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 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

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 <dpee’s + osha Belleut sb. Jy Dewees Seo ae 3

Peraeopod 7 markedly sexually dimorphic in form, basis (o) strongly expanding postero-

distally, margin deeply serrate; gnathopod 2( 0), propod distinctly widening distally; uropod 3,

peduncle with (4) 5 postero-distal spines T. ditmari (Derzhavin)

Peraeopod 7 slightly sexually dimorphic, basis (¢’ ) not expanding distally, margin weakly

serrate; gnathopod 2(<o), propod margins subparallel, not distinctly widening distally; uropod 3,

peduncle 3-(4) spinose T. ochotensis (Brandt) (p. 15)

Antenna | very short, flagellum 3-4 segmented, shorter than peduncle; peraeopod 4, segment 5

very short and stout, about '/, length of segment 6; gnathopod | (’), segment 4 lacking posterior

“blister’’; uropod | sexually dimorphic, outer ramus (o’) with 5-6 close-set marginal comb-

spines (fig. 6) T. georgiana (Bousfield) (p. 13)

Antenna | normal, flagellum 5-7 segmented, longer than peduncle; peraeopod 4, segment 5

about % length of segment 6; gnathopod | (o’), segment 4 with posterior “‘blister’’ (fig. 5);

uropod | not sexually dimorphic, outer ramus (o¢) with 3-4 (5) broadly spaced spines .......

T. traskiana (Stimpson) (p. 10)

eoeoeceveeeeeeee eee eee ee ee ee we ee ow

oeoeeeeeeer eee eee ee ee we ee eh hl Ohh Oh Oh Oh HO He eo eH ee em hm em em em hh hh

Ce ee }

Traskorchestia traskiana (Stimpson 1857)

Figure 5

Orchestia traskiana Stimpson 1857a, p. 90

:‘Stebbing, 1906a, p. 534

‘Bousfield, 1958, p. 885, fig. 10d; 1975, p. 363,

We, 230, 1961, p, 35, le. 1.2. TORI p. 83.

fig. 17

Orchestia sp.: O’Clair, 1977, p. 446

non Orchestia taskiana: Bulycheva, 1957, p. 166,

fig. 60

Material examined

Alaska: Amchitka, Aleutian Islands; C.E. O’Clair

Plot 5, upper 1A2, 22 May 1974, 1 imm., ibid.,

Plot 9, 1A2, 27 August 1974, 2 imm., ibid. (no

data). 2 9 0... br. HH.

Southeastern Alaska: 865 specimens from Bous-

field & McAllister 1961 stns. (Seward to Dixon

Entrance) Al, A4, A7, A8, Al4, A1l8, A22, A25,

A29, A31, A37, A38, A55, A57, A68, A75, A87,

A&88&, A118, A121, A131, A140, A151, A159,

A162, A166, A175; Bousfield 1980 stns. (Sitka

FEPION) "SZ. Soy Sic LL.

10

British Columbia: Queen Charlotte Islands; 200

specimens from Bousfield 1957 stns. H2a, H8a,

H13, W2, W4a, W16, E5, E13, El4a, E17, E20,

E24, E25; north mainland coast; 120 specimens

from Bousfield 1964 stns. H2, H4, H9, H11, H20,

H23, H34, H38, H40, H41, H42, H44, H45, H47,

H51, H60, H61; central mainland coast; 40

specimens from Bousfield 1959 stns. N3, N4, N6,

N11, N15, N21, N22; Vancouver Island, north

outer coast, 35 specimens from Bousfield 1959

stns. Ol, O2a, O03, O04, O16, and 12 specimens

from Bousfield 1975 stns. P23a and P32; inner

coast and Johnstone Strait, 470+ specimens from

Bousfield 1959 stns. V4a, V14, V19, V20, V21,

V22, V25, V26; Strait of Georgia: re-examination

of Bousfield 1955 material from G4, G9, G13,

G21, M1, M8, M11; Vancouver Island south end

and outer coast: 100 specimens from Bousfield

1970 stns. 706, 712, 716, 719; Barkley Sound

region, 70 specimens from Bousfield 1975 stns.

P5, P6, Pl6a, P17b, P19b; 28 specimens from

Bousfield 1976 stns. B3, B14; 23 specimens from

Bousfield 1977 stns. B4a, Bl2a, B6a.

4

a

L|

a

A

a

v

\)

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Figure 5. Traskorchestia traskiana (Stimpson). Piper’s Lagoon, Vancouver Island, B.C. of - 17.0 mm,

2 ov. - 13.0 mm.

British Columbia: Miscellaneous collections:

J.F.L. Carl, Victoria region, 1956: Ross Bay,

2 February, 6 odd, 5 9 @ br. II, NMNS No.

5220; ibid., 18 February, 5 o'd', 1 9 ov., NMNS

No. 5219; Shoal Bay, 19 February, 2 ob od',2 9 Q,

NMNS No. 5223; Willows Beach, 25 February,

1 o, NMNS No. 5217; Gonzales Bay, 27 Febru-

ary, 6 dod do (to 19.0 mm), 1 2 ov., NMNS No.

222; Cadboro Bay, 8 June, 2 dd, 2 2 9 ov.,

NMNS No. 5116; ibid., 15 September, 4 do,

NMNS collections; ibid., 1959: China Beach,

4 July, 4 od dv, 1 9, 1 imm., NMNS 5877; ibid.,

Chatham Island, 12 July, 4 od oh, 1 9, NMNS

No. 5869. C.G. Carl coll.: Pt. Grey, south entrance

point to Burrard Inlet, Lot. No. 55, 10 August

1933, 39 oo, 10 29 F ov., NMNS collections;

Savary Island, Lot No. 47, 30 March 1929,

10 ov, 5 2 F ov., NMNS collections; Anthony

Island, Queen Charlotte Islands, 17 October

1956, 14 oo ,9 2 F (2 ov.), 1 imm., NMNS No.

5886; Grassy Island, B.C., 29 June 1958, 19 do,

1 2 ov., NMNS No. 5873; Hohuae Island, B.C.,

2 July 1958,2 do ,2 & Y ov., NMNS No. 5879.

S. Nash coll.: Nanoose Bay, 11 January 1979,

Lot No. 1, 16 large &o o& (to 15.0 mm), 5 small

oo (to 7.0mm), 4 2 9,9imm.:; ibid., Lot No. 2,

200,1 2 br. I; NMNS collections; R.M. O’Clair

coll.: stn. 760060, Muchalat Inlet, Vancouver

Island, opposite Gold River, July 1976,2 do,

2 292 ov., NMNS collection.

Washington State: 152 specimens from Bousfield

1966 stns. W2, W4, W6, W8, W9, W11, W20,

W21, W22, W26, W28, W29, W34, W35, W36,

W47, W48.

Oregon coast: 38 specimens from Bousfield 1966

stns. W51, W52, W53, W55, W56, W57, WS8.

California: Santa Barbara, beach intertidal,

25 October 1965, W.L. Klawe coll.,3 o 0,1 co

transf. (at 10.0 mm), NMNS No. 13992; ibid.,

18 August 1969, A. Wenner coll.,9 oo (to

18.0 mm), 4 992, NMNS No. 13993; ibid.,

29 May 1970,8 0d ov, 1 2 ov., NMNS No. 13968.

Re-examination of all material from central

California published by Bousfield (1961).

Mexico: La Mission, Baja California, March 1952,

T.E. Bowman coll., 3 0&0 o&', USNM No. 102423;

Turtle Bay, B.C., 7 June 1956, W.L. Klawe coll.,

13 large o ob (to 18.0 mm), 2 small oo’, 4 small

22 ov., (to 10.0 mm), NMNS collections; El

Socorro Beach, near El Rosario, HW sand,

June 1980, R. Appy coll., 31 oo (to 17.0 mm),

21 9299, several ov., NMNS collections.

Re-examined all material from Bahia San

Quintin, in Barnard (1964).

Diagnosis

Male (14-18 mm). Medium large, plesiomorphic

species characterized by: antenna 1, flagellum

5-6 segmented; antenna 2, flagellum about

15-segmented, distinctly longer than unmodified

peduncle. Mouthparts normal; mandible, left

lacinia 4'/,-dentate; maxilliped outer plate

relatively tall, apex subacute. Gnathopod 1,

segment 4 with small but distinct posterior blister

or tumescent lobe. Gnathopod 2, propod longer

than deep, palm smooth, oblique. Coxa 3, lower

margin nearly straight, deepest posteriorly.

Peraeopod 4, segment 5 not markedly short or

thick. Peraeopods 5-7, dactyls short; bases

smoothly rounding behind, margins with

numerous evenly spaced basally “‘beaded”’ small

spines. Peraeopod 7, segment 6 with transverse

apical row of 5-6 fine setae, lacking distal

longitudinal facial row of brush setae. Abdom-

inal side plates 2 and 3, hind corners weakly

produced. Pleopods normal, little reduced, rami

7-12 segmented. Uropod 1, peduncular margins

with 3-5 short spines; inner ramus with 3-4 inner

marginal and one or two outer marginal spines,

outer ramus with 3-4 (occasionally 5 or 6) spaced

outer marginal spines; uropod 2, peduncle with

3 stout inner and outer marginal spines, inner

ramus with 2 inner and outer marginal spines,

outer ramus with 2 short outer marginal spines.

Uropod 3, peduncle with 5 posterodistal spines;

ramus shorter, tapering, with 2 groups of posterior

marginal spinules and slender apical spines.

Telson lobes each with transverse row of medio-

dorsal spines, a single subapical dorsal spine and

3-4 apical spines. Coxal gill on peraeopod 6

large, about twice length of gill on peraeopod 5.

Female (13 mm ov.). Gnathopod 1, segment 6

with small posterodistal tumescence overlapped

by part of unguis only. Gnathopod 2, basis widest

proximally, segment 4 broadly and shallowly

tumescent. Brood plate broadest proximally.

Brood plate (P5) about half length of that of

peraeopod 2, margins with 16-18 curl-tipped setae.

Remarks

T. traskiana bears several characters in common

with T. ochotensis, especially in uropods, gnatho-

pods, and overall size range; however, the lack of

sexually dimorphic peraeopod 7, unreduced

pleopods, and less strongly dorsally spinose

telson, relate it more closely to T: georgiana, the

other North American species. ‘‘Orchestia”

solifuga Iwasa 1939 appears distinct from T. tras-

kiana in the more elongate peduncular segment 3

of antenna 1, 4-dentate mandibular left lacinia,

weakly lobate maxilliped palp segment 2, short

palm of gnathopod 1 (@), narrow basis of

gnathopod 2 (@), unlike sparsely crenulate basis

of peraeopods 5-7, marginally setose pleopod

peduncles, and sublinear uropod 3 peduncle.

Until the condition of the peraeopod dactyls,

coxal gills, maxilliped palp, and brood plate

setae can be determined, the generic status of

O. solifuga is uncertain.

Distributional ecology

A widely eurytopic species, occurring mainly

under drift debris of rocky and stony beaches, but

also on sand (co-occurring with Megalorchestia

spp.) and in estuaries and salt marshes (often

with Paciforchestia and T. georgiana) of both

exposed and protected coasts, from the Aleutians

and western Alaska (northern and western limits

not determined) south to northern Baja Cali-

fornia; some populations terrestrial (sympatric

with Porcellio) in leaf litter near shore in the

Santa Barbara region. Females ovigerous during

spring and summer; bearing one brood per year

in the north, and in the south (south of Prt.

Conception) two generations per year, with the

spring brood maturing and breeding during the

late summer and early fall (Wenner, 1974). Mature

males of two size classes occur in warm-water

shores (southern California and Strait of

Georgia, B.C.).

Beach fleas and sandfleas were found by Murie

(1959) and Sheffer (1959) to be the most com-

monly available food item of blue foxes (Alopex

lagopus (L.)) on 22 islands of the western

Aleutians, from the Near Islands (including

Attu), through the Rat Islands (including

Amchitka and Kiska), eastward to the Andreanof

Islands (including Adak). Murie’s statements

that ‘“‘... (beach fleas) swarm on the beaches

where winnows of dead kelp furnish a favourable

habitat. They lurk under bits of wood or any-

thing else that may lie on the sand and preserve

the required moist shelter underneath.’’, imply

the presence of both beach fleas and sandhoppers.

Scheffer (1959) specifies in his records, Traskor-

chestia traskiana, the only species identified with

certainty in material of Talitridae from this

region (above). Murie’s further statements that

“it is easy for a fox to pick up a full meal of

sand fleas’, and, ‘‘an island with extensive

beaches, sand or gravel, is favourable for foxes’”’,

could reasonably apply to the occurrence of

Megalorchestia spp., or even to Trinorchestia.

Moreover, his description (p. 290) of “‘sand fleas”’

from Unimak Island (Fox Islands, eastern

Aleutians) as food of red foxes (Vulpes vulpes L.)

almost certainly would include 7. traskiana, the

dominant species in preserved material from

southern Alaska.

Traskorchestia georgiana (Bousfield 1958)

Figure 6

Orchestia georgiana Bousfield 1958, p. 887, fig. 3;

ror, p. 3; 1975, p. 363; 1981, fig. 17

: Staude et al., 1977, p. 12, couplet 21b and figs.

Material examined

British Columbia: Queen Charlotte Islands,

Bousfield stn. E25, 8 August 1957, north end

merspring Is., Moresby Is., 1 ¢, 1 2 -ov.,

13

NMC-C-1981-476. Southern Mainland; E.L.

Bousfield stn. EB 10, Stearman Beach, West

Vancouver, 19 June 1976, 1 imm. 9 NMC-C-

1981-483. Vancouver Island. E.L. Bousfield stns.

V25, V27, Little Tribune Bay, Hornby Is., drift

debris under Sargassum, sand beach, pan traps,

30-31 July 1959, 700+ specimens ( obo’, 2 Q,

imm.), NMC-C-1981-490. Savory Island, 30

March 1929, G.C. Carl coll., 1 @ subad.,

NMC-C-198 1-482. Sydney, B.C. gravel beach at

HW, June 1976, R. Long coll., 1 o& subad.,

18 99 br. I & Il, NMC-C-1981-487; Brady’s

Beach, near Bamfield, B.C., tide pools at HW,

29 July 1976, E.L. Bousfield coll., 7 &éo& (to

13.5 mm), 3 9 9 ov., 1 imm., NMC-C-1981-484.

Washington: E.L. Bousfield stn. M10b, Boundary

Bay, Salicornia flats, 2 September 1957, 1 o,

NMC-C-198 1-477.

California: Re-examined material from San Diego

region published by Bousfield (1961).

Mexico: Turtle Bay, near Ensenada, Baja Cali-

fornia, HW drift, 7 June 1956, W.L. Klawe coll.,

14 ¢oic' (to 15.0 mm); 1 o transfico Vy oe

(to 9.0 mm), 5 imm., NMNS Cat. No. 5123,

slide mount.

Diagnosis

Male (12-14 mm). Small to medium-sized

apomorphic species characterized by: antenna |

short, flagellum 3-4 segmented; antenna 2 short,

flagellum barely longer than peduncle, 12-14

segmented; maxilliped outer plate small, apex

rounded, barely exceeding inner plate; coxa 2

large, rounded below, nearly masking coxa 1;

gnathopod 1, segment 4 lacking posterior blister,

dactyl shorter than palm; gnathopod 2, propod

relatively short and deep, paim short, gently

convex, nearly vertical. Peraeopod 4, segment 5

very short and thick (relative to peraeopod 3);

dactyl short, strongly pinched, unguis short.

Peraeopods 5-7 relatively short and small (for

body), bases less broadly expanded than in

T. traskiana, marginal spinules fewer and more

widely spaced, peraeopod 7 not sexually dimor-

phic, hind lobe of basis distally obtuse, not

evenly rounded, segment 6 with transverse

apical row of 4-5 setae. Abdominal side plates 2

and 3 deep, distally broad, hind corners acute;

pleopods slender, somewhat reduced, Ist slightly

the shorter, rami 5-6 segmented. Uropod l,

peduncle with 8 inner and outer marginal spines,

inner ramus with 4-6 inner and outer marginal

spines; outer ramus with 5-6 tightly spaced outer

Figure 6. Traskorchestia georgiana (Bousfield). Cape Lazo, Vancouver Island, B.C. o& - 13.5 mm,

Q - 10.5 mm.

marginal ‘“‘comb-spines” (distally); uropod 2

peduncle with 4 outer marginal spines, strongest

distally, inner ramus with 2-3 inner and outer

marginal spines, outer ramus with 2-3 outer

marginal spines. Uropod 3, peduncle short, deep,

with 3 posterodistal spines; ramus short, tapering,

with distal and apical short spines. Telson short,

lobes each with dorsolateral medial groups of

3 spines, a subapical single spine, and 2-3

unequal apical spines. Coxal gills largest on

peraeopod 2 and peraeopod 6, gill of peraeopod 6

about twice length of gill on peraeopod S.

