Serials National Museums (QQH91 National Museum of Canada AN of Natural Sciences P82 Ottawa 1982 Publications in Biological Oceanography, No. 11 THE AMPHIPOD SUPERFAMILY TALITROIDEA IN THE NORTHEASTERN PACIFIC REGION. I. FAMILY TALITRIDAE: SYSTEMATICS AND DISTRIBUTIONAL ECOLOGY. E.L. Bousfield Senior Scientist National Museum of Natural Sciences National Museums of Canada Ottawa, Canada K1A 0M8 Publications d’Océanographie biologique, n° 11 Musées nationaux Musée national du Canada des Sciences naturelles National Museums National Museum of Canada of Natural Sciences Ottawa 1982 Publications in Biological Oceanography, No. 11 THE AMPHIPOD SUPERFAMILY TALITROIDEA IN THE NORTHEASTERN PACIFIC REGION. l. FAMILY TALITRIDAE: SYSTEMATICS AND DISTRIBUTIONAL ECOLOGY. E.L. Bousfield Senior Scientist National Museum of Natural Sciences National Museums of Canada Ottawa, Canada K1A 0M8 Publications d’Océanographie biologique, n° 11 Musées nationaux Musée national du Canada des Sciences naturelles National Museum of Natural Sciences Publications in biological oceanography, no. 11 Published by the National Museums of Canada © National Museums of Canada 1982 National Museum of Natural Sciences National Museums of Canada Ottawa, Canada Catalogue No. NM 95-7/11 Printed in Canada ISBN 0-662-11269-5 ISSN 0068-7995 Musée national des Sciences naturelles Publications d’océanographie biologique, n° 11 Publié par les Musées nationaux du Canada © Musées nationaux du Canada 1982 Musée national des Sciences naturelles Musée nationaux du Canada Ottawa, Canada N° de catalogue NM 95-7/11 Imprimé au Canada ISBN 0-662-11269-5 ISSN 0068-7995 Contents Abstract, v Résumé, vi Introduction, 1 Systematics 3 Superfamily Talitroidea Bulycheva 1957, emend Bousfield 1979a, 3 Family Talitridae Bulycheva 1957, emend Bousfield 1979a, 1982, 3 Section I. Beach Fleas (Non-substrate modifiers, sensu MacIntyre, 1963), Protorchestia new genus, 7 Protorchestia nitida (Dana 1852), 8 Traskorchestia new genus, 9 Key to species of Traskorchestia, 10 Traskorchestia traskiana (Stimpson 1857), 10 Traskorchestia georgiana (Bousfield 1958), 13 Traskorchestia ochotensis (Brandt 1851), 15 Paciforchestia new genus, 17 Key to species of Paciforchestia, 17 Paciforchestia klawei (Bousfield 1961), 18 Transorchestia new genus, 19 Key to described species of Transorchestia, 20 Transorchestia chiliensis (Milne-Edwards 1840), 20 Transorchestia enigmatica (Bousfield & Carlton 1967), 22 Orchestia Leach 1813-14, 22 Section I!. Sandhoppers (substrate modifiers, sensu MacIntyre 1963), 24 Platorchestia new genus, 26 Key to known species of Platorchestia, 27 Platorchestia chathamensis new species, 27 Megalorchestia Brandt 1851, 29 Key to species of Megalorchestia, 30 1. | Megalorchestia californiana Brandt 1851, 30 2. Columbiana group, 32 Megalorchestia columbiana (Bousfield 1958), 32 Megalorchestia minor (Bousfield 1957), 33 Megalorchestia dexterae new species, 35 3. Pugettensis group, 36 Megalorchestia pugettensis (Dana 1853-55), 37 Megalorchestia corniculata (Stout 1913), 39 Megalorchestia benedicti (Shoemaker 1930), 40 Trinorchestia new genus, 41 Trinorchestia trinitatis (Derzhavin 1937), 43 Orchestoidea Nicolet 1849, 43 Orchestoidea tuberculata Nicolet 1849, 45 Talitrus Latreille 1802, 45 Talitrus saltator (Montagu 1808), 46 Pseudorchestoidea new genus, 47 Key to species of Pseudorchestoidea, 48 Pseudorchestoidea brito (Stebbing 1891), 48 Pseudorchestoidea gracilis (Bousfield & Klawe 1963), 49 Pseudorchestoidea meridionalis (Schuster 1954), 51 Pseudorchestoidea mexicana new species, 51 ill Pseudorchestoidea biolleyi (Stebbing 1908), 53 Pseudorchestoidea spp., 54 Section III. Landhoppers, 55 Arcitalitrus sylvaticus (Haswell 1880), 55 Talitroides topitotum (Burt 1934), 55 Discussion and Conclusions, 57 Acknowledgments, 62 References, 63 Tables: Table I. Major taxonomic characters of Family Talitridae and their plesiomorphic and apomorphic states, 66 Table II. Non Substrate Modifiers and Character States Considered, 69 Table III. Substrate Modifiers and Character States Considered, 70 Table IV. All Talitridae; Selected Character States, 71 Table V. Distribution of North Pacific Beach Fleas and Landhoppers, 72 Table VI. Distribution of North Pacific Talitroidean Amphipoda, 73 Date of submission: August 28, 1981 Date of acceptance: November 19, 1981 Abstract Based on material obtained by field expeditions of the National Museums of Canada since 1955, and from other sources, this study provides new information on the distribution and ecology of semiterrestrial and terrestrial amphipods of the North American Pacific and adjacent coastal marine regions. It revises the component species within formal new generic and informal systematics-ecological groupings, based mainly on newly recognized taxonomic characters and employing numerical taxonomic methodology. The study also outlines primary factors that contribute to the anomalous regional combination of primitive and advanced morphological types within beach fleas and sandhoppers, and the total absence of native terrestrial species in the North American Pacific coast. Herewith recorded and/or described in the region from Alaska to Baja California, are the following beach flea species: Traskorchestia new genus, and species 7. ochotensis (Brandt), T. traskiana (Stimpson), and T. georgiana (Bous- field); Paciforckestia new genus, and species P. klawei (Bousfield); Transorchestia new genus, and species T: enigmatica (Bousfield & Carlton); and Platorchestia new genus, and P. chathamensis new species; the following sandhopper species: Megalorchestia Brandt, and species M. pugetten- sis (Dana), M. corniculata (Stout), M. benedicti (Shoemaker), M. californiana Brandt, M. colum- biana (Bousfield), M. minor (Bousfield) and M. dexterae new species; and the introduced landhopper species Arcitalitrus sylvaticus (Has- well) and Talitroides topitotum (Burt). From adjacent regions, similarly treated are the sand- hopper genera Trinorchestia new genus, and species T. trinitatis (Derzhavin) from the north- western Pacific region; and from the Pacific coast of Mexico and Central America the sandhoppers Pseudorchestoidea new genus, with species P. gracilis (Bousfield & Klawe), P. meridionalis (Schuster), P. mexicana new species and P. biolleyi (Stebbing). Taxonomically pertinent descriptions and redescriptions are provided for Protorchestia new genus and species P. nitida (Dana), and Transorchestia chiliensis (Milne-Edwards) from southern Chile; Orchestoidea tuberculata Nicolet from central Chile; Talitrus saltator Latreille from the eastern Atlantic, and Orchestia (sens. str.) from the Atlantic-Caribbean region. Résumé De nouvelles données, fondées sur des spécimens recueillis lors d’expéditions du Musée national des Sciences naturelles depuis 1955 ou provenant d’autres sources, apparaissent dans cette étude sur la répartition et l’écologie des amphi- podes terrestres et semi-terrestre de la céte du Pacifique en Amérique du Nord et des régions cOtiéres adjacentes. Les espéces étudiées sont révisées et reclassées formellement dans de nouveaux genres et officieusement en groupe- ments écosystématiques, surtout en fonction de caractéres taxinomiques nouvellement reconnus et suivant une méthodologie numérique de taxinomie. L’étude décrit aussi les principaux facteurs favorisant l’"anomalie que constitue la présence, dans une méme région, de types morphologiquement primitifs et avancés de talitres des plages et de talitres des sables, et l’absence totale d’espéces terrestres indigénes sur la céte Pacifique de l’Amérique du Nord. Pour la région s’étendant de |’Alaska jusqu’en Basse Californie, on signale et (ou) décrit dans l’étude les espéces suivantes de talitres des plages : nouveau genre Traskorchestia et espéces T. ocho- tensis (Brandt), 7. traskiana (Stimpson) et T. georgiana (Bousfield); nouveau genre Paciforchestia et espéce P. klawei (Bousfield); nouveau genre Transorchestia et espéce T. enig- v1 matica (Bousfield & Carlton); et nouveau genre Platorchestia et nouvelle espéce P. chathamensis ; les espéces suivantes de talitres des sables : genre Megalorchestia Brandt et espéces M. pugettensis (Dana), M. corniculata (Stout), M. benedicti (Shoemaker), M. californiana Brandt, M. colum- biana (Bousfield), M. minor (Bousfield) et nouvelle espéce M. dexterae; et les espéces intro- duites de talitres terrestres Arcitalitrus sylvaticus (Haswell) et Talitroides topitotum (Burt). Pour les régions adjacentes, on étudie de maniére semblable les talitres des sables suivants : nou- veau genre Trinorchestia et espéce T. trinitatis (Derzhavin) de la région du Pacifique nord- ouest ; et nouveau genre Pseudorchestoidea, avec espéces P. gracilis (Bousfield & Klawe), P. meri- dionalis (Schuster), P. mexicana (nouvelle espéce) et P. biolleyi (Stebbing), de la céte Pacifique du Mexique et de l’Amérique centrale. Font l’objet de descriptions et redescriptions taxinomiques pertinentes : nouveau genre Protorchestia et espéces P. nitida (Dana) et Transorchestia chilien- sis (Milne-Edwards) du sud du Chili; Orchestoidea tuberculata Nicolet du centre du Chili, Talitrus saltator Latreille de lest de l’Atlantique et Orchestia (sensu stricto) de la région Atlantique- Antilles. ABBREVIATIONS FOR FIGURES (Sree antenna | er Mandible ee antenna 2 Lips pedis left a calceolus a right ae head es See upper lip Pianist » « pleopod BEPh. 3.054 brood plate PMS 0.5 0s urosome BrSet ....brood setae See uropod 2h: epimeral plate ae lower lip Gal ty: 3% gnathopod 1 I ee maxilla | a gnathopod 2 a maxilla 2 Bgiad sets peraeopod Mxpd ....maxilliped CX. oi+ys sCORal plate ee male ANATUN ererscare immature Rta ics « female Ras ad oid juvenile subad ....subadult Oot wise dactyl Vii Introduction The long, much-indented North American Pacific coastline is characterized by diverse rocky, sandy, and estuarine intertidal habitats. The region is washed by cool nutrient-rich ocean currents that support lush growths of attached marine algae and sea grasses, and offers ample environmental scope for members of the shore-dwelling gammaridean superfamily Talitroidea. Until very recently, however, only 23 species in six tali- troidean families had been recorded in the coastal marine region from Alaska to southern Cali- fornia (Bousfield, 1975, 1979d; Barnard, 1975) of which 11 species were members of the semi- terrestrial and terrestrial family Talitridae. Early records of North Pacific species, especially of the far-eastern seas of the USSR, had been sum- marized admirably by Gurjanova (1951) and Bulycheva (1957). Contributing mainly to taxo- nomic, distributional and ecological information on regional talitrids during the post-World War II period were the studies of Barnard (1954, 1955, 1964) and Bousfield (1957, 1958, 1961, 1970). Despite these noteworthy advances, several taxonomic problems of shore-dwelling talitrids (e.g., Bousfield, 1957; Bousfield & Klawe, 1963) remained unresolved and distributional records were spotty and incomplete. Characterization of genera and species based almost solely on characters of the gnathopods, an inheritance from 19th century taxonomists, has only recently been replaced or supplemented by more reliable and phyletically significant characters from all body regions (e.g., Bowman (1977), Friend (1980), and Bousfield (in prep.))'. Also, much more extensive amphipod material has been obtained in the Canadian Pacific and adjacent coastal regions since 1955 and especially since 1966 (see station lists of Bousfield and Jarrett, 1981). Recent analysis of this material has filled in many of the early distributional hiatuses and permitted more accurate comparison of the eastern and western North Pacific faunas and of the total North Pacific rim fauna (lat. 35°N-65°N) with the corresponding North Atlantic talitrid fauna (see Bousfield, 1979d, 1981). The richer and more diverse North Pacific fauna undoubtedly reflects the much greater age of its coastline and pre- 'Bousfield, E.L. A revised classification of Talitrid amphipods (Crustacea: Amphipoda: Talitridae). (In prep.) sumed more prolonged evolutionary opportuni- ties. Despite changing configurations through accretion of microcontinents during the Palaeozoic, this coastline has remained an essentially open surf coast since the early Mesozoic (see Howarth, 1981; Ben-Abraham, 1981) Based mainly on material obtained during field expeditions of the National Museums of Canada in the Alaska to California coastal marine regions, especially since 1966, the present study provides detailed records of the distribution and ecology of regional species of Talitridae; it furthermore attempts to group the species more realistically and formally on a systematics- ecological basis, and relates these components to faunas of adjacent and world-wide regions that have previously been described and/or are currently under revision. ze : : P5-7 ae Sod Figure 1. TALITRIDAE. Major Taxonomic Characters. a, b: peraeopod dactyls; c, d: mandibular left lacinia mobilis; e-—h: maxilliped; i-l: uropod 3; m-p: telson. Systematics Recent advances in talitroidean systematics (e.g., Bousfield, 1979a, 1981, in prep.'; Friend, 1980) have underscored the family level distinc- tiveness of the Talitridae and facilitated recog- nition of more natural (phyletic) species groupings. Newly recognized taxonomic charac- ters especially of mouthparts, peraeopods, and surface ultrastructure (see especially Bousfield in prep.'; Bousfield and Halcrow, in prep.) are used herewith to supplement and clarify previous taxonomic diagnoses of the family and com- ponent genera. Superfamily Talitroidea Bulycheva 1957, emend Bousfield 1979a Family Talitridae Bulycheva 1957, emend Bousfield 1979a, 1982 Body smooth, rarely rugose; pleon rarely dorsally toothed; urosome short, segments 2 and 3 usually telescoping dorsally into 1; animal capable of saltation (in air). Eyes medium to very large, often nearly dorsally contiguous, rarely small or lacking. Antenna | very short, rarely longer than peduncle of antenna 2 which is often elongate and sexually dimorphic. Buccal mass directly beneath head or slightly prognathous. Upper lip rounding below; lower lip lacking inner lobes, varying little. Mandible, left lacinia 4 or 5- (occasionally 6-) dentate, right lacinia usually tri- or quadricuspate. Maxilla 1, palp minutely 2-segmented or lacking; maxilla 2, inner plate shorter, single stout proximal plumose seta, varying little; maxilliped plates often small and weakly armed, palp segment 4 much reduced or lacking, rarely unguiform. Coxa | reduced, not posteriorly cuspate; coxae 2-4 deeper, each with distinct posterior marginal cusp, which may be small or lacking. Gnathopods unlike, strongly sexually dimorphic except in some terrestrial genera and one intertidal genus; gnathopod | weakly subchelate or simple (lacking palm), distal 'Bousfield, E.L. A revised classification of Talitrid amphipods (Crustacea: Amphipoda: Talitridae). (In prep.) *Records from Pacific coast of Central America and Galapagos include those of Stebbing (1906b) and Monod (1970) for **Orchestia’’ costaricana, and Bousfield (in prep.). segments often tumescent behind (especially in male); gnathopod 2 minutely subchelate or chelate and propod mitten-shaped in females, immatures and many terrestrial males, often powerfully subchelate and of ‘‘amplexing”’ form especially in semi-terrestrial males. Peraeopods slender, spinose, occasionally fossorial; dactyls cuspid- actylate and unlike in peraeopods 3 and 4, or non-cuspidactylate and alike. Peraeopod 5 usually much shorter than 6 and 7, all trending to dissim- ilarity of form and size in advanced groups. Pleopods slender, variously modified, reduced, or vestigial, rarely sexually dimorphic. Uropods | and 2 well developed, rami moderately long, spinose apically and usually marginally, uropod 1 often set on short “prepeduncle’; uropod 3 uniramous. Telson lobes variously fused, usually slightly separated distally, and armed apically and often dorsally with spines. Coxal gills short, plate-like or sac-like (marine intertidal species), or large, pleated, convoluted or lobate (most terrestrial species); sternal gills lacking. Brood plates variable; usually subovate, marginally setose, setae hook-tipped in plesiomorphic species, but often very reduced, linear, with marginal setae few