A A A P (BC), At (STL) P(BER) P (SEAL-ORE) P(BC-CAL) A-At-AC) PB(BC-CAL) At (AC) A A At (AC, deep) At (ST L, deep) A (Barrow), At (ST L) A A At-A (G, N, deep) A At-A (ST L) A PA (BER) At (ST L - LABR, deep) P (BC, deep), At (ST L, slope) P(AL-CAL) P (BC-CAL), At (ST L, deep) P (BER-CAL) P-A (BER) P-A (BER) P-A (BER) P (BER) A-P-At (ST L-LABR) A-P-At (STL) Superfamily Oedicerotoidea Bousfield, 1979 Family Oedicerotidae Lilljeborg, 1865. Acanthostepheia behringiensis (Lockington, 1877) A. malmgreni (Goes, 1866) A (BER) P (BER), A-At (ST L, deep) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 91 Aceroides distinguendus (Hansen, 1888) A (BAR) A. edax J. L. Barnard, 1967 P (CAL, deep) A. goesi Just, 1980 A A , . latipes (Sars, 1882) P (SE AL-BC), A-At (ST L deep) A. sedovi Gurjanova, 1946 A Americhelidium americanum (Bousfield, 1973) G(FL)) A. millsi Bousfield & Chevrier 1996 P(WA) A. pectinatum Bousfield & Chevrier, 19% P(BC-ORE) A. micropleon (Barnard, 1977) P(S CAL) A. setosum Bousfield & Chevrier, 19% P(SE AL-BC) A. variabilum Bousfield & Chevrier, 19% P(BC-WA) A. shoemakeri (Mills, 1%2) P (BER-CAL) A. rectipalmum (Mills, 1%2) P (BER-CAL) Ameroculodes edwardsi (Holmes, 1903)(Ledoyer, 1972) A. kolmesi Bousfield 1996 Arrhinopis longicornis Stappers, 1911 Arrhis lutkeni Gurjanova, 1936 A. phyllonyx (M. Sars, 1858) Bathymedon antennarius Just, 1980 B. covilhani J. L. Barnard, 1961 B. flebilis Barnard, 1967 B. kassites Barnard, 1966 B. longimanus (Boeck, 1871 ) B. nanseni Gurjanova, 1946 B. pumilis Barnard, 1962 B. obtusifrons (Hansen, 1887) B. roquedo Barnard, 1962 B. saussurei (Boeck, 1871) B. vulpeculus Barnard, 1971 Deflexilodes enigmaticus Bousfield & Chevrier, 1996. D. intermedius Shoemaker 1930 D. norvegicus (Boeck 1871) D. similis Bousfield & Chevrier, 1996 D. simplex Hansen, 1887 D. tesselatus Schneider, 1884 D. tuber culatus Boeck, 1871 Finoculodes omnifera Barnard, 1971 Hartmanodes hartmanae (Barnard, 1962) H. nyei (Shoemaker, 1933) Kroyera carinata Bate, 1857 Machaironyx muelleri Coyle, 1980 Monoculodes brevirostris Bousfield & Chevrier, 1996 M. castalskii Gurjanova, 1951 M. diamesus Gurjanova, 1936 M. demissus Stimpson, 1853 M. emarginatus J. L.Bamard, 1962 M glyconicus Barnard, 1967 M latissimanus (Stephensen, 1931) M. latimanus (Goes, 1861 M. longirostris (Goes, 1866) M. murrius Barnard, 1962 At (AC) (not FL!) At (V) G (FL?) A-At (ST L) P(AL?) A-At (ST L, slope) A P (ORE, deep) P (ORE-CAL, deep) P (CAL-deep) At (G, N, slope) P (BER-BC)A-At (ST L) P (ORE-S CAL) A-At (STL) P (CAL) At (ST L) P (ORE-S CAL, deep) P(SE AL-BC) A-At (AC) (not FL!) P(S CAL), At -(ST L) P(AL-BC) A-At (ST L, slope) At (ST L) A-At (ST L, slope) P (ORE, deep) P (S. CAL) G (FL) (see Ortiz, 1979) P (BC?) P(BER) P (BC) P(BER) P (BER?-BC) AT (AC) P (ORE-CAL) P (CAL, deep) P (ORE-BC?), At (STL) P (SE AL-WA)-A-At (ST L, slope) A-At (N, slope) P (CAL) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 92 M. necopinus Barnard, 1967 M. packardi Boeck, 1871 M. perditus J. L. Barnard, 1966 M. recandesco Barnard, 1967 M. sudor Barnard, 1967 M. tenuirostratus Boeck, 1871 Monoculopsis longicornis (Boeck, 1871) Oediceroides trepadora (Barnard, 1961) Oediceros borealis Boeck, 1871 O. saginatus Kroyer, 1842 Pacifoculodes spinipes (Mills, 1962) p bruneli Bousfield & Chevrier 1996 P. barnardi Bousfield & Chevrier, 1996 P. levingsi Bousfield & Chevrier, 1996 P, crassirostris (Hansen, 1887) P. zernovi (Gurjanova, 1936) Paroediceros behringiensis Lockington, 1877 P. lynceus (M. Sars, 1858) P. propinquus (Goes, 1866) Perioculodes cerasinus Thomas & Barnard, 1985 P. longimanus (Bate & Westwood, 1868) Rostroculodes borealis (Boeck, 1871) R. hanseni Stebbing, 1894 R. kroyer i (Boeck, 1871) R. longirostris (Goes, 1866) R. schneider i (Sars, 1895) R. vibei (Just, 1980) Synchelidium tenuimanum Norman 1895 Westwoodilla brevicalcar (Goes, 1866) W. megalops (Sars, 1882) (syn with caeculcft ) P (CAL, deep) A-At (AC) P (BC-S CAL) P (ORE, deep) P (Cal, deep) At (N, slope) P (BER)-A-At(AC) P (ORE-CAL, deep) A-At (AC) A-At (G) P(BC-ORE) P(SEAL) P (CAL) P(BC) P(AL) P(BER-BC) P (BER)-A P (ALEUT), A-At (ST L, slope) A-At ST L, slope) G (FL-BL) At (STL) P-A (BAR), At-A (G, N) A A-At (ST L, slope) P-A (BAR) P-A (BAR), A-At (ST L) A-At (LABR) At (ST L, shelf) P (BC-CAL), A-At (STL) A Superfamily Leucothoidea Bousfield, 1979 Family Leucothoidae Dana, 1852 Anamixis cavitura Thomas, 1997 G (NE) A. hanseni Stebbing 1899 G (MI) A. linsleyi Barnard, 1955 P (S CAL) Leucothoe alata J. L. Barnard, 1959 P (S CAL) L. spinicarpa (Abildgaard, 1789) A-At (ST L) Leucothoides pacifica Barnard, 1955 P (S CAL) L. pottsi Shoemaker, 1933 G (FL) (-Anamixis linsleyi J. L. Barnard, 1955 Family Pleustidae Buchholz, 1874 Subfamily Pleustinae Bousfield & Hendrycks, 1994a Pleustes (Pleustes) panoplus (Kroyer, 1838) Pleustes ( Pleustes ) panoplus var 4 Bousfield & Hendycks, 1994b Pleustes (P.) panoplus var. 5 Bousf. & Hendrycks, 1994b At (A-AC) P (BER)-A P(BER) AMPHI PACI FI C A VOL. 3 NO. 1 MAY 16, 2001. 93 Pleustes ( P .) tuberculaius (Bate, 1858) Pleustes (Catapleustes) victoria e Bousfield. & Hendrycks, 1994b P. ( C.) constantinus Bousfield & Hendrycks, 1994b P. ( C.) constantinus var., Bousf. & Hendrycks. 1994b Thorlaksonius amchitkanus Bousfield & Hendrycks, 1994b T. borealis Bousfield & Hendrycks, 1994b T. depressus (Alderman, 1936) T platypus (Barnard & Given, 1960) T. brevirostris Bousfield & Hendrycks, 1994b T. subcarinatus Bousfield & Hendrycks, 1994b T. grandirostris Bousfield & Hendrycks, 1994b Thorlaksonius carinatus Bousfield & Hendrycks, 1994b T. truncatus Bousfield & Hendrycks, 1994b P(BER) P(BC) P(BER) P(BC) P(BER) P (SEAL-ORE) P (ORE-CAL) P (CAL) P(SE AL-CAL) P (SEAL-ORE) P(BC-CAL) P(SEAL-BC) P(BC) Subfamily Mesopleustinae Bousfield & Hendrycks, 1994a Mesopleustes abyssorum (Stebbing, 1888) P (ORE deep) Subfamily Pleustoidinae Bousfield & Hendrycks, 1994a Pleustoides carinatus (Gurjanova, 1972) P (BER?) Subfamily Atyiopsinae Bousfield & Hendrycks, 1994a, emend Cadien & Martin, 1999 Myzotarsa anixiphilius Cadien & Martin, 1999 P (S CAL) Subfamily Eosymtinae Bousfield & Hendrycks, 1994a Eosymytes minutus Bousfield & Hendrycks, 1994a P (BC) Subfamily Stenopleustinae Bousfield & Hendrycks, 1994a Arctopleustes glabricauda (Dunbar, 1954) Stenopleustes gracilis (Holmes, 1905) S. inermis Shoemaker, 1949 S. latipes M. Sars, 1858) Sympleustes olricki Hansen, 1887 A-At (UNG) At (AC-DEL) G (FL?) At (AC-DEL) At (ST L, slope) A Subfamily Pleusymtinae Bousfield & Hendrycks, 1994a Pleusymtes coquillus Barnard, 1971 p (ORE-CAL) p . glaber (Boeck, 1861)? P (CAL), A-At (AC) p. glabroides (Dunbar, 1954) A-At (LABR) p. pulchella (G. O. Sars ,1876) A-At (AC?) p. subglaber (Boeck, 1871) P (CAL) Pleustomesus medius (Goes, 1866) P?, A-At (ST L, slope) Subfamily Pleusirinae Bousfield & Hendrycks, 1994a Pleusirus secorrus Barnard, 1969 P (AL-CAL) AMPH1PACIFICA VOL. 3 NO.l MAY 16, 2001. 