Contents

Dickinson, John J. The systematics and distributional ecology of the family Ampeliscidae (Amphipoda:

Gammaridea) in the Northeastern Pacific region. I. The genus Ampelisca.

(Submitted for publication: April 23, 1981)

(Accepted for publication: June 18, 1981) |

Conlan, K.E. and E.L. Bousfield. The amphipod superfamily Corophioidea in the Northeastern Pacific

region. Family Ampithoidae: Systematics and distributional ecology.

(Submitted for publication: May 20, 1981)

(Accepted for publication: September 24, 1981) 4]

Conlan, K.E. and E.L. Bousfield. The superfamily Corophioidea in the North Pacific region. Family

Aoridae: Systematics and distributional ecology.

(Submitted for publication: May 20, 1981)

(Accepted for publication: September 24, 1981) WF

Jarrett, N.E. and E.L. Bousfield. Studies on the amphipod family Lysianassidae in the Northeastern

Pacific region. Hippomedon. and related genera: Systematics and distributional ecology.

(Submitted for publication: June 12, 1981)

(Accepted for publication: September 24, 1981) 103

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The Systematics and Distributional Ecology of the family

Ampeliscidae (Amphipoda: Gammaridea)

in the Northeastern Pacific Region

1. The genus Ampelisca

John J. Dickinson

National Museum of Natural Sciences, Ottawa, Canada, K1A OM8

ABSTRACT

The genus Ampelisca is represented by 14 species on the North American continental shelf in the region

between the Bering Sea and Northern California. Taxonomic analysis of the extensive collections in the

Canadian National Museum of Natural Sciences extends the ranges of nine species from Oregon or

California to northern British Columbia or southeast Alaska. Three species are recorded from near

St. Lawrence Island in the Bering Sea. A new species Ampelisca hessleri is described from northern British

Columbia waters. A bathymetric subspecies Ampelisca macrocephala unsocalae Barnard 1960 is elevated

to species rank. Two geographic variants are elevated to separate species: Ampelisca fageri from A. schel-

lenbergi Shoemaker 1933; Ampelisca careyi from A. macrocephala Liljeborg 1852. A key to the genus for

the northeast Pacific region is included. A cluster analysis is performed on the northeast Pacific Ampelisca

fauna. Sub-generic phyletic relationships are pointed out, but no attempt is made to divide the genus

Ampelisca due to the mosaic distribution of characters among the species.

RESUME

Le genre Ampelisca est représenté par quatorze espéces sur le plateau continental nord-américain, dans la

région qui s’étend de la mer de Béring au Nord de la Californie. L’analyse taxinomique des vastes collec-

tions du Musée national des sciences naturelles du Canada permet d’attribuer a neuf espéces un territoire

plus grand, qui s’étend de l’Oregon ou de la Californie au nord de la Colombie-Britannique ou au sud-est

de |’Alaska. Trois espéces provenant du voisinage de l’ile Saint-Laurent, dans la mer de Béring, sont

signalées. Une nouvelle espéce, provenant des eaux septentrionales de la Colombie-Britannique est

décrite : Ampelisca hessleri. Une sous-espéce bathymétrique est reconnue comme espéce distincte :

Ampelisca macrocephala unsocalae Barnard 1960. Deux variantes géographiques sont aussi reconnues

comme espéces distinctes : Ampelisca fageri (précédemment A. schellenbergi Shoemaker 1933) et A. careyi

(précédemment A. macrocephala Liljeborg 1852). L’auteur fournit une clé des espéces appartenant a ce

genre dans le nord-est du Pacifique et une analyse par populations des espéces d’Ampelisca du nord-est

du Pacifique. Les relations phylétiques entre les espéces sont indiquées, mais il n’est pas question de sub-

diviser le genre Ampelisca, vu lentrelacement des caractéres entre les espéces.

Introduction

Species of the genus Ampelisca in the family

Ampeliscidae are broadly and abundantly

distributed on sand and mud bottoms of con-

tinental shelves (Barnard 1969a; Dickinson et al.

1980). The majority of about 100 worldwide

species are sublittoral forms, but about 20% are

bathyal and abyssal. Species in this genus are

exclusively marine and spend most of their lives

as infaunal tube dwellers feeding on organic

detritus (Enequist 1949; Mills 1967a). The breed-

ing behaviour of Ampelisca is unknown in most

species, but the terminal stage male is known to

be pelagic in shallow water species, and mature

females are thought to swim to the surface when

they are ready to mate (Mills 1967a). It seems

likely that bathyal and abyssal members of the

genus deviate from this pattern, but their breeding

behaviour is completely unknown. The life cycle

in most species of Ampelisca is unknown, but

studies by Mills (1967a) and Kanneworff (1965)

have demonstrated that the life cycle can be as

short as a few months (Ampelisca abdita) or as

long as 2 years (Ampelisca macrocephala). Species

of Ampelisca can be as small as 4-5 mm (Ampelisca

milleri, Ampelisca hancocki) or as large as

25-35 mm (Ampelisca macrocephala, Ampelisca

eschrichti). Zoogeographically, species of

Ampelisca have been reported to have very broad

distributions both geographically and _ bathy-

metrically. However, many of the species which

have been previously treated as having geo-

graphic or bathymetric variants (i.e., Ampelisca

macrocephala) may turn out to be series of closely

related species when examined more critically.