Female (5-8 mm). Much smaller than male at

maturity. Gnathopod 1, propod slender, palm

short, exceeded fully by unguis of dactyl, postero-

distal angle not (or very slightly) tumescent.

Gnathopod 2, basis slightly expanded proximally,

margins nearly parallel, segment 4 broadly

tumescent behind, 5 shallow, 6 short; brood plate

narrowing distally, with about 40 long hook-

tipped setae, mostly along anterior margin.

Uropod 1, outer ramus with 2-3 well spaced

marginal spines. Brood plate of peraeopod 5

smaller, margin with about 20 setae.

Remarks

Although related to T. traskiana more closely

than to western Pacific species, 7. georgiana is

unique in possessing sexually dimorphic uropod 1,

short pleopod 1, and females that are often barely

'/, the length of males at maturity.

Distributional ecology

Occurs mainly at the drift line of protected

stony and pebbly beaches, occurring on sand with

windrows of eelgrass and Sargassum, usually

ult

i

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:

\\il

Figure 7. Traskorchestia ochotensis (Brandt). Orca Island, Alaska. co - 15.0 mm. St. Paul Island,

Bering Sea. 92 ov. - 14.0 mm.

co-occurring with T. traskiana, in summer-warm Traskorchestia ochotensis (Brandt 1851)

bays and lagoons and inland seas, occsionally Figure 7

in warm tidal pools and warm springs near shore,

from southern Queen Charlotte Islands and Strait Orchestia ochotensis Brandt 185lc, p. 94, t. 6,

of Georgia, sporadically south to Pt. Conception, fig. 18-26

and commonly southward to Baja California. :‘Stebbing, 1906a, p. 543

Animals are slower moving, less saltatory than :Derzhavin, 1937, p. 88, pl. 1,(2)

T. traskiana. Females ovigerous in spring and :Gurjanova, 1951, p. 804, fig. 559

summer, probably with two or more generations :Bulycheva, 1957, p. 166, figs. 61a, b

per year in the south. :Bousfield, 1981, p. 86, fig. 17

non Orchestia ditmari Derzhavin 1923, p. 187

Is

Material examined

Southeastern Alaska: Bousfield & McAllister stn.

A153, Mummy Is., Orca Inlet, 16 July 1961, 1 ¢

(16.0 mm), NMNS Cat. No. NMC-C-1981-580,

slide mount; St. Paul Is., Bering Sea, July 1914,

Gu Parker-coll., 5 ocd (to 14:5 mm), 2: O.9

ov., NMNS Cat. No. NMC-C-1981-578, slide

mounts.

Western Pacific: Southern Sakhalin Is., USSR,

4 September 1946, E.F. Gurjanova coll.,3 vo

(to 18.5 mm), 4 2 2 (br. II, to 13.0 mm), Zool.

Inst. No. 32/32313, slide mounts; Nosappu,

Hokkaido, Japan, 21 June 1971, H. Morino coll.,

3° oo (to 20:5 mm); 2-2 2..0v: @o- 13,0 mm),

NMNS Cat. No. NMC-C-198 1-576, slide mounts.

Diagnosis

Male (15-20 mm). Medium to large, moderately

apomorphic species characterized by: antenna 1,

flagellum nearly equal to peduncle, 6-7 seg-

mented; antenna 2 stout (not incrassate),

flagellum 15-18 segmented. Mandibular left

lacinia 4+-dentate, Sth tooth minute, occasion-

ally lacking (especially in female); maxilliped palp

normal, outer plate medium, apically rounded.

Coxa | about '/, masked by coxa 2; coxae 2-4

wider than deep, rounded below. Gnathopod 1,

segment 4 lacking posterior tumescence or blister;

dactyl slightly shorter than palm of propod;

gnathopod 2, propod large, slightly broadening

distally, palm gently convex, oblique. Peraeopod

4, segment 5 short (about 7/, segment 6), dactyl

strongly pinched, nail long. Peraeopods 5-7,

bases slightly dissimilar in shape, hind margin of

5 and 6 rounded, of 7 straight, weakly serrate;

peraeopod 7 longest and slightly sexually

dimorphic, segment 6 distally with longitudinal

facial row of brush setae increasing in length

distally, and short transverse apical row of 5-6

short setae. Abdominal side plates 2 and 3 deep,

hind corners acute, hind margins weakly serrulate

and setulose; pleopods much reduced, third

shortest, rami 2-5 segmented, distinctly shorter

than peduncles. Uropod 1, peduncular margins

with 3-4 spines, inner ramus with 5-6 inner

marginal and 3 outer marginal spines; outer

ramus with 2-3 widely spaced marginal spines,

not sexually dimorphic; uropod 2, outer margin

of peduncle with 3-4 spines, heaviest distally;

inner ramus with 2-3 inner and outer marginal

spines; outer ramus with 2-3 outer marginal

spines. Uropod 3, peduncle relatively shallow,

with 3-4 mostly short posterior marginal spines;

ramus much shorter, with short posterior

marginal spines and longer apical spines. Telson

lobes each with several dorsolateral spine groups

and 3-4 apical spines of unequal size. Coxal gills

2 and 6 much the largest, 6 elongate, more than

twice the length of gill 5.

Female (12-15 mm ov.). Gnathopod -1, palm

distinct, with small posterodistal tumescence,

exceeded by most of unguis of closed dactyl,

posterior marginal spines short. Gnathopod 2,

segment 2 widest proximally, segment 4 broadly

tumescent behind, segment 5 deeply tumescent,

6 slightly shorter than 5. Brood plate margins

subparallel, with about 40 long hook-tipped setae,

mainly anteriorly. Peraeopod 5, length of brood

plate about half that of gnathopod 2, margins

20-setose. Females are distinguishable from

females of 7. traskiana not only by the greatly

reduced pleopod rami, but also by the longer

antenna 1, smaller palm of gnathopod 1, longer

and narrower brood plates, relatively longer

peraeopods 5-7, with longer dactyls and less

strongly convex posterior margins of bases.

Distributional ecology

Under drift-line debris, on sandy, rocky and

stony beaches of exposed and semi-protected

shores of the entire North Pacific rim region from

Japan (north Honshu and Hokkaido) and

Sakhalin in the west through Kamchatka, St. Paul

Island to southeastern Alaska (Prince William

Sound) in the east. Since the distribution of

Traskorchestia ochotensis brackets the Aleutian

Island chain, the records of Murie (1959) for

‘“‘beach fleas’? and “‘sand fleas’ from that region

very probably include this species also. Females

ovigerous in May-July; apparently only one

brood per year.

Remarks

Minor morphological variations occur in mature

males and females throughout the range, but a

15.0 mm male from Orca Island, southeastern

Alaska compares closely with a 15.0 mm subadult

male from Noppura, Hokkaido. Mature males

have a sexually dimorphic peraeopod 7 in which

segment 4 broadens distally, the hind margin is

arcuate, and segment 6 is bowed rather markedly,

not straight. This species is not to be confused

with the somewhat smaller T: ditmari Derzhavin

(1923) that occurs along the shores of the Sea of

Okhotsk in Kamchatka and the Kurile Islands

(Derzhavin, 1937). Lack of material of T. ditmari

prevents critical evaluation of differences in

females and mouthparts, pleopods, etc.; however,

comparison on material of 7. ochotensis (on hand)

with the rather good drawings of Derzhavin

(1923) has revealed species differences believed

significant.

Paciforchestia new genus

Diagnosis

Medium to medium-large beach fleas, character-

ized by: smooth, unmodified bodies; eyes lateral,

medium, not dorsally contiguous; inferior

antennal sinus distinct, shallow; antenna 1

extending beyond peduncular segment 4 of

antenna 2, peduncle 3 longest, flagellum

upturned distally; antenna 2 slender, elongate,

geniculate at flagellum, not incrassate (dc).

Buccal mass directly beneath head, not very

deep; mandible, left lacinia fully 5-dentate;

right lacinia tricuspate; maxilliped palp obscurely

4-segmented (4th minute, masked by distal spines

of 3), segment 2 broad, inner distal lobe large.

Coxae 2-4 broad, rounded ventrally, cuspate

posteriorly. Gnathopod 1 (o'), segments 4, 5

and 6 tumescent behind, palm not exceeded by

dactyl; gnathopod 1 (@) blister vestigial or

lacking on segment 4; dactyl not exceeding palm.

Gnathopod 2 (0) powerfully subchelate, palm

smoothly oblique, unguis of dactyl attenuate.

Gnathopod 2 (9), basis slightly to moderately

expanded anteriorly, segment 3 slightly elongate,

segments 4 and 5 with weak or low posterior

blisters. Peraeopods 3 and 4 elongate, unequal,

cuspidactylate, dactyls short, that of peraeopod 4

weakly pinched; peraeopod 4, segment 5 shorter

than in peraeopod 3; peraeopods 5-7 homo-

podous, increasing posteriorly, bases rounded

behind; coxa 5 anterolobate; peraeopod 7

sexually dimorphic in length but not in form.

Abdominal side plates deep, weakly setulose

behind, hind corners minutely acuminate.

Pleopods more or less reduced, pleopod 3

shortest, rami shorter than peduncles, peduncular

margins smooth, retinacula 2-8. Uropods | and 2

slender, rami and peduncles marginally spinose,

peduncle | with laterodistal (inter-ramal) spine.

Uropod 3, peduncle very deep, strongly spinose

posteriorly, ramus shorter, spinulose posteriorly

and apically. Telson long, narrowing, lobes

separated near tip, with a few short apical spines,

occasionally singly inserted dorsolateral spines.

Coxal gills reduced, modified, that of peraeopod

6 elongate, sinuous. Brood plates large, subovate,

marginal setae numerous, long, simple-tipped.

Type species: Parorchestia klawei Bousfield

1961

Additional species:

Paciforchestia pyatakovi (Derzhavin 1937),

p. 89, fig. II(i)

See also Bulycheva, 1957, p. 172, figs. 62a,

b, and Morino, 1975, p. 178, figs. 9-11

Paciforchestia tenuimana (Iwasa 1939), p. 268,

pl. XIII, figs. 10-11

Etymology

The generic name is a combining form of

Orchestia and the regional geographic name

Pacific. Gender: Feminine.

Remarks

‘““Orchestia’”’? kokuboi Ueno 1929 has many

features of this genus, but the laterally plumose

peduncles of the pleopods, short telson, and

absence of a distolateral spine on uropod | are

atypical. Paciforchestia is extremely plesio-

morphic, little advanced beyond Protorchestia

and members of the palustral group and primitive

terrestrial Talitridae of the southern hemisphere.

(Tables II, IV; figs. 26, 28)

Key to species of Paciforchestia

All pleopods much reduced, rami distinctly shorter than peduncle, 1-3 segmented; uropod 3,

ramus slender, cylindrical, about */, peduncle length; telson lobes with apical spines only (rarely

Ser MA ORSONILET Al SPEMESY™§ "ee ssteheete a dt so as wk ORR, coe a ae be wee cee Do 2

Only pleopod 3 much reduced, rami shorter than peduncle, 4-8 segmented; uropod 3, ramus

short, subconical, about '/, length of peduncle; telson lobes with 2 dorsolateral spine groups,

PME EA RSIa mcs Wits. oe, Ses ee Me es 2s x os alsa ta es ee ee eke we P. klawei (Bousfield) (p. 18)

Antenna 1, peduncular segment 3 distinctly ('/,) longer than peduncle 2, hind margin with small

spines; uropod tPouter ramus marginally smooth ... 62.3. 2.00.05. ee. P. tenuimana (Iwasa)

Antenna |, peduncular segment 3 slightly longer than peduncle 2, hind marginsmooth; uropod 1,

Surce rants distally with marginal spines .%..... 02 ee a P. pyatakovi (Derzhavin)

Figure 8. Paciforchestia klawei (Bousfield). San Juan Islands, Washington, USA. of - 14.5 mm,

@ ov. - 13.0 mm.

Paciforchestia klawei (Bousfield 1961)

Figure 8

Parorchestia klawei Bousfield 1961, p. 3, fig. 1, 2

(2) Barnard, 1969b, p. 76, 221

Material examined

British Columbia: E.L. Bousfield 1955 stn. Pé6a,

Wickininnish Bay, south end, fine sand at HW

level, 2 August 1955, 1 imm. (5.0 mm); stn. M2,

Emmond’s Beach, near Powell River, pebble

beach, HW level, 25 August 1955, 1 @ br. II

(11.5 mm), NMC-C-1981-555, slide mount;

Portland I. (48°43.5’N,123°22’W), intertidal, 15

June, 1927, 1 & (14.5 mm), BCPM collections

(courtesy P. Lambert).

Washington: San Juan Co.; Kiket Is., intertidal,

26 October 1968, Houghton coll., 2 ¢éo& (to

15.0 mm), 1 @ br. II (brood plate II stage)

(to 14.0 mm), Friday Harbor Marine Laboratory

collection, slide mounts. Goose Island, sand

beach under gravel and Fucus mat (0-30 cm

level), 26 April 1977. E. Kozlof student team B.,

1 9 br. II, Friday Harbor Marine Laboratory

collections.

Mexico: Los Coronados, off Baja California,

25 January 1957, W.L. Klawe coll. 2 oo

(to 10 mm), 2 2 9, 1 ov. (to 10.0 mm). NMNS

Cat. No. 6167, slide mount.

Diagnosis

As described originally. The smaller North

American Pacific species differs from the two

known large western Pacific species in a number

of characters that in toto may be of subgeneric

significance, viz: Antenna 1, peduncular segments

2 and 3 short, little longer than segment 1;

gnathopod 2 (c'), palm with weakly defined

hinge tooth; gnathopod 2 ( 2) basis very slightly

expanded proximally, weakly arcuate posteriorly;

coxae 2-4 medium deep, hind margin nearly

vertical (versus sloped forward); coxa 5, anterior

lobe moderately larger and deeper than posterior

lobe; peraeopods 3-7, dactyls short, unguis not

attenuated. Peraeopod 7 slightly longer than

peraeopod 6 (versus much longer) in male. Pleo-

pod rami multi-articulate, peduncles with 3-5

retinacula. Uropod 2, inner ramus with inner and

outer marginal spines; uropod 3, ramus short

and subconical versus slender-cylindrical. Telson

broadly tapering, lobes with dorso-lateral and

apical groups of spines, apical notch shallow.

Coxal gills smaller, less strongly lobate or

attenuated. Brood plates broader, marginal

setae more numerous.

Remarks

This species complex encompasses plesiomorphic

features of palustral and some terrestrial talitrids

in the homopodous peraeopods; fully subchelate

and palmate gnathopod 1 (both sexes); short

dactyls, weakly “pinched” or incised peraeopod 4

dactyl; uropod | with distolateral peduncular

spine; and elongate, weakly armed telson, but is

distinguished by the 5-dentate left lacinia,

cuspidactylate peraeopods, and deep peduncle of

uropod 3.

Distribution ecology

Under drift debris on protected but bermless

coarse sand and pebbly beaches, often with

Traskorchestia traskiana and occasionally with

T. georgiana, but usually much less abundant;

throughout the Strait of Georgia, British

Columbia (in summer-warm, slightly brackish

areas only); also Santa Barbara, Southern

California, to northern Baja California.

Transorchestia new genus

Diagnosis

Medium to medium-large smooth-bodied beach

fleas, almost exclusively southern hemisphere,

characterized by: eyes medium, lateral; antenna 1,

peduncular segment 3 longest; antenna 2 sexually

dimorphic, peduncle incrassate in male; inferior

antennal sinus distinct, medium deep; buccal

mass directly ventrally; mandibular left lacinia

cleanly 4-dentate (rarely with Sth vestigial tooth),

right lacinia tricuspate; maxilliped palp obscurely

4-segmented (4th minute, masked by spines of

3rd), segment 2 broad, with distinct medio-distal

lobe. Gnathopod | (0) deeply subchelate, palm

much exceeding dactyl, segments 5 and 6 strongly,

and 4 weakly tumescent behind; gnathopod |

(2) fully subchelate, dactyl slightly (or not)

exceeding transverse palm, segments 4-6 lacking

tumescent process; gnathopod 2 (o'), propod

powerfully expanded, palm oblique, usually

with stout (hinge) tooth, dactyl sinuous; gnatho-

pod 2 (9), basis little expanded, segment 3 short,

4 and 5 shallowly tumescent behind. Coxae 2-4

rounded ventrally, strongly cuspate behind.