94 Subfamily Dactylopleustinae Bousfield & Hendrycks, 1994a Dactylopleustes echinoides Bousf. & Hendrycks, 1995b P (BC-CAL) Subfamily Neopleustinae Bousfield & Hendrycks, 1994a Neopleustes pulchellus (Kroyer, 1846) At-A (ST L, slope) “ Parapleustes " bicuspis (Kroyer, 1838) A-At “P. " assimilis (Sars, 1895) A-At “P. ” gracilis Buchholz, 1874 At (G) Pleustostenus displosus Gurjanova, 1972 P (BER?) "Sympleustes” cornigerus Shoemaker, 1964 P-A (BAR) Subfamily Parapleustinae Bousfield & Hendrycks, 1994a Chromopleustes johanseni Bousfield & Hendrycks, 1995ab C. oculatus (Holmes, 1908) . C. lineatus Bousfield & Hendrycks, 1995ab Incisocalliope aestuarius (Watling & Maurer, 1973) I. karstensi J. L. Barnard, 1959 . Micropleustes nautilus (Barnard, 1969) M. nautiloides Bousfield & Hendrycks, 1995b Parapleustes americanus Bousfield & Hendrycks, 1995b Gnathopleustes pugettensi (Dana, 1853) G. serratus Bousfield & Hendrycks, 1995b G. pachychaetus Bousfield & Hendrycks, 1995b G. trichodus Bousfield & Hendrycks, 1995b G. simplex Bousfield & Hendrycks, 1995b G. den (Barnard, 1969) Trachypleustes trevori Bousfield & Hendrycks, 1995b . T. vancouverensis Bousfield & Hendrycks, 1995b Commensipleustes commensalis (Shoemaker, 1952) Incisocalliope aestuarius (Watling & Maurer, 1973) I. newportensis Barnard, 1959 /. bairdi (Boeck, 1871) I. makiki (Barnard, 1970) P (BER) P(AL-CAL) P (SEAL-NCAL) At(V);G(FL?) A P (AL-CAL) P (BC-CAL) P(AL-BC) P(SE AL-CAL) P(SE AL-CAL) P (SEAL-ORE) P(BC) P (BC) P (CAL) P(AL-BC) P (BC) P (CAL) G (FL?) P(S CAL) P(S CAL) P (HAW) Superfamily Stenothoidea Bousfield ( 2001) Family Amphilochidae Boeck, 1871 Subfamily Amphilochinae Barnard & Karaman, 1991 Amphilochoides odontonyx (Boeck, 1871) A-At (shelf) Apolochus barnardi Hoover & Bousfield, 2001 P(CAL) A. casahoya (McKinney, 1978) G (FL-TEX) A. delacaya (McKinney, 1978) G. A. litoral is (Stout, 1912) P(SE AL-CAL) A. manudens (Bate, 1862) At (ST L) A. picadurus (Barnard, 1962) P (CAL) A. staudei ( Hoover & Bousfield, 2001) P(BC-WA) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 95 A. pillai Barnard & Thomas, 1983 A. tenuimanus Boeck, 1871 Gitana abyssicola Sars, 1892 G. calitemplado Barnard, 1962 G. ellisi Hoover & Bousfield, 2001 Gitanopsis arctica Sars, 1892 G. bispinosa (Boeck. 1871) G. inermis (Sars, 1882) Hourstonius vilordes (Barnard, 1962) H. laguna (McKinney, 1978) H. tortugae (Shoemaker, 1933) G(FL) A-At (ST L, slope) A-At (ST L, deep) P (CAL) P(BC) A-At ST L) A-At (ST L) A-At (ST L, slope) P(SEAL-CAL) G(FL-TEX) G(EL) Subfamily Cyproideiinae Barnard & Karman, 1991 Haplopheonoides obesa Shoemaker, 1956 G (FL) Family Stenothoidae Boeck, 1871 Mesometopa esmarki (Boeck, 1871) P (CAL) M. neglecta Barnard, 1966 P (CAL) M. sinuata Shoemaker, 1964 P(ORE-CAL) Metopa alderi (Bate ,1857) A-At M. abyss alls Stephensen, 1931. A (G-EM) M. boecki Sars, 1892 A-At M. borealis Sars, 1882 A-At M. bruzelii (Goes, 1866) A-At (ST L) M. cistella Barnard, 1969 P (CAL) M. clypeata (Kroyer, 1842) A-At M. dawsoni Barnard, 1962 P (CAL) M. glacialis (Kroyer, 1842) P(BER), A-At (STL) M. groenlandica (Hansen, 1887) A-At (ST L) M. invalida G. O. Sars, 1892 At (STL) M. leptocarpa G. O. Sars, 1882 A-At (ST L) M. longicornis Boeck, 1870 A-At (ST L) M. norvegica (Lilj, 1950?) At ST L) M. propinqua G. O. Sars, 1892 A-At (ST L) M. pusilla G. O. Sars 1892 At (ST L) M. robusta Sars, 1892 A-At (STL) M. samsiluna Barnard, 1962 P (CAL) M. sinuata Sars, 1892 A-At-(ST L) M. solsbergi Schneider, 1884 At (ST L) M. spinicoxa Shoemaker, 1955 A (AC) M. spitzbergensis Briiggen, 1909 A-At (St L) M. sporpis Barnard, 1969 P (CAL, deep) M. tenuimana Sars, 1892 A-At (ST L) Metope Ha aporpis Barnard, 1962 P(CAL) M. carinata (Hansen, 1887) A-At M. longimana (Boeck 1871) A-At M. nasuta (Boeck, 1871) A-At Metopelloirtes micropalpa (Shoemaker, 1930) At (AC) AMPHIPACIFICA VOL. 3 NO. 1 MAY 16, 2001. 96 M. tattersalli Gurjanova, 1938 A (BAR) Parametopa alaskensis Holmes, 1904) P(AL) P. crassicornis Just, 1980 A-At (STL) Parametopella cypris (Holmes 1905) At (V -C) G (W FL) P. inquilina Watling, 1976 At (C-V)G(FL) P. nitiis Barnard, 1962 P (CAL) P. texensis McKinney et al, 1978 G (W FL ) P. cf. texensis McKinney, Kalke & Holland, 1978 G(FL) Proboloides holmesi Bousfield, 1973 At (V) P. nordmanni (Stephensen, 1931) A-At P. pacifica (Holmes, 1908) P (CAL, deep) P. tunda Barnard, 1962 P (CAL) Raumajara carinata (Shoemaker, 1955) P, A (BAR) Stenothoe alaskensis Holmes, 1904 P (BER) S . brevicornis Sars, 1882 A-G? S. estacola Barnard, 1962 P (CAL) S. frecanda Barnard, 1962 P (CAL) S. georgiana Bynum & Fox, 1977 At (C); G (FL) S. gallensis Walker , 1904 G (FL) S. marina Bate, 1857 P (CAL?) -At? S. minuta Holmes, 1905 G(FL) S. monoculoides Montagu, 1815 At (STL) S. symbiotica Shoemaker, 1956 G(FL) S. valida Dana, 1852 P (CAL) Stenothoides bicoma Barnard, 1962 P (CAL) S. burbancki Barnard, 1969 P (CAL) Stenula incola Barnard, 1969 P (CAL) S. modosa Barnard, 1962 P (CAL) 5. nordmanni (Stephensen, 1931) P (BAR), A-At (STL) S. peltata (S. I. Smith, 1874) A-At (ST L, slope) Zaikometopa erythrophthalmus (Coyle & Mueller, 1981) P(AL) Superfamily Iphimedioidea Lowry & Myers, 2000 Family Epimeriidae Boeck, 1871 (=Paramphithoidae Sars, 1895) Epimeria cor a Barnard, 1971 P (deep) E. longispinosa K.H. Barnard, 1916 At (E FL deep) E. loricata G.O Sars, 1879 A-At (G-BF) E. obtusa Watling, 1981 At (C -E FL) E. yaquinae McCain, 1971 P(ORE, deep) Paramphithoe hystrix Ross, 1835 P-A-At (G, N, slope) P. polyacantha (Murdoch, 1885) A Ushakoviella echinophora Gurjanova, 1955 P(BER-SEAL) Family Iphimedidae Boeck, 1871 Acanthonotozoma inflatum (Kroyer, 1842) A. monodentatus Kudrjaschov, 1965 A. rusanovae Bryazhgin, 1974 A-At P (BER?) P (BC-AL), At (STL) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 97 Acanthonotozoma serratum (Fabricius, 1780) A- At (ST L) A. sinuatum Just, 1978 A-At (ST L) Coboldus hedgpethi (Barnard, 1969) P (CAL) Curidia debrogania Thomas, 1983 G (FL) Iphimedia rickettsi Shoemaker, 1931 P (AL) I. zora Thomas & Barnard, 1991 G (FL) Family Odiidae Coleman & Barnard, 1991 Cryptodius kelleri (Bruggen, 1907) C. unguidactylus Moore, 1992 Imbrexodius oclairi Moore, 1992 P(BC-CAL) P(BC) P (BC) Incerta sedis Family Lafystiidae Sars, 1895 Lafystius acuminatus Bousfield, 1987 L. frameae Bousfield, 1987 L. morrhuana Bousfield, 1987 L. sturionis Kroyer, 1842 Paralafystius mcallisteri Bousfield, 1987 Protolafystius madillae Bousfield, 1987 AT (V), G (FL?) AT (V), G (FL?) A-At (AC) A-At (AC) P(SEAL-BC) P (BC) Superfamily Dexaminoidea Bousfield, 1979 Family Atylidae G. O. Sars, 1882 Subfamily Atylinae Boeck, 1871; revised Bousfield & Kendall 1994 Atylus carinatus (Fabricius, 1793) P (BER)-A-At (STL) A. atlassovi (Gurjanova, 1951) P (BER)-A A. borealis Bousfield & Kendall, 1994 P(SEAL-WA) A. bruggeni (Gurjanova, 1938) P (BER)-A A. collingi (Gurjanova, 1938) P (BER)-A A. georgianus Bousfield & Kendall, 1994 P(BC-ORE) A. melanops (Oldevig, 1959) A A. nordlandicus Boeck, 1871 A A. rylovi (Bulycheva, 1952) P (W PAC) A. tridens (Alderman, 1936) P(BC-CAL) Subfamily Nototropinae Bousfield & Kendall, 1994 Aberratylus aberrantis (J. L. Barnard, 1962) P (CAL?, deep) Nototropis minikoi (Walker, 1905) At (V-C), G(ER?) N. smithi Goes, 1866 A, At? N. swammerdamii (Milne-Edwards, 1830) AT (AC-V), G? N. urocarinatus McKinney, 1980 G (FL-TEX) Subfamily Lepechinellinae Schellenberg, revised Barnard & Karaman 1991 Lepechinella bieri Barnard, 1957 P (CAL, deep) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 98 Subfamily Anatylinae Bulycheva, 1955; revised Bousfield & Kendall, 1994 Kamehatylus nani Barnard, 1970 P (HAW) Family Dexaminidae Leach, 1813/14 Subfamily Polycheriinae Bousfield & Kendall, 1994 Polycheria osborni Caiman, 1898 P (SE AL-CAL) P. carinata Bousfield & Kendall, 1994 P (BC) P. mixillae Bousfield & Kendall, 1994 P (BC) Subfamily Prophliantinae Nicholls, 1939 Guernea nordenskioldii (Hansen, 1887) A-At (AC) G. reduncans (Barnard, 1958) P (BC-CAL) Subfamily Dexamininae Leach, 1813/14; revised Bousfield & Kendall, 1994 Dexamine thea Boeck, 1861 At (AC) Superfamily Ampeliscoidea Bousfield, 1979 Family Ampeliscidae Costa, 1857 Ampelisca abdita Mills, 1964 At (ST L, V-C), G (?) A. aequicornis Bruzelius, 1859 At-A (AC) A. agassizi (Judd, 1896A) (= A. vera Barnard, 1954) P (CAL)-At (V), G (E-FL) A. amblyops Sars, 1891 At (FL, deep) A. amblyopsoides J. L. Barnard, 1960 P (CAL) A. bicarinata Goeke & Heard, 1983 G (FL-MI) A. birulai Briiggen, 1909 P (BER), A A. brachycladus Roney, 1990 P (CAL?) A. brevisimulata Barnard, 1954 P(ORE-BC) A. burkei Barnard & Thomas, 1989 G (FL) A. carey i Dickinson, 1982 P(BC-ORE) A. ciego Barnard, 1966 P (CAL) A. coeca Holmes, 1908 P (S CAL) A. cristata Holmes, 1908 P (BC-ORE) A. cristoides Barnard, 1954 P (S CAL) A. declivitatus Mills, 1967 At (deep) (STL) A. eoa Gurjanova, 1951 P (BER) A. erythrorhabdota Coyle & Highsmith, 1989 P (BER) A. eschrichti Kroyer, 1842 P (BER?)- A- At (ST L) A. fageri Dickinson, 1982 (= A. schellenbergi Shoemaker, 1933 P (ORE) A. furcigera Gurjanova, 1936 P (BER) A. gibba Sars, 1882 At (ACdeep) A. hancocki Barnard, 1954 P (BC-ORE) A. hessleri Dickinson, 1982 P(ORE) A. holmesi Pearse, 1908 G (FL-MI AMPHIPACIFICA V0L.3N0.1 MAY 16, 2001. 99 A. indentata Barnard, 1954 P(CAL) A. latipes Stephensen, 1925 At (STL-AC) A. lobata Holmes, 1908 (- A. articulata Stout, 1913) P (AL) A. macrocephala Liljeborg, 1852 P (BER)-A-At (AC) A. mexicana Barnard, 1954 P (S CAL) A. milleri Barnard, 1954 P (CAL) A. pacifica Holmes, 1908 P (CAL) A. plumosa Holmes, 1908 P (AL) A. pugetica Stimpson, 1864 P(BC-WA) A. rornigi Barnard, 1954 P (CAL) A schellenbergi (see Coyle & Highsmith, 1989) P(BER) A. shoemakeri Barnard, 1954 P (CAL) A. typica (Bate, 1856) AT (AC) A. uncinata Chevreux, 1887 At (AC, deep) A. unsocolae Barnard, 1960 P(ORE) A. vadorum Mills, 1963 At (G, V)(E FL?) A. venetiensis Shoemaker, 1916 P (CAL) A. verrilli Mills, 1967 At (V-C?)(E FL?) Byblis barbarensis Barnard, 1960 P (CAL) B. bathyalis Barnard, 1966 P(CAL) B. brevirama Dickinson, 1983 P (ORE),- A B. crassicornis Metzger, 1875 P (BER?) B. frigidis Coyle & Highsmith, 1989 P(BER) B. gaimardii (Kroyer, 1846)? P (BER?)- A- At (AC) B. longispina Dickinson, 1983 P(BC) B. medialis Mills, 1971 At (AC, deep) B. mill si Dickinson, 1983 P (BC) B. mulleni Dickinson, 1983 P(ORE) B. pearcyi Dickinson. 1983 P (BER),- A B. robustus Coyle & Highsmith, 1989 P (BER) B. serrata S. I. Smith, 1873 At (V), G (E FL?) B. tannerensis Barnard, 1966 P (CAL) B. teres (see C. & H., 1989) P (BER) B . thyabilis Barnard, 1971 P(ORE) B. veleronis Barnard, 1954 P(BC-CAL) Haploops fundiensis Wildish & Dickinson, 1982 At (AC) H. laevis Hoek, 1882 P (CAL)-A-At (ST L) H. sibirica Gurjanova, 1929 A H. lodo Barnard, 1961 P (CAL?) H. setosa Boeck, 1871 P (ER), At (AC) H. similis Stephensen, 1925 At (AC, shelf to deep) H. spinosa Shoemaker, 1931 At (AC) H. tubicola Liljeborg, 1856 P (BER)-A-At (ST L) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 100 Superfamily Melphidippoidea Bousfield, 1979 [= cheirocratids Barnard & Barnard, 1983 (part)] Family Melphidippidae Stebbing, 1899 Casco bigelowi (Blake, 1929) Melphisana bola Barnard, 1962 Melphidipella macer (Norman, 1869) Melphidippa amorita Barnard, 1966 M. borealis Boeck, 1971 M. goesi Stebbing, 1899 M. macrura G. O. Sars, 1894 At (ST L-AC-DEL) P(AL-CAL) P(BC) P (CAL)? P-A?-At (ST L) A-At (AC) At (ST L) Family Hornelliidae Bousfield, 1982 Horne Ilia (Metaceradocus) tequestae Thomas & Barnard, 1986 G (FL) H. occidentalis (Barnard, 1959) P (S CAL) Family Megaluropidae Thomas & Barnard, 1986 Megaluropus longimerus Schellenberg, 1925? Gibber os us devaneyi Thomas & Barnard, 1986 G. myersi (McKinney, 1980) G. visendus (Barnard, 1969) Resupinus coloni Thomas & Barnard, 1986 P (BC-CAL) P (S CAL?) P (CAL?); G (FL-TEX) P (B CAL) P (CAL) Superfamily Liljeborgioidea, Bousfield, 1979 Family Liljeborgiidae Stebbing, 1899 Idunella aequicornis (Sars, 1876) A-At (ST L) /. bowenae Karaman, 1979 At (V, shelf) I. smithi Lazo-Wasem, 1985 At (V), G (E FL?)) Liljeborgia bousfieldi McKinney, 1979 G (FL-TEX) L. cota Barnard, 1962 P (ORE-CAL, deep) L. fissicornis M. Sars, 1858?) A-At (N, slope) L. geminata Barnard, 1969 P (CAL?) L pallida (Bate, 1857) P (CAL)? G (FL) Listriella albina Barnard, 1959 P (ORE-CAL, deep) L. barnardi Wigley, 1966 At (V-C), G (W FL) L. carinata McKinney, 1979 G (FL-TEX) L. clymenellae Mills, 1962 At (V-C), G (FL?) L. diffusa Barnard, 1959 P (CAL) L. eriopisa Barnard, 1959 P (S CAL) L. goleta Barnard, 1959 P (ORE-CAL) L. melanica Barnard, 1959 P (CAL) L. quintana McKinney, 1979 G (TEX) Family Sebidae Walker, 1908 Subfamily Sebinae Holsingerl980 Seba aloe Karaman, 1971 G (W FL) S. profunda Shaw, 1989 P (BC, deep) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 101 Subfamily Seborgiinae Karaman, 1992 Relictoseborgia hershleri (Holsinger, 1992b) FW (TEX) 1R. relicta (Holsinger, 1980) FW (TEX) Family Colomastigidae Stebbing, 1899 Colomastix bousfieldi LeCroy 1995 G (FL-TEX) C.. camura LeCroy, 1995 At (C), G (FL-TEX) C. cornuticauda LeCroy, 1995 G (W FLA) c. denticornis LeCroy, 1995 G(WFL) c. falcirama LeCroy, 1995 G(FL) c. gibbosa LeCroy, 1995 G (FL) c. halichondriae Bousfield, 1973 At, G (FL-TEX) c. heardi LeCroy, 1995 AT (C), G (FL-YUC) c . irciniae LeCroy, 1995 G(FL) c. janiceae Heard & Perlmutter, 1977 G (FL-YUC), At (C) c. tridentata LeCroy, 1995 At (C),G (FL-YUC) Superfamily Crangonyctoidea Bousfield 1973 [= crangonyctoids Barnard & Barnard, 1983 (part)] Family Crangonyctidae Bousfield 1973 (revised Holsinger 1977) Bactrurus brachycaudus Hubricht & Mackin,1940 FW B. hubrichti Shoemaker, 1945 FW B . mucronatus (Forbes, 1876) FW Crangonyx aberrans D. Smith, 1983 FW C. alpinus Bousfield, 1963 FW (P) C. anomalus Hubricht, 1943 FW C. antennatus Packard, 1881 FW C. dearolfi Shoemaker, 1942 FW C. floridanus Bousfield, 1963 FW C. forbesi (Hubricht & Mackin, 1940) FW C. gracilis Smith, 1871 FW C. grandimanus Bousfield, 1963 FW C. hobbsi Shoemaker, 1941 FW C. minor Bousfield, 1958 FW C. obliquus (Hubricht & Mackin, 1940) FW C. packardi S. I. Smith, 1888 FW c. pseudo gracilis Bousfield, 1958 FW c. richmondensis richmondensis