Previous major studies of Ampelisca from the

Pacific coast of North America are those of

Holmes (1908) and Barnard (1954a, 1960, 1967a,

197la) from California and Oregon. Barnard

(1954a, 1960) found 26 species of Ampelisca in

his extensive studies of the southern California

region, and 8 of these species are known from his

limited studies of the Oregon coast (Barnard

1971la). Only scattered records exist north of

Oregon, and only one species of Ampelisca

agassizi has been previously reported from British

Columbia (Mills 1967b). This study on the

systematics and distributional ecology of

Ampelisca in the region between the Bering Sea

and Oregon will attempt to fill in this large gap

in our knowledge of this important genus. The

systematics and distributional ecology of the

other two genera of Ampeliscidae (Byblis and

Haploops) represented in the northeast Pacific

coastal waters will be treated in a subsequent

paper by Dickinson (in prep.)

Materials

This study was based primarily on collections in

the National Museum of Natural Sciences. These

collections range from southeast Alaska to

Oregon, but concentrate on British Columbia.

The primary source of this material was a series

of field expeditions conducted by Dr. E.L. Bous-

field and colleagues at the National Museum of

Natural Sciences. Station data for expeditions

to southern Vancouver Island (1955), Queen

Charlotte Islands (1957), northern Vancouver

Island (1959), southeastern Alaska (1961) and

northcentral British Columbia (1964) were

published previously (Bousfield 1957; Bousfield

1963; Bousfield and McAllister 1962; and Bous-

field 1968). Data for expeditions to Washington

and Oregon (1966), southern Vancouver Island

and Burrard Inlet (1970, 1975 and 1978), and

southeastern Alaska (1980) are available in

manuscript form from the National Museum of

Natural Sciences and will be published in the

near future.

Other major sources of Ampelisca material

used in this study were collections donated to the

National Museum of Natural Sciences by Dr.

Colin Levings of the Pacific Environmental

Institute, Dr. Derek Ellis of the University of

Victoria, and material borrowed from Dr. A.G.

Carey at Oregon State University, Dr. R. Brusca

of the Allan Hancock Foundation, Dr. J.L.

Barnard of the U.S. National Museum, and

Dr. Mary Nerini of the N.O.A.A. lab in Seattle.

Preserved material and prepared slide mounts

of specimens are housed in the National Museum

of Natural Sciences, and representative specimens

are also deposited in the collections of the United

States National Museum, Washington, D.C.

During National Museum of Natural Sciences

field trips to Vancouver Island and southeastern

Alaska in 1977-1980, Dr. E.L. Bousfield (NMNS)

and Mr. Ron Long, Photography Section,

Biology Department, Simon Fraser University,

made a series of colour photographs of living

amphipods as a means of recording their natural

pigmentation patterns. These colour transpar-

encies are catalogued in the National Museum of

Natural Sciences collections and to date include

representatives of A. careyi, A. cristata, A. lobata,

A. pugetica, and A. unsocalae. Regrettably, the

transparencies are too few to reveal inter- and

intraspecific variation in colour pigmentation

for Northeast Pacific Ampelisca; the inclusion of

colour differences in the species diagnoses would

therefore be premature. However, a few brief

notes on colour pattern are included for photo-

graphed species in the hope that future workers

will be stimulated to pursue the investigation of

these patterns in Ampelisca and other coastal

amphipod groups.

SYSTEMATIC SECTION

Ampeliscidae Bate 1857

Diagnosis: Body smooth, pleonites 1-3 smooth or

occasionally weakly carinate, urosomite | usually

carinate, often strongly, especially in male.

Urosomites 2-3 fused. Head usually longer than

deep, lacking rostrum. Eyes 0-6, (when present)

dorsofrontal pair always with lenses, ventro-

frontal pair sometimes with lenses, and dorso-

caudal pair always lacking lenses. Antenna |

usually short, lacking accessory flagellum.

Antenna 2 usually long, peduncular segments 4

and 5 elongated. In terminal male stage, brush

setae present on ventral surface of peduncular

segments 1-2 and basal flagellar segments of

antenna | and on dorsal surface of peduncular

segments 3-5 of antenna 2. Mouthparts slightly

modified. Upper lip with median apical notch.

Lower lip broad, inner lobe well developed,

mandibular lobes weak or lacking. Mandibular

molar and palp well developed, left lacinia with

4-5 cusps. Maxilla | inner plate weakly setose,

outer plate with 11 apical spine teeth. Maxilla 2,

inner plate moderately setose. Maxilliped plates

well developed, palp strong and dactylate.