and simple in apomorphic species. Recent revision of the 200+ known world species (Bousfield, in prep.)' has informally subdivided family members on a systematics- ecological and behavioural basis to encompass (1) palustral talitrids (semi-aquatic in salt marshes and mangrove swamps)’, (2) beach fleas (mainly intertidal and coastal leaf-litter, non-substrate- modifying talitrids), (3) sandhoppers (intertidal substrate-modifying talitrids of sandy beaches), and (4) landhoppers (truly terrestrial, supra-tidal non-substrate-modifying talitrids). Since each group contains at least two obviously convergent generic morphotypes and is therefore polyphy- letic, no formal recognition of these four groups is proposed. However, the system has some practical taxonomic application and is utilized herewith. Figure 2. TALITRIDAE. Major Taxonomic Characters; Gnathopods. a-d: gnathopod | o&; e-h: gnathopod | Q; i-l: gnathopod 2 o&'; m-p: gnathopod 2 9. Key to principal systematics-ecological groupings of Talitridae 1. | Dactyls of peraeopods 3-7 non-cuspidactylate; peraeopods 3 and 4 dactyls alike; segments 5 SUR CHIA VCs i UR) FST a See eh OO OBS BADE lac 6 WE 0s ocala 0 Dactyls of peraeopods 3-7 cuspidactylate; dactyl of peraeopod 4 variously thickened and “pinched” (notched behind), unlike that of peraeopod 3; peraeopod 4, segment 5 distinctly shorter that in peracopod S(iibs. Why Si wins Ae Os ic. Ss se WEE Se ls ee 2. Peraeopods 5-7 subsimilar in form, increasing in length posteriorly (fig. 31); coxal gills plate-like, subsimilar on peraeopods 2-6; gnathopod | distinctly subchelate in male and female, palm not exceeded by dactyl (figs 28. ©). a haxnavsave chyna nated eu swtodu h5% 6 20 Mla tee eas Peraeopods 5-7 more or less unequal in form and size; coxal gills various, usually convoluted and complexly lobate, unlike on peraeopods 2-6; gnathopod 1, palm weak or lacking (especially 9 ), Gaceeded by dactyl (itgs, 26,1) 05.4.8 wer non-cuspidactylate landhoppers (incl. Arcitalitrus, p. 55) 3. Mandibular left lacinia mobilis 4-dentate (fig. 1d); semi-aquatic in salt marshes, mangrove swamps, and some fresh-waters (New Caledonia) ......... palustral talitrids (not treated hereby) Mandibular left lacinia mobilis 5-dentate (fig. 1c); intertidal on rocky shores .............. Dir 56 SA ees Laden Wala many oleh eo ha sacs ete non-cuspidactylate beach fleas (Protorchestia, p. 7) 4. Appendages (antennae, gnathopod |, peraeopods, uropods) stout, more or less strongly spinose and modified for burrowing (figs. 3g, 1); pleopod peduncles marginally spinose (figs. 3p, q, r); RUS CRATE INS UT EO WETS Aisi eve 6 + cack o abinals die ele x alk oe se sandhoppers «) Appendages slender, with slender spines, not modified for burrowing (figs. 3d, n); pleopod peduncles very weakly or not marginally spinose (fig. 30); non-substrate modifiers of rocky frarerysandy) beaches and forest leaf litter’... 41252. Tua ee ae eats ci es BMC Oe ae Seva ae bs Paced ce ere ke ote eA beach fleas and cuspidactylate landhoppers 6 5. Mandible, left lacinia 5- or 6-dentate; uropod 3, ramus relatively long (figs. 1k, 1); gnathopod 2 V2 )..bas® broadened anteriorly (ip; 2p) o2) . . . NA ROE 5-dentate sandhoppers (p. 24) Mandible, left lacinia usually 4-dentate; uropod 3 short, ramus < peduncle (figs. li, j); gnathopod 2 Par) Dasis narrow (ie, 2m) 5 i Se wes se ee A ee 4-dentate sandhoppers (p. 24) 6. Maxilliped palp, segments 2 and 3 broadly expanded, segment 2 with medio-distal lobe and strongly spinose medial margin (figs. If, g); coxal gills 3-5 relatively small, smaller than 2 and 6; gnathopod 2 (°) of functionally amplexing form, powerfully subchelate (fig. 21); peraeopod 4 dactyl usually strongly pinched or incised posteriorly, different from peraeopod 3 (fig. 1b) ... Me Pee ot enieSgas «ar areas ute wed Sena ee acai ak tae ok beach fleas (‘‘Orchestia”’ groups) (p. 5) Maxilliped palp, segments 2 and 3 usually spinose, not broadly expanded, segment 2 never with inner distal lobe (fig. le); coxal gills 3-5 (as well as 2 and 6) large, convoluted or modified; gnathopod 2(<') usually mitten-like (asin 9 andimm.) (fig. 2n), ifsubchelate, dactylattenuated; meracopod 3 and 4 dactyls not strongly different (fig Sh) re) . deg yc cats ~ ithe soe ens ces Aa ee een ase De ae Ep cuspidactylate landhoppers (incl. Talitroides topitotum) (p. 55) Section I. Beach Fleas (Non-substrate modifiers sensu MacIntyre, 1963) Key to genera of beach fleas of the North Pacific rim region and selected world-wide genera (includes Traskorchestia, Paciforchestia, Transorchestia, Orchestia, Protorchestia, Platorchestia)' 1. Mandibular left lacinia 5-dentate; antenna 2 not, and peraeopods 6 and 7 usually not noticeably sexually dimorphic in form (except Traskorchestia ditMmari) ......cecevceeceneecosseucnes Mandibular left lacinia 4-dentate; antenna 2 and peraeopods 6 and 7 moreor less strongly sexually dimorphic, often incrassate or thickened or of different form in of (figs. 3c, k) .........++4-- Mandibular left lacinia 5-dentate; antennae and peraeopod 7 (often also 6) incrassate (co) .... Pee OOS os ee Bhs a ie vil bo ae Oe eae Platorchestia n.g. (p. 26) ‘Genus Platorchestia is included also in the sandhoppers. Figure 3. TALITRIDAE. Major Taxonomic Characters. a-c: head & antennae; d-g: urosome & uropods; h, i: peraeopods 3 & 4; j, k: peraeopods 6 & 7; l-n: peraeopods 5, 6, & 7; o-r: pleopods. Maxilliped palp distinctly 4-segmented (fig. le); gnathopod 2 ( 9 ), basis linear, not expanded (fig. 2m); coxal gills 2-6 plate-like, subsimilar; telson lacking dorsal spines (fig. lm) ......... a or el ee Ce eee Be ee 9 Protorchestia n.g. (Southern Hemisphere) (p. 7) Maxilliped palp obscurely 4-segmented, 4th masked by spines of 3 (figs. If, g); gnathopod 2 (o'), basis more or less expanded anteriorly (fig. 20, p); coxal gills modified, 3, 4 and 5 reduced; telson with dorsal; marginal; and apical spines (figs; In, 0)... F500. oi Sie es Vea ws bap sd odes 3 Uropod | with strong distolateral (inter-ramal) spine (fig. 3d); telson longer than broad, narrow- ing distally, with single dorsolateral spine (fig. 8); gnathopod | ( 2 ), segments 5 and 6 posteriorly tumescent, dactyl not exceeding palm (fig. 2e); brood plate setae simple (fig. 20) ............ ET Tee ee ee ee ae ee ee ee ee PE, Paciforchestia n.g. (p. 17) Uropod | lacking distolateral spine (fig. 3e); telson short, broader than long, with groups of dorsal and marginal spines (fig. lo); gnathopod 1 ( 9), segments Sand 6not tumescent posteriorly, dactyl slightly exceeding palm (fig. 2f); brood plate setae hook-tipped (fig. 2m) ............. Pathe ba eile Ease kW ghas Sd Pade Bae Seana Gaetaiee oc Te ee ey ate arene eae ee Traskorchestia n.g. (p. 9) Gnathopod | ( ? ), dactyl barely or not exceeding distinct palm (fig. 2f); telson elongate, narrow- ing distally, lobes with dorsolateral row of spines (fig. 1n); brood plate setae hook-tipped .... ov. Bow Mak si egret lw pee ae Transorchestia n.g. (p. 19) Gnathopod | ( ? ), palm short, distinctly exceeded by dacty]l (fig. 2g); telson short, with dorsal, marginal, and apical groups of spines (fig. lo); brood plate setae simple ......... Orchestia Leach Protorchestia new genus Diagnosis Small, littoral and supralittoral talitrids characterized by Hyale-like, unmodified bodies; eyes dorsolateral, nearly contiguous; inferior antennal sinus very shallow; antenna | equal to or slightly exceeding peduncle 4 of antenna 2; antenna 2 not elongate nor sexually dimorphic in form. Buccal mass not prognathous; mandibular left lacinia 5-dentate, right lacinia quadricuspate (?); maxilliped palp distinctly 4-segmented, 4th not unguiform, segment 2 expanded medially but lacking inner distal lobe. Coxae 2-4 rounded below, hind margin with cusp. Gnathopod | (both sexes) fully subchelate (stronger in &), dactyl not exceeding (usually shorter than) palm, segments 4, 5 and 6 tumescent posteriorly; gnathopod 2 (o') powerfully subchelate, propod short, deep, palm smoothly convex; gnathopod 2 (2) basis linear, not expanded, segment 3 short, segments 4 and 5 tumescent behind, 6 short, mitten-shaped. Peraeopods 3-7 non cuspidacty- late, unguis short; peraeopods 3 and 