Ellis, 1940 FW c. r. occidentalis Hubricht & Harrison, 1941 FW (P) c r. laurentianus Bousfield, 1958 FW c rivularis Bousfield, 1958 FW c. serratus (Embody, 191 1) FW c. setodactylus Bousfield, 1958 FW c. shoemakeri (Hubricht & Mackin, 1940) FW Stygonyx courtneyi Bousfield & Holsinger, 1989 FW (P) Stygobromus abditus Holsinger, 1978 FW 5. ackerlyi Holsinger, 1978 FW s. alabamensis alabamensis (Stout, 1911) FW AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 102 s . a. occidentalis (Hoi singer, 1967) FW s. allegkeniensis (Holsinger, 1967) FW s. araeus (Holsinger, 1969) FW s. arizonensis Holsinger, 1974 FW s. balconis (Hubricht, 1943) FW s. baroodyi Holsinger, 1978 FW s. barri (Holsinger, 1967) FW s. barryi Holsinger, 1978 FW s. bifur catus (Holsinger, 1967) FW s . bigger si Holsinger, 1978 FW 5. borealis Holsinger, 1978 FW 5. bowmarti (Holsinger, 1967) FW 5. canadensis Holsinger, 1980 FW 5. Caroline ns is Holsinger, 1978 FW 5. clantoni (Greaser, 1934) FW 5. coloradensis Ward, 1977 FW S . conradi (Holsinger, 1967) FW 5. cooperi (Holsinger, 1967) FW 5. cumberlandus Holsinger, 1978 FW 5. dejectus (Holsinger, 1967) FW 5. dicksoni Holsinger, 1978 FW 5. elatus (Holsinger, 1967) FW 5. elliotti Holsinger, 1974 FW 5. emarginatus (Hubricht, 1943) FW 5. ephemerus (Holsinger, 1969) FW 5. estesi Holsinger, 1978 FW S . exilis Hubricht, 1943 FW 5. fecundus Holsinger, 1978 FW S. ferausoni Holsinger, 1978 FW S. finleyi Holsinger, 1978 FW 5. flagellatus (Benedict, 18%) FW 5. franzi Holsinger, 1978 FW 5. gracilipes (Holsinger, 1%7) FW 5. gradyi Holsinger, 1974 FW 5. grahami Holsinger, 1974 FW S. grandis Holsinger, 1978 FW S . hadenoecus (Holsinger, 1966) FW 5. harai Holsinger, 1974 FW 5. hayi (Hubricht & Mackin, 1940) FW 5. heteropodus Hubricht, 1943 FW 5. hoffmani Holsinger, 1978 FW 5. holsinger i Ward, 1977 FW S. hubbsi Shoemaker, 1942 FW s. indentatus (Holsinger, 1%7) FW s. inexpectatus Holsinger, 1978 FW s. interitus Holsinger, 1978 FW S' iowae Hubricht, 1943 FW S' kenki Holsinger, 1978 FW S. lacicolus Holsinger, 1974 FW S' leensis Holsinger, 1978 FW S. longipes (Holsinger, 1966) FW Co Co CO OO AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 103 S. lucifugus (Hay, 1882) S. mackenziei Holsinger, 1974 S. mackini Hubricht, 1943 S. minutus Holsinger, 1978 S. montanensis Holsinger, 1974 S montanus (Holsinger, 1967) S. morrisoni (Holsinger, 1967) S. mundus (Holsinger, 1967) S. my stic us Holsinger, 1974 S. nanus Holsinger, 1978 S. nortoni (Holsinger, 1969) S. obrutus Holsinger, 1978 S. obscurus Holsinger, 1974 S. onondagaensis (Hubricht & Mackin, 1940) S oregonensis Holsinger, 1974 S. ozarkensis (Holsinger, 1967) S. parvus (Holsinger, 1969) S. pecki (Holsinger, 1967) S. pennaki Ward, 1977 S. phreaticus Holsinger, 1978 S. pizzinii (Shoemaker, 1938) S. pollostus Holsinger, 1978 S . pseudospinosus Holsinger, 1978 S. putealis (Holmes, 1909) S. puteanus Holsinger, 1974 S. quatsinensis Holsinger & Shaw, 1987 S. redactus Holsinger, 1978 S. reddelli (Holsinger, 1966) S. russelli (Holsinger, 1967) S. secundus Bousfield & Holsinger, 1981 S. sheldoni Holsinger, 1974 S. sierrensis Holsinger, 1974 S. mithi Hubricht, 1943 5. sparsus Holsinger, 1978 (1969?) S. spinatus (Holsinger, 1967) S. spinosus (Hubricht & Mackin, 1940) S. stegerorum Holsinger, 1978 S. stellmacki (Holsinger, 1967) S . subtilis (Hubricht, 1943) S . tahoensis Holsinger, 1974 S. tenuis tenuis (S. I. Smith, 1874) S. t. potomacus (Holsinger, 1967) S. tritus Holsinger, 1974 S. vitreus Cope, 1872 S wengerorum Holsinger, 1974 Synpleonia pizzini Shoemaker, 1941 Synurella chamberlaini Shoemaker, 1936? bifurca (Hay, 1882) chamberlaini (Ellis, 1941) dentata Hubricht, 1943 johanseni Shoemaker, 1920 AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 104 Superfamily Talitroidea Bulycheva, 1957 Family Hyalidae Bulycheva, 1957 Apohyale pugettensis (Dana, 1853) A . anceps (Barnard, 1969) A. californica (Barnard, 1969) Hyale media (Dana, 1853) H. nilssoni Rathke 1843 H. oculata Bousfield, 1981 H. perieri (Lucas, 1846) Leptohyale longipalpa Bousfield 1981 Parallorchestes ochotensis (Brandt, 1851) P. brevicornis Bousfield, 1981 P. minor Bousfield, 1981 P. spinosa Bousfield, 1981 P. subcarinata Bousfield, 1981 P. supracarinata Bousfield, 1981 P. trispinosa Bousfield, 1981 P. nuda Bousfield, 1981 P. americana Bousfield, 1981 P. minima Bousfield, 1981 P. occidentalis Bousfield, 1981 P. subcarinata Bousfield, 1981 Par hyale hawaiensis (Dana, 1853) P. fascigera Stebbing, 1897 Plumulohyale plumulosa (Stimpson, 1857) Protohyale frequens (Stout, 1913) P. canalina Barnard, 1979 P. nigra (Haswell, 1879) P, lagunae (Stout, 1913) P. intermedia (Bousfield, 1981) P. seticornis (Bousfield, 1981) P. oclairi (Bousfield, 1981) P. spinosa (Bousfield, 1981) P (SE AL-CAL) P(CAL-BC) P(BC-CAL) G(FL) At (AC) P(BC) G(FL) P(BC) P (AL-BC) P (AL-BC) P(BC) P(BC)? P(SEAL-WA) P(BER) P(BC) P(BC) P(AL) P (BC) P(BC) P (SE AL-WA)C G(FL) G(FL) P (BC-CAL), At (V), G (FL?) P(BC-CAL) P (S CAL?) P(CAL) P(S CAL) P(SE AL-ORE) P(SE AL-CAL) P(SE AL-WA) P(SE AL-BC) Family Hyalellidae Bulycheva, 1957 Subfamily Hyalellinae Bousfield, 1996 Allorchestes angusta Dana, 1853 P (AL-CAL) A. bella bella Barnard, 1974 P (BER-CAL) A.. pacifica Bousfield, 1981 P(BC) A. parva Bousfield, 1981 P(BC) A. subcarinata Bousfield, 1981 P(AL) A. urocarinata Bousfield, 1981 P(SE AL-BC) A. carinata Iwasa, 1939 (Bousfield, 1981) P(BER) Hyalella (Hyalella) azteca (Saussure, 1858) FW H. (H.) inermis S. I. Smith, 1974 FW H. (H.) longicornis Bousfield, 1996 FW H. (H.) muerta Baldinger, Shepard, & Threloff 2000 FW AMPHIPACIHCA VOL. 3 NO. 1 MAY 16, 2001. 105 Hyalella (H.) montezuma Cole & Watkins, 1977 FW H. (H.) sandra Baldinger, Shepard, & Threloff 2000 FW H. (H.) texana Stevenson & Peden, 1973 FW Parhyalella whelpleyi (Shoemaker, 1933) G (FL) Family Dogielinotidae Gurjanova, 1953 Proboscinotus loquax (Barnard, 1968) P (ORE-WA) Family Najnidae J. L. Barnard, 1972 Najna consiliorum Derzhavin, 1937 P (BER) N. kitimati Barnard, 1979 P (CAL) N. lessoniophilum Bousfield, 1981 P(CAL) N. rugosum Bousfield, 1981 P(AL-BC) N. setosum Bousfield, 1981 P (BC-ORE?) N. plumulosum Bousfield, 1981 P (BC-ORE?) Family Eophliantidae Sheard, 1936 Lignophliantis pyrifera Barnard, 1969 P (S CAL) Family Phliantidae Stebbing, 1899 Pariphinotus (Heterophlias) escabrosus (Barnard, 1969) P (BC-CAL) P. (H.) seclusus (Shoemaker, 1933) At (C), G (FL) Family Talitridae Raflnesque, 1815 (a) Palustral subgroup (pragmatic subfamily group, Bousfield, 1984) Uhlorchestia uhleri (Shoemaker, 1930) At (C-FL)), G (FL-TEX) U. spartinophila Bousfield & Heard, 1986 At (V-C), G (FL) (b) Beachflea subgroup (Bousfield, 1984) Orchestia gammarella (Pallas, 1766) Orcheslia grillus Bose, 1802 Paciforchestia klawei (Bousfield, 1959) Platorchestia chathamensis Bousfield, 1982 P. plate nsis (Kroyer, 1845) Tethorchestia sp 1 (- tropica Shoemaker MS) Tethorchestia brevipleopoda (Bousfield MS) Traskorchestia traskiana (Stimpson, 1856) T. georgiana (Bousfield, 1958) T. ochotensis (Brandt, 1851) Transorchestia