Anterior coxae usually longer than broad. Coxa 4

strongly excavate posteriorly. Coxae 5-6 anteriorly

lobate. Coxa 7 posteriorly lobate. Gnathopod 1

simple or weakly subchelate occasionally sexu-

ally dimorphic. Gnathopod 2 elongate, and

usually simple. Peraeopods 3 and 4 glandular,

segment 5 short, dactyls elongate, simple, with

ducts. Peraeopods 5-7, bases broad. Peraeopods

5 and 6, similar in form and size, segment 5

usually with comb teeth on posterior margin,

dactyls short. Peraeopod 7 short, different in

form than peraeopods 5 and 6. Coxal gills on

peraeopods 2-6, usually pleated, sexually dimor-

phic, mature females having flatter, narrower,

and less pleated gills than mature males. Brood

plates linear, marginal setae simple. Pleopods

strong; retinacula paired, short stemmed, curved

tip, one margin toothed. Uropods 1-2, rami

usually subequal and narrowly lanceolate.

Uropod 3 varies in form, but rami usually long,

broadly lanceolate, and setose, outer ramus one-

segmented. Telson bilobate, sometimes sexually

dimorphic in size and form, lobes usually deeply

and narrowly separated, apices often notched

and spinose.

Ampelisca Kroyer 1842

Diagnosis: Head longer than deep, narrowing

anteriorly with anteroventral corner unproduced.

Eyes (when present) have lenses in both dorso-

frontal and ventrofrontal pairs. Antenna 2,

flagellum with more than 5 flagellar segments.

Mandibular palp, segment 3 usually much

shorter than segment 2. Maxilliped, inner plate

short and broad. Anterior coxae much longer

than broad. Peraeopods 3-4, segment 5 very short

and often partially overhung by segment 4.

Peraeopods 5-6, dactyls very short and hook-

like. Basal lobe of peraeopod 7 expanded distally,

posterior margin oblique, and lacking setae on

anterior margin near junction with segment 3.

Peraeopod 7 with seven distinct segments, dactyl

broad at base. Telson lobes elongated and fused

only basally in both sexes.

Remarks: The diagnosis of the family Ampeli-

scidae closely parallels Bousfield’s (1978)

diagnosis of the superfamily Ampeliscoidea, but

encompasses more precisely the generic vari-

ability within the family. The diagnosis of the

genus Ampelisca attempts to strike a middle

ground between the narrow diagnosis of Barnard

(1969a) and the broad diagnosis of Bousfield

(1973). The chief problem in any definition

involving the family Ampeliscidae is the poor

state of knowledge regarding the monotypic

genus Triodos K.H. Barnard 1916. Barnard’s

original description and illustrations are very

incomplete, and subsequent work by Griffiths

(1976) adds little new taxonomic information.

Key to Ampelisca of the Northeast Pacific Region

feet eon'sideplate 3 with.an acute tooth on lower posterior corner. ....-.........0. 60.2 Ss eee 2

Eicon sideplate 3 without anacute tooth on lower posterior cormer:. ...02) 0 0200. So eee 9

2. Segment 5 of peraeopod 7 with notch in anterior margin and uropod | not reaching beyond

nm OA RCUEL CNT, CTD CT MAO 2 550 dsr eles canner cveraeas Abs ich hay sandepnanard tue «5 RRM ey LA 3

Segment 5 of peraeopod 7 without notch in anterior margin and uropod | reaching end of

Tens WNT Ie EUS MUNN lect Bh kk ON on are otchn icawinn ax adem rarer eer oe ammriagan div DAD SAA RSG eee Micanne Nol eyat 4

3. Dorsal carina of urosomite | saddle shaped; posterior lobe of segment 4 of peraeopod 7

broad, extending % length of segment 5; dorsal surface of telson with long spines ...........

we he os OH Be ee Ce veee@esceseanvcecuwevetscnenseesaeeves

a Sale cersade aie aunts RNR He Ampelisca pugetica p.17

Dorsal carina of urosomite | not saddle shaped; posterior lobe of segment 4 of peraeopod 7

slender, extending less than '/, length of segment 5; dorsal surface of telson with short

RR regal eis Satara aa om Rw oo

Dina ete ire ahs Cenere Gene Ampelisca eschrichti p. 18

10.

bf,

14.

15.

Uropod 2 lacking subapical spine on outer ramus; posterior margin of pleon sideplate 3

neany straight: dactyl of peracopod 7 short and thick i. .s0.4.....58 2% Ampelisca hancocki

Uropod 2 with long subapical spine on outer ramus; posterior margin of pleon sideplate 3

convex dactylort peracopod 7 longandslender. .....uslivesS Gxeeiaiek. Lactose

Lower front margin of head concave and parallel to upper margin; pleon sideplate 2 witha

tooth onilowerposteriorcomen i <aweursdh......cawe) wets lessee. 6h eee

Lower front margin of head convex or slightly concave but never parallel to upper margin;

pleanisideplate 2:without a tooth on lower posterioncorner «2.1942 .a’. dawn 20k. eee

Pleon sideplate 3 produced into a large process above tooth; telson, apex of each lobe narrow,

lacking large notch; urosomal carina massive not lamellar ........ Ampelisca brevisimulata

Pleon sideplate 3 weakly convex, lacking large process above tooth; telson lobe, apex with

larsemotchsuresomal caringdamellar ces... 3... ..dceeslave teh. Ampelisca cristata

Dorsal surface of telson with short blunt spines concentrated in a central row..............