4, all segments closely subsimilar, segments 5 subequal; peraeopods 5-7 similar in form (homopodous), increasing in length posteriorly, bases rounded behind, segment 5 not short, peraeopod 7 not incrassate in o'; coxa 5 aequi- or slightly anterolobate; coxa 6, hind lobe steeply oblique. Abdominal side plates 1-3 nearly smooth behind, hind corners subacute; pleopods normal, linear, peduncles each with 2 retinacula, margins not spinulose. Uropods | and 2 short, outer ramus smooth, peduncle with distolateral (inter-ramal) spine; uropod 3 short, ramus tapering, with apical and marginal spines. Telson lobes narrow- ing and separated distally, with apical spines only. Coxal gills plate-like, subsimilar. Brood plates ( 9 ) large, subovate, smallest on peraeopod 5, marginal setae long, numerous, hook-tipped. Type-species: Orchestia nitida Dana 1852 (Cape Horn region, South America) Additional species: Protorchestia campbelliana (Bousfield 1964) (Campbell Island, New Zealand) Parorchestia americana Bousfield 1964 (nomen nudum) Uruguay. Etymology The generic name signifies its very primitive morphology, embodying exclusively plesio- morphic characters, that is probably close to the first or ancestral talitrid morphotype. Gender: Feminine. Remarks The following species could be included in this genus except that the left mandibular lacinia is 4-dentate (a presumed apomorphic condition), antenna | nearly equals the peduncle of antenna 2, uropod 3 ramus lacks marginal spines, and peraeopod dactyls are more elongate, among other differences: Parorchestia rectipalma K.H. Barnard 1940 (South Africa), and a series of undescribed estuarine species from Tasmania Figure 4. Protorchestia nitida (Dana). Puerto Robalo, Chile. o& - 10.5 mm; @ ov. - 6.5 mm. and New Zealand (Bousfield, unpublished infor- mation). Remarkably, Protorchestia is _plesio- morphic in all characters considered in this study (Tables II, IV, figs. 26, 28) and little advanced beyond its presumed aquatic hyalid ancestors. Protorchestia nitida (Dana 1852) Figure 4 Orchestia nitida Dana 1852, p. 204; 1853, p. 868, t. 58, fig. Sa-f ‘Stebbing, 1906a, p. 539 Material examined Stn. F.1, Puerto Robalo, Isla Navarino, Chile, MW-LHW levels, 29 January, 1970, E.L. Bous- field and J.W. Markham colls., 1 &, 1 2 (ov.) (slide mounts); approximately 54 ood’, 38 9 9 (many ov.), and 80 juv. identified from 12 other stations in the Cape Horn region, Chile, February 1970. Hudson 70 Expedition, E.L. Bousfield and J.W. Markham colls., NMNS collections. Distributional ecology Known only from southern Chile and the Cape Horn region, mainly under algae and stones, occasionally in tide pools, from MW to lower HW levels (rarely under drift debris at H W level), and below main zone of Transorchestia chiliensis, mainly on protected and semi-protected beaches, occasionally in brackish water. Females ovigerous in summer (January-February), presumably annual. Remarks The female is very similar to P. campbelliana (Bousfield) but differs in the larger eye, longer telson, more elongate uropod 3, and smaller size at maturity. Although the differences are slight, species distinction is maintained pending des- cription of the male of P. campbelliana. Traskorchestia new genus Diagnosis Small to medium-large beach fleas characterized by: smooth, unmodified bodies; eyes lateral, not dorsally contiguous; antenna | equal to or slightly exceeding peduncle 4 of antenna 2, peduncle 3 longest; antenna 2 not elongate, peduncle not incrassate in male; inferior antennal sinus distinct, medium deep; buccal mass directly beneath head, not significantly prognathous; mandible, left lacinia 4 '/, (-5) -dentate, right lacinia tricuspate; maxilliped palp medium, obscurely 4-segmented (4th minute, masked by spines of 3), segment 2 broadly expanded medially, with distinct medio- distal lobe. Coxa 1 medium, half-overlapped by 2; coxae 2-4 broader than deep, lower margins nearly straight, hind margins cuspate. Gnatho- pod | (co), dactyl shorter than palm, segments 5 and 6 (occasionally 4) tumescent behind; gnathopod | (@), palm short, slightly exceeded by dactyl, segment 4, 5 not, 6 slightly postero- distally tumescent behind; gnathopod 2 (co) powerfully subchelate, propod short, deep, palm smoothly convex; gnathopod 2 (@), basis moderately expanded anteriorly, segment 3 short, segments 5 and 6 shallow-tumescent posteriorly. Peraeopods 3-7 cuspidactylate, dactyls (espe- cially unguis) short; peraeopod 4, segment 5 distinctly shorter, and dactyl strongly pinched, differing from respective segments of peraeopod 3; peraeopods 5-7 more or less homopodous, increasing in length posteriorly, bases rounded behind, that of peraeopod 7 may be sexually dimorphic (e.g., 7. ditmari); coxa 5 aequi- or slightly antero-lobate, coxa 6 hind lobe steeply oblique, anterodistally rounding. Abdominal side plates medium deep, hind corners subacute, posterior margins weakly spinulose; pleopods slender, rami normal or variously reduced, peduncles with 2 retinacula, all peduncles with a few outer marginal spines. Uropods | and 2 short to medium, outer ramus marginally spinose, inner ramus both margins spinose, peduncles lacking laterodistal (inter-ramal) spine; uropod 3, peduncle deep, posterior margin spinose, ramus tapering, margins and apex short-spinose. Telson short to medium, with apical and dorsal groups of spines, apex notched. Coxal gills lobate, 3-5 much smaller than 2 and 6. Brood plates ( ¢ ) large, subovate, margins with numerous long hook-tipped setae. Type-species: Orchestia traskiana Stimpson 1857 (present selection) Additional species: Traskorchestia ochotensis (Brandt 1851) Traskorchestia ditmari (Derzhavin 1923) Traskorchestia georgiana (Bousfield 1958) Etymology A combining form of the generic name Orchestia and the species name traskiana. Gender: Feminine. Remarks The genus Traskorchestia is closest to the austral and south Pacific new genus Transorchestia (Tables II, IV, figs. 26, 28), especially in the elongate segment 3 of antenna 1; relatively deep inferior antennal sinus; obscurely 4-segmented maxilliped palp; deep subchelate and distaliy tumescent segments of gnathopod 1 (¢c); moderately expanded basis of gnathopod 2 ( 2); large brood plates with curl-tipped marginal setae; short peraeopod dactyls; short, spinose uropods lacking pronounced distolateral (inter- ramal) spine; and deep, posteriorly spinose peduncle of uropod 3. Traskorchestia is more plesiomorphic than Transorchestia .in the essentially 5-dentate condition of the mandibular left lacinia; non-incrassate antenna 2 (cc); smooth-palmed propod and non-sinuous dactyl of gnathopod 2 (o'); weakly (or not) incrassate condition of peraeopods 6 and 7 (0); and oblique, rounded hind lobe of coxa 6; but is more apomorphic in the more reduced palm of gnatho- pod | (?) with overlapping dactyl; reduced rami and more strongly spinose peduncles of the pleopods; more heavily spinose uropods; and shorter, broader, more dorsally spinose telson. Traskorchestia is also more apomorphic than Paciforchestia in nearly all generic characters, except in the form of the brood plates and marginal setae. On the other hand Traskorchestia is generally more plesiomorphic than Plator- chestia, except in the reduced dentition of the mandibular left lacinia, and the more strongly spinose rami of uropod 1, but is not more apo- morphic than Orchestia in any character, except the more spinose pleopods. Key to species of Traskorchestia Peraeopod 7 ( o’), posterior margin of basis straight, or basis distally widening; segment 6 distally with longitudinal facial row of brush setae; telson with 3-4 dorsolateral spine groups ....... 2 Peraeopod 7 (’), posterior margin of basis evenly rounded, basis narrowing distally; segment 6 with 5-6 transverse apical brush setae only; telson lobes each with 1-2 dorsolateral spine BEOIDS. rear ts Mao. «Hi. «resin 1981, fig. 18: 19755 p.