Peraeopods 3-7 cuspidactylate, dactyls short;

peraeopod 4 shorter than peraeopod 3, segment 5

shorter, dactyl weakly pinched; peraeopods 5-7

weakly heteropodous, peraeopod 7 longest,

peraeopods 6 and 7 sexually dimorphic in size

and form; coxa 5 anterolobate, coxa 6 postero-

lobate, anterior margin of hind lobe nearly

vertical. Abdominal side plates 2 and 3 weakly

serrate behind, hind corners subacute; pleopods

slender, normal (rami usually longer than

peduncle), outer margin of peduncles very weakly

spinose and/or lined with fine ‘“‘fuzz’’ setae.

Uropod 1, peduncular distolateral (inter-ramal)

spine not developed, rami weakly marginally

spinose; uropod 2, inner ramus, both margins

spinose. Uropod 3, peduncle deeply broadened,

posterior margin short-spinose; ramus short.

Telson elongate, tongue-shaped, lobes distally

narrowing, separated apically, each with long

dorsolateral marginal row of short stout spines,

and apical spines. Coxal gills 3-5 somewhat

reduced, those of peraeopod 2 and peraeopod 6

large and bladder-like. Brood plates ( 9 ) broadly

subovate, margins with numerous long hook-

tipped setae.

Type-species: Orchestia chiliensis Milrie-

Edwards 1840

Additional species: Transorchestia serrulata

(Dana 1852)

Transorchestia enigmatica (Bousfield & Carlton

1967)

Transorchestia chiliensis gracilis (Chilton 1920);

Hurley, 1957, p. 160

Transorchestia miranda (Chilton 1916); Hurley,

1957, po 162; figser4 75

Transorchestia sp. (Australian species, Bous-

field, (in prep.)!

(?)Transorchestia bollonsi (Chilton 1909);

Hurley, 1957, p. 160, fig. 3; Bousfield,

1964, p. 54, fig. 5

'Bousfield, E.L. A revised classification of Talitrid amphipods

(Crustacea: Amphipoda: Talitridae). (In prep.)

Etymology

The generic name reflects its phyletically inter-

mediate position between the most primitive

and most advanced members of the beach flea

genus Orchestia (sens. lat.) Gender: Feminine.

Remarks

The four species and subspecies of the chiliensis

complex are herewith maintained as separate

entities pending study of more extensive material

from South America, Australia, New Zealand,

and other land masses of the southern hemi-

sphere. 7. bollonsi from the subantarctic islands

of New Zealand is tentatively included in this

group despite its special morphological features

(reduced pleopod rami, knob-like brood plate

setae, short palm of gnathopod | (2), and more

heavily spinose uropod rami). The full species

groups may be separated according to the

following key. The genus appears intermediate

between the plesiomorphic Traskorchestia and

the more apomorphic Orchestia sens. str., and

may be antecedent to the latter, hence the name

Transorchestia. (Tables II, IV; figs. 26, 28)

Key to described species of Transorchestia

1. | Uropod 3, ramus sublinear, nearly equal to peduncle; gnathopod 1 ( 2), palm short, distinctly

exceeded by dactyl; pleopods reduced, rami distinctly shorter than peduncle; brood plate setae

club-tipped Transorchestia bollonsi (Chilton)

Uropod 3, ramus subconical, much shorter than peduncle, gnathopod 1 ( 9 ), palm little (or not)

exceeded by dactyl; pleopods normal, rami not shorter than peduncles; brood plate setae

hook-tipped

2. | Gnathopod 2 (<'), palm smooth, convex, lacking well-defined hinge tooth; peraeopod 7 (’)

segment 5 strongly broadened posteriorly, width nearly equal to length

T. miranda (Chilton)

Gnathopod 2(<'), palm concave posterior to strong hinge tooth; peraeopod 7(0')segment Sless

strongly inflated, width about ’/, length

T. chiliensis group (p. 20)

(comprises T. chiliensis (Milne-Edwards), T. serrulata (Dana), T. enigmatica (Bousfield &

Carlton), and T: chiliensis gracilis (Chilton)).

Transorchestia chiliensis (Milne-Edwards 1840)

Figure 9

Orchestia chiliensis Milne-Edwards 1840, p. 18

:‘Stebbing, 1906a, p. 537

‘Ruffo, 1949, p. 53, fig. 18

non Orchestia chiliensis: Hurley, 1957, p. 157,

fie. 2

Material examined

Cape Horn region, South America, Hudson 70

Expedition: E.L. Bousfield stn. F.1, Puerto

Robalo, Isla Navarino, Chile (54°57’S,67°40’W),

29 January 1970, 7 &o& (to 14.5 mm), 11 9 9

(7 ov.) 3 juv.; stn. F.14, Playa Aaron, Isla

Navarino, 5 February 1970, 2 oc; stn. F.17,

Cabo Maria, Isla Navarino, 7 February 1970,

36.0,5 3 2 (ov.),3 juv.,/sth..F.20, Peninsula

Scott, west beach, Isla Navarino, 10 February

1970, Jig S312) 9 2. 164quy.asin, F 34: Punta

Zegers, Tierra del Fuego, Chile, 22 February

1970,.19 o do (to:.16.0: mm), 119 9 (ev) (to

13.0 mm), 3 juv. NMNS collections, slide mounts.

20

Diagnosis

Male (14.0-16.0 mm). A medium-sized, stout-

bodied species, closely agreeing with the descrip-

tion and figures of Ruffo (1949), but somewhat

less well with ‘‘O. chiliensis” (probably O. serru-

lata Dana) as described and figured in detail by

Hurley, 1957 (above). Distinctive features are:

antenna 1, peduncular segment 3 often with 2

posterior marginal spines, and segment | with

posterodistal spine (rather than setal group);

antenna 2, segments 4 and 5 stoutly incrassate,

flagellum 17-segmented. Gnathopod |, segment 4

with weak but distinct posterior blister; gnatho-

pod 2, basis not noticeably expanded or serrate

antero-distally; propod, palmar hinge tooth very

prominent, posterior sinuous bulge of dactyl

about mid-point, closing on palm just below

hinge tooth; peraeopod dactyls very short,

relatively thick proximally, outer margin convex,

unguis very short; dactyl of peraeopod 4 very

short, nail about '/, dactyl body length. Peraeo-

pods 6 and 7, segments 4 and 5 strongly

incrassate, width more than '/, length, broader

peieeepaiiaedieenas

.

ZY

Y

\ MX2

S

Q?

oe,

cag anne JA

gees.

Figure 9. Transorchestia chiliensis (Milne-Edwards). Puerto Robalo, Chile.

12.0 mm.

on peraeopod 7. Pleopods 1, 2 and 3, peduncles

with 1, 2 and 3 outer marginal short spines, and

fine fuzz setae, barely visible on 1. Uropod 2,

inner ramus with 2 medial marginal and | lateral

marginal spines respectively. Uropod 3, basis

with 5 posterodistal spines, middle spine the

largest. Telson lobes each with about 6 dorso-

lateral spines, proximal Ist and 3rd largest, and

2 unequal apical spines.

Female (12.0 mm, ov.). Gnathopod 2, basis

relatively short and broad, anterior margin very

slightly indented distally, lined with 6-7 well-

spaced spinules. Brood plate of peraeopod 2

broadest near base, anterior margin with about

30, and posterior margin distally with about 10,

long hook-tipped setae.

21

LFT

MD

& - 15.0 mm, 9 ov. -

Remarks

Mature males of O. chiliensis differ from similar-

sized rocky-shore specimens from Portobello,

New Zealand, in the shorter but more spinose

antenna 1, longer flagellum of antenna 2, more

proximal sinuosity of the dactyl of gnathopod,

relatively short, bowed dactyls of peraeopods,

slightly more spinose pleopod peduncles, and

slight but consistent differences in armature of

uropods and telson. In comparable female

material from New Zealand, the basis of gnatho-

pod 2 is relatively longer, the anterior margin

more strongly incised, and the brood plates of

peraeopod 2 have about 25 and 15 hook-tipped

setae on anterior and posterior margins

respectively.

Transorchestia enigmatica (Bousfield & Carlton

1967)

Figure 10

Orchestia enigmatica Bousfield & Carlton 1967,

Doves. ties: “1,*2

Orchestia_ chiliensis:

fig. 234a, b

Bousfield, 1975, p. 363,

Material examined

Lake Merritt, Oakland, Alamedo Co., California,

among tubes of Mercierella enigmatica, 17 June

1965, J. Carlton coll., 1 & (15.5 mm), Paratype,

USNM File No. 260765, slide mount (in NMNS,

Ottawa). Type and paratype material from Lake

Merritt, NMC-C-1981-53,532.

Distributional ecology

Intertidally under damp rotting wood, leaves,

and other organic debris, and occasionally

among the tubes of Mercierella enigmatica

Fauvel, around the brackish-water margins of

Lake Merritt, the levels of which fluctuate

according to the flood gate. Females ovigerous

in summer (June to August).

Remarks

The species has been recorded to date only from

the shores of Lake Merritt, within the city of

Oakland, California. After further examination

of fully mature males, the species was assigned

to O. chiliensis by Bousfield (1975). However,

careful comparison of all material of T. enigmatica

from Lake Merritt with extensive series of T: chi-

liensis from the Cape Horn region (the general

type-locality) reveals consistent differences,

especially in the longer, more slender peraeopod

dactyls, more narrowly ovate female brood

plates with more numerous posterior marginal

setae, more distal location of posterior marginal

bulge of the dactyl of gnathopod 2 (co), and

minor differences in the form and armature of

uropod 3 and telson. These differences indicate

that 7. enigmatica is distinct from rocky shore

material of 7. chiliensis sens. str., especially from

the Cape Horn region. However, comparison of

mature male and female specimens of T. enig-

matica with similar-sized material from a salt-

marsh near Miranda, North Island, New Zealand,

E.L.. Bousfield; collector, October 1978 (cf.

Paviour-Smith, 1956), yields closer agreement in:

yi)

less incrassate antennae and peraeopods 6 and 7;

more attenuated, slender peraeopod dactyls;

more strongly spinose peduncles of pleopods

2 and 3, and more similar armature of uropod 3

and telson. In Miranda marsh females, however,

the brood plate of gnathopod 2 is less broad, and

the anterior and posterior margins have 23 and:

15 hook-tipped setae respectively. T. chiliensis

gracilis Chilton from Juan Fernandez, is a

terrestrial species, shows several distinctive

morphological features of gills and gnathopods,

and is almost certainly a valid full species.

Although the probability is high that T. enigma-

tica is conspecific with one of these (or an

undescribed) species or forms from the southern

hemisphere, the species name is herewith retained

pending study of more extensive materials from

all pertinent regions.

Orchestia Leach 1813-14

Diagnosis

The genus Orchestia has been more narrowly

redefined by Bousfield (in prep.)' to encompass

the type group of shore-dwelling and immediate

“coastal terrestrial” species of the Atlantic-

Mediterranean region (e.g., of Karaman, 1970),

with the following primary characteristics: body

smooth, medium to medium-large. Eyes lateral,

not enlarged or dorsally contiguous. Inferior

antennal sinus distinct, medium deep. Antenna |

short, peduncular segment 3 not longer than 2,

bare; antenna 2 usually sexually dimorphic.

Buccal mass slightly prognathous, mandibular

left lacinia cleanly 4-dentate, rarely with vestigial

Sth tooth; maxilliped palp 3-segmented, segment 2

broad, with distinct, medio-distal lobe. Gnatho-

pod | (co) fully subchelate, palm little exceeding

dactyl, segments 5 and 6 (but not 4) tumescent;

gnathopod 1 (@), segment 5 slender, dactyl

distinctly exceeding short palm, segments 4-6

lacking tumescent process. Gnathopod 2 (c),

propod powerful, palm oblique, usually with

tooth, dactyl usually sinuous; gnathopod 2 (9),

basis broadly expanded anteriorly, segment 3

short, 4 not tumescent posteriorly. Peraeopods

3-7 cuspidactylate, peraeopod 4 shorter than 3,

segment 5 shorter, dactyl distinctly pinched.

'Bousfield, E.L. A revised classification of Talitrid amphipods

(Crustacea: Amphipoda: Talitridae). (In prep.)

Figure 10. Transorchestia enigmatica (Bousfield & Carlton). Lake Merritt, Oakland, California. o& -

11.5 mm; 2 @ ov. - 11.0 mm. (Recomposed from Bousfield & Carlton, 1967).

Peraeopods 5-7 weakly heteropodous, dactyls

slender; peraeopod 7 longest, usually sexually

dimorphic in form and size; coxa 5 anterolo-

bate, 6 posterolobate, hind lobe oblique, antero-

23

distally rounded. Abdominal side plates 2 and 3

weakly crenulate behind, posterior corner acute,

lower face not pitted. Pleopods slender, normal

or slightly reduced; peduncles 2 and 3 very weakly

spinose, lacking fine setae, each with 2 retin-

acula. Uropods 1 and 2, peduncle with disto-

lateral spine weak or lacking, rami marginally

spinose, both margins of inner ramus spinose.

Uropod 3, peduncle not strongly broadened;

ramus short, tapering. Telson not elongate, lobes

with dorsolateral and apical spine groups, weakly

notched or separated apically. Coxal gills 3-5

reduced. Brood plates large, narrowly ovate;

marginal setae numerous, simple.

Type-species: Oniscus gammarellus Pallas 1766

(monotype and subsequent synonymy)

The group comprises about 15 mostly

Mediterranean-Atlantic species including O. mon-

tagui (Audouin 1826), O. mediterranea Costa

1857, O. cavimana Heller 1865, O. gambierensis

Chevreux 1908, O. magnifica Vecchi 1931,

O. remyi Schellenberg 1950, + subspecies

roffensis Wildish 1968, O. grillus Bosc 1802,

O. stephenseni Cecchini 1928, O. kosswigi Ruffo

1949 and O. microphtalma Amanieu & Salvat

1963, (See also Karaman, 1970).

Section II. Sandhoppers

(substrate modifiers, sensu MacIntyre, 1963)

Recent revision of world sandhoppers (Bousfield,

in prep.)' has subdivided the group primarily

on the basis of dentition of the mandibular left

lacinia mobilis. This character is somewhat

variable within the two groups and not absolutely

taxonomically critical, reflecting the extreme

difficulty of defining natural groups on a rigid

taxonomic basis, and advisability of the character

consensus approach to natural groupings. Thus,

dentition of the left lacinia mobilis tends to be

strongly correlated with certain conditions of

other morphological characters, as follows:

4-dentate group

Body generally stout, surface occasionally rugose

or ridged. Eyes medium to moderately enlarged,

not contiguous mid-dorsally. Antenna 2 short,

especially in female, seldom sexually dimorphic

in form; buccal mass directly ventral; mandibular

left lacinia 4-dentate, rarely 5-dentate. Coxae

2-4 deep, weakly or not cuspate behind. Gnatho-

pod 1 (c) often fully subchelate, dactyl exceeding

palm. Gnathopod 1 (2), palm small or lacking.

Gnathopod 2 ( 9 ), basis little expanded anteriorly,

segment 4 rarely tumid or processiferous behind.

'Bousfield, E.L. A revised classification of Talitrid amphipods

(Crustacea: Amphipoda: Talitridae). (In prep.)

24

Coxa 5 aequilobate, coxa 6 strongly and deeply

posterolobate, anterior margin vertical. Uropod 3

short, ramus tapering (seldom compressed),

rarely longer than peduncle or sexually dimor-

phic. Telson very short and broad, lobes fused

at apex, rarely with dorsal spines (marginal only).

Coxal gills relatively large, 3-5 not markedly

smaller than 2 and 6; brood plates broadly ovate,

large on peraeopod 5.

Composition

Orchestoidea Nicolet (sens. str.), ““Orchestoidea”’

brasiliensis (Dana 1853), Talitrus saltator

Montagu 1808, and most sand beach hoppers of

southern Australia-New Zealand and South

Africa (e.g., of Hurley, 1956).

5-dentate group

Body stout or slender, surface usually smooth.

Eyes medium to very large, often nearly conti-

guous mid-dorsally, often sexually dimorphic.