I ee, Gia. d Makerere sale. « . . ee eee a ee | Ampelisca macrocephala

Dorsal surface of telson with long slender spines scattered randomly over lobes ............

Lower front margin of head concave; antenna | reaching end of peduncle of antenna 2; tooth

Cmpleomsidepinte: Sshonyand qhick woe ws ... . aul Piadvamiowetes ceees Ampelisca careyi

Lower front margin of head convex; antenna | not reaching end of peduncle of antenna 2;

toothon pleomsideplate 3 long.and slender... . icc. 20 sb. Ampelisca unsocalae

Rami of uropod 3 short and flattened; basal lobe of peraeopod 7 short only reaching end of

segment 3; spines lacking on posterior margin of segment 5 of peraeopods 5and6 ..........

Rami of uropod 3 long and not flattened; basal lobe of peraeopod 7 extending beyond

segment 3; spines present on posterior margin of segment 5 of peraeopod5 and6...........

Antenna | long, extending beyond peduncle of antenna 2; outer ramus of uropod 2 lacking

Cine aee eine eee eee RE a bapiey.. Sede... . SRR See caer Ampelisca hessleri

Antenna | short, not extending beyond peduncle of antenna 2; outer ramus of uropod 2 with

SHOa Pied PilcmNnnne CTU. . Seine Tee... . Saslesnaest oo chee ween aR oe

Basal lobe of peraeopod 7, lower posterior margin strongly oblique, eyes present ...........

Ser ae Meee re ane. SI ey a Gas weewiew Tod, Ampelisca birulai

Basal lobe of peraeopod 7, lower posterior margin weakly oblique, eyes absent ............

ee eee a. ORE ga. ges... Seals wierd fee Bs Ampelisca plumosa

Segment 3 of peraeopod 7 longer than segment 4, mandibular palp segment 2 inflated .......

Pe er eR Se er eee eed... . Lowa mivevs well aiaers Ampelisca milleri

Segment 3 of peraeopod 7 subequal to segment 4 in length; mandibular palp segment 2

slenderno mim larcdssublinear 4). .sicase.... SORE les wee kee. Jee al Re

Urosomite | produced dorsally into a well developed carina; uropod | with rami much longer

than short stout peduncle; antenna | not reaching end of peduncle of antenna2 ............

pr amse Sara e tears Sh hes, 182194 3s. ORRIN eee es Ampelisca agassizi

Urosomite | not produced into a carina, uropod | with rami shorter than peduncle; antenna |

hedemine cudor peduncle of antenna 2° lene... . . Se SPSTR Ae CSS. Le db 2ee ad ee

Segment 5 of peraeopod 7 lacking spine bearing notch on anterior margin; outer ramus of

uropod | spinulose on inner margin; inner margin of inner ramus of uropod 3 with six spine-

DGaeTe Se reatIOMst, 45, UNSn 8. teehee eaten, swe Bye Geet oath) Ampelisca lobata

Segment 5 of peraeopod 7 with spine bearing notch on anterior margin; outer ramus of

uropod | unarmed on inner margin; inner margin of inner ramus of uropod 3 lacking spine-

Cates e mr Acros te, fates HL a STIRS oct nssisvn ted snesa nae oO RE nas ee ee

Uropod 3, inner ramus distinctly longer than outer ramus, inner margin of inner ramus

Dearie asetics of largeteetht U....% Mi Mie Ri ind SARS ae Ampelisca schellenbergi

Uropod 3, inner ramus subequal to outer ramus, inner margin of inner ramus microdentate

Oly 3s... ROS RI es eas {eee eerie eet Ampelisca fageri

2hS

. 10

Ji?

Ampelisca agassizi (Judd) 1896

Figure 1.

Byblis agassizi Judd 1896, pp. 599-603, figs. 9-11

Ampelisca compressa Holmes 1903, p. 273;

1905, pp. 408-481, fig. (no number).

Ampelisca agassizi Holmes 1905, pp. 418-482,

fig. (no number).

:Mills 1967b, pp. 643-645, fig. 3.

‘Bousfield 1973, p. 135, pl. 38.

Ampelisca vera Barnard 1954a, pp. 23-26, pls.

14-16.

Material examined: British Columbia — Queen

Charlotte Islands: Bousfield stns. 1957, H1

(6 spec.). J.W. Scoggan stns., 1965, JWS 89, 104,

105, 106 (30 spec.). North central coast: Bous-

field stns. 1964, H8, H34, H36, H37, H48, H49

(about 60 spec.). C. Levings, Swanson Bay stns.,

1975, 51B-001, 51B-002, 51B-003, 51B-004, 51B-005,

eer

LL |

51B-007, 51B-008, 51B-009, 51B-010, 51B-011,

51B-012, 51B-013 (about 400 spec.). Vancouver

Island, Shoal Bay, Victoria: J.F.L. Hart, 1949

(2 spec.); Bousfield stns. 1959, 06, V22 (4 spec.).