enigmatica (Bousfield & Carlton, 1968) At (AC) At (AC-V); G (FL-TEX) P(S CAL-B.C) P(BC) At (AC-V), G (FL-TEX) G(FL) G(FL) P(Al-CAL) P(CAL-BC) P (ALEUT) P (CAL, intr.) (c) Sand hopper subgroup (Bousfield, 1984) Americorchestia longicornis (Say, 1818) A. barbarae Bousfield, 1992 At (AC-V), G G (TEX) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 106 Americorchestia heardi Bousfield, 1992 A. megalophthalma (Bate, 1862) A. salomani Bousfield, 1992 Megalorchestia calif orniana (Brandt, 1851) M. M. M. M. M M. columbiana (Bousfield, 1958) minor (Bousfield, 1957) dexterae Bousfield, 1982 pugettensis (Stimpson, 1856) corniculata (Stout, 1912) benedicti (Shoemaker, 1936) G (FL-LA) At(AC-C) G (FL-LA) P P P(S CAL) P (S-B CAL) P P (CAL) P (CAL) (d) Landhopper subgroup (Bousfield, 1984) Arcitalitrus sylvaticus (Haswell, 1879) Talitroides topitotum (Burt, 1934?) T. alluaudi (Chevreux, 1896) P (CAL, intr.) P (CAL, intr.), G (FL-MI, intr.) P (BC - CAL), At (intr.), G (FL) Superfamily Pontoporeioidea Bousfield, 1979 [= Haustorioidea Barnard & Drummond, 1982 (part)] Family Bathyporeiidae Bousfield, 1978 Amphiporeia gigantea Bousfield, 1973 A. lawrenciana Shoemaker, 1929 A. virginiana Shoemaker, 1933 Bathyporeia parkeri Bousfield, 1973 B. quoddyensis Shoemaker, 1949 At (AC) At (AC) At AC), G (H FL?) At (V-C)G(EFL) At (AC-V) Family Pontoporeiidae Dana, 1855 Diporeia brevicornis (Segerstrale, 1937) FW D. erythrophthalma (Waldron, 1953) FW D. filicornis (Smith, 1974) FW D. hoyi (Smith, 1874) FW D. intermedia (Segerstrale, 1977) FW D. kendalli (Norton, 1909) FW Monoporeia ajfinis (Lindstrom, 1885) A-At (ST L), P (AL) Pontoporeia femorata Kroyer, 1842 P (AL-BC)-A-At (ST L-AC) Priscillina armata (Boeck, 1861) A-At (STL -AC) Family Haustoriidae Stebbing, 1906 Acanthohaustorius bousfieldi Frame, 1982 At (V) A. cf. bousfieldi Frame, 1980 G(EFL) A. intermedius Bousfield, 1965 At (V-C) A. nr. intermedius Bousfield, 1965 G (FL?) A. mill si Bousfield, 1965 At (V-C) G (E. FL) A. uncinus Foster, 1988 G (FL-MI) A. pansus Thomas & Barnard, 1984 G(FL) A. shoemakeri Bousfield, 1965 At (V-C) A. cf. shoemakeri Bousfield 1965 G (NW FL) A. similis Frame, 1980 At (V-C) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 107 A. spinosus (Bousfield, 1962) At (AC-Del) A. uncinus Foster, 1989 G (FL-MI) Eohaustorius brevicuspis Bosworth, 1973 P(BC-ORE) E. ecus (Gurjanova, 1951) P(BER) E. sencillus Barnard, 1962 P (CAL) E. washingtonianus (Thorsteinsen, 1941) P ( AL-CAL?) E. estuarius Bosworth, 1973 P (BC-ORE) E. sawyer i Bosworth, 1973 P (BFC-ORE) Haustorius canadensis Bousfield, 1962 At (SW G-V) (G (FL?) H. jayneae Foster & LeCroy, 1991 G(NE) Lepidactylus dytiscus Say, 1818 G (EFL) L. triarticulatus Robertson & Shelton, 1980 G (FL-TEX) Neokaustorius biarticulatus Bousfield, 1965 At (V-C), G (E FL?) N. schmitzi Bousfield, 1965 At (V-C), G (EFL) Parahaustorius attenuatus Bousfield, 1965 At (V) P. holmesi Bousfield, 1965 At (AC) G (FL?) P. longimerus Bousfield, 1965 At (V-C)G(FL?) P. cf. longimerus Bousfield, 1965 G (W. FL) P . obliquus Robertson & Shelton, 1978 G (FL-TEX) Protokaustorius bousfieldi Robertson & Shelton, 1978 G (FL-TEX) P. deichmannae Bousfield, 1965 At (V) G (FL?) P. wigleyi Bousfield, 1965 At (V) G (FL?) Pseudohaustorius americanus (Pearse, 1908) G (FL-MI) P. borealis Bousfield, 1965 At (V) P. caroliniensis Bousfield, 1965 At V-CAR-E FL?) Superfamily Gammaroidea Bousfield, 1977 [= gammaroid group Barnard & Barnard, 1983 Family Gammaridae Leach, 1813 Chaetogammarus stoerensis (Reid, 1938) At (AC) C. ischnus (Sars, 1896) FW (intr.)(Witt, etai, 1998)) Eulimnogammarus obtusatus (Dahl, 1938) At (AC-ST L) Gammarus acherondytes Hubricht & Mackin, 1940 FW G. annulatus S. I. Smith, 1874 At (STL- AC) G. bousfieldi Cole & Minckley 1961 FW G. daiberi Bousfield, 1969 P (CAL, intr.)-At (V-C)-G? G. desperatus Cole, 1981 FW G. duebeni Liljeborg, 1851 At (AC) G. fasciatus Say, 1818 FW G. hyalelloides Cole, 1976 FW G. jenneri Bynum & Fox, 1977 At (V-C) G. lacustris lacustris Sars, 1864 FW G. lawrencianus Bousfield, 1956 A-At G. limnaeus S. 1. Smith, 1874 FW G. minus minus Say, 1818 FW G. minus pinicollis Cole, 1976 FW G. paynei Delong, 1992 FW G. pecos Cole & Bousfield, 1970 FW G. pseudohmnaeus Bousfield, 1958 FW G. tigrinus Sexton, 1939 At (AC-C)G (FL-LA) AMPHIPACIHCA VOL. 3 NO. 1 MAY 16, 2001. 108 G. troglophilus Hubricht & Mackin, 1940 G. (Lagunogammarus) oceanicus (Segerstrale, 1947) G. (L.) setosus Dementieva, 1931 G. (L.) wilkitzkii (Birula, 1897) G. ( Mucrogammarus ) mucronatus (Say, 1818) G. (M.). palustris Bousfield, 1969 Marinogammarus finmarchicus Dahl, 1938 FW At(A-AC) P (AL-BC)-A-At A At (AC-V-C), G(FL), P (Salton Sea) G(FL?) At (AC-V-C) Family Anisogammaridae Bousfield, 1977 Anisogammarus pugettensis pugettensis (Dana, 1853) P(AL-CAL) A. amchitkana Bousfield, 2001 P(AL-) A. epistomus Bousfield, 2001 P(BC) A. slattery i Bousfield, 2001 P(BER-WA) Barrow gammarus mcginitiei (Shoemaker, 1955) P-A (BAR) Carineogammarus makarovi (Bulycheva, 1952) P (SE AL) Eogammarus oclairi Bousfield, 1979 P(BC-ORE) E. confervicolus (Stimpson, 1856) P (SEAL- CAL) E. psammophilus Bousfield, 1979 P (ALEUT) Locustogammarus levingsi Bousfield, 1979 P(SEAL-BC) L. locustoides (Brandt, 1851) P(AL-BC) Ramellogammarus campestris Bousfield & Morino, 1992 FW P (ORE) R . californicus Bousfield & Morino, 1992 FW P (CAL) R . columbianus Bousfield & Morino, 1992 FW P (BC-ORE) R. oregonensis (Shoemaker, 1944) FW P (ORE) R. ramellus (Weckel, 1907) FWP(CAL) R . similimanus (Bousfield, 1961) FW P (ORE) R. setosus Bousfield & Morino, 1992 FW P (ORE) R. littoralis Bousfield & Morino, 1992 FW P (ORE) R. vancouverensis Bousfield, 1979 FWP(BC) Spinulogammarus subcar inatus (Bate, 1862) P(AL-BC) Spasskogammarus tzvetkovae Bousfield, 1979 P(BER) FamUy Gammaroporeiidae Bousfield, 1977 Gammaroporeia alaskensis (Bousfield & Hubbard, 1968) P (SE AL) FamUy Mesogammaridae Bousfield, 1977 Paramesogammarus americanus Bousfield, 1979 P (SE AL) Superfamily Hadzioidea Bousfield, 1977 [= hadzioids Barnard & Barnard, 1983] FamUy Allocrangoncytidae Holsinger, 1989 Allocranqonyx hubricht i Hoi singer, 1971 FW A. pellucidus (Mackin, 1935) FW FamUy Hadzudae S. Karaman, 1933 Wecke lu d sub g roup (Ho l sin g e r . 1 99 2) (= weckeliids of Barnard & Barnard, 19831 Allotexiweckelia hirsuta Holsinger, 1980 FW (TX) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 109 Holsingerius samacos (Hoi singer, 1980) FW H. smaragdinus Holsinger, 1992b FW Mexiweckelia hardeni Holsinger, 1992b FW Paramexiweckelia ruffoi Holsinger, 1993 FW Texiweckelia texensis (Holsinger, 1973) FW (TEX) Texiweckeliopsis insolita (Holsinger, 1980) FW (TEX) Harlziid si i bar oun (— hadziids of Barnard & Barnard* 1983) Dulzura sal J. L. Barnard, 1969 P (CAL) Protohadzia sp. Zimmerman & Barnard, 1977 G (FL?) P. sckoenerae (Fox, 1973) At (C); G (FL) Metaniphargus beattyi Shoemaker, 1942 G (FL?) Netamelita barnardi McKinney et al, 1978 G (TEX) N. brocha Thomas & Barnard, 1991c G(FL) N. cortada Barnard, 1962 P(S CAL) Spathiopsis looensis Thomas & Barnard, 1985 G (FL) Gammarellas l=ntiuanids Barnard & Barnard. 