\355,0f1g. 4232 ‘Bowers,..1963,; p: 3172 figs. 3e, 4; 1975. pe357; fig. 228 ‘Staude et al., 1977, p. 12, fig. 20a Orchestoidea corniculata: Thorsteinson, 1941, Dead Talorchestia tridentata: Stebbing, 1899, p. 398, t. 30b (male) Material examined Alaska: 290 specimens from Bousfield and McAllister 1961 stns. A22, A43, A61, A66, A68, A70, A127, A140 (Prince William Sound to Baranof Is.). British Columbia: Queen Charlotte Islands; 260+ specimens from Bousfield 1975 stns. W1, W9a, W11, W16, H13, E18b, E20; north main- land coast; 270 specimens from Bousfield 1964 stus:- tit. bic, 4, At, 20, 23; 734, 35, H38, H40, H42, H44, H48, H51, H60; central mainland coast, 445+ specimens from Bousfield 1959 stns. N1, N3, N4, N6, N11, N16, N22; Vancouver Island, north outer coast, 715+ specimens from Bousfield 1959 stns. Ol, O2c, 03, O05, O7, O12, O16, O18, 58 specimens from Bousfield 1975 stns. P23, P29; north inner coast and Johnstone Strait, 545+ specimens from Bousfield 1959 stns. V1, V2, V3, V4b, V22; Strait of Georgia: re-examination of Bousfield 1955 station material from G1, G4, G9, G13, M1, M8, M11; southern Vancouver Island, 190 specimens from Bousfield 1970 stns. P710, P714, P716, P717, P720, and 5 specimens from Bousfield 1975 stns. P6, P14 (Barkley Sound region). Washington State: 145 specimens from Bousfield 1966 stns. W2, W4, W8, W24, W26, W30, W31, W33, W35, W36, W38, W41, W42, W44, W47, W48. Oregon coast: 50 specimens from Bousfield 1966 stns. W53, W55, W56, W58, W60. California: Re-examination of material of Bowers and author from Monterey region, previously published by Bousfield (1961). Miscellaneous material: British Columbia, Vancouver Island region: Ucluelet, July 1909, J. Macoun coll., 18 &o& (to 16.0 mm), 3 9 9, 1 imm. NMNS No. 913; J.F.L. Carl collections: head Departure Bay, 28 July 1938, 1 0, 1 Q, 2 imm., Cabbage Is., Strait of Georgia, 30 August 1956, Il oo, 18 9 29, NMNS No. 5109; Gonzales Bay, 27 February 1956,3 9 9,4 imm., NMNS No. 5108; Cadboro Bay, 25 February 1956,2 ooh, 3 292, 1 imm. NMNS 5107; ibid., 8 June, 5 ood, 2 9 9, NMNS No. 5116; ibid., 15 September, 6 9 9 subad., (with sternal mites); Saanich Spit, 21 August 1956 (in company with M. californiana), 2 ob ,2 9 9, NMNS No. 5110; China Beach, 4 July 1959, 22 oo (to 17.5 mm), 26 29 2, NMNS No. 5877; Chatham Island (near Victoria), 12 July 1959, 1 ¢o&, 3 99, NMNS No. 5869. Diagnosis Megalorchestia pugettensis varies in size (mature males 13.0 to 16.0 mm) and somewhat in morphology. Southernmost populations tend to have less spinose abdominal side plates, and in gnathopod 2 (oc), the posterodistal palmar 38 process is very short or lacking, almost as in M. corniculata. However, the dark lateral bars on coxae 5-7, are distinctive at all pigmentation intensities throughout the range (Bowers, 1963). Remarks Megalorchestia pugettensis is the most plesio- morphic known species of the genus, particularly in the small eyes, relatively short propod of gnathopod 1 that (ind) is postero-distally tumescent (forming a “pseudopalm”’), the relatively weakly and short-spinose uropods, weakly armed telson, and least broadly expanded pleopod peduncles. These characters are more or less shared with two similar-sized but more plesiomorphic western Pacific sandhoppers, Talorchestia sinensis Tattersall and 7. nipponensis Morino and to a lesser extent with western Pacific sandhoppers of the even more plesiomorphic genus Platorchestia (see Table III, and fig. 27). Distributional ecology Occurs on nearly all coarse to fine sand beaches of both surf-exposed shores and protected embayments, including estuaries, washed by summer-cold to warm, high salinity to brackish waters, from southern Alaska (western and northern limits not determined) to central California (Carmel Point). M. pugettensis co-exists with M. californiana on open fine sand beaches, burrowing mainly slightly seaward of it at the drift debris (HW) level, and co-occurs frequently with M. columbiana at or slightly above it, under drift debris. M. pugettensis is the only sandhopper occurring on poorly sorted or very coarse sands of small, steeply sloping beaches, frequently with Traskorchestia traskiana in such wave-protected situations. Megalorchestia pugettensis (and possibly also M. columbiana) almost certainly occurs on the Alaskan peninsula and adjacent Fox Islands, probably westward at least to Unalaska (Orchestoidea of Sheffer 1959, p. 377). Murie (1959, p. 290) states that ‘‘sand fleas were present in unbelievable numbers under boulders and in rotting kelp on Unimak Island, where they composed almost the entire droppings of red foxes (Vulpes vulpes L.).” The species is univoltine, breeding once per year; females ovigerous in late winter and early spring, adults dying out during the summer. Figure 17. Megalorchestia corniculata (Stout). Shell Beach, California. Megalorchestia corniculata (Stout 1913) Figure 17 Orchestoidea corniculata Stout 1913, p. 647, fig. C spoustield, 1961, p. 7, fig. 4a-d; 1981, fig. 17 :Craig, 1973a, b. geewers, 1963, p. 317, fig. 2, 5-d; 1975, ip. 356, fig. 3d. ‘Straughan, 1981, pp. 375, 426. non Orchestoidea corniculata: Thorsteinson, 1941, p. 55 39 o - 21.0 mm, ov. - 18.0 mm. Material examined California: Duxbury Reef, Salinas beach, 28 August 1960. B. Neal coll., 24 specimens NMC-C-1981-474; Santa Barbara Beach at HW, 14 June 1961, McGrath coll., 24+ speci- mens, Pacific genus Talorchestia. Key to species of Pseudorchestoidea 1. Antenna 2relatively stout, some distal flagellar segments toothed behind; coxa 6, anterior margin of posterior lobe vertical, rounding; uropod 3 short, marginal and apical spines few, large; gnathopod 2 (o'), palm smoothly merging with posterior margin; eastern Atlantic ......... oeeoeeveveevreeevee eee eee ee ee we we eee OO Om OH OH oO ee ew wo Pe ines y wndia Sedat coe eee P. brito (Stebbing) (p. 48) Antenna 2 slender, distal flagellar segments not toothed (except P. mexicana); coxa 6, anterior margin of posterior lobe oblique, gently convex; uropod 3 elongate, marginal and apical spines small, numerous; gnathopod 2(<'), propod with distinct prominence at posterior palmar angle; GAStERI aCINIG Sc. Se hee ele vhee 0c a dite booed 2. Peraeopods 6 and 7, bases relatively small, unlike in size and form; uropod 2, outer ramus distinctly shorter than inner; gnathopod 2 (mature co’), propod small, short, weakly developed, posterior angle with prominent tumescent lobe (fig. 2,L). ............ P. biolleyi (Stebbing) (p. 53) Peraeopods 6 and 7, bases relatively large, subsimilar in size and form; uropod 2, rami subequal in length; gnathopod 2(c’), propod large, ovate, strongly developed, posterior angle with spinose prominence (fie, 2K) s+... a eee es 3. | Pleopod rami well developed, 12-15 segmented, little shorter than respective peduncles; uropod 1, outer ramus with marginal spines; antenna 1, flagellum 6-7 segmented, longer than FOE EG Ley Ninel as I aig olde aa Segara Be ceeey ate ee P. gracilis (Bousfield & Klawe) (p. 49) Pleopod rami reduced, 4-10 segmented, much shorter than peduncles; uropod 1, outer ramus marginally bare; antenna 1, flagellum 4-5 segmented, about equal in length to peduncle 2 .... 4 4. _ Dactyls of peraeopods 6 and 7, anterior margin pectinate (with several slender spines); peraeo- pods 6 and 7, bases rounded behind; uropod 1, inner ramus with medial (inner) marginal spines only; antenna 2 distal flagellar segments not toothed ........... P. meridionalis (Schuster) (p. 51) Dactyls of peraeopods 6 and 7, anterior margin withsingle distal slender spine; peraeopods 6and 7, posterior margin of basis nearly straight; uropod 1, inner ramus with both inner and outer marginal spines; antenna 2 distal flagellar segments toothed....... P. mexicana new species (p. 51) Pseudorchestoidea brito (Stebbing 1891) Talorchestia brito Stebbing 1891, p. 324, pl. 15 :Chevreux & Fage, 1925, p. 279, fig. 279, 289 ‘Vader, 1970, p. 83, figs. 2-6 non Orchestoidea brito: Bulycheva, 1957 Material examined England: Blyth, Northumberland county, sandy beach, March 1955, D.I. Williamson coll., 2 oo, 1 Q ov., (to 11.5 mm), NMNS Cat. No. 5106, slide mounts. Portugal: Sagres, sand beach, E. Mateus coll., (no other data), 5 obo, (to 12.5 mm), NMNS Cat. No. 10465, slide mount. Remarks Vader (1970) has provided detailed figures of most of the important taxonomic characters omitted in the original description and by sub- sequent workers on material from the Atlantic- Mediterranean region. Compared to eastern Pacific species of the genus, P. brito is a heavier bodied animal, with stouter antennae, and stouter Figure 22. Pseudorchestoidea gracilis (Bousfield & Klawe). Cabo San Lucas, Baja California, Mexico. o - 18.0 mm, 9 ov. - 14.0 mm. spines on uropods and peraeopods; the pleopods are less reduced; gnathopod 2, palmar margin merges smoothly with the posterior margin; and the ramus of uropod 3 is short, with a long terminal spine. Distributional ecology Occurs mainly on exposed, flat, sandy, high salinity, warm-water beaches, from southwestern Norway, along the European-Atlantic coast to Morocco and in the Mediterranean eastward to the Black Sea. The species is active on the beach by day, mostly lower on the shore than 49 associated beach-hopper species, feeding mainly on unicellular algae, diatoms, and scavenging on animal carcasses, rather than on cast-up plant debris. Apparently cannibalism is common in P. brito. Females ovigerous in summer, May- September. Pseudorchestoidea gracilis (Bousfield & Klawe 1963) Figure 22 Orchestoidea gracilis Bousfield & Klawe 1963, pil, figs: 12 Figure 23. Pseudorchestoidea meridionalis (Schuster). 