Antenna 2 relatively long, usually strongly

sexually dimorphic, peduncle often incrassate in

male; mandibular left lacinia usually cleanly

5-6 dentate, seldom with Sth (proximal) tooth

vestigial or lacking. Coxae 2-4 often as broad

as deep, usually strongly cuspate behind.

Gnathopod 1 (<') seldom fully subchelate, palm

usually vestigial or lacking. Gnathopod 1 (92)

palm usually lacking, very short when present.

Gnathopod 2 (¢), basis more or less strongly

expanded anteriorly, segment 4 often with

posterior process. Coxa 5 anterolobate; coxa 6

posterolobate, lobe often shallow, anterior

margin often oblique. Uropod 3 various, ramus

usually longer than peduncle, often laterally

compressed, occasionally sexually dimorphic.

Telson lobes more elongate, often separated

apically, bearing dorsal and apical spines. Coxal

gills strongly reduced, those of peraeopods 3-5

small. Brood plates elongate ovate, variable in

length, occasionally lacking, that of peraeopod 5

short, weakly setose.

Composition

Platorchestia new genus (partim), Megalorchestia

Brandt, Trinorchestia new genus, Pseudorches-

toidea new genus, Talorchestia Dana, most

species of the Indo-Pacific and northern

hemisphere and new generic groups proposed by

Bousfield (in prep.)'.

Key to genera of sandhoppers of the North Pacific rim region

and selected world-wide genera

Mandibular left lacinia basically 4-dentate (fig. 1d); gnathopod 2 ( 9 ) basis sublinear, little

expanded anteriorly (fig. 2n); uropod 3 short, ramus cylindrical or tapering, not longer than

peduncle (figs. 1i, }); telson short, broad, entire, with marginal spines only (fig. Ip) ......... 2

Mandibular left lacinia basically 5-dentate; gnathopod 2 ( 2 ) basis more or less strongly

broadened anteriorly (fig. 2p); uropod 3 medium to elongate, ramus usually laterally compressed,

longer than peduncle; telson as long as broad, with dorsal and marginal spines (fig. lo) ...... 4

Gnathopod | distinctly subchelate in male, often very weakly so in female (figs. 2a,g) .......

Be eter ate, Care ee waren e a's Sa Most “‘Talorchestia”’ species of the southern hemisphere

Gnathopod | lacking distinct palm in male and female (Figs. 2b,c,d,h) ............0e00ee 3

Gnathopod 2 (oc) powerfully subchelate; pleopods reduced, peduncles widened, outer margin

strongly long-spinose (fig. 3q); uropod 3, apical spines numerous, short ............+ee0ee-

ete Setar eS ee AOE wists ate aig KGW 0k! 6 w «dea ws RO oR Orchestoidea Nicolet (p. 43)

Gnathopod 2, propod small, mitten-shaped in male (as in female and imm.); pleopods normal,

rami long, peduncles slender, outer margin short-spinose (fig. 3p); uropod 3, ramus with single

REGIE BEIGE Decides Fhe ee erieih ss Winnll Pe dw sow RORY sae atae Sea Talitrus Latr. (p. 45)

All pleopod peduncles spinose along entire outer margin (figs. 3n, p); uropod 1, peduncle and

outer famus always stoutly marginally spinose (fig. 3g)... ...6:0 0% 2 hace sive alee Oe 5

Pleopod peduncles, especially pleopod 1, outer margin not spinose or only partly so; uropod 1,

peduncle and outer ramus weakly or not spinose marginally (fig. 3f) ..............0 cee eeee 6

Pleopod rami very reduced, much shorter than peduncles; peduncles basally broadened,

marginal spines long, strong (fig. 3r); gnathopod 1 (co and 9), propod long, tapering distally,

ieneth greater than .'/, anterior margin of carpus (figs. 2d, h). so < 6 cccdenestet.cvieas sees

nh oh cs Sa ae re ech hee elt cE via a brine RR CRN Megalorchestia Brandt (p. 29)

Pleopod rami not reduced, multi-segmented; peduncles linear, marginal spines short (fig. 3p);

gnathopod 1(¢' and 9), propod short, stout, length about '/, anterior margin of carpus (fig. 2b)

EGE is. cs eee Sotieiciae Ws eitaadlaw » « ss ewan «cs Pep ds See Trinorchestia new genus (p. 41)

Peraeopod 5 much smaller than peraeopods 6 and 7 and differing markedly in form of basis and

segment 4 (fig. 3n); peraeopod 5 dactyl short, thick, nail vestigial; gnathopod | (co), propod

totally lacking palmar margin between posterodistal blister and unguis (fig. 2c); uropods land 2,

rami usually with terminal spade spines (fig. 3f). Mandibular left lacinia, Sth (proximal) tooth

ene OE BRCM Stoica. « ila uate: « «ss we Ree Ee Pseudorchestoidea new genus (p. 47)

Peraeopod 5 smaller than, but similar in form to peraeopods 6 and 7 (fig. 3m); peraeopod 5,

dactyl normal, nail elongate; gnathopod | (o’) more or less subchelate, propod with short,

distinct palm (fig. 2a); uropods | and 2, terminal spines regular, acute-tipped (fig. 3c). Mandibular

fereiacinia Gleanly. S-ClENLALE lenses. sadenres «s+» Baluwanrs eepet< eo ewes chaos ee ene 7

Gnathopod 1, propod with distinct palm, dactyl of female exceeding, dactyl of male little or not

exceeding, palm; coxa of peraeopod 6, posterior lobe, anterior margin vertical, often with distinct

distal process (fig. 3j); antenna 2 and peraeopod 7 strongly sexually dimorphic in form, some

segments incrassate or broadened in male (figs. 3c,k) ............ Platorchestia new genus (p. 26)

Gnathopod 1, propod with small but distinct palm only in male, always exceeded by dactyl; coxa

of peraeopod 6, hind lobe rounding anterodistally, margin usually oblique; antenna 2 and

peraeopod 7 (in male) elongate but not differing in form from female .......... Talorchestia Dana

25

Platorchestia new genus

Diagnosis

Medium-sized, smooth-bodied, semi-fossorial

talitrids, little evolved from the “beach flea”

facies (relatively weakly spinose appendages),

characterized by: antenna | short, not exceeding

peduncle 4 of antenna 2, peduncular segments

subequal; antenna 2 short, sexually dimorphic

(peduncle usually strongly incrassate in male);

inferior antennal sinus shallow, distinct; eyes

medium to small, vertically subrectangular,

quadrate to round. Buccal mass directly beneath

head; left mandibular lacinia cleanly 5-dentate;

maxilliped palp relatively short, broad, obscurely

4-segmented, segment 2 strongly spinose medially,

with mediodistal lobe, segment 3 rounded

apically, spines masking minute 4th segment.

Coxa | shallow, shorter than cuspate coxae 2-4,

anterodistal border rounded, not sharply acute.

Gnathopod 1 (o') distinctly subchelate, dactyl

equal to or slightly exceeding palm, propod not

narrowing distally, carpus not elongate, seg-

ments 5 and 6 with posterior tumescence;

gnathopod 1 (9), propod with short palm,

greatly exceeded by dactyl, segments 4-6 not

tumescent behind; gnathopod 2 (o') powerfully

subchelate, palm nearly vertical, notched or

sinuous, dactyl stout; gnathopod 2 (9), basis

broadened anteroproximally; segment 3 short,

segment 5 (not 4) shallow-tumescent posteriorly,

segment 6 shorter than carpus. Peraeopods 3-7

cuspidactylate, nails short. Peraeopods 3 and 4

unequal; in peraeopod 4, segment 5 often very

short (width nearly equal length), body of dactyl

usually strongly pinched or sharply constricted;

peraeopods 5-7 more or less similar in form,

increasing in length posteriorly, bases rounded

behind, peraeopod 7 (and often peraeopod 6)

sexually dimorphic in form, segments 4 and 5

incrassate in male; coxa 5 anterolobate; coxa 6,

hind lobe nearly vertical, antero-distal corner

right-angled, or with short distal process.

Abdominal sideplates 1-3, hind margin weakly

serrulate, hind corners acute; pleopods normal,

linear, outer margins of peduncles 2 and 3 weakly

spinulose, inner with 2 retinacula. Uropods short,

not heavily spinose; uropod 1, distolateral (inter-

ramal) spine not developed, outer ramus

marginally bare or nearly so, terminal spines

long; uropod 2, inner ramus spinose on inner

and outer margins. Uropod 3, peduncle moder-

ately broad, spinose posteriorly; ramus shorter

26

than peduncle, with short posterior and apical

spines. Telson short, lobes distally rounded, with

dorsal and apical spine groups. Coxal gills

reduced, especially on peraeopods 3-5, 2 and 6

longest. Brood plates (9), elongate-ovate,

smallest on 5, marginal setae long, simple-tipped.

Type-species: Orchestia platensis Kréyer 1845;

Bousfield, 1973, p. 160, plate 46(2)

Additional species!

Platorchestia crassicornis (Derzhavin) 1937

‘Bulycheva, 1957, p. 159, figs. 56a, b, (as

O. platensis)

:Morino, 1975, p. 127, figs. 4-5 (as O. platensis)

Platorchestia pachypus (Derzhavin) 1937, p. 91,

pl 2y sae? Soipku3; fie2

:Bulycheva, 1957, p. 146, fig. 52

:Morino, 1975, p. 179, figs. 4-6

(?) Platorchestia zachsi (Derzhavin) 1937, p. 90,

pho 2) fie 2.

:Gurjanova, 1951, p. 811, fig. 565

:Bulycheva, 1957, p. 146, fig. 53

(?) Platorchestia chathamensis new species (p. 27)

(2) Platorchestia japonica Tattersall 1922

:Morino, 1975, p. 175

Etymology

The generic name, Platorchestia, is a combining

form of Orchestia and the type-species name

platensis. Gender: Feminine.

Remarks

This genus differs from Orchestia (sens. str.)

mainly in the generally more spinose appendages;

powerfully incrassate antenna 2 (o’); 5-dentate

left mandibular lacinia; very short segment 5 of

peraeopod 4; right-angled and processiferous

hind lobe of coxa 6; marginally unarmed (or

weakly armed) outer ramus of uropod 1; more

heavily spinose pleopod peduncles, and smaller,

less marginally setose brood plates. Members of

the genus are restricted to the North Pacific

(especially western shores); however, P. platensis

(Krdyer) is nearly cosmopolitan along tropical-

temperate coastlines, but has not yet been

recorded from the eastern Pacific. Most members

of the group demonstrate some fossorial

behaviour and frequently occur on sandy beaches,

often as pioneer species, before more specialized

‘““Talorchestia’’ fritzi Stebbing 1903 from the Pacific coast of

Costa Rica and Galapagos (see Shoemaker, 1932, and

Monod, 1970) is closely allied here, but is tentatively retained

in the Caribbean group of Talorchestia Dana.

burrowing talitrids take over. The Platorchestia

group appears to be a north-temperate counter-

part of the tropical Indo-Pacific plesiomorphic

Talorchestia group, both of which burrow

shallowly, and are likely precursors of more

specialized substrate-modifying groups. Plator-

chestia is relatively apomorphic when grouped

with the beach fleas (allied to Orchestia), but

very plesiomorphic relative to the sandhoppers

(near Talorchestia) (Tables II, III, IV; figs. 26,

21; 28).

Key to known species of Platorchestia

1. Coxal plate 5, hind lobe with distinct anterodistal process (fig. 3j); uropod 3, ramus nearly equal

in length to peduncle, with 1-4 posterior marginal spines as well as apical spines; gnathopod 2

(2), basis strongly broadened anteriorly

Coxa 5, posterior lobe lacking distinct anterodistal process, lower margin meeting anterior

margin nearly directly at right angle; uropod 3, ramus distinctly shorter than peduncle, with

apical spines only; gnathopod 2 ( 2), basis moderately expanded anteriorly (fig. 20)

2. Eyes vertically subrectangular; gnathopod | (o’), dactyl nearly equal to total palm (including

blister); gnathopod 2 (o"), palm medially notched; peraeopod 7 (0), segment 5 moderately

expanded, width not greater than half length; telson apically notched

Eyes subovate; gnathopod 1 (0°), dactyl distinctly shorter than total palm; gnathopod 2(<),

palm sinuous; peraeopod 7 (o’), segment 5 very broadly expanded, width equal to or greater than

length; telson lobes separated in distal '/, to '/,

P. pachypus (Derzhavin)

3. | Gnathopod | (o’), carpus elongate, anterior margin about 1'/, times propod; gnathopod 2( <0)

palmar notch medial; uropod 1, outer ramus usually with | marginal spine

P. crassicornis (Derzhavin)

Gnathopod | (o'), carpus normal, anterior margin slightly longer than propod; gnathopod 2

(do), palmar notch nearer posterior angle; uropod 1, outer ramus lacking marginal

spine(s)

P. platensis (Kréyer)

4. _ Pleopods reduced, rami 3-4 segmented, distinctly shorter than peduncle; eye small,

rounded

eoeeeeeeeveeeeee eee eee ee ee ee we we ee eh Ohl Oh Oh Oh Oh Oh Hh OHO OH Hh Ow

P. chathamensis new species (p. 27)

Pleopods normal, rami multi-segmented, only slightly shorter than peduncle; eye normal,

subquadrate

5. | Gnathopod 1 (’) neotenically arrested at half transformed stage (fig. 21); antenna 2(<),

peduncle strongly incrassate; uropod 1, outer ramus with marginal spines; seashore

species

Ce |

P. zachsi (Derzhavin)

Gnathopod | (<') fully powerfully subchelate, palm convex (fig. 21); antenna 2 (o’), peduncle

only slightly broader than in female; uropod 1, outer ramus marginally bare; terrestrial

species

Platorchestia chathamensis new species

Figure 11

Material examined

Chatham Island, near Victoria, B.C. among logs,

at HW drift line, 4 July 1959, E.L. Bousfield and

G.C. Carl coll, 1 9 ov. Holotype NMC-C-

1981-509.

Diagnosis

Female (9.0 mm, ov.). A relatively small,

apomorphic, somewhat aberrant species, tenta-

tively assigned to this genus (in absence of male

specimens to complete diagnosis). Eye small,

4

P. japonica (Tattersall)

rounded. Antenna 1, flagellum 3'/,-segmented,

much shorter than peduncle. Antenna 2 very

short, appearing damaged or regenerating in

single female specimen available. Buccal mass

appearing slightly prognathous. Mandibular

left lacinia, proximal (Sth) tooth small. Maxilli-

ped palp relatively short, broad, segment 3

short, rounded distally, 4th segment minute but

not fused to 3rd. Gnathopod 1, carpus not

elongate; propod palm very short, oblique,

dactyl with single stout posterior seta; gnatho-

pod 2, basis relatively little expanded anteriorly,

broadest medioproximally; segments 4 and 5

shallow-tumescent behind. Peraeopod 4, dactyl

Figure 11. Platorchestia chathamensis new species. Chatham Island, Victoria, Vancouver Island, B.C.

2 ov. - 9.0 mm. P. platensis (Kréyer). Nova Scotia.

weakly constricted, nail long. Peraeopods 5-7,

slender, bases subsimilar, posterior margins

convex, weakly crenulate; dactyls relatively long,

slender; coxa 6, hind lobe without anterodistal

process, lower margin slightly sinuous. Abdomi1-

nal side plates 1-3, hind corners acuminate.

Pleopods slender, distinctly reduced, rami 3-4

segmented, much shorter than peduncles;

peduncles: 1, 2 and 3 with 0, 2 and 4 ‘outer

marginal short spines respectively. Uropods |

and 2 weakly spinose; uropod 1, outer ramus

marginally bare, inner ramus with 2-3 inner and

outer marginal short spines. Uropod 3 short,

peduncle with 3 stout posterior marginal spines;

ramus distinctly shorter than peduncle, sub-

conical, with apical spines only. Telson lobes

fused almost to apex, each with apical spines

and 2 groups of dorso-lateral spines. Coxal gills

relatively large, lobate (?). Brood plates 2-4

28

o - 12.0 mm gnathopods.

broadly sublinear, each with about 12 marginal

simple setae, brood plate of peraeopod 5 very

short, spade-shaped with 3-4 apical simple setae.