1983) Tabatzius muelleri (Ortiz, 1976) G (FL-YUC) T. copillius (McKinney & Barnard, 1977) G (FL7-YUC) Family Melitidae Bousfield, 1973 [= melitids + ceradocids sensu Barnard & Barnard, 1983 (part)} Abludomelita obtusa (Monatagu, 1813) P? (WA?), At (ST L) Ammaera hixorti Thomas & Barnard, 1985 G(FL) Bathyceradocus tore Hi (Goes, 1966) P (bathyal), A-At (ST L, deep) Ceradocus colei (Kunkel, 1910) At (V) C. paucidentatus Barnard, 1952 P (CAL) C. rubromaculatus (Stimpson, 1856) P C. sheardi Shoemaker, 1948 G (W FL) C. shoemakeri Fox, 1973 At (C); G (FL) C. spinicauda (Holmes, 1908) P(BC-CAL) Denticeradocus sp. (see Barnard, 1952) P (CAL) Desdimelita barnardi Jarrett & Bousfield, 1996 P (BC) D. desdichada (J. L. Barnard, 1962) P(SEAL-CAL) D. calif ornica (Alderman, 1936) P(AL-CAL) D. microdentata Jarrett & Bousfield, 1996 P(SE AL-ORE) D. microphthalma Jarrett & Bousfield, 1996 P(SE AL) D. transmelita Jarrett & Bousfield, 1996 P (BC) Dulichiella appendiculata (Say, 1818) P (SCAL);At (C); G(FL-LA) Elasmopus antermatus (Stout, 1913) P(SEAL-CAL) E. balcomanus Thomas & Barnard, 1988 G(FL) E. bampo Barnard, 1979 P (CAL) E. holgurus Barnard, 1962 P(CAL) E . lemaitrei Ortiz, 1994 G (FL? CUBA) E. levis (S. I. Smith, 1873) At (V-C), G (FL) E. mutatus Barnard, 1962 P (WA -CAL) E. pectenicrus (Bate, 1862) G(FL) E. pocillimanus (Bate, 1862) G (FL) E . serricatus Barnard, 1969 P(S CAL) E. thomasi Ortiz, 1994 G(FL-CUBAt) Eriopisa elongata Bruzelius, 1859 P (CAL) -At (V-C, shelf), G (FL?) AMPHIPACIFICA V0L.3N0.1 MAY 16, 2001. 110 E incisa McKinney, Kalke & Holland, 1978 E. schoenerae Fox, 1973 Eriopisa sp. (Barnard, 1952) Jerbarnia americana Watling 1981 Lupimaera lupana (Barnard, 1969) Maera danae (Stimpson, 1853) M. cf. danae Krapp-Schickel & Jarrett. 2000 M. loveni (Bruzeiius, 1859) M. fusca (Bate, 1864) M. nelsonae Krapp-Schickel & Jarrett, 2000 M. bousfieldi Krapp-Schickel & Jarrett, 2000 M.. jerrica Krapp-Schickel & Jarrett, 2000 M. similis Stout, 1913 Maera diffidentia J. L. Barnard, 1969 M.. rathbunae Pearse, 1908 Maera sp. (nr. rathbunae ) Krapp-Schickel & Jarrett, 2000 M. grossimana (Montagu, 1808)? M. prionochira Bruggen, 1907 M. quadrimana (Dana, 1853) M. re is hi Barnard, 1979 M. serrata Schellenberg, 1938 M. sulca (Stout, 1913) M. williamsi Bynum & Fox, 1977 Megamoera arnoena (Hansen, 1887) M. bowmani Jarrett & Bousfield, 1996 M. borealis Jarrett & Bousfield, 1996 M. dentata (Kroyer, 1842) M. glacialis Jarrett & Bousfield, 1996 M. kodiakensis (Barnard, 1964) M. mikulitschae (Gurjanova, 1953) M. rafiae Jarrett & Bousfield, 1996 M. subtener (Stimpson, 1856) M unimaki Jarrett & Bousfield, 1996 Melita alaskensis Jarrett & Bousfield, 1996 M. intermedia Sheridan, 1980 M. elongata Sheridan, 1979 M. longisetosa Sheridan, 1979 M. nitida (S. I. Smith, 1874) M. oregonensis Barnard, 1954 M. shoemakeri (= M . nitida Shoemaker, 1936) M. sulca (Stout, 1913) Melitoides makarovi Gurjanova, 1934 M. valida (Shoemaker, 1964) Quadrimaera carla Krapp-Schickel & Jarrett, 2000 ?Q- vigota Barnard, 1969 Quasimelita quadrispinosa (Vosseler, 1889) Q. formosa (Murdoch, 1885) Spathiopus looensis Thomas & J. L. Barnard, 1985 G (TEX) G(FL?) P(S CAL) At (C-E FL)-,G (FL) P (CAL) P-At (AC) P(AL-SEAL) P (Al-WA)-A-At (N) P(AL-WA) P (BER-CAL) P(BC-CAL) P (SEAL- ORE) P(BC-MEX) At (NC-G(FL) G(FL-MI) P (BC - At (NC) P(BC-ORE) P(AL) G (CUBA-FL ) P (CAL?) G (CUBA-FL?) P(S CAL) At (C); G (FL) A P(SEAL) P(SEAL) P-A-At P(SEAL) P(SEAL) P(BER) P(SEAL) P(BC-CAL) P (ALEUT) P(AL) G (W FL) G (W FL) At (V-C). G (W FL) P (intr.)-At, G (E FI) P(BC-CAL) G (YUC) P(S CAL) P(BER P(BER) P (BC - CAL) P (CAL ) PA (SE AL)-A-At (ST L) A (AL)-At (ST L) G(FL) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. Ill Superfamily Bogidielloidea Bousfield, 1977 [= bogidiellids Barnard & Barnard,, 1983 (part)] Family Atesiidae Holsinger & Longley, 1980 Artesia welbourni Holsinger, 1992b A. subterranea Holsinger, 1980 FWTEX FW TEX Family Bogidiellidae Hertzog 1936 Parabogidiella americana Holsinger, 1980 FWTEX Superfamily Corophioidea Barnard & Barnard, 1983 (revised) Family Ampithoidae Stebbing, 1899 Ampithoe dalli Shoemaker, 1938 A. divisura Shoemaker, 1933 A. kussakini Gurjanova, 1955 A. longimana (S. I. Smith, 1873) A. lacertosa Bate, 1858 A. plumulosa Shoemaker, 1938 A. ramondi Audoin, 1828 (= A. divisura?) A. rubricata (Montague, 1808) A. rubricatoides Shoemaker, 1938 A. sectimanus Conlan & Bousfield, 1982 A. simulans Alderman, 1936 A. valida S. I. Smith, 1873 A. volki Gurjanova, 1938 Cymadusa compta (S. I. Smith, 1873) C. filosa Savigny, 1816 C. uncinata (Stout, 1912) Peramphithoe eoa (Barnard, 1954) p femorata (Kroyer, 1845) P. burner alis (Stimpson, 1864) P. mea (Gurjanova, 1938) P. lindbergi (Gurjanova, 1938) p, stypotrupetes Conlan & Chess, 1992 P. plea (Barnard, 1965) p. tea (Barnard, 1965) Pleonexes aptos Barnard, 1969 Pseudamphithoides bacescui Ortiz, 1976 Sunamphitoe pelagica (Milne-Edwards, 1830) P(AL-ORE) G (FL Keys) P(AL-BC)) P?-At(V),G(WFL) P(AL-SCAL) P (BC-S CAL) P?, G (FL) At (AC) P(BER) P (SEAL-ORE) P(AL-ORE) P (BC-CAL), At (V)), G (FL) P (BER?) AT (V), G (W FL) G(FL) P (BC-CAL) P (BER?) P-At? P (SE AL-S CAL) P (ALEUT) P (BER-CAL) P(SEAL-CAL) P (BC-CAL) P (SE AL-S CAL) P (S CAL) G (FL?, CUBA) At (offshore), G Family Biancolinidae J. L. Barnard, 1972 Biancolina brassiacephala Lowry, 1974 G Family Aoridae Stebbing, 1899 Arctolembos arcticus (Hansen, 1887) Acuminodeutopus heteruropus Barnard, 1959 Aoroides columbiae Walker, 1898 A P (CAL) P (BER-CAL) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 112 A. exilis Conlan & Bousfield, 1982 A. inermis Conlan & Bousfield, 1982 A. intermedius Conlan & Bousfield, 1982 A. spinosus Conlan & Bousfield, 1982 Bemlos audbettius Barnard, 1962 B. concavus (Stout, 1913) B. mackinneyi Myers, 1978 B. macromanus Shoemaker, 1925 B. sanmartini Ortiz, Lalana & Lopez, 1992? Columbaora cyclocoxa Conlan & Bousfield, 1982 Grandidierella bonnieroides Stephensen, 1948 G. notoni Shoemaker, 1935 G. japonica Stephensen , 1938 Lembos (Arctolembos) arctica Hansen, 1887 L. ( Globosolembos) francanni Reid, 1951 L. (Globosoolembos) smithi ( Holmes, 1905) L. borealis Myers, 1976 L. bruneomaculatus brunneomaculatus Myers, 1977 L. brunneomaculatus mackinneyi Myers, 1978 L. dentischium Myers, 1977 L. hypacanthus (K. H. Barnard, 1916) L. kunkelae Myers, 1977 L. minimus Myers, 1977 L. ovalipes Myers, 1979 L. rectangulatus Myers, 1977 L. setosus Myers, 1978 L. smithi (Holmes 1905) L. spinicarpus spincarpus (Pearse, 1912) L. spinicarpus inermis Myers, 1979 L. tigris Myers, 1981 L. tigrinus Myers, 1979 L. tempus Myers, 1981 L. unicornis Bynum & Fox, 1977 L. unifasciatus unifasciatus Myers, 1977 L. unifasciatus reductus Myers, 1979 L. websteri Bate, 1856 Leptocheirus pinguis (Stimpson, 1853) L. plumulosus Shoemaker , 1932 L. rhizophorae Ortiz, 1981 Liocuna caeca Myers, 1981 Microdeutopus anomalus (Rathke, 1843) Af. gryllotalpa Costa, 1853 myersi Bynum & Fox, 1977 Neohela