8.0 mm. Material examined Re-examination of Type and Allotype male and female and Paratype female material from a beach at Cabo San Lucas, Baja California, Mexico, NMNS Cat. Nos. 6634 and 6635, and slide mounts. Diagnosis Close to P. meridionalis Schuster and P. mexi- cana n. sp., in the following characters: eyes large, nearly dorsally contiguous; gnathopod 2 (co) powerfully subchelate and amplexing in form; propod subovate, palm very oblique, with distinct tooth and depression hear hinge, and rounded spinose protuberance at posterior angle; posterior margin very short; peraeopods 6 and 7 very slender and elongate, 7 distinctly longer 50 San Diego, El Salvador. MD 3 - 11.0 mm, @ ov. - \ than 6 (especially in male), bases more or less rounded behind; uropods 1 and 2, spade spines strongly developed; uropod 3 elongate and sexu- ally dimorphic; and telson tongue-shaped, narrowing distally. Differing from P. meridionalis and P. mexicana in the more elongate antenna | (flagellum 6-7 segmented), smaller, more quad- rate eye; more strongly spinose peraeopods, shorter (non-pectinate) dactyls of peraeopods 6 and 7; larger basis and dactyl of peraeopod 5; more elongate pleopods; more heavily spinose uropods | and 2, in which the outer ramus of uropod | is marginally spinose; and the shorter, less sexually dimorphic uropod 3. The group of P. gracilis, P. meridionalis and P. mexicana is slightly more plesiomorphic than P. biolleyi Stebbing, but more apomorphic than the larger European P. brito in which the eye is smaller, peduncle of antenna 2 (o') somewhat thickened, gnathopod 2 (0°) palm less strongly toothed and less oblique; peraeopods 6 and 7 not very elon- gate or differing markedly in length; uropod rami shorter but more strongly spinose; uropod 3 very short; and telson subquadrate. Distributional ecology Occurs on open and semi-protected beaches of southern Baja California, burrowing in the upper tidal sand by day, and actively foraging at night over the entire beach, especially on the wet sand near the water’s edge. Animals are occasionally bioluminescent, presumably derived from lumin- escent bacteria in the food. Pseudorchestoidea meridionalis (Schuster 1954) Figure 23 Orchestoidea meridionalis Schuster 1954, p. 103, fig. 1 :Bousfield & Klawe, 1963, p. 2 (key) Material examined San Diego, Department La Libertad, El Salvador, sandy beach, upper tidal zone, 24 November 1957, O. Schuster coll., 1 o& (11.0 mm), 5 sub- adult do, 3 2 F subadult, 1 imm. 9. NMNS Cat. No. 5851, slide mount. Diagnosis As diagnosed in the key, in morphological comparison with P. gracilis and P. mexicana (above) and as illustrated. Also, sexual dimorphism was noted in the form of the abbreviated dactyl of peraeopod 5, which is more elongate and less swollen in the male. Gnathopod 2 (2), basis is strongly expanded anteriorly, and the propod is distinctly longer than the carpus, the minute dactyl located about '/, from the apex. Antenna 2 flagellum distal segments simple, not shortened or toothed. Distributional ecology Known only from surf-exposed sandy beaches of El Salvador, Pacific coast of Central America (Schuster, 1954), in the upper third of the tidal zone. Pseudorchestoidea mexicana new species Figure 24 51 Material examined Mexico: Mazatlan, Sinaloa Province, on sand beach near HW level, G.A. Cole, coll., 6 August 1969. Female Holotype, br. II, 10.0 mm, subadult male Allotype, 8.5 mm; female, br. II, Paratype; adult male (anterior peraeonal region only) Paratype. NMC-C-1981-507. Ibid, lot No. 2. In crab burrows at HW level, 2 2 9 (br. I and II); Paratypes NMC-C-1981-507. Santiago Bay, Department Colima, sand beach, 29 January 1964, W.L. Klawe coll., | o& subad. (9.0 mm), 1 9 br. II (11.0 mm), | imm., NMC-C-1981-508. Diagnosis Female (br. II, 10.0 mm). Slender-bodied, slender-legged, sandhopper characterized by: eyes very large, rounding above and nearly contiguous mid-dorsally; antenna | very short, flagellum 4-5 segmented, about equal to seg- ment 2 of peduncle; antenna 2 slender, distal 7-8 segments somewhat shortened, sharply toothed behind. Mandible, left lacinia cleanly 4-dentate; maxilliped palp, segment 3 slightly longer than wide. Coxa 1, anterodistal angle subacute. Gnathopod 1, propod elongate, posterior margin about ’/, length of anterior margin. Gnathopod 2, basis broadly expanded anteromedially, segment 4 with prominent posterior tumescent lobe. Peraeopods 3-7 very slender, relatively weakly spinose; dactyls slender, those of peraeopods 6 and 7 with single antero- distal slender spine; peraeopod 5, dactyl very abbreviated and stout, basis asymmetrically broadly rounding behind; peraeopods 6 and 7, basis relatively large and subsimilar, posterior margins nearly straight, weakly spinose. Abdominal side plates smooth below and nearly unarmed behind; pleopods very slender, | shortest; rami much reduced, 4-6 segmented; peduncle of 1, outer margin spinose proximally, of 2 and 3 short-spinose distally. Uropods | and 2 slender, weakly armed, terminal spade spines well developed; uropod 1, outer ramus _ lacking marginal spines, inner ramus with inner (medial) marginal spines and a few proximal medial marginal spines; uropod 2, rami subequal, inner ramus with both margins spinose, outer ramus with outer marginal spines only. Uropod 3 elongate, peduncle weakly short-spinose distally, ramus longer, laterally compressed, with short facial and apical spines. Telson short, tapering broadly, apex notched, lobes with a few dorso- lateral and apical short spines. Coxal gills Figure 24. Pseudorchestoidea mexicana new species. Mazatlan, Mexico. 8.5 mm, o& adult (gnathopods only)- small, normal, 6 not elongate. Brood plates not fully developed, lacking setae in material available. Male (8.5 mm) Allotype. Very like the female except gnathopod 1, carpus and propod tumescent behind, blister of propod subterminal. Gnatho- pod 2 in transforming stage, posterior margin of basis with 6-8 scattered spines, segment 4 with posterior process; propod small, palm smoothly convex, with small depression near hinge; dactyl 52 ? br. II. - 10.0 mm, o& subad. - short, tip reaching base of posterodistal palmar tumescence. Presumed mature male Paratype (length unknown). Gnathopod 2, basis with antero- distal lobe; propod large, powerful, palmar margin very oblique, spinose, with large hinge tooth and depression and spinose posterodistal prominence; dactyl hind margin weakly spinulose, with prominent hinge tooth. Figure 25. Pseudorchestoidea biolleyi (Stebbing). Punta Arenas, Costa Rica. o& -12.5mm, 9? - 8.5mm. Remarks Closely allied morphologically to Pseudorches- toidea meridionalis and P. gracilis, as described above, in the key (pp. 71-73) and as illustrated; more closely similar to P. meridionalis in size; eye shape; short flagellum of antenna 1; very reduced dactyl of peraeopod 5; reduced rami of pleopods (4-6 segmented vs. multi-segmented); unarmed outer ramus of uropod 1; shorter telson, and less heavily spinose