Remarks

The single female specimen is unlike all other

known 5-dentate ‘‘Orchestia”’ species in charac-

ters above and in the key, and is herewith

considered a distinctive new species. The overall

facies is reminiscent of Orchestia microphtalma

Amanieu & Salvat 1963, a log-burrowing species

from the Atlantic coast of France; these simi-

larities include the small rounded eye, short

antenna 2 and reduced pleopods. However, the

latter species is a true 4-dentate Orchestia, with

more strongly spinose appendages, including

marginally spinose outer ramus of uropod 1,

posteriorly spinose ramus of uropod 3, and

shorter peraeopod dactyls, among other differ-

ences. The single female of P. orientalis bears

a superficial resemblance to ecologically asso-

ciated females of Traskorchestia spp., but is

excluded from that genus by the more minute

maxilliped palp segment 4, the expanded basis

of gnathopod 2 (@), the short, narrow brood

plates with simple (vs. hook-tipped) marginal

setae, deep right-angled coxal lobe of peraeopod

6, the more slender elongate dactyls of the

peraeopods, and the weakly spinose uropods.

Confirmation of generic assignment awaits

discovery of the male of P. chathamensis.

Etymology

The species is known only from Chatham Island,

near Victoria, B.C., in the eastern North

Pacific region.

Megalorchestia Brandt 1851

Orchestoidea: Stebbing, 1906a, p. 527 (partim)

:Bulycheva, 1957, p. 130 (partim)

‘Bousfield, 1957, p. 120 (partim)

‘Barnard, 1969a, p. 471 (partim)

Diagnosis

Medium to large, smooth-bodied, heavily spinose

substrate-modifying talitrids of the eastern North

Pacific rim region, characterized by: head short,

deep, inferior antennal sinus wide, shallowly

incised, but deep to accommodate large segment |

of antenna 2 peduncle; buccal mass directly

ventral; eye medium to very large. Antenna |

shorter than peduncle 4 of antenna 2, flagellum

shorter than peduncle. Antenna 2 sexually

dimorphic, often elongate and/or peduncle

incrassate in male, peduncle 4 ( 9 ) not shortened.

Mandibular left lacinia cleanly 5-6 dentate, right

lacinia tricuspate; maxilliped palp 3-segmented,

‘tall’, segment 2 with pronounced spinose

mediodistal lobe, segment 3 subconical, narrow-

ing distally. Coxa 1 deep, anterodistal angle

subacute; coxae 2-4 subquadrate, variously

cuspate behind. Gnathopod 1 (co) powerfully

fossorial, spinose, carpus elongate (anterior

margin nearly equal to basis) with posterodistal

tumescence; propod elongate, narrowing distally,

lacking true palm, postero-distal blister very

shallow or lacking; gnathopod | (¢) segments 5

and 6 lacking blister, 6 without palm. Gnatho-

pod 2 (o') powerfully subchelate, palm of propod

variously oblique and toothed, dactyl often with

posterior marginal tooth near hinge; gnathopod 2

29

(2), basis more or less strongly expanded antero-

medially; segment 3 short; 4 with elongate

posterior process; 5 shallow-tumescent behind;

6 shorter than 5. Peraeopods 3-7 cuspidactylate,

dactyls slender; peraeopod 4, segment 5 and

dactyl shorter than in peraeopod 3, dactyl

constricted distally, overhanging base of unguis.

Peraeopods 5-7 tending to dissimilarity (in

southern species); peraeopod 6 slightly longer

than peraeopod 7, all strongly spinose, dactyls

moderately long. Coxa 5 anterolobate; coxa 6

posterolobate. Abdominal side plates nearly

smooth behind and below, hind corners weakly

acuminate, 2 deepest, lower margin convex.

Pleopods strongly reduced, 3rd shortest;

peduncular outer margin long-spinose, inner

margin with 2 retinacula. Uropods | and 2,

peduncles and both rami marginally strongly

spinose; uropod 3 medium, short spinose; ramus

longer than peduncle, laterally compressed, apex

rounded. Telson short, broad, lobes nearly

totally fused, with dorsal and apical short spines.

Coxal gills small, longest on peraeopod 6. Brood

plates subovate, lacking on peraeopods 2-4 in

some species, margins with simple setae, small on

peraeopod 5.

Type-species: Megalorchestia californiana

Brandt 1851 (original designation)

Additional species: Megalorchestia pugettensis

(Dana 1853)

Megalorchestia corniculata (Stout 1913)

Megalorchestia benedicti (Shoemaker 1930)

Megalorchestia minor (Bousfield 1957)

Megalorchestia columbiana (Bousfield 1958)

Megalorchestia dexterae new species

Remarks

The phyletic separation of heavily-bodied

4-dentate sandhoppers of the southern hemi-

sphere from heavy-bodied 5-dentate sandhoppers

of the northern hemisphere has necessitated

removal of the North American Pacific complex

from Orchestoidea Nicolet 1849 and resurrection

of the genus Megalorchestia Brandt 1851 to

accommodate them. Megalorchestia presently

contains three distinctive sub-groups: the

californiana group (monotypic); the columbiana

group (3 species); and the pugettensis group

(3 species), as diagnosed herewith (pp. 29-40, and

figs. 12-18, 27). Possible formal recognition of

these subgroups awaits study of more extensive

material from southern California, the Gulf of

California, and western Alaska.

Key to species of Megalorchestia

Eyes medium, vertically subquadrate, not bulging from head surface; antenna 2 short, flagellum

distinctly shorter than peduncle, peduncular segments 4 and 5 short, stout, strongly incrassate in

male; peraeopod 4, segment 5 very short, width nearly equal to length pugettensis group 2

Eyes large, subrotund, more or less bulging from sides of head in dorsal view; antenna 2,

flagellum equal to or longer than peduncle, elongate in males, peduncle moderately

incrassate in males; peraeopod 4, segment 5 width about //, to 7/, length

Uropod 2, outer ramus distinctly shorter than inner; peraeopod 5, segments 3 and 4 very

broad (3 broader than long), differing in form from those of peraeopod 6 and 7; gnatho-

pod 2(<'), palm with large distal tooth and deep depression near hinge ................005

eRe bikes eRe ste a eudie a age Pia’ nah eet will aifoheity aie ais so 4a RRO M. benedicti (Shoemaker) (p. 40)

Uropod 2, rami subequal; peraeopod 5, segments 3 and 4 not unusually broad, not markedly

different in form from those of peraeopod 6 and 7; gnathopod 2 (c’) with strong palmar tooth

near hinge

Coxa 5 very deep, anterior lobe as deep as coxa 4; coxa 6, hind lobe very deep, anterior margin

vertical; gnathopod 2 (o’), posterodistal angle of palm rounded

M. corniculata (Stout) (p. 39)

Coxa 5, anterior lobe distinctly shallower than coxa 4; coxa 6, hind lobe with oblique anterior

margin; gnathopod 2 (0), posterodistal angle of palm with paired short protuberances between

which tip of dactyl closes M. pugettensis (Dana) (p. 37)

Peraeopod 5, segments 3 and 4 very broad, differing in form from those of peraeopod 6 and 7

(fig. 3f); coxa 5 shallow, anterior lobe notas deep as coxa 4; uropod 1, outer ramus with only outer

marginal row of spines; gnathopod 2 (<0), palm with acute tooth near hinge, dactyl sinuous or

with posterior tooth near hinge columbiana group 5

Peraeopod 5, segments 3 and 4 normal, differing little in form from those of peraeopod 6 and 7;

coxa 5, anterior lobe about as deep as coxa 4; uropod 1, outer ramus with inner and outer

marginal row of spines; gnathopod 2 (o), palmar hinge tooth broad, blunt, dactyl regular ...

eae eee RRS. oarah, LeieGinslsle dee 6 vo owas Ri ORS Mapu Hal’ M. californiana Brandt (p. 30)

Peraeopod 5, dactyl very short and stout, nail (unguis) lacking; uropod 2, outer ramus with only

outer marginal spine row; gnathopod 2 (<'), palm very oblique, posterior margin of propod

distinctly less than half the anterior margin M. dexterae n. sp. (p. 35)

Peraeopod 5, dactyl short but not stout, nail developed; uropod 2, outer ramus with both inner

and outer marginal row of spines; gnathopod 2( 6’), palm normally oblique, posterior margin of

propod about half anterior margin

Peraeopods 6 and 7 slender, elongate, segment 6 longer than 5; gnathopod 2 ( ), posterior

tumescent process of segment 4 very large, deeper than segment; animals slender, small (to

15.0 mm) M. minor (Bousfield) (p. 33)

Peraeopods 6 and 7 stout, heavy, segment 6 not longer than 5; gnathopod 2( @ ), posterior process

of segment 4 short, not deeper than segment; animals heavy bodied, large (males to 22.0 mm)

eoeoeoeveveeeeeereeeeeeeeeeeeee eee ec ee we ewe we ewww eee Ho HP eH ee wo we ew ee he ew

eeoeeeeveereeeeeeeeeeeeeeeeeee eee ee ee ee eo

eeoeeeeeeveeeeeeeee eee eeeeeee eee ee ee ee ee wee eh Ohl hl Ohh Ohh Oh Oh Ohl hh Oh hh

eoeeeeeeeeee eee eee eee ee ee we wee ee hm Oh Oh Oh Oh Hh Oh Oh Hh Oh Ohh Oh Oh Oh PO HH Oh ee em hh how

Megalorchestia californiana Brandt 1851

Figure 12

Megalorchestia californiana Brandt 1851b, p. 311;

Brandt 185lc, p. 142, pl. 6

Orchestoidea californiana: Stebbing, 1906a, p. 528

*Boustield, 1957, p.. 122, pl, 24;.1958. p. 891,

fis 10); 1961, p.. 1b 1975. 0. 355., ite.229

‘Bowers, 1963, p. 316, figs. 1, 3a; 1975, p. 355

fig. 224

30

M. columbiana (Bousfield) (p. 32)

Material examined

British Columbia: E.L. Bousfield stations, 1959,

Vancouver Island, outer coast, Cape Scott south-

ward, 174 specimens from stns. O1, O2c, O3, O5,

O7c, O12, O16, O18; inner coast, north end, 55

specimens from V1, V2; south end 22 specimens

from Chatham Island; ibid. 1964, Vancouver

Island, near Bamfield, 42+ specimens from H44,

H45; ibid. 1970, P707 (Pachena Bay), Vancouver

Island, 40 specimens; ibid. 1975, P14b, (Keeha

Figure 12. Megalorchestia californiana Brandt. Brady’s Beach, Vancouver Island, B.C. o& - 23.0 mm,

9 ov. - 20.0 mm.

Bay), 84 specimens; ibid. 1976, B6 (Brady’s

Beach), 3 oo, 39 Q (fig’d specimens); ibid.

1977, 13 specimens from BSe (Witty’s Lagoon),

Bllc (Wreck Bay).

Washington State, USA: Bousfield 1957 stn.

M14 (Fidalgo Island), 60+ specimens; ibid. 1966,

120 specimens from W14, W16, W17, W20, W34,

W39, W41, W45, W46, W47.

Oregon State (outer coast) ibid. 1966: 68

specimens from W50, W53, W55, W56, W59,

W61, W63, W66, National Museum of Natural

Sciences collections.

Miscellaneous collections: British Columbia,

Vancouver Island, Young and Spreadborough

collections, 1909: (1) Wreck Bay, 11 June,7 oo’,

3 29 ov. (2 lots), NMNS No. 1047; Ucluelet

31

(Long Beach), July,9 i ov, 1 9, 8imm. NMNS

No. SITS: “Long. Bay’ (Uciuelet), ot. lio. 7.

20 July 1931, C.G. Carl coll., 10 care © «.

NMC-C-1981-570. JFL Carl collections, Van-

couver Island 1934, stns. 2231-14S, May; 2241-12,

28 June, 10 imm.; Cadborough Bay, Victoria,

8 June 1956, 10,8 2 9 (3 ov.) NMNS No. 5116;

Sydney Spit, coarse sand at HW, 26 August 1956,

2 oo subad. NMNS No. 10348.

California: Santa Barbara, sand beach, 25 Octo-

ber 1965, W.L. Klawe coll., 4 o& subad., 1 2 br. I,

12 imm., NMNS No. 13974; Santa Barbara,

August, 1969, A.Wenner coll., 8 ooh, 5 9 9,

NMNS No. 13975. Re-examined all materials

from British Columbia, California, and Bahia

San Quintin, Mexico, previously recorded by the

author (1957, 1961, 1964), NMNS collections.

Remarks

M. californiana is close to the apomorphic

columbiana group in overall form of body and

appendages, but differs in the larger body size,

less shortened urosome, undifferentiated peraeo-

pod 5, deeper coxal plates (especially of peraeo-

pod 5), more spinose abdominal side plates, and

more heavily long-spinose uropods 1 and 2. Also,

the eyes are not as large, relative to head size,

the maxilliped palp segment 2 is broader, gnatho-

pod 1 (@) carpus broadens distally, dactyl of

gnathopod 2 (co) is untoothed behind; the

peraeopod dactyls are relatively short and stout,

the gills less markedly reduced, and the mature

female has fully developed brood plates on

peraeopods 2 to 5. The dorsal abdominal mark-

ings are less striking than in M. columbiana (see

Bowers, 1975), but the brilliant orange-red colour

of antenna 2 of mature males is diagnostic.

Distributional ecology

Occurs only on smooth sand beaches of open

coasts exposed to surf action and cool, high

salinity waters; it ranges from the north end

of Vancouver Island, B.C., southward to Point

Conception, California, and sporadically south

to northern Baja California. The species is

univoltine, breeding once per year; females

ovigerous in spring and early summer. Like most

semi-terrestrial talitrids of Pacific rim shores,

M. californiana is frequently parasitized on gills

and sternum by mites (Thinoseius sp.).

Columbiana group

This highly apomorphic sandhopper group is

characterized by relatively slender bodies and

very short urosome; large, bulging eyes; elongate

antenna 2; shallow coxal plates; slender peraeopod

dactyls; markedly reduced and different form of

peraeopod 5; unequal rami of uropod 2; and

reduced gills and brood plates. All occur on fine

to coarse sand beaches but often on protected,

steeper shores, subject to brackish or even

freshwater influence.

Megalorchestia columbiana (Bousfield 1958)

Figure 13

Orchestoidea columbiana Bousfield 1958, p. 890

fig. 4; 1961, p. 9, fig. 4f-h; 1975, p. 355, fig. 233

-Bowens;. 1975. Dp, 333, 112. 220, 1903, .p. 317,

fig. 3d

32

Material examined

Alaska: 260 specimens from Bousfield &

McAllister 1961 stns. A20, A22, A58, A60, Aél,

A65, A66, A70, A71, A73, A87, A155 (Day

Harbour, near Seward, to Dixon entrance).

British Columbia: Queen Charlotte Islands,

4 specimens from Bousfield 1957 stns. H13, E1;

north mainland coast; 90 specimens from E.L.

Bousfield 1964 stns. H35, H38, H40, H44, H45,

H48, H60; E-L. Bousfield 1959 stn. N3, 200+

specimens; Vancouver Island, north end: 19 spe-

cimens from E.L. Bousfield 1959 stns. O2a, O2c,

V1; Vancouver Island, south end; 36+ specimens

from E.L. Bousfield 1970 stns. P713 (Clo-oose

Beach) and P720 (Port Renfrew).

Washington State: 70 specimens from E.L.

Bousfield 1966 stns. W16, W20, W21, W24, W26,

W38, W39, W45, W46.

Oregon State: 45 specimens from E.L. Bousfield

1966 stns. W50, W52, W56, W59, W63, W64.

Re-examined all material from British Columbia

and California including type material from

Wickininnish Bay, and figured material from

Carmel Beach, Monterey Co., California,

previously published by the author (1958, 1961).

NMNS collections, Ottawa.

Remarks

Megalorchestia columbiana is the most plesio-

morphic member of the group in having a

relatively large, stocky body; short, heavy

peraeopods; and (with the closely related

M. minor) a more vertical palm of gnathopod 2

(o); normal dactyl of peraeopod 5; and both

margins of the outer ramus of uropod 2 spinose,

On surf-exposed fine sand beaches the animal

occasionally co-occurs with M. californiana

from which it is distinguished by the more

conspicuous dorsal abdominal markings (at all

stages) and shorter urosome, and by the bluish-

white antennae of the mature male (Bowers,

1963). In northern and central regions, especi-

ally on protected coarse sand beaches, it co-occurs

frequently with M. pugettensis; the dorsal colour

patterns are often difficult to distinguish, but

M. columbiana is usually less darkly pigmented;

moreover, the body of M. columbiana is broader

anteriorly, and the eyes appear to bulge from the

head margin in dorsal view. M. columbiana is

superficially similar to Trinorchestia trinitatis

Derzhavin in the large eyes, elongate antenna

(co), and form of gnathopod 2 (0), but is readily

distinguished by the elongate narrowing propod

Figure 13. Megalorchestia columbiana (Bousfield). Carmel Point, California.

of gnathopod 1 (both sexes), the specialized

constriction overhang of the dactyl of peraeopod

4, and much more reduced and modified pleo-

pods, and the specialized form of peraeopod 5S.