monstrosa Boeck, 1861 N. intermedia Coyle & Mueller, 1981 N. pacifica Gurjanova, 1953 Paramicrodeutopus schmitti (Shoemaker, 1942) Pseudunciola obliquua (Shoemaker, 1949) Pterunciola spinipes Just, 1977 Rildardanus laminosa (Pearse, 1912) Rudilemboides naglei Bousfield, 1973 P (SE AL-CAL) P(BC-CAL) P(BC) P (SEAL-ORE) P (CAL) P (CAL-BC?) G(FL) P(S CAL) G (FL - CUBA ) P (SE AL-S CAL) G (W FI -TEX) G? (YUC?) P (CAL-BC, intr.) A-P(BER) G(FL) G (FL-YUC) At (G-S) G (FL) G (FL-TEX) G(FL) G(EFL) G(FL) G(FL) G (W FL) G(FL) G (W FL) At (V-C) G(FL) G(W FL) G (W FL) G(W FL) G (W FL) At (C), G (FL) G(FL) G (W FL) At (STL; V), G (E FL?) A-At (AC) At (V), G (E FI) G (FL - CUBA) G (W FL) At (V) At (V) At (C-FL), G (FL) A-At (ST L) P(W AL) P (CAL?) P (S CAL) At (AC-V, shelf) At (off NC, deep) At (HAT)-G (W FL-AL) At (V-C), G (FL) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 113 Unicola crassipes Hansen 1887 U. dissimilis Shoemaker 1945 U. inermis Shoemaker 1945 U. irrorata Say, 1818 U. laticornis Hansen, 1887 U. leucopis (Kroyer, 1845) U. serrata Shoemaker, 1945 U. spicata Shoemaker, 1945 A-At (N, slope) At (V-C), G (E FL?) A-At (AC-CHES) At (AC-V) A-At (AC-CV, deep) A-At (AC) G (FL-AL) At,- G Family Cheluridae Allman, 1847 Chelura terebrans Philippi, 1839 Tropichelura gomezi Ortiz, 1976 T. insulae Barnard, 1959 Family Isaeidae Stebbing, 1906 P(CAL, intr.). At (AC), G G. (FL- CUBA) G (FL) Ampelisciphotis podophthalma (J L. Barnard, 1958) Audulla chelifera Chevreux, 1901 Cheirimedia macrocarpa america Conlan, 1983 C. macrodactyla Conlan, 1983 C. similicarpa Conlan, 1983 C. zotea (Barnard, 1962) Cheirophotis megacheles (Giles, 1885) Chevalia aviculae Walker, 1904 C. carpenteri Barnard & Thomas, 1987 C. inaequalis (Stout, 1913) C. mexicana Pearse, 1913 Gammaropsis atlantica Stebbing, 1888 G. effrena (Barnard, 1964) G. ellisi Conlan, 1983 G. inaequistylis Shoemaker, 1930 G. maculatus (Johnston, 1827) G. mamola (Barnard, 1962) G. martesia (Barnard, 1964) G. melanops G. O. Sars, 1882 G. nitida (Stimpson, 1853) G. ocellatus Conlan, 1994 G. ociosa (J. L. Barnard, 1962) G. shoemakeri Conlan, 1983 G. sophiae (Boeck, 1861) G. spinosa (Shoemaker, 1942) G. sutherlandi Nelson, 1981 G. thompsoni (Walker, 1898) Microprotopus raneyi Wigley, 1966 M. shoemakeri Lowry, 1972 Pareurystheus alaskensis (Stebbing, 1910) p. dentatus (Holmes, 1908) p. tzvetkovae (Conlan, 1983) Photis bifurcata Barnard, 1962 P. brevipes Shoemaker, 1942 P (CAL) G (FL) P(BC-ORE) P(BER) P(AL-BC) P(BC-CAL) P (CAL) P (BC); G (FL) G (FL) G (FL) G (FL-LA) G (FL) P (CAL) P(C-CAL) A-At (STL shelf) A-At (N) P (CAL) P (CAL)? At (G) A-At (AC) P (CAL, deep) P (CAL) P(BC-S.CAL) A-At (AC slope) P(BC-S.CAL) AT (N C), G (SE FL) P(SEAL-SCAL) At (C), G (W FL) At (V-C), G (E. FL- LA) P (AL) P(BER-BC) P (AL) P(WA-CAL) P(AL-CAL) AMPHIPACIF1CA VOL. 3 NO. 1 MAY 16, 2001. 114 P. californica Stout, 1913 P. chiconola Barnard, 1962 P. conchicola Alderman, 1936 P. dentata Shoemaker, 1945 P. elephantis Barnard, 1962 P. fischmanni Gurjanova, 1938 P. kurilica Gurjanova, 1955 P. lacia J. L. Barnard, 1962 P. linearmanus Conlan, 1994 P • longicaudata (Bate & Westwood, 1862) P. macromana McKinney et al, 1978 P. macinerneyi Conlan, 1983 P. macrocoxa Shoemaker, 1945 P. macrotica Barnard, 1962 P ■ melanica McKinney, 1980 P. oligochaeta Conlan, 1983 P. pachydactyla Conlan, 1983 P ■ parvidons Conlan, 1983 P ■ pugnator Shoemaker, 1945 P. reinhardi Kroyer, 1842 P ■ spasskii Gurjanova, 1951 P. spinicarpa Shoemaker, 1942 P- tenuicornis G. O. Sars, 1882 P. trapherus Thomas & Barnard, 1991b P. typhlops Conlan, 1994 P. viuda J. L. Barnard, 1962 Podoceropsis amchitkensis Conlan, 1983 p - angustimana Conlan, 1983 (= G. ociosa ?) P - barnardi (Kurjaschov & Tzvetkova, 1975) P- chionoecetophila Conlan, 1983 P- setosa Conlan, 1983 Protomedeia articulata Barnard, 1962 P • fasciata Kroyer, 1842? P- grandimana Bruggen, 1905 P • penates Barnard, 1966 P- prude ns Barnard, 1966 P • stephenseni Shoemaker, 1 955 P (CAL) P (deep) P(WA-CAL) At (V) G (FL) P (CAL) P(BER) P (BER-ORE) P(BC-CAL) P (CAL) G (FL) G (FL-W TEX) P(BC-WA) At (AC-V) P (CAL) G (FL-TEX) P(SEAL-BC) P(SE AL-BC) P (BC -WA) At (C-FL), G (FL) P?-A-At (AC-BF) P (AL-BC) P (CAL)? A-At (G-I, C) G(FL) P (CAL, deep) P(S CAL) B (AL) P (BC) P(BER-BC) P (ALEUT-ORE) P (AL) P (ORE-S CAL) P? (WA), A-At P (BER-BC)A-At P(BCCA) P (BC-S CAL) A-At (ST L) Family Neomegamphopidae Myers, 1981 Neomegamphopus heardi Barnard & Thomas, 1987 N. hiatus Barnard & Thomas, 1987 N. kalanii Barnard & Thomas, 1987 N. pachiatus Barnard & Thomas, 1987 N. roosevelti Shoemaker, 1942 G (FL)? G(FL) G(EFL) G (S FL)? G(FL) Family Ischyroceridae Stebbing, 1899 (contains subfamilies Cerapiinae Budnikova and Ischyrocerinae Stebbing) Bonnierella linearis californica Barnard, 1966 P (OR-deep) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 115 Cerapus tubularis Say, 1818 At (V-C) C. benthophilus Thomas & Heard, 1979 G(FL) C. cudjoe Lowry & Thomas, 1991 G(FL) Erictkonius brasiliensis (Dana, 1853) P(intr.?),G(FL) E. difformis Milne-Edwards, 1830 P(WA?), A- At E . fasciatus (Stimpson, 1853 ) AT (ST L-V) E. rubricornis (Stimpson, 1853) P (BER-CAL),- At E. tolli Bruggen, 1909 A- At (ST L) Ischyrocerus anguipes (Kroyer, 1838) P-A-At (to DEL) (not FL!) /. claustris (Barnard, 1969) P (CAL) L commensal is Chevreux, 1900 A-At (STL) /. gurjanovae Kudrjaschov, 1975 P(BER) I. latipes Kroyer, 1842 A-At (ST L, slope) I. malacus Barnard, 1964 P (CAL, deep) /. megalops G. O. Sars, 1894 At (G-EM) L nanoides (Hansen, 1887) P (WA?), At (ST L) /. parvus Stout, 1913 P(S CAL) /. pegalops Barnard, 1962 P (CAL) I. serratus Gurjanova, 1938 P(AL?) I. tuberculatus (Hoek, 1882) Gurjanova P(BER) /. tzvetkovae Kudrjaschov, 1975 P(BER) Jassa borowskyae Conlan, 1990 P(AL-CAL) J. carltoni Conlan, 1990 P (CAL) J. marmorata Holmes, 1903 P-At (ST L -V-C)- G (E FL), J . morinoi Conlan, 1990 P(BC-CAL) J. myersi Conlan, 1990 P(CAL) /. oclairi Conlan, 1990 P ( AL-BC J. shawi Conlan, 1990 P(BC-CAL) J. slattery i Conlan, 1990 P(BC-CAL) J. staudei Conlan, 1990 P (SE A1 - BC, ORE) Microjassa bahamensis Conlan, 1995 At (E FL?) M. boreopacifica Conlan, 1995 P(SE AL-BC) M barnardi Conlan, 1995 P (ORE-CAL) Af. bousfieldi Conlan, 1995 P (CAL) Af. floridensis Conlan, 1995 G(FL) Af. litotes Barnard, 1954 P(BC-CAL) Af. macrocoxa Shoemaker (1942) AT (AC-G?) Af. micropalpa Shoemaker (1942) At (V-C?) A/. tetradonta Conlan, 1995 P (CAL?) G (FL) Parajassa angularis Shoemaker, 1942 P (CAL?) Ventojassa ventosa (Barnard, 1962) P (CAL, deep) Neoischyrocerus claustris (J. L.Bamard, 1969) P (CAL) Family Corophiidae Dana, 1849 Subfamily Corophiinae Bousfield & Hoover, 1997 Americorophium spinicorne (Stimpson, 1957) P(AL-CAL) A. aquafuscum (Heard & Si kora, 1972) At (C), G (FL-MI) A. brevis (Shoemaker, 1949) P (SE AL-CAL) A. ellisi Shoemaker, 1943 G (FL-LA) AMPHIPACIFTCA VOL. 3 NO. 1 MAY 16, 2001. 