appendages. P. mexicana differs from P. meridionalis mainly in the smooth (vs. pectinate) dactyls of peraeopods 6 and 7; the more nearly straight posterior margins of the 53 basis of peraeopods 6 and 7; the more strongly spinose pleopod peduncles; and more heavily spinose rami of uropods | and 2 (inner ramus of uropod | with both inner and outer marginal spines). Sexual dimorphism in uropod 3 could not be determined from the material available. Etymology The specific name refers to the Pacific mainland coast of Mexico, the type locality of the species. Pseudorchestoidea biolleyi (Stebbing 1908) Figure 25 Orchestoidea biolleyi Stebbing 1908, p. 241, fies. 1,52 ‘Bousfield & Klawe, 1963, p. 2 (key) Material examined Costa Rica: Airport Beach, Limon (98°58’N, 83°01’W) 19 March 1971, D.M. Dexter coll., IG, D vue ov. (to- 110 mim), 3 ime. 1 juv., NMC-C-1981-514; Playita Blanca, Playas del Coco, (10°34’N, 85°42’W), 9 February 1971, DEM Dexter coll. 3 cc, 8 9 9, 9 imm. NMC-C-1981-512, Naos Is., sand beach, 19 June 1973, Davi. “Dexter coll, 2 cod sto. 12.0 mm), 4 992, 13 juv., NMC-C-1981-516; Puerto Columbia, sand beach, 12 April 1971, D.M. Dexter coll., 1 co transf. NMC-C-1981-513; Punta Arenas, sandy beach, under logs and driftwood, 26 November 1959, W.L. Klawe coll.; Lot No. f, 2,6 ¢ (to 11.5.mm),. 1! of transi., 1 9: Lot Nov g7 ) i. lo transf., 3 2 9. subad., 3 imm., 2 juv., NMNS No. 5862. Diagnosis Small, slender bodied, long-legged, weakly spinose sandhoppers, closely allied to the meridionalis group in having large, nearly dorsally contiguous eyes; slender antenna 2, flagellar segments simple (not toothed behind); elongate peraeopods (7 distinctly longer than 6); uropods 1 and 2, slender, elongate; prepeduncle elongate; ramus of uropod 3 elongate, short- spinose and laterally compressed; and telson distally narrowing. P. biolleyi is distinctive in the following characters; maxilliped palp, segment 3 as long as wide, rounding apically; mandibular left lacinia with small proximal (Sth) tooth; gnathopod 1 (@), propod relatively short, posterior margin of carpus about half anterior; gnathopod 2 (mature o') weakly sub- chelate, propod small, neotenically arrested at transforming stage; gnathopod 2( 9), basis moderately expanded anteriorly; segment 4, posterior process short; peraeopod 5 very small, short, basis rounding behind; peraeopods 6 and 7, bases relatively small, more or less unlike in size and form, hind margin of 7 nearly straight; dactyls slender, not pectinate anteriorly; pleopods slender, rami multi-segmented, little reduced; peduncle of 1 not markedly shorter than 2 and 3, outer margins of | smooth. Uropods | and 2 weakly spinose, terminal spade spines sub- linear; uropod 1, outer ramus lacking marginal spines; uropod 2, outer ramus distinctly shorter 54 than inner (especially in male), inner ramus with inner marginal spines only. Uropod 3 not markedly sexually dimorphic. Telson short spade-shaped, with scattered small dorsal spines. Distributional ecology Known only from sandy beaches of the Pacific coast of Costa Rica and Panama, in the upper tidal zone. Females ovigerous in late winter and spring; probably two or more broods per year. Pseudorchestoidea spp. Single immature specimens probably referable to this genus, but not identifiable to species, have been examined from the following localities: 1. Isla Grande, Mexico: Stn. 5 (no further data), 1 subad. & (10.0 mm). NMC-C-1981-460, slide mount. The specimen corresponds with the type material of P. mexicana in the weakly spinose uropods (uropod | outer ramus marginally bare), weakly armed distal segments of peraeopod 5, non-pectinate peraeopod dactyls, reduced rami and proximally spinose margins of peduncle 1 of the pleopods, but differs in the larger number of segments in the pleopod rami (6-10, vs. 4-7), the lack of teeth on the distal segments of the flagellum of antenna 2, and the more distal position of the posterior tumescent lobe of the propod of gnathopod 1. 2. Ensenada, Baja California, Mexico: Under seaweed at HW line, sand beach, 3 June 1956, W.L. Klawe coll., 1 imm. 9 (5.5 mm), NMC-C- 1981-464, slide mount. This very immature specimen is clearly a sandhopper close to Pseudorchestoidea in overall facies, including slender peraeopods 6 and 7 of which 7 is the longer; slender pleopods; short palmless propod of gnathopod 1; slender, weakly armed uropods 1 and 2; 4-dentate mandibular left lacinia; and spade-shaped, distally notched telson, among other features. However, the relatively unmodi- fied peraeopod 5, the dactyl of which is of normal length, but with ‘“‘fuzz’’ setae on the posterodistal margin, the nearly unarmed margins of the pleo- pod peduncles, and the short ramus of uropod 3, are more like Caribbean species of Talorchestia. The presence of these single enigmatic specimens, and the distributionally limited material of this study, underscore the need for more intensive collecting of amphipods along the beaches of Mexico and Central America in order to delimit more fully the diversity and phyletic relationships of the regional Talitridae. Section III. Landhoppers Truly terrestrial amphipods are not native to the North American Pacific coastal or inland con- tinental regions. Native terrestrial species are known, however, from Central America, includ- ing Costa Rica and Panama (Bousfield, in prep.)', from the Galapagos (Bowman, 1977), from Hawaii and the north central Pacific Islands (Bousfield & Howarth, 1976) and from western Pacific coastal regions, especially Japan (Iwasa, 1939; Tamura and Koseki, 1974). Absence of native cool-temperate landhopper species in coastal rainforests of the North American Pacific coastal regions contrasts with the rich landhopper faunas of climatically similar forested land regions of the southern hemisphere such as South Island, New Zealand, and Tasmania (see Hurley, 1955, 1968; Friend, 1980). These popu- lations have been explained on the basis of a probable Gondwanaland origin of primary ter- restrial groups, and coincidental evolution of angiosperm leaf litter, subsequent continental drift, and evolution of other regional landhopper groups from immediate seashore progenitors (Bousfield, in press)'. Thus, native landhoppers of the Central American, Hawaiian, and Japanese coastal continental rain forests are of the cuspidactylate (modern) type and related morphologically to local seashore (beach flea) types. Synanthropic populations of a few land- hopper talitrids have been recorded in parks, botanical gardens, and zoological gardens of coastal cities of California (records summarized in Bousfield, 1975; Biernbaum, 1980). Non-cuspidactylate (ancient) types, such as Arcitalitrus dorrieni (Hunt), and A. sylvaticus (Haswell) have relatively low eurytopicity and when introduced by man to new habitats of the northern hemisphere, the amphipods spread slowly or remain in place (Richardson, 1980). On the other hand, cuspidactylate types such as Talitroides alluaudi Chevreux, and T. topitotum (Burt) seem much more hardy and may spread rapidly from their points of introduction, especially in coastal, winter-mild continental areas and on tropical islands (Bousfield & Howarth, 1976; Biernbaum, 1980). Key to introduced terrestrial amphipods of North America 1. | Peraeopods 3-7 cuspidactylate; peraeopod 4 shorter than 3, segment 5 shorter than and dactyl different from those of peraeopod 3 respectively; coxal gill of peraeopod 6 reverse L-shaped or curved forward Talitroides 2 Peraeopods 3-7 non-cuspidactylate (simplidactylate); peraeopods 3 and 4 subequal, segments 5 ) and dactyl similar in size and form; coxal gill of peraeopod 6 goose-necked, attenuated posteriorly oeeeeeeeeeeeeeeeeeeeee eee ee we we we ew Arcitalitrus sylvaticus (Haswell) (p. 55) 2. Antenna | elongate, nearly reaching tip of peduncle 5 of antenna 2; uropod 1, distolateral (inter- ramal) spine with complex tip oeeeeeeeeeeeee ee eee ee we we ee eee ee eh ee ee T. topitotum (Burt) (p. 55) Antenna | shorter, tip reaching about mid-point of peduncle 5 of antenna 2; uropod 1, distolateral spine with simple tip Arcitalitrus sylvaticus (Haswell 1880) Talitrus sylvaticus Haswell 1880, p. 246, pl. VIII, fig. 1 :Bousfield & Carlton, 1967, p. 282 :Bousfield, 1975, p. 353 (key), 364 Distributional ecology Recorded in North America only from leaf litter (mainly Eucalyptus) and damp debris in Golden Gate Park, and adjacent areas of San 'Bousfield, E.L. A revised classification of Talitrid amphipods (Crustacea: Amphipoda: Talitridae). (In prep.) oeeeeeeeeeeeee eee ee ee ee eee hl hl Ohl Oh OHO OO hh hh hh Hh ee ee ee aye) T. alluaudi Chevreux Francisco, California; endemic to southeastern Australia (New South Wales and Victoria). No additional regional material has been examined since the records of Bousfield & Carlton (1967). Talitroides topitotum (Burt 1934) Talitropsis topitotum Burt 1934, p. 184, fig. 11 :Vader, 1972 Talitrus decoratus Carl 1934, p. 134, p. 742, figs. 1-6 Talitrus pacificus Hurley 1955, p. 155, fig. 3 Talitrus sylvaticus Shoemaker, 1936, p. 60, fig. 1 **Talitrus’’ sylvaticus: Bousfield, 1975, p. 353 (key), 364, fig. 231 Material examined California: Balboa Park, San Diego Co., mixed litter, Eucalyptus and Pittisporum, near zoo, 23 February 1978, K.K. Bohnsack coll., 20 2 9. br. I and II (to 9.5 mm). NMC-C-1981-547. 