Bousfield stns. 1975, P29b (1 spec.). Bousfield

stns. 1976, B-26 (4 spec.). Bousfield stns. 1977,

B8 (1 spec.). Verdier Pt., Saanich Inlet, K. Conlan,

1976-77 (about 250 spec.), D.V. Ellis, 1980,

Sidney (20 spec.). Southern mainland, Bousfield

stns. 1959, N23b. Washington: Bousfield stns.

1955, M10, Bousfield stns. 1966, W13 (2 spec.).

Oregon: OSUBI 01280. 46°6’'N,124°8’W, 70 m,

4 Dec. 1974 (1 spec.) OSUBI 01267. 44°26'3N,

124°14.7'W, 60 m, 21 June 1976 (2 spec.)

Distributional ecology: Geographic range: North

Atlantic, Nova Scotia to Caribbean (Barnard

1954b; Mills 1967b, 1971). North Pacific, Queen

Charlotte Islands to Ecuador (Barnard 1954a,

Figure 1. Ampelisca agassizi (Judd). Swanson Bay, B.C., 9 9 mm ov. Shoal Bay, B.C., ¢& 10 mm

197la). Bathymetric range: North Atlantic,

5-450 m (Mills 1971; Dickinson et al. 1980).

North Pacific 0-266 m (Barnard 1971la). NMNS

coll. 0-300 m, primarily 40-100 m. Abundance:

North Atlantic, up to 15,000/m? (Dickinson et al.

1980). North Pacific, rarely over 500/m’.

Life cycle: Ovigerous females found in B.C.

waters during July (1), November (1). Terminal

males: April (2).

Diagnosis: (Female). A medium sized Ampelisca

(8-11 mm) characterized by: head short, lower

front margin oblique nearly straight. Antenna |

short, not reaching end of peduncle of antenna 2,

peduncular segment 2 twice length of segment 1

and with 8-10 long setae on ventral surface,

flagellum setae long. Antenna 2, about as long

as body, flagellum setae long. Mandibular palp,

segment 3 over % length of segment 2. Coxae 1-3

without slit or tooth on lower posterior corner.

Gnathopod 1, nearly simple. Gnathopod 2, seg-

ment 5 less than twice the length of segment 6.

Peraeopod 3, plumose setae along entire posterior

margin of segment 4. Peraeopods 5-6, 2-3 rows of

weakly developed comb spines along posterior

margin of segment 5. Peraeopod 7, basis broad,

segment 3 subequal to 4 in length, segment 4

produced posteriorly into a slender, setose lobe

extending most of the length of segment 5,

segment 6 longer than 5, segment 7 slender and

subequal to 5 in length. Uropod 1, peduncle

short and deep, rami long reaching end of

uropod 2, inner ramus spinulose, outer ramus

unarmed. Uropod 2, rami shorter than peduncle,

inner ramus spinulose on both margins, outer

ramus spinulose on outer margin only. Uropod 3,

rami lanceolate, outer ramus bearing more

plumose setae than inner. Telson lobes, broad,

apices blunt and armed with 4-5 setules, dorsal

surface bearing 2-3 pairs of spines along cleft.

Pleon sideplate 3, lower posterior corner rounded.

Urosomite 1, well developed dorsal carina which

is rounded posteriorly. Urosome compressed in

length. Gill flattened in mature females, dendritic

folds short, narrow and curved downward.

(Male). Very rare in collections. Differ from

female in: presence of brush setae on peduncles of

antenna 1-2; gills less flattened and dendritic for

entire length; urosomite 1, deep notch cut in

anterior dorsal surface, massive hood-like carina

projecting dorsally over urosomite 2; uropod

bearing plumose setae on both margins of

both rami.

Remarks: Barnard (1960), Mills (1967b) and

the author have compared specimens from the

Atlantic and Pacific coasts of North America,

and have been unable to find a morphological

basis for separating the populations on either

side of the Isthmus of Panama. This suggests that

evolution in this species must be relatively slow

since these populations have been disjunct for at

least one million years. Ampelisca agassizi is

relatively distinct morphologically from other

species of the northeast Pacific region (see pheno-

gram Fig. 21) and is easily identified by its

combination of rounded pleon side plates, blunt

telson, lanceolate uropod 3, and compressed

urosome (see Key p. 3).

Ampelisca hessleri n. sp.

Figure: 2.

Material examined: British Columbia — Queen

Charlotte Islands: J.W. Scoggan coll. 1965;

JWS103 (54°37’N,133°55’W), 225 m (3. speci-

mens), JWS 104 (54°32’N,133°25’W), 325 m

(1 specimen). North central coast: C. Levings

1963 coll; Swanson Bay (53°0.25’N,128°30.9’W),

152 m, 4 April 1973; 2 (6.0 mm) holotype

(NMC-C.1981-378), 2 @9 paratypes (NMC-

C.1981=377).