Distributional ecology

Occurs at the level of drift debris and slightly

below, mainly on protected, steeper-sloping, fine-

to-coarse sand beaches of estuaries and embay-

ments, often in brackish areas, from southern

Alaska (northern and western limit not deter-

mined) to Carmel Point, Monterey Bay, central

California. The species is univoltine, breeding

33

o& - 22mm, 2 - 16.0mm.

once per year, females ovigerous in the spring and

early summer. Animals occur on the same beaches

as, but slightly seaward of, M. californiana,

burrow not quite as deeply, and tend to be more

diurnal (especially the juveniles and immatures).

Megalorchestia minor (Bousfield 1957)

Figure 14

Orchestoidea minor Bousfield 1957, p. 126, fig. 25

‘Straughan, 1981, p. 375

Orchestoidea columbiana: Straughan, 1981, p. 375

Figure 14. Megalorchestia minor (Bousfield). Pt. Dume, California.

Material examined

California: D.E. Bowers collections, April 1960

from: San Simeon, San Luis Obispo Co.,9 oo

(to 14.0 mm), 9 9 9,5imm., NMNS No. 5430;

Santa Barbara, Santa Barbara Co., 16 oo,

16 92 9, br. P5 only, NMNS No. 5431; west side,

Pt. Magu, Ventura Co.,3 ooh, 2 292, NMNS

No. 5428; between Pt. Magu and Sycamore

beach, 11 oho, 28 9 9, NMNS No. 5435; just

east Seguit Pt., Los Angeles Co., 7 9 9, NMNS

No. 5427; Dume Cove, Los Angeles Co.,7 2 9,

NMNS No. 5433; just west Pt. Dume, Los

Angeles Co., 14 oi oh, 17 2 9, NMNS No. 5432;

34

o - 14.0 mm, Q ov. - 15.0 mm.

Municipal Beach, Los Angeles Co., 4 oo,

14 9 9, NMNS No. 5434. Santa Barbara, Santa

Barbara Co., sand beach at HW Level, 25 Octo-

ber. 1965, W:L. _Klawe coll.,..3° o o,..subage

4 99 br. P5 (to 14.5 mm), 2 imm. NMC-C-

1981-503, slide mounts; ibid. August 1969,

A. Wenner coll., 3 9 9. br. Pl, PS only Ge

11.0 mm), NMC-C-1981-505.

Mexico: Bahia San Quintin, HW stn. SQ104,

5 August 1960, J.L. Barnard coll., 1 (peraeopod

6 only)?. Re-examination of all specimens from

California (Playa del Ray, Pt. Dume) and

Mexico (Ensenada, B.C.), including type slide

mounts (basis of fig. 14), reported in previous

author publications (1957, 1961).

Distributional ecology

Occurs on surf-exposed flat sand beaches,

burrowing at or near the HW drift debris, from

just north of Pt. Conception (San Luis Obispo

Co.) to northern Baja California (Ensenada and

Bahia San Quintin). The species is probably

univoltine; ovigerous females have setose brood

plate only on peraeopod 5.

Remarks |

The species has been seldom collected and little

studied since the original description based

on material from Los Angeles county, California

(Bousfield, 1957). The species is small and

slender-bodied (mature males to about 15.0 mm),

and is distinguished from the closely related

M. columbiana by the relatively shorter antenna 1;

more slender and less heavily spinose peraeopods

and uropods; more markedly modified peraeo-

pod 5 (nail of dactyl shorter); less strongly

developed hinge tooth of palm and dactyl of

gnathopod 2 (o'); more elongate posterior

process of segmient 4 of gnathopod 2 (9); and

the more spinose armature of the outer ramus of

uropod 2 which (in M. minor) extends further

apically along the inner margin than along the

outer margin. The species occurs mainly on

beaches that are exposed to relatively heavy

contamination from offshore oil wells and urban

pollution in the southern California region (e.g.

Straughan, 1981) and might be considered an

endangered crustacean species.

Megalorchestia dexterae new species

Figure 15

Material examined

San Juanico, Baja California, sand beach at HW

level, 21 July 1975, D.M. Dexter, coll. Male

holotype (17.0 mm), female allotype (br. L.,

13.5 mm), male subadult paratype (11 mm).

NMC-C-1981-506, slide mounts.

Diagnosis

Male (17.0 mm). A medium-sized, slender-bodied,

finely spinose species of the columbiana group

characterized by: eyes very large, nearly conti-

guous dorsally, and posteriorly nearly reaching

peraeon |; antenna 1, flagellum 5-6 segmented;

35

antenna 2, peduncle long, segment 4 long;

flagellum about 25-segmented, subterminal

segments weakly toothed. Maxilliped palp rela-

tively short, segment 3 short. Gnathopod 1,

propod long, slender, arcuate, nearly equal in

length to carpus, without posterodistal tumes-

cence; carpus very elongate, with small postero-

distal blister. Gnathopod 2, propod large, ovate,

palm very oblique, stoutly spinose, especially at

posterior angle, with large rounded tooth near

hinge; posterior margin short, about one-quarter

anterior margin; dactyl long, sinuous. Peraeopods

3-7, dactyls relatively slender, dactyl of 4 not

strongly pinched, barely overhanging base of

nail; peraeopod 5, anterior marginal peg-spines

of segment 6 short; dactyl very short, thick, not

‘fuzzy’ behind, apex blunt, nail lacking;

peraeopods 6 and 7 slender, elongate, bases

subsimilar in shape, segment 6 about equal to 5.

Abdominal side plates very weakly setulose

behind, smooth below. Pleopods 1-3, rami very

short, 1-2 segmented, scarcely half length of

respective peduncles. Uropods | and 2, peduncles

and rami slender, armed with numerous relatively

long slender simple spines; outer ramus of

uropod 2 distinctly shorter than inner ramus,

with outer marginal spines only. Uropod 3 rela-

tively long, not markedly sexually dimorphic,

ramus about equal in length to peduncle. Telson

short, small, apex subacute, entire, dorsal

spines slender.

Female (13.5 mm., br. I). Antenna 2, flagellum

as long as peduncle, about 25-segmented,

subterminal segments toothed behind. Gnatho-

pod 2, basis broadest medioproximally; hind

lobe of segment 4 short, not deeper than

segment, segment 5 hind lobe with small distal

process. Brood plates not developed in single

specimen available.

Etymology

The species is named in honour of Dr. Deborah

M. Dexter, the collector, who has specialized in

the world-wide study of sand-beach crustaceans.

Distributional ecology

Known only from the type locality which is the

most southerly record of the genus (sens. str.)

on the North American Pacific coast.

Remarks

Megalorchestia dexterae bears a superficial

resemblance to some members of Pseudorches-

Figure 15. Megalorchestia dexterae new species. San Juanico, Baja California, Mexico.

0. pr. 1 -= 133 mm:

toidea, especially P. gracilis Bousfield & Klawe,

with which it may abut distributionally in

Baja California. These similarities include the

relatively slender body, large eyes, elongate

antenna 1, relatively shallow coxal plates, form

of gnathopod 2 (co), distinctive peraeopod 5

(especially dactyl), and slender uropods. The

generic differences, however, include in Megalor-

chestia a less elongate body, slightly incrassate

peduncular segments of antenna 2 (c), taller

maxilliped palp, more elongate and more power-

36

3o - 17.0 mm,

fully fossorial gnathopod 1, smaller coxal gills,

and differing form of the pleopods, uropod 3

and telson.

Pugettensis group

This relatively plesiomorphic complex is charac-

terized by a short, broad, stocky body; relatively

small eyes; short antennae with stoutly incras-

sate peduncular segments (0); deep coxal plates,

especially the lobes of peraeopods 5 and 6;

Figure 16. Megalorchestia pugettensis (Dana). Brady’s Beach, Vancouver Island, B.C. o& - 17.0 mm,

we- 13.5 mm.

strongly toothed, normally oblique palmar

margins of gnathopod 2 (<’), dactyl not toothed

behind; relatively short, stocky, short-dactylate

peraeopods; subsimilar form of peraeopods 5-7;

segment 4 very short in peraeopod 4; and short

uropods and telson. The two larger species occur

more frequently on coarse sand and fine gravel,

often on protected shores, whereas the smaller

M. benedicti is an open surf, fine sand beach

dweller.

Megalorchestia pugettensis (Dana 1853-55)

Figure 16

Orchestia (Talitrus) pugettensis Dana 1853 and

e595 p. 859, t. 57, fig. 3a-d

Orchestoidea pugettensis:Stebbing, 1906a, p. 528

:Thorsteinson, 1941, pl. 1, figs. 1-9

a7

:Bousfield, 1958, p. 890, fig. 101; 1961, p. 7,

fig. 3> 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, <ibid., .26.June,sll dicks (to 26.0) mm);

ibid., 18 August, A. Wenner coll.,3 ood, 4 2 &,

3 imm.; Corona del Mar, 28 May 1953, J.L.

Barnard coll., 5 specimens, NMNS Cat. No.

5128; Pt. Loma, San Diego, February 1956,

W.L. Klawe coll., 5 imm. NMC-C-1981-466.

Re-examination of all central Californian material

listed in Bousfield (1961, p. 7), including stn. C8,

Shell Beach, 10 July 1959, 37 specimens NMNS

Cat. No. 5819, and slide mounts.

Mexico: El Socorro Beach near El Rosario,

Baja California, fine sand and cobbles, under

wave cast Phyllospadix at HHW level, June 1980,

R.G. Appy coll., 10 & o& (to 19.0 mm), 22 9 9

(mostly ov.), 10 imm. NMC-C-1981-463.

Remarks

Megalorchestia corniculata is the largest and most

spinose species of the pugettensis group (mature

males to 25.0 mm), but is otherwise very similar

to M. pugettensis. The species is distinguished by

relatively large eyes, especially in southern popu-

lations; weakly tumescent gnathopod 1 (0’); non-

protruding paired spinous ridges at the posterior

palmar angle of gnathopod 2 (0); very broadly

expanded basis of gnathopod 2 and small brood

plate (2); very deep anterior and posterior lobes

of coxae 5 and 6 respectively; relatively short

and powerfully spinose uropods; more broadly

expanded pleopod peduncles; and more dorsally

spinose telson. Dorsally, the pigmentation pattern

is similar to that of M. pugettensis, but laterally,

M. corniculata lacks a dark horizontal bar on

peraeon and/or coxa 7. Antenna 2 (mature male)

is usually salmon-pink, rarely bluish or brownish,

as in M. pugettensis.

Distributional ecology

Occurs mainly on surf-exposed, high salinity,

cold-temperate, coarse sand and fine gravel

beaches, beneath cliffs of the California coast,

from north central region (Mission Head) to

northern Baja California (El Socorro Beach).

Animals burrow at HW level in the zone of

rotting kelp, above freshly deposited kelp,

females with eggs seaward of those with young,

but above most juveniles. The burrows of adults

(especially females with eggs and especially in

the winter) may reach depths of 50-60 cm

(maximum 76 cm). Adults are exclusively

nocturnal but juveniles emerge earlier in the

evening and remain at the surface later in the

morning, after sunrise (Craig, 1973b). The species

tends to orient landwards under a wide range of

stimuli by day; lunar position is the dominant

night-time orienting stimulus (Enright, 1961).

The species occasionally co-occurs with M. puget-

tensis in the northern part of its range, and with

Paciforchestia klawei and Traskorchestia traskiana

in the south, but rarely with other sandhoppers

(e.g. Straughan, 1981).

40

Megalorchestia benedicti (Shoemaker 1930)

Figure 18

Orchestoidea benedicti Shoemaker 1930, p. 112,

fig. 3

:‘Bousfield, 1961, p. 9; 1975, p. 355, 363; 1981,

p. 86, fig. 17

:Bowers, 1963, p. 318, fig. 3c; 1975, posaaee

fig. 227

:‘Straughan, 1981, pp. 375, 423

Material examined

California: Piedras Blancas, San Luis Obispo

Co., sand beach at HW level, 30 April 1947,

J.L. Barnard coll., 1 2, br. pl. PS only (8.0 mm);

Santa Barbara, Santa Barbara Co., sand beach,

intertidal, 25 October 1965, W.L. Klawe coll.,

2o00,1 @ (br. pl. P5 only), NMC-C-198 1-530;

Santa Barbara, sand beach, HW level, 18 August

I969, A. Wenner coll. 5 oo (to 10.0 mm

NMC-C-1981-529. All specimens from slide

mounts, previously published upon from Cali-

fornia (1957, 1961) and Mexico (Ensenada, 1957;

Bahia San Quintin, 1964) were re-examined;

fig. 18 is based on material from Morro Beach,

San Luis Obispo Co.

Remarks

This is the smallest and most apomorphic species

of the pugettensis complex. It differs from the

other species not only in smaller size and more

strongly contrasting checkerboard pigmentation

pattern, but also in the more elongate propod and

the deeper proximal palmar incision of gnatho-

pod 2 (o); the relatively narrow basis of gnatho-

pod 2 (9); the somewhat unlike form of peraeo-

pod 5 (with respect to peraeopods 6 and 7); the

more elongate and slender peraeopods 6 and 7,

especially dactyls; the more expanded pleopod

bases and shorter rami; the unequal rami of

uropod 2; and the shorter, broader telson. In

these features, M. benedicti links the pugettensis

group with the columbiana group.

Distributional ecology

Occurs mainly on surf-exposed, fine sand, berm

beaches from northern California (Eureka) to

central Baja California. The species burrows

shallowly under drift debris at HW level, just

below the main population of M. californiana

with which it frequently co-occurs.

Figure 18. Megalorchestia benedicti (Shoem.). Morro Beach, California.

Trinorchestia new genus

Orchestoidea: Bulycheva, 1957, p. 130 (partim)

Diagnosis

Medium-sized, slender-legged, spinose beach-

hoppers characterized by: body smooth, broad,

narrowing abdominally; eyes very large (especi-

ally in o&), nearly contiguous dorsally. Inferior

antennal sinus medium deep, short. Antenna |

short, peduncle 2 longest, flagellum shorter than

peduncle. Antenna 2 elongate, peduncular seg-

ments moderately incrassate (o'). Buccal mass

4]

o -9.5mm, 9 ov. - 8.0 mm.

relatively shallow, directly ventral to head;

mandible left lacinia cleanly 5-dentate, right

lacinia tricuspate; maxilliped palp tall, segment 3

subconical, 4 fused to 3. Coxal plate 1 deep,

anterodistal angle acute; coxae 2-4 subquadrate,

cuspate posteriorly. Gnathopod 1,- carpus elon-

gate, powerful, propod much shorter, broadest

medially, spinose, lacking palm; dactyl strong;

carpus and propod with posterior tumescent lobe

in male only. Gnathopod 2 (co) powerfully sub-

chelate; palm with notch and tooth near hinge,

dactyl strong, with posterior tooth near hinge;

segment 4 with posterior tumescent process.

Figure 19. Trinorchestia trinitatis (Derzh.) Seto, Japan.

Gnathopod 2 (¢) basis moderately expanded

anteriorly; segment 4 with posterior tumescent

process. Peraeopods 3-7 slender, cuspidactylate;

peraeopod 4 slightly smaller than 5, segment 4

smaller, dactyl pinched, body not overhanging

base of nail; coxa 5 aequilobate, coxa 6 postero-

lobate, anterior margin of hind lobe convex,

oblique; peraeopods 5-7 homopodous in form,

basis and segment 4 similar, peraeopods 6 and 7

much longer than 5, 6 longest. Abdominal side

plates smooth and convex below. Pleopods sub-

equal, nearly normal, rami little reduced, outer

ramus the shorter; peduncles sublinear, outer

42

So - 18.0 mm, 9 br. II. - 18.0 mm.

margins short-spinose. Uropods 1 and 2, all rami

with inner and outer marginal spines, those of

uropod 2 subequal in length, terminal spines

simple. Uropod 3, ramus longer than peduncle,

short-spinose, laterally compressed. Telson short,

lobes fused distally, with dorsal and apical spines.

Coxal gills 2-5 short, sinuous, 6 elongate. Brood

plates undescribed.