56 Distributional ecology In leaf litter and under moist garden debris, along the Gulf coast of the United States from Louisiana east to Florida and north to South Carolina; in California, authentically recorded only from Balboa Park, and Pasadena, but probably occur- ring in the San Francisco Bay area also. Bierbaum (1980) has summarized records in North America and world-wide. Discussion and Conclusions The present generic revisionary concepts within the family Talitridae are based mainly on analysis of character states within North Pacific and selected world-wide species and higher groupings, as summarized in Tables I, II, III, and IV. The cluster analysis and resulting phenograms (figs. 26, 27, 28) complement the results of a revision! of world-wide Talitridae recently completed by the writer (Bousfield, in prep.).? The phyletic (evolutionary) conditions of morphological characters used in this analysis are differen- tiated within each table by the symbol 0 (plesio- morphic) or | (apomorphic). These conditions (subject to confirmation in some instances) were derived by comparison with related out- groups in presumed ancestral families Hyalidae, Hyalellidae, Najnidae and palustral Talitridae (see Bousfield 1981, in prep.).? ? Thus, an index of plesio-apomorphy is obtained by totalling the numbers for all character states for each species; low total values characterize primitive or plesiomorphic, and high numbers the advanced (derived) or apomorphic taxonomic groupings. An attempt was made to select or define characters that would minimize instances of the intermediate condition across the taxonomic spectrum; such cases were slotted into either the primitive or advanced category (a taxonomic judgment) in order to utilize a simplified two- stage average linkage cluster analysis (Sneath & Sokal 1973). The total body analysis more clearly reveals evolutionary trends in talitrid subgroups, morphological relationships that were generally masked by previous reliance on a few characters (especially of the gnathopods, some mouthparts, etc.) for generic (Bulycheva 1957), or subgeneric (Hurley 1975) delineation. Thus the transcending ‘Elements of this revision were presented orally at a Symposium on Biogeography, Annual Meeting of the Cana- dian Society of Zoologists, University of Waterloo, May 1981. *Bousfield, E.L. A revised classification of Talitrid amphipods (Crustacea: Amphipoda: Talitridae). (In prep.) *Bousfield, E.L. The amphipod superfamily Talitroidea in the northeastern Pacific region: 2. Family Hyalidae. Systematics and distributional ecology. (In prep.) Bousfield, E.L. The amphipod superfamily Talitroidea in the northeastern Pacific region: 3. Family Najnidae. Systematics and distributional ecology. (In prep.) 2 significance of condition of peraeopod dactyls and dentition of the mandibular left lacinia mobilis, the more precise definition of character states within the mouthparts, gnathopods, uropod 3 and telson, and recognition of signifi- cant new character states within the antennae, coxal plates, peraeopods, pleopods, uropods | and 2, gills and brood plates have collectively necessitated reassessment of generic groupings within the family. Most previously accepted full genera have been validated herewith, but in a more restricted sense, and based on the type species in each case. Some generic names, previously synonymized, have been resurrected, a few subgeneric names have been elevated to full generic status, several new generic names have been created, and all species have been reassigned within the old and new generic concepts. Taxonomic characters proposed here- with are at a level of taxonomic significance similar to those utilized in recent revisions of amphipod family and superfamily groups of the North Pacific region (e.g., Bousfield, 1979b (Gammaroidea); Conlan & Bousfield, 1982, (Ampithoidae)); and of other regions (Friend, 1980 (terrestrial Talitridae)). Formal recognition of the broad systematics- ecological units of Talitridae employed here is compromised by morphological convergences on the one hand, and ecological overlap on the other. Now required is a refinement of the system that will satisfactorily deal with these limitations and avoid formalization of polyphyletic group- ings, as in the past. Also required is much more information on the functional morphology and behaviour of these animals. Even with these limitations, the present system provides a basis for more natural grouping of species and genera, and new insights into the probable evolutionary history of the Talitridae. Thus, within the beach flea group (Table II, and fig. 26), the North Pacific rim region encompasses native genera (Traskorchestia, Paciforchestia) that are primitive relative to those of the North Atlantic and tropical-warm-temperate regions (Orchestia sens. str., “‘Orchestia’’ floresiana group) but are comparably apomorphic to those of antiboreal (southern hemisphere) regions (e.g., Protorchestia, Transorchestia). The North Pacific also encom- BT 4SaYydIO TZ4TdJ BTSOYOI0ZTd BIZSaYO104BTd TABUYIP BTISeyoIONSBU STSUdaPOYOO BTYsaydJONSeIy BUBTSIOIS BTASayoONSBI], BUBTHYSBIZ BIYSaYOIONSBI] BT ASayo1OSuBI], SOPTOLZTTeL SNITTE4TOLy BTSAZVA ,BTASeYoIO, BUBOTIBISOD ,BT4SIYoI0,, BUBUTNUS BTFSeydIOJTOBY TAO¥B4BAd BTYSaYILOJ TOBY TOABTH BTYSaYIOJTOBY BT4SayoI04OIg % Apo- 100 morphy ve) ~ AyLueLLWLS 4% 50 Figure 26. Phenogram of Species of Beach Fleas, Palustral Talitrids, and Landhoppers, North Pacific Rim region, and other related groups. 58 Pseudorchestoidea Platorchestia Talorchestia Orchestoidea Trinorchestie meridionalis nn un ] 2 al § oo ° ed ° ed » o on Z ae] e S| & a 100 rey oT 5. co ae 5) = -— ny se 50 beph ters Megalorchestia ugettensis corniculata benedicti columbiana dexterae minor californiana Figure 27. Phenogram of Species of Sandhoppers, North Pacific Rim Region, and other related groups. passes the genus Platorchestia, most species of which trend to a fossorial mode of life on sandy beaches; thus Platorchestia may be close to the presumed ancestor of most 5-dentate, heavy-bodied, cold-temperate sandhoppers of the northern hemisphere (see also Table III and figs. 27, 28). Paradoxically perhaps, North Pacific rim sandhoppers, especially those of the North American coast, are generally apomorphic, and component members (Megalorchestia, Pseudorchestoidea) are among the world’s most specialized sandhoppers. The phenograms (figs. 27, 28) confirm the need for formal recognition of the new generic categories and more natural 59 realignment of key genera (e.g., Talitrus with Orchestoidea; Megalorchestia with Trinorchestia, Talorchestia, and Platorchestia; Pseudorchestoidea with Caribbean members of Talorchestia), as out- lined also by the author elsewhere (Bousfield, in prep.)’. Formal recognition of subgroups within sandhopper genera (e.g., within Megalorchestia)* 'Bousfield, E.L. A revised classification of Talitrid amphipods (Crustacea: Amphipoda: Talitridae). 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