Distributional ecology: Known only from

northern B.C. waters at upper bathyal depths

(150-325 m).

Diagnosis: (Female). A small Ampelisca

(6-8.0 mm) characterized by: head, eyes present,

lower front margin oblique and convex. Antenna |

long, extending well beyond peduncle of

antenna 2, peduncular segment 2 only slightly

longer than 1, flagellum setae of medium length.

Antenna 2 about % body length, peduncular

segment 4 about twice length of antenna |

peduncular segment 2, flagellum setae of medium

length. Mandibular palp, segment 3 about half

length of segment 2. Coxae 1-3, lower posterior

corner toothed. Gnathopod 1, palm elongate

extending 7/, posterior margin of segment 6.

Gnathopod 2, segment 5 less than twice the

length of segment 6. Peraeopod 3, plumose setae

confined to distal third of posterior margin of

segment 4. Peraeopods 5-6, posterior margin of

segment 5 unarmed. Peraeopod 7, basal lobe

relatively slender and lower posterior margin

obliquely rounded, segment 3 subequal to 4 in

length, segment 4 with posterior lobe weak and

sparsely setose, segments 5 and 6 subequal in

length, segments 3-7 slender and stick-like.

Uropod 1, peduncle stout, rami shorter than

Figure 2. Ampelisca hessleri n.sp. Swanson Bay, B.C., 2 holotype, 6 mm

peduncle, inner margin of inner ramus with 4

spines, outer ramus unarmed. Uropod 2, peduncle

longer than rami, outer ramus shorter than inner,

both rami spinulose, no apical spine on outer

ramus. Uropod 3, rami flattened, broad and

foliaceous, distal margins of outer ramus ser-

rated with a plumose seta set in each serration,

apex of inner ramus notched medially with 6-8

plumose setae set in notch. Telson lobes, tapering

to a blunt tip bearing 3-4 setules, dorsal surface

with 3-4 short spines along each side of the cleft.

Pleon side plate 3 rounded, without tooth at

lower posterior corner. Urosomite | elevated into

carina which is upturned posteriorly. Gills flat-

tened and unpleated in mature females.

(Male). Unknown.

Remarks: This new species is very similar to

A. plumosa Holmes and A. birulai (see pheno-

gram Fig. 21), but these species can be distin-

guished by the relative lengths of antennae, shape

of the basal lobe of peraeopod 7, stoutness of

rami of uropod 3, and the presence or absence of

a tip spine on uropod 2 (see key p. 3 and

Figs. 2-4).

Etymology: This species is named in honour of

Dr. Robert Hessler who has contributed greatly

to our knowledge of deep sea benthic ecology,

and who also has a special interest in the evolu-

tion of peracarid crustaceans.

Ampelisca plumosa Holmes 1908

Figure: 3;

Ampelisca plumosa Holmes 1908, pp. 509-510,

figs. 18.

‘Barnard 1960, pp. 30-31, fig. 8

Material examined: Southern California —

North Coronado Island: Off San Diego, 1130-

1190 m, USNM cat. no. 38542, holotype (13.0

mm) sub. o.

Distributional ecology: Known only from

/ Z

Z |

i ci oe

Figure 3. Ampelisca plumosa Holmes. North Coronado Island, California. & holotype, 13 mm, subadult

bathyal depths off southern California (Holmes

1908, Barnard 1960). Dickinson and Carey’s

(1978) records in abyssal depths off Oregon are

another new species in this group.

Diagnosis: The diagnosis of this species is

based solely on a subadult male (holotype

13.0 mm) which usually differs from the female

only in the structure of the gills and the presence

of penal papillae. This species is characterized by:

head, eyes absent, lower front margin of head

nearly vertical. Antenna | just reaching end of

peduncle of antenna 2, peduncular segment 2

about one and half times length of segment 1,

flagellum setae medium length. Antenna 2 about

body length, peduncular segment 4 long, more

than twice length of antenna | peduncular

segment 2, flagellum setae medium length.

Coxae 1-3, lower posterior corner toothed.

Gnathopod 1, palm short, extending only '/,

length of posterior margin of segment 6. Gnatho-

pod 2, segment 5 elongate, more than twice length

of segment 6. Peraeopod 3, plumose setae along

entire posterior margin of segment 4. Peraeopods

5-6, posterior margin of segment 5 unarmed.

Peraeopod 7, basal lobe relatively broad and

lower posterior margin gently rounded, segment

3 subequal to 4 in length, segment 4, posterior

lobe weakly developed and sparsely setose, seg-

ments 5 and 6 subequal in length, segments 3-7

slender, stick-like. Uropod 1, peduncle stout,

rami slightly shorter than peduncle, inner margin

of inner ramus with 2 spines, outer ramus

unarmed. Uropod 2, peduncle longer than rami,

outer ramus smaller than inner ramus bearing a

medium length apical spine, both rami spinulose.