116 A. salmonis (Stimpson, 1857) A. stimpsoni (Shoemaker, 1941) Apocorophium acutum (Chevreux, 1908) A . lacustre (V anhoffen, 1911) Apocorophium simile (Shoemaker, 1934) A. louisianum (Shoemaker, 1934) Corophium volutator (Pallas, 1776) Crassicorophium crassicorne (Bruzelius, 1859) C. clarencense (Shoemaker, 1949) C, bonelli (Milne Edwards, 1830) Laticorophium baconi (Shoemaker, 1934)] Monocorophium insidiosum (Crawford, 1937) M. acherusicum (Costa, 1 857) M. californiaruim (Shoemaker, 1934) M. carlottensis Bousfield & Hoover, 1997 M. oaklandense (Shoemaker, 1949) M. steinegeri (Gurjanova, 1951) M. uenoi (Stephensen, 1932) M. tuberculatum (Shoemaker, 1934) Sinocorophium alienensis (Chapman, 1988) P(SEAL-WA) P (CAL) P (CAL, intr). At (V-C). G (FL) At (V).G(E FL) At (C-EFL) G (FL-LA) At (AC) P-A-At (tAC-CHES) P-A (BER) P-A-At (AC)(not FL!) P (AL-CAL) (not FL!) P (BC-CAL, intr?). At (ST L; V) (FL?) P (AL-CAL) A-At (CHES), G (FL) P (BC-CAL) P(BC-SEAL) P (CAL) P (BER) P (CAL, intr.) At (V-C), G (FL) P (CAL, intr.) Subfamily Siphonoecetinae Just, 1983 Siphonoecetes smithianus Rathbun, 1905 At (V, shelf) Caribboecetes crassicornis Just, 1 984 G (FL?) Family Podoceridae Leach, 1814 Dulichia rhabdoplastis McLoskey, 1970 D. tuberculata Boeck, 1870 Dulichiopsis remis (Barnard, 1964) Dyopedos arcticus (Murdoch, 1885) D. bispinus (Gurjanova, 1930) D. falcata (Bate, 1857) D. monacanthus (Metzger, 1875) D. porrectus Bate, 1857 D. spinosissima Kroyer, 1845 D. unispinus (Gurjanova, 1951) Paradulichia typica Boeck, 1870 Podocerus brasiliensis (Dana, 1853) P. chelonophilus Chevreux & DeGueme, P. cristatus (Thomson, 1879) P. fulanus Barnard, 1962 P. kleidus Thomas & Barnard, 1992b P. spongicolus Alderman, 1936 P(SE AL-CAL) P (WA?)-A-At (ST L) P (AL?) P (WA, CAL)-A-At (ST L) P (AL-BC)-At A-At (ST L, slope) A-At (ST L - CHES) A-At (ST L) A-At (AC, slope) P (BER) PA (Barrow), At (ST L, slope) P (S CAL), G (FL) 1888 G (FL)(see Thomas & Barnard, 1992a) P (CAL) P(S CAL) G (FI) P (BC?-CAL) AMPHIPACIRCA VOL. 3 NO.l MAY 16, 2001. 117 SUBORDER CAPRELLIDEA Leach, 1814 Superfamily Caprelioidea Laubitz, 1993 Family Caprogammaridae Kudrjaschov & Vassilenko, 1966, emend McCain, 1970 Subfamily Caprogammarinae K. & V., 1966 Caprogammarus gurjanovae Kudrjaschov & Vassilenko, 1966 WNP Family Caprellidae White, 1847, emend McCain, 1970 Subfamily Caprellinae Leach, 1814 Caprella angusta Mayer, 1903 P(BC-ORE) C. alaskana Mayer, 1903 P (BER-ALEUT-CAL)) C. andreae (Mayer, 1890) At (AC)- G c. borealis Mayer, 1903 P(AL-WA) c. brevirostris Mayer, 1903 P (CAL) c. calif ornica Stimpson, 1857 P (BC-CAL) c. carina Mayer, 1903 A c. ciliata G.O. Sars, 1880? P ( AL)-N At c. constantina Mayer, 1903? P(BER) c. cristibrachium Mayer, 1903? P (BER-ALEUT) c. danielevskii Czem. 1868 At (FL)- G c. drepanocheir Mayer, 1890 P(AL-WA) c. dubia Hansen, 1888 At -A c. equilibra Say, 1818 At (V) - G - P (BC-WA intr?) c. gracilior Mayer, 1903 P(AL-CAL) c. greenleyi McCain, 1969 P (ORE-CAL) c. incisa Mayer, 1903 P(SEAL-CAL) c. irregularis Mayer, 1890 P(AL-WA) c. kincaidi Holmes 1904? P(BER) c. laeviuscula Mayer, 1890? P(AL-ORE) c. linearis L. 1758 A-At-N P c. mendax Mayer, 1903 P (BC) c. mutica Schurin, 1935 P (CAL) c. natalensis Mayer, 1903 P (BC-CAL) c. paulina Mayer, 1903 P (BER-ALEUT) c. penantis Leach, 1814 At-G; P (CAL intr?) c. pilidigita Laubitz, 1970 P (BC-WA) c. pilipalma Dougherty & Steinberg, 1953 P (CAL) c. pustulata Laubitz, 1 970 P (SEAL-ORE) c. radiuscula Laubitz, 1970 P(SEAL-WA) c. rinki Stephensen, 1933 At (deep) c. scabra Holmes, 1904 P (SE AL) c. scaura Templeton, 1836 P (CAL)-At c. septentrionalis Kroyer, 1842? P (BER?), A-At c. striata Mayer, 1903 P (AL-WA?)-A c. trispinus Honeyman, 1889 At (deep) c. ungulina Mayer, 1903 P (BC, deep) c. unica Mayer, 1903 At AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 118 C. uniforma La Follette, 1915 p (CAL) C. verrucosa Boeck, 1872 P (BC-CAL) Metacaprella anomala (Mayer, 1903) P (AL-CAL) M. ferresa Mayer, 1903 P (A-CAL) M. horrida G. O. Sars, 1880 At- A M. kennerlyi (Stimpson, 1864) P (AL-CAL) Subfamily Aeginellinae Vassilenko, 1968 Aeginella spinosa Boeck, 1861 N At (deep) Aeginina longicornis (Kroyer, 1842) A-At Family Pariambidae Laubitz, 1993 Deutella californica Mayer, 1890 P ( SE AL-CAL) D. abracadabra Steinberg & Dougherty, 1952 At-G D. incerta Mayer, 1903 C Hemiaeginina minuta Mayer, 1890 At, G Luconacea incerta Mayer, 1903 At (V) Paracaprella tenuis Mayer, 1903 At (V) P. pusilla Mayer, 1890 At G P. cf. temir (fide Nelson, 1995) O Family Protellidae McCain, 1970, emend Laubitz, 1993 Mayerella limicola Huntsman, 1915 At M. banksia Laubitz, 1970 P (AL-CAL) M. acanthopoda Benedict 1997 P (S CAL) Protellina ingolfi Stephensen (1942?) N At (deep) Proaeginina norvegica (Stephensen, 1931) N At Protoaeginella sp. Laubitz & Mills, 1972 N At Tritella pilimana Mayer 1903 P (AL-ORE) T, laevis Mayer, 1903 P (BC-S CAL) T. tenuissima Doughty & Steinberg 1953 P (CAL, deep) Family Paracercopidae Vassilenko, 1968 Cercops holbolli Kroyer, 1842 A C. compactus Laubitz, 1970 P Paracercops setifer Vassilenko, 1972 P (BER?) AMPHIPACIHCA VOL. 3 NO.l MAY 16, 2001. 119 Family Caprellinoididae Laubitz, 1993 Pseudaeginella biscaynensis (McCain, 1968) At (FL) G Pseudoliropus vanus Laubitz, 1970 P (BC, deep) Family Phtisicidae Vassilenko, 1968 Subfamily Phtisicinae Vassilenko, 1968 Phtisica marina Slabber, 1769 At-G Perotripus brevis (La Follette, 1915) P (AL-S CAL) Hemiproto wig ley i McCain, 1968 At (G -E FL) Infraorder Cyamida Bousfield, 1979 Family Cyamidae Rafininesque, 1817 (revised Margolis, McDonald, & Bousfield 2000) Cyamus (Cyamus) ceti (L.) Lamarck, 1801 C. ( Cyamus ) erraticus R. de Vauzeme, 1834 C. ( Cyamus ) ovalis R. de Vauzeme, 1834 C. ( Cyamus ) gracilis R. de Vauzeme, 1834 C. ( Cyamus ) monodontis Lutken, 1873 C. (Cyamus) nodosus Lutken, 1860 P (AL-CAL), At P (BC)-At P(SEAL) P (SE AL)-At P (BER)-A-At A on Balaena mysticetus on Balaena glacialis on Balaena glacialis on B. gracilis on beluga, narwhal on narwhal Cyamus (Paracyamus) balaenopterae K. H. Barnard, 1931 C. (Paracyamus) boopis Lutken, 1870) P-At P (AL-CAL) Atl on balaenopteae (blue, fin) on Megaptera Cyamus (Mesocyamus) catodontis Margolis, 1954 C. (Mesocvyamus) orubraedon Waller, 1989 C. (Mesocyamus) mesorubraedon Margolis et al., 2000 P ('BC)-At P P on Physeter on Berardius bairdi on Physeter Cyamus(Apocyamus) scammoni Dali, 1872 C. (Apocyamus) eschrichtii Margolis et al, 2000 C. (Apocyamus) kessleri Brandt, 1872 P (AL-CAL) P P (AL-CAL) on Eschrichtius on Eschrichtius on Eschrichtius Orcinocyamus orcinus (Leung, 1870) P (BC+) on Orcinus orca Isocyamus delphini (Guerin-Meneville, 1836) /. globicipitis Lutken, 1973 I. kogiae Sedlak-Weinstein, 1992 P (AL-CAL) At P (CAL?) on porpoises, dolphins on Globicephalus on Kogia (pygmy sperm) Neocyamus physeteris (Pouchet, 1888) P(SE AL-BC) on Physeter, Globicephalus Platycyamus flaviscutatus Waller, 1989 Platycyamus thompsoni (Gosse, 1855) ampullatus P At on Berardius bairdi on Hyoperodon Syncyamus pseudorcae Bowman, 1955 At on Pseudorca Scutocyamus parvus Lincoln & Hurley, 1974 At on white-beak dolphin ( Cephalorhynchus ) AMPHIPACIFICA VOL. 3 NO.l MAY 16, 2001. 120 NOTES