Remarks

This monotypic genus has no single distinctive

characteristic and appears not closely related to

other sandhopper genera. However, shared

synapomorphies appear to relate it to Pseudo-

orchestoidea, Talorchestia, and possibly Megal-

orchestia on the one hand (figs. 27, 28) and with

the western Pacific ‘*Talorchestia’’ sinensis-

nipponensis group on the other (see Morino,

1972). The possibility that more than one species

of Trinorchestia occurs in the western Pacific

region merits careful study.

Type-species: Orchestoidea trinitatis Derzhavin

1937 (monotypic)

Etymology

The generic name is a combining form of

Orchestia and the type species name frinitatis.

Gender: Feminine.

Trinorchestia trinitatis (Derzhavin 1937)

Figure 19

Orchestoidea trinitatis Derzhavin 1937, p. 88, fig. 1

:Vader, 1970, p. 134, fig. 49

:Morino, 1972, p. 57, figs. 9-11

:Bousfield, 1981, fig. 17

Orchestoidea brito: Bulycheva, 1957, p. 134, fig. 49

Talorchestia brito: Iwasa, 1939, p. 273, figs. 13-15

:Stephensen, 1944, p. 65, figs. 22-23

:Gurjanova, 1951, p. 809, fig. 563

Material examined

Japan: Seto, Honsu, sand beach, 1957, A. Yamazi

coll., 1 o& (18.0 mm), NMC-C-1981-536, slide

mount; Ishikari, Hokkaido, 21 August 1968,

H. Morino coll., 1 & (18.0 mm), 1 ¢& (transform-

ing stage), 2 2 9 (subadult, 12.5 mm), NMC-C-

1981-537, slide mounts.

Distributional ecology

Occurs mainly on surf-exposed cold-temperate

fine sand beaches of the open western Pacific

coast from Commander Islands, Kamchatka

Peninsula (including Bering and Myedny Islands)

in the north, through the Kuriles, Sakhalin and

Hokkaido to southern Kyushu (Japan). Life

cycle is presumed univoltine, females probably

Ovigerous in late spring and summer. Animals

are essentially nocturnal, occasionally diurnal

according to availability of food (caprellids,

large flies) (Morino, 1972).

Remarks

The animals are somewhat variable morphologi-

cally by region as well as sex and instar.

Stephensen (1944) first drew attention to the

43

presumed cuspidactylate condition of the

peraeopods.

Orchestoidea Nicolet 1849

Orchestoidea Nicolet 1849, p. 229

:‘Stebbing, 1906a, p. 527 (partim)

‘Barnard, 1969a, p. 471 (partim)

Diagnosis

Large, stout bodied, sculptured, strongly fossorial

sandhoppers of the Pacific coast of South

America, characterized by: head deflexed slightly

near frons. Inferior antennal sinus shallowly

incised, but deep to accommodate large pedun-

cular segment 1. Eyes large, bulging, not conti-

guous dorsally. Antenna | short, flagellum short,

peduncular segments | and 2 longest; antenna 2

medium, elongate, not incrassate in o’. Buccal

mass relatively shallow, directly beneath head;

mandibular left lacinia cleanly 4-dentate, right

lacinia tricuspate (?); maxilliped palp 3-segmented,

‘tall’, segment 2 with spinose medio-distal lobe,

segment 3 elongate, subconical, segment 4

totally lacking. Coxal plate 1 deep, anterodistal

angle subacute; coxal plates 2-4 deep, hind

marginal cusps vestigial or lacking. Gnathopod 1

powerful, spinose; segment 5 elongate; segment 6

elongate, narrowing distally, lacking palm; dactyl

strong; segments 4 and 5 devoid of tumescent

process (oh and 9). Gnathopod 2 (<') powerfully

subchelate, palm incised near hinge; dactyl strong,

with posterior marginal swelling near hinge;

gnathopod 2 (¢), basis long, little expanded,

anterior margin sinuous; segment 3 short; 4

spinose behind, lacking posterior tumescent

process; 5 shallow posteriorly; 6 nearly equal to

5. Peraeopods 3-7 stout, spinose, cuspidactylate,

dactyls slender; peraeopod 4 shorter than 3,

segment 5 distinctly shorter; dactyl pinched,

body not overhanging nail. Coxa 5 aequilobate,

6 very deeply posterolobate, anterior margin

vertical, rounding; peraeopods 5-7 weakly

heteropodous, basis and segment 4 sub-similar

in form, 6 and 7 much longer than 5, 6 longest.

Abdominal side plates 1-3 deep, convex and

smooth below, hind margins spinose; pleopods

reduced, decreasing posteriorly, rami shorter

than peduncle; peduncles broadly sublinear,

outer margin strongly long-spinose, overhanging

outer ramus, inner margin with 2 retinacula.

Uropods 1 and 2 stout; peduncles and rami,

inner and outer margins strongly spinose,

terminal spines simple; rami of uropod 2 sub-

Wao

Figure 20. Orchestoidea tuberculata Nicolet. Chachagua, Chile.

equal in length. Uropod 3 very short, peduncle

not expanded, short-spinose; ramus compressed,

rounding apically, short spinose. Telson short,

broad, lobes fused distally to form a single

plate, with marginal and apical spines only.

Coxal gills not greatly reduced, shallow-

acetabulate, 2 and 6 longest. Brood plates

subovate, subequal on 2-5, marginal setae simple.

Type-species: Orchestoidea tuberculata Nicolet

1949 (monotypic)

Remarks

The diagnosis (above) is based on the type-

species which has never been fully described. The

44

& - 21.0 mm, Q br. II. - 17.0 mm.

genus belongs to a southern hemisphere group

of sandhoppers in which the mandibular left

lacinia is primarily 4-dentate, the frons is slightly

deflexed; antenna 2 (@), especially peduncular

segment 4, very short; coxal plates deep; hind

lobe of coxa 6 very deep, anterior margin

vertical; gnathopod 2 (¢), basis narrow, seg-

ment 4 rarely tumescent behind; brood plates

broadly subovate, 5th subequal, little reduced;

coxal gills 3-5 little reduced; pleopod peduncles

spinose, broadly sublinear; uropod 3 small; and

telson lobes rarely with dorsal spines. Except

for ‘“‘Orchestoidea”’ brasiliensis (Dana), all other

southern continental sandhoppers have a more or

less subchelate gnathopod | (co), and antenna 2

and peraeopods 6-7 tend to be variously sexually

dimorphic, and the bodies are smooth and eyes

smaller. ‘‘Orchestoidea”’ brasiliensis is placed in

a monotypic genus by Bousfield (1982 in prep.)

because of its distinctive peraeopod 5 with

reduced dactyl, and differing structure of gills,

pleopods, and uropods.

Orchestoidea tuberculata Nicolet 1849

Figure 20

Orchestoidea tuberculata Nicolet 1849, p. 229,

2a1, t. 2, fig. 4

:‘Stebbing, 1906a, p. 527

Material examined

Chile: Cachagua, ocean beach at HW level,

13 November 1973, D.E. Dexter coll., 22 dod

(to 21.0 mm), 45 2 Q (1 ov. several br. I and II)

plus 5 imm., NMC-C-1981-519, slide mounts;

Corral (near Valdivia), ocean beach, Lamb &

Holz coll. (no data), 1 & (19.0 mm), 2 9 9 ov.

(16.0 mm); Playa Grande, Mehuin, Province

Valdivia, June 1976, C. Varela coll., several

imm. oo and 9 Q.

Peru: Mejia, near Mollendo (approx. 15°S, 72°W),

28 July 1957, H. Heller coll., several imm. 0d ov

and 2 9, NMC-C-1981-518.

Remarks

The species bears a superficial resemblance to

large, heavy-bodied sandhoppers of the North

American Pacific coast. However, the numerous

generically distinctive features (noted above and

in the key) necessitate the removal of the boreal

North American sandhopper complex from the

genus Orchestoidea and resurrection of Megalor-

chestia Brandt 1851, to encompass them.

Distributional ecology

Occurs under drift debris at the HW level of

surf-exposed, high salinity, cool temperate

beaches of the Pacific coast of South America

from south of Valparaiso to southern Peru (Mejia

near Mollendo). No animals were taken on the

beaches of the Cape Horn region during the

Hudson 70 expedition (unpublished informa-

tion); in that region, the dominant sand-beach

amphipod is “‘Orchestia” scutigerula Dana.

45

Talitrus Latreille 1802

Talitrus Latreille 1802, vol. 1, p. 78, vol. 2, p. 148

‘Stebbing, 1906, p. 524 (partim)

‘Hurley, 1955, p. 145 (partim)

‘Bulycheva, 1957, p. 127 (partim)

‘Barnard, 1969a, p. 472 (partim)

Talitrus (Talitrus) Hurley 1975, p. 161

Diagnosis

Medium sized, heavy, smooth-bodied sand-

hoppers characterized by: head and buccal mass

broadened below. Interior antennal sinus

shallowly incised, deep to accommodate peduncle

segment 1. Eyes medium, not dorsally contiguous.

Antenna | short, peduncle 2 longest; flagellum

shorter than peduncle. Antenna 2 elongate,

peduncle longer in male but not incrassate.

Mandibular left lacinia 5-dentate, proximal (5th)

tooth small, right lacinia tricuspate. Maxilliped

palp “‘tall’’, appearing 3-segmented, segment 2

broad, with spinose mediodistal lobe, segment 3

subconical, 4th (terminal) segment minute,

masked by spines of segment 3. Coxa 1 deep,

anterodistally acute. Coxae 2-4 subquadrate,

cuspate behind. Gnathopod | simple, powerfully

fossorial, larger in male; carpus elongate and

heavily spinose; propod elongate, arcuate,

narrowing distally, lacking palm; dactyl strong;

propod and carpus lacking posterior tumescent

process (oh, @). Gnathopod 2, propod mitten-

shaped in both sexes; basis sublinear, somewhat

more expanded antero-medially in female;

segment 4 lacking posterior tumescent process;

segment 5 tumescent postero-proximally;

segment 6 shorter than 5. Peraeopods 3-7

cuspidactylate, segments stout, spinose, not

elongate; dactyls medium. Peraeopod 4 shorter

than 3, segment 5 distinctly shorter; dactyl short,

prominently pinched, but not overhanging unguis.

Coxa 5 aequilobate, lobes deep; coxa 6, hind

lobe very deep, anterior margin vertical, rounding

below. Peraeopods 5-7 weakly heteropodous in

form, basis and segment 4 of short peraeopod 5

differing slightly from those of 6 and 7; peraeo-

pod 6 longest. Abdominal side plates 1-3 deep,

smooth below, weakly spinulose posteriorly;

pleopods about normal, rami little reduced,

multisegmented; peduncle sublinear, not expanded,

outer margin short-spinose, inner with 2 retin-

acula. Uropods | and 2 stout, inner and outer

margins of peduncles and rami strongly spinose,

terminal spines simple; uropod 2, outer ramus

shorter than inner. Uropod 3 short, peduncle not

broadened but spinose posterodistally; ramus

cylindrical, tapering, short spinose behind,

apical spine long. Telson short, broad; lobes

nearly completely fused distally, with apical

and marginal spines only. Coxal gills relatively

large, somewhat sac-like, little reduced on

peraeopods 3-5. Brood plates large, subovate,

little reduced on peraeopod 5, marginal setae

simple.

Type-species: Talitrus saltator Montagu 1808.

(monotypic)

Remarks

The genus Talitrus is a true sandhopper, most

closely allied with Orchestoidea Nicolet, and

46

other 4-dentate genera; it is remote morpholo-

gically, as well as ecologically and behaviourally

from the true landhoppers (Table IV, fig. 28).

Despite the neotenic loss of amplexing type

gnathopod 2 in the male (an advanced character

that is convergently and superficially similar to

that of many landhopper genera), Talitrus is

plesiomorphic in several other taxonomic fea-

tures, especially relative to those of large, stout-

bodied sand hopper genera of North American

coasts. (see Table III and fig. 27).

Talitrus saltator (Montagu 1808)

Figure 21

Talitrus saltator: Sars, 1895, p. 23, pl. 9

:Chevreux & Fage, 1925, p. 271, fig. 282

Material examined

England: Northumberland; sand beach, March

1955,.D.1,.. Williamson. coll., .5 od;. 6.9.9

(3 ov.), NMNS Cat. No. 5014, slide mounts.

Portugal: Vila de Conde, A. Mateus coll. (no

data), 2 o'c'; 4 99, 2 imm., NMNS Cat.

No. 10464.

Remarks

Although remote geographically from the North

American Pacific coastal fauna, this northeastern

Atlantic species is illustrated herewith to supple-

ment the illustrations of Sars (1895), Bulycheva

(1957) and others, in fully diagnosing the genus

Talitrus and establishing more precisely its

phyletic position among the Talitridae.

Pseudorchestoidea new genus

Orchestoidea: Bulycheva, 1957, p. 130 (partim)

:Bousfield, 1957, p. 120 (partim)

‘Barnard, 1969a, p. 471 (partim)

Diagnosis

Small to medium, smooth, shallow-bodied,

slender-legged sand hoppers characterized by:

head and buccal mass shallow, inferior antennal

sinus short, sharply incised. Eyes very large,

bulging, nearly contiguous dorsally (especially

male). Antenna 1 short, peduncular segments 2

and 3 subequal, flagellum shorter than peduncle.

Antenna 2 slender, elongate (especially male),

peduncular segments slender, not incrassate in

male. Mandibular left lacinia 4-dentate, or with

very weak 5th (proximal) tooth, right lacinia

tricuspate; maxilliped palp tall, appearing

3-segmented, segment 2 with small mediodistal

spinose lobe, segment 3 rounded-subconical,

segment 4 minute (masked by spines of 3) or

lacking. Coxa 1 shallower than 2-4, antero-

distally subacute; coxae 2-4 shallow, wider than

deep, hind margins cuspate. Gnathopod | (¢&

and @) strongly fossorial, spinose; carpus

elongate; propod distinctly shorter, lacking true

palm; dactyl strong; carpus and propod with

small posterior marginal blister in male only.

Gnathopod 2 (co) more or less powerfully

subchelate (or neotenically arrested at partly

transformed stage); propod ovate, palm usually

oblique; dactyl strong, with posterior marginal

process near hinge; gnathopod 2 ( 2 ), basis short,

expanded anteriorly; segment 3 short; 4 with

47

small posterior tumescent process (occasionally

also in male); 5 deeply tumescent behind; 6 mitten-

like, subequal to 5, lobe well beyond dactyl.

Peraeopods 3-7 slender, lightly spined, cuspi-

dactylate, dactyls slender; peraeopod 4 slightly

shorter than 3, segment 4 not greatly shortened,

dactyl weakly pinched, body not overhanging

base of unguis. Coxa 5 aequilobate, elongate,

lobes shallow; coxa 6, hind lobe shallow, anterior

margin usually oblique. Peraeopods 5-7 hetero-

podous, peraeopod 5 markedly shorter than, and

basis and segments 3-6 of different form from,

those of peraeopods 6 and 7; dactyl of 5 very

short, thick, with ‘“‘fuzz’’ setae posteriorly, nail

vestigial; peraeopods 6 and 7 elongate, 7 longer.

Abdominal side plates 1-3 medium deep, nearly

smooth below and behind, hind corners acu-

minate. Pleopods slender; peduncles sublinear,

elongate, inner margin with 2 retinacula, 2 and 3

always with outer marginal spines; rami subequal,

multisegmented or partly reduced, shorter than

respective peduncles. Uropods 1 and 2 slender,

weakly spinose, with terminal spade _ spines;

uropod 1, outer ramus bare, or with outer

marginal spines only; prepeduncle of urosome 1

elongate; uropod 2, rami usually subequal, outer

ramus with outer marginal spines only. Uropod 3

usually elongate, occasionally sexually dimorphic;

peduncle not expanded but weakly spinose

behind; ramus equal to or longer than peduncle,

laterally compressed, margins and apex short-

spinose. Telson usually narrowing distally,

apex slightly notched or entire, lobes with short

apical and dorsal spines. Coxal gills small,

sinuous, those of peraeopods 3-5 smallest, that

of peraeopod 6 not elongate. Brood plates small,

narrowly ovate, smallest on peraeopod 5, margins

distally and sparsely simple-setose.