Uropod 3, rami flattened, relatively slender, and

foliaceous; distal margins of outer ramus ser-

rated with a plumose seta set in each serration,

apex of inner ramus notched medially with 6-8

plumose setae set in notch. Telson lobes tapering

to a blunt tip bearing 3-4 setules, dorsal surface

with 4 pairs of long spines along cleft. Pleon

sideplate 3, posterior corner rounded, without a

tooth. Urosomite 1 elevated into a carina which

is upturned posteriorly. Gills highly pleated in

subadult male.

(Pelagic terminal male). Unknown.

Remarks: This species is unknown from study

area, but it is included to help distinguish it

from A. hessleri n.sp. and A. birulai Bruggen.

It was also thought a more critical diagnosis and

more detailed illustrations would be useful to

future workers since representatives of this species

group of Ampelisca seem to be common at bathyal

and abyssal depths of the northeast Pacific.

Ampelisca birulai Bruggen 1909

Figure 4. |

Ampelisca birulai Gurjanova 1951, p. 310,

me 173.

‘Shoemaker 1955;p. 10, fig. 2,n-q, 3,a-c

Material examined: Alaska — Beaufort Sea

(71°21.9N,152°33.4W), 99 m, 3 November 1976,

Oms.U.B.1. 00985 (1 ov. 9); Bering Sea: St.

Lawrence Island (64°N,169°W), July 1980,

J. Oliver coll., (3 specimens).

Distributional ecology: Geographic range:

Arctic Ocean and Bering Sea (Shoemaker 1955).

Bathymetric range: 10-100 m (Shoemaker 1955).

Life cycle: Ovigerous females found in Beaufort

Sea during November (1).

Diagnosis: A medium sized Ampelisca (8-10 mm)

characterized by: head, eyes present, lower front

margin oblique, nearly straight. Antenna |

reaching just beyond peduncle of antenna 2,

peduncular segment 2 subequal to segment 1,

flagellum setae medium length. Antenna 2 about

half body length, peduncular segment 4 more

than twice length of antenna 1, peduncular

segment 2, flagellum setae medium length.

Mandibular palp, segment 3 about ’/, length of

segment 2. Coxae 1-3, lower posterior corner

strongly toothed. Gnathopod 1, segments 5 and 6

stout, palm short and deep, almost vertical,

Figure 4. Ampelisca birulai Bruggen.

Pitt Point, Beaufort Sea, Alaska. 9 8 mm, Ov.

dactyl short and thick. Gnathopod 2, segment 5

less than twice length of segment 6. Peraeopod 3,

plumose setae sparsely distributed along entire

posterior margin of segment 4. Peraeopods 5-6,

posterior margin of segment 5 unarmed. Peraeo-

pod 7, basal lobe relatively broad with lower

posterior margin steeply oblique, segment 3

subequal to 4 in length, segment 4 posterior lobe

weak and sparsely setose, segments 5 and 6 sub-

equal in length, segments 3-7 slender and stick-

like. Uropod 1, rami and peduncle subequal in

length, inner mfargin of inner ramus armed with

3 spines, outer ramus unarmed. Uropod 2,

peduncle longer than rami, outer ramus shorter

than inner and bearing a sub-apical spine, both

rami spinulose. Uropod 3, rami flattened,

relatively slender, and foliaceous; distal margins

of outer ramus serrated with plumose seta set in

each serration, apex of inner ramus notched

aD PS

SS — =——

oF —

Figure 5. Ampelisca lobata Holmes. Stn. H21

medially with 4-6 plumose setae set in notch.

Telson lobes each tapering to a blunt tip bearing

3-4 setules, dorsal surface with 1-2 pairs of short

spines along cleft. Pleon sideplate 3, lower

posterior corner rounded without a tooth.

Urosomite 1, elevated into a carina which is

upturned posteriorly. Gills flattened and

unpleated in mature female.

(Pelagic terminal male). Unknown.

Remarks: Ampelisca birulai is extremely similar

to A. plumosa (see phenogram fig. 21), but is

easily distinguished by the shape of its basal lobe

on peraeopod 7 (see key p. 3).

Ampelisca lobata Holmes 1908

Figure 5.

Ampelisca lobata Holmes 1908, pp. 517-518,

fie. i25

‘Barnard 1954a, pp. 11-14, pls. 5-6

(1964), North Bank Island, B.C. 9 7 mm, ov.

Ampelisca articulata Stout 1913, pp. 639-640

Material examined: British Columbia — Queen

Charlotte Islands: Bousfield stns. 1957; H2a,

H14, W9, W12 (4 specimens). J.W. Scoggan stns.,

1965; JWS 106 (4 specimens). Northcentral coast:

Bousfield stns., 1964; H10, H21, H26, H28, H31,

H34, H36, H50 (11 specimens). L. Marhue stns.,

1966; LMI17 (1 specimen). Vancouver Island:

NMNS coll. 1909; Ucluelet (5 specimens), NMNS

coll. 1931; Nanoose Bay, Nanaimo (1 specimen).