Type-species: Orchestoidea biolleyi Stebbing

1908 (present selection)

Additional species:

Pseudorchestoidea brito (Stebbing 1891),

piv324pl.,.15

Pseudorchestoidea meridionalis (Schuster

1954)

Pseudorchestoidea gracilis (Bousfield &

Klawe 1963)

Pseudorchestoidea mexicana new species

(p. 50)

Etymology

The generic name is derived from the Greek

‘“‘pseudes”’ meaning “‘false’”’ and the root name

Orchestoidea, and alludes to the mainly super-

ficial similarity between the new genus Pseudor-

chestoidea and the true Orchestoidea. Gender:

Feminine.

Remarks

The genus Pseudorchestoidea presently comprises

three subgroups; the plesiomorphic brito group

(monotypic) of the northwestern Atlantic and

Mediterranean regions; the intermediate meri-

dionalis group, containing three species, and the

apomorphic monotypic biolleyi group, the latter

two groups endemic to the Pacific coast of

Central America and Mexico. The brito group is

rather distinct from the other two groups (Table

III and fig. 27) and links the genus to other sand-

hopper complexes, especially within the Indo- >

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). (In prep.)

*Similarly, the 8 species of ‘‘Talorchestia” sandhoppers

recorded from New Zealand by Hurley (1956) comprise three

or four distinctive morphological and ecological-behavioural

groups that require further analysis and probable generic

recognition.

“Orchestia" costaricana

Talitroides

"Orchestia" vaggala

Transorchestia

Traskorchestia

Protorchestia

Paciforchestia

Arcitalitrus

100

o

~

@

nO

oO

Ly?)

oO

e =

Le)

yi

@

>

rw)

-—_-———

Similarity

v3

50

Platorchestia

-

ie)

-—_<-<—=- —\ 8 = = ween &- = = = eas see

o

©

= o

‘ on x @

S| .31 BS st ®

« + ee ee) (ee fe

» 5 <x & (3) » be

wn & rs) ° & " °

<= od °o =] i=] £ @

a. sa a oa fl gf Ss

o i] (=) a. & § x

% Apo-

56 64 67 72 72 80 90 morphy

Figure 28. Phenogram of Genera of all Groups of Talitridae, North Pacific Rim Region, and other

related genera.

awaits more intensive study of functional

morphology, behaviour, and the life histories of

component species (e.g., Morino 1978, Bowers,

1964).

The distributions of 37 species of Talitridae in

the North Pacific region have been summarized

in Tables V and VI. The gross Pacific rim

coastline has been subdivided into 15 sub-regions

that more or less reflect existing coastal marine

biogeographic provinces (see Brusca & Waller-

stein, 1979, Tzvetkova, 1975). Distributional data

for beach fleas and landhoppers reveal that

highest numbers of species occur on the North

60

American coast in zones 11 and 12, on either side

of 35°N latitude, and on the Asiatic coast in

zones | and 2, at comparable latitudes, and in

comparable numbers; numbers of species decrease

both north and south of those zones. For the

sandhoppers, a similar pattern emerges, for

similar numbers of species. Total numbers

(including introduced species in North America)

also confirm this trend. Thus, fewest species of

Talitridae (4/37) have been recorded from the

vast 6000 km coastline from Kamchatka to

Northern British Columbia. Low species numbers

in the Cortez and Mexican provinces undoubtedly

reflect inadequate sampling and suggest that

further collecting along the Baja and mainland

Mexican coasts would yield much new distribu-

tional, and probably also taxonomical, infor-

mation.

As noted previously (Bousfield, 1979c; 1981),

the northernmost species (e.g., 7. traskiana,

M. pugettensis, and M. columbiana) tend to be the

most plesiomorphic within their generic and

subgeneric groupings, and also tend to be the

widest ranging and most eurytopic species.

Conversely, the most apomorphic species within

these same groupings tend to be the most

southerly in distribution and have the narrowest

latitudinal ranges (e.g., 7. ditmari and T. geor-

giana within the genus Traskorchestia, and

M. benedicti in the Megalorchestia pugettensis

group).

Of primary significance in the evolution of

terrestrial amphipods may be the total lack of

native terrestrial Talitridae in the North

American coastal region, despite their presence

in Central America, Cocos Is., Galapagos Is.,

Hawaii, and Japan (Table VI). This hiatus is not

plausibly explained solely in terms of centres of

origin and continental drift dispersal (see also

Bousfield 1982, in press). Land amphipods in all

peripheral Pacific regions above are ‘‘modern”’

cuspidactylate types, closely related to regional

seashore species groups (e.g., P. japonica to

P. platensis in Japan, ‘“‘Orchestia”’ pickeringi to

“Orchestia’” floresiana in Hawaii, etc.); thus,

terrestrial evolutionary potential is presumably

present in North America also.

Land amphipods may have evolved in concert

with the evolution of angiosperm leaf litter, the

preferred food in continental deciduous rain

forests of Tasmania and Victoria (Friend, 1980).

However, the rain forests of the North American

Pacific coast at comparable latitudes and with

comparable winter-mild climates are today

essentially climax-coniferous, and their forest

floor organic debris is singularly deficient in

deciduous (angiosperm) leaf litter. In this impor-

tant respect they contrast markedly with rain

forests of other peripheral North Pacific regions,

61

and with those of Tasmania and Victoria, where

conifers are relatively rare or lacking, and

deciduous leaf litter is predominant and renewed

seasonally as food for the amphipods. Paratropi-

cal rain forests, including feather palms, fan

palms, and mangroves apparently existed along

the Pacific Northwest coast (northeastern Cali-

fornia to northwestern Washington and S.E.

Alaska) during warm moist Eocene climates and

became predominantly broad-leaved deciduous

during the cooler Oligocene (Wolfe, 1978).

Penetration of Palaeocene leaf litter habitats

by terrestrial amphipods may never be known

with certainty; the cooling climates and decreas-

ing deciduous forests of the Neogene to Recent

eras would have been unfavourable for them, and

none occur there today. Indirect evidence of

possible former occurrence might be obtained

through analysis of feeding types among the

regionally endemic isopods, myriapods, and

other leaf litter arthropods that normally occur

in association with terrestrial amphipods.

Finally, palustral amphipods having immediate

common ancestry with non-cuspidactylate

“‘ancient’”’-type landhoppers (Table II and fig. 26)

are known only from the Central American and

northern South American coasts of the eastern

Pacific, presumably northward to the limit of

mangrove swamps in Mexican and Cortez pro-

vinces, and unknown north of those regions

where native Spartina salt marshes are also rare

or totally lacking. Pertinently, ‘‘Orchestia’”’

vaggala Bowman 1977, the only known terrestrial

amphipod from the evolutionarily isolated

Galapagos Islands in the eastern Pacific, is

very closely related to the chelate (apomorphic)

palustral talitrids (e.g., O. costaricana Stebbing)

endemic to salt marshes and mangrove swamps

of the Caribbean region (Table II, fig. 26), but

not terrestrial there. Perhaps the North American

Pacific palustral or beach flea counterpart with

terrestrial potential is the very plesiomorphic

Paciforchestia klawei (fig. 26) whose western

Pacific analogues (P. tenuimana and _ possibly

“P.” kokuboi) have invaded truly terrestrial

habitats in the southern islands of Japan.

Acknowledgments

The accumulation of extensive material required

for this study has required the generous coopera-

tion of many organizations and _ individuals.

Appreciation to the many agencies and colleagues

who collaborated in the field has been accorded

in several pertinent publications (see especially

Bousfield & Jarrett 1981) and is herewith again

gratefully acknowledged. Several colleagues

have contributed valuable suggestions and

insights into talitrid taxonomy and biogeography

over the years, some of which is embodied

inextricably herewith; I refer especially to past

exchanges with Drs. T.E. Bowman, J.L. Barnard,

and the late C.R. Shoemaker, Smithsonian Insti-

tution, Washington; Dr. Wim Vader, Tromsg¢,

Norway; Dr. A.M.M. Richardson and Dr. A.J.

Friend, Hobart, Tasmania; Dr. Hiroshi Morino,

Mito, Japan; Dr. N.L. Tzvetkova and the late

Dr. E.F. Gurjanova, Leningrad, USSR; Dr. J.

62

Carlton, Woods Hole, Mass.; Dr. D.M. Dexter,

San Diego State College, California; Dr. D.E.

Hurley, Wellington, New Zealand; and museum

colleagues Dr. J.J. Dickinson and Mrs. D.R.

Laubitz. Loan or donation of additional study

lots has been noted in the accounts of material

examined, but for much adventitiously collected

material a note of special appreciation is made to

Dr. W. Klawe, La Jolla, California, Mr. Craig P.

Staude, Friday Harbor Laboratories, Drs.

Stewart and Jarmila Peck, Carleton University,

Ottawa, Dr. Daryl Bowers, Oakland, Calif.;

Mrs. J.F.L. Carl, Victoria; B.C., and Dra

Dexter. Most of the line drawings were rendered

by Mrs. F.E. Zittin, Vancouver, B.C., supple-

mented with a few illustrations by Mr. C. Paquette,

Ottawa, Mr. C. Douglas, NMNS staff artist, and

the author.

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Table 1. Major Taxonomic Characters of Family Talitridae and Their Plesiomorphic and Apomorphic States*

No.

r

10.

Be

pe

24.

ae:

Taxonomic Character

Eye size and position

Antenna 1, length

Antenna 1, flagellum

Antenna 2, dimorphism

Antenna 2, peduncle

Antenna 2, peduncular

segment 4

Mandible, left lacinia

Maxilliped palp

Maxilliped palp, segment 4

Maxilliped palp, segment 2

Coxa 1

Coxa 5, anterior lobe

Coxa 6, posterior lobe

Coxa 6, posterior lobe, anterior

margin

Gnathopod | (0) subchelation

Gnathopod | (co), segment 4

Gnathopod 1 (&)

Carpus (segment 5)

Gnathopod | (<o)

Propod (segment 6) length

Gnathopod | (c)

Propod tumescence

Gnathopod | ( 9) subchelation

Gnathopod 1 ( 9) propod palm

Gnathopod | (@)

Propod tumescence

Gnathopod 2 (o'), subchelation

Gnathopod 2 (oc), propod

palm

Gnathopod 2 (co), dactyl

Phyletic State

Plesiomorphic

Medium, lateral

Elongate, equal to or exceeding

A, peduncle 4

Equal to or longer than

peduncle

Not (or slightly) sexually

dimorphic

Not (or slightly) thickened

(incrassate)

Slightly shorter than peduncle

5 (especially in 9)

More or less 5-dentate

Short, broad

Present; distinct or masked by

spines of 3

Lacking mediodistal lobe

Rounded anterodistally

Shallow, distinctly less than

coxa 4

Shallow, longer than deep

Rounded distally

Strong, dactyl not exceeding

palm

Posterior tumescent lobe pre-

sent, occasionally very small

Short, posterior margin about

half anterior margin

Short (straight), length about

twice maximum width

Postero-distal tumescent lobe

large, distinct

Strong, dactyl not (or very

slightly) exceeding palm

Present, distinct

Postero-distal tumescent lobe

present (may be vestigial)

Strongly subchelate, amplexing

form

Smoothly convex

Regular, smoothly arcuate

*Note: Second, or alternate apomorphic state is italicized.

66

Apomorphic

Large, dorsally contiguous;

very small or lacking.

Short, not reaching end of A,

of peduncle 4

Shorter than peduncle

Distinctly sexually dimorphic

in form

Distinctly incrassate in &

Much shorter than peduncle 5

Gn 2)

4-detate; 6-dentate

Tall, segment 3 subconical

Fused variously to 3 or totally

lacking

With mediodistal lobe

Acute or subacute antero-

distally

Deep, subequal to depth of

coxa 4

Deeper than long

Right-angled or with antero-

distal process

Weak, dactyl exceeding palm;

simple, palm lacking

Tumescent lobe totally lacking

Elongate, posterior margin

*/, anterior margin

Elongate (arcuate), length 3-5

times maximum width

Posterior lobe weak, flat, barely

visible or lacking

Weak, dactyl exceeding palm

Lacking, or trace only

Postero-distal tumescent lobe

totally lacking

Neotenically arrested at trans-

forming stage; mitten-like (as

in Q)

Toothed, incised; Sinuous,

concave

Toothed behind; sinuous

Table 1 Cont'd.

No.

26.

Bi.

28.

29.

30.

a1.

32.

ao.

34.

35.

36.

ST.

38.

39:

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

Taxonomic Character

Gnathopod 2 (@), basis

Gnathopod 2 (9), segment 4,

posterior process

Peraeopods 3 and 4

Peraeopod 4, segment 5

Peraeopods 3 and 4, dactyl

form

Peraeopods 3-7

Peraeopods .3 and 4, dactyl

length

Peraeopods 5-7, form

Peraeopod 5, form

Peraeopod 5, dactyl

Peraeopods 6 and 7, length

Peraeopocs 6 and 7, dimor-

phism

Peraeopods 5-7, dactyls

Pleopods

Pleopod rami

Pleopod peduncles, form

Pleopod peduncles, armature

Uropod 1, prepeduncle

Uropod 1, armature

Uropod 1, peduncle

Uropod 1, outer ramus

Uropod 2, rami

Uropod 2, inner ramus

Uropod 2, apical spines

Phyletic State

Plesiomorphic

Sublinear or slightly broadened

anteriorly, margins subparallel

Broadly rounded

Similar in size and form

Normal, length 2-4 times maxi-

mum width

Subequal in form and size

Simplidactylate; non-cuspidactyl-

ate (or cusps minute)

Short, length about twice width

Homopodous; bases similar in

form, or nearly so (9)

Not unusually small, or differ-

ing (segments 3 & 4)

Normal, as in peraeopods 6

and 7

Peraeopod 7 longer

Not sexually dimorphic in form

Short, length 2-3 times maxi-

mum width

Subsimilar; all of normal length

Normal, not (or little) shorter

than peduncle

Linear or sublinear

Outer margin smooth or nearly

Ye)

Short, depth greater than

length

Weakly spinose; spines few

and/or short

Disto-lateral (inter-ramal) spine

well developed

Margin lacking spines (or

nearly so)

Subequal

Marginal spines in single row

Normally slender, tapering tips

acute

67

Apomorphic

Distinctly broadened antero-

medially, anterior margin

arcuate

Long, acute; lacking

Peraeopod 4 distinctly shorter

Very short, length about equal

to width

Dactyl 4, body pinched, shorter

and stouter than dactyl 3

Cuspidactylate; cusps distinct,

well developed

Long, length 3-4 times maxi-

mum width

Heteropodous; bases distinctly

unlike in shape

Very much smaller and of

different form from peraeopods

6 and 7

Very short, modified, stout,

nail often vestigial

Peraeopod 6 longer

Sexually dimorphic in form

(especially peraeopod 7)

Elongate, slender, length more

than 3 times width

Unequal in size; one or more

reduced

One or more pleopods with

rami distinctly shorter than

peduncle

Broadened or expanded later-

ally and/or basally

All peduncular outer margins

spinose throughout

Elongate, length greater than

depth

Strongly spinose; spines numer-

ous (6-10 per segment) and/or

long

Disto-lateral spine very small

or lacking

Marginally spinose (3—many)

Outer ramus distinctly the

shorter

Marginal spines in 2 rows

Some spines with spade-like

tips (spade spines)

Table 1 Cont'd.

No.

50.

51.

52.

33.

54.

55.

56.

a1.

58.

39.

60.

Taxonomic Character

Uropod 3, peduncle form

Uropod 3, ramus length

Uropod 3, ramus form

Uropod 3, ramus apical spines

Telson, form

Telson, apex

Telson, armature

Brood plates, size and form

Brood plate, peraeopod 5

Brood plates, marginal setae

Coxal gills, form

Phyletic State

Plesiomorphic

Not markedly expanded, length

greater than depth

Shorter than peduncle

Cylindrical, tapering

One or more spines stout,

elongate

Elongate, spade-shaped

Variously cleft, lobes distally

separated

Apical and/or lateral spines

only

Large, broadly subovate

Large, only slightly smaller

than 2-4

Tips hooked or clavate

All plate-like, or sac-like, little

differing in size, little modi-

fied

68

Apomorphic

Strongly expanded behind,

depth (width) about equal to

length

Equal to or much longer than

peduncle; extremely _ short

(length = width)

Laterally compressed, mar-

gins subparallel

All spines weak, short

Short, broad, length not

greater than width

Lobes distally fused to single

plate

Dorsal, plus apical and lateral

spines

Small to medium, narrowly

ovate or sublinear

Much smaller than 2-4

(except when 2-4 lacking)

Tips simple, tapering

Peraeopods 3-5 (especially)

reduced, acetabulate, or very

small; enlarged, convoluted,

and/or elongate

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