Bousfield stns., 1955; Fl (3 specimens). Bous-

field stns., 1959; V4b, V6, 01, 03, 05, 07b (50

spec.). N.A. Powell, NMNS coll. 1969; Wiffen

Spit (2 specimens). Bousfield stns., 1970; P702,

P713, P719 (3 specimens). Bousfield stns., 1975;

P5b, PSc, Pl7e (30 specimens). Bousfield stns.,

1976; B26, B27 (100.specimens). USNM coll.

Round Island, Nanoose Bay, Ucluelet (6 speci-

mens). D.V. Ellis, Sidney Harbour stns., 1979;

1601, 2401, 2602, 3401 (4 specimens). Southern

mainland coast: Bousfield stns., 1977; E3, P3,

(32 specimens).

Distributional ecology: Geographic range —

Queen Charlotte Islands to Baja California

(Barnard 1954a,b, 1967a, 1971la). Records from

Caribbean Sea (Barnard 1954b) and South

America (Barnard 1954a) were unillustrated and

are suspect until examined by more recent critical

standards. Bathymetric range: 0-183 m_ with

southern submergence (Barnard 1969b). 0-144 m

off British Columbia but most commonly 0-40 m.

Sediment preference: Strong association with

plants (Barnard 1969b). Mixed bottoms of rock

and sand off British Columbia.

Life cycle: Ovigerous females found in B.C.

waters during May (1), July (7), August (1),

November (1).

Diagnosis: (Female). A medium sized Ampelisca

(7-9 mm) characterized by: head, lower front

margin oblique and convex. Antenna | extending

just beyond end of peduncle of antenna 2,

peduncular segments | and 2 subequal, flagellum

with short setae. Antenna 2 about two thirds

body length, flagellum with medium length setae.

Mandibular palp segment 3 about four-fifths

length of palp segment 2. Gnathopod 1, palm

elongate. Gnathopod 2, segment 5 less than twice

length of segment 6. Coxae 1-3 slender, slit at

lower posterior corner. Peraeopod 3 slender,

segment 4 bearing plumose setae only on distal

third of the posterior margin. Peraeopods 5 and

6, three sets of comb spines on posterior margin

of segment 5. Peraeopod 7, basal lobe deep and

relatively slender, segment 4 longer than 3, witha

small setose posterior lobe, segments 5 and 6 sub-

equal in length. Uropod | reaching to end of

uropod 2, both rami spinulose. Uropod 2, outer

ramus slightly shorter than inner ramus, both

margins of both rami spinulose. Uropod 3, inner

ramus longer than outer ramus, inner ramus

strongly serrate along inner margin, with six

blunt spines each set in a serration, inner ramus

without setae, outer ramus with a few terminal

setae. Telson, lobes notched laterally at tip, each

notch bearing a short blunt spine, dorsal surface

of each lobe with 3-5 short blunt spines in a

central row. Pleon sideplate 3 usually quadrate at

lower posterior corner, occasionally minutely

toothed in large specimens. Urosomite | raised

only slightly dorsally, posterior edge flush with

urosomite 2. Gills flattened, with dendritic folds

short, narrow and curved downward.

(Male). None in NMNS collections. According

to Barnard (1954a), mature males differ in having:

brush setae on peduncles of antenna | and 2,

flagellum of both antenna elongated; gnathopod

1, segment 6-narrowed distally (see A. pugetica

figure); uropod 3, rami subequal in length, inner

ramus lacking serrations, both margins of both

rami setose. Subadult males have less flattened

gills with more dendritic folds. Presumably,

adult males would have even more complexly

folded gills.

Remarks: Ampelisca lobata is most similar to

A. fageri (see discussion and phenogram), especi-

ally in structure of urosome, gills, and telson.

However, the two species are easily separated on

differences in peraeopod 7 and uropod 3 noted

in the key.

A single photograph of A. lobata (HAAA lab. 1)

shows the colour patterns in the female specimen

from the Burrard Inlet region. The majority of

the body is translucent with faint traces of red

pigment on the antennae and peraeopods. The

anterior portion of the head is highly pigmented

in cardinal red and yellowish white. The red

pigment is concentrated around the eyes and is

probably the subcuticular non-retinular screening

pigment referred to in Halberg et al. 1980. The

yellowish white pigment is located peripheral to

the major concentrations of red pigment. It

appears closer to the cuticular surface, and is

more granular in appearance. The distal segments

of the anterior peraeopods also have scattered

spots of white pigment.

Ampelisca fageri n. sp.

Figures 6-7.

Ampelisca schellenbergi Barnard 1954a in part,

pp. 14-16; pls 7ay'ey pls8a;) by dj..e; f, ‘ge.

Material examined: British Columbia

Whiffen Spit, Sooke Basin, Vancouver Island;

(48°21’N,123°44’W), Bousfield stns., 1955; FI,

gravelly sand among boulders, 31.3°/ 9, 10.9°C,

intertidal, 17 August 1955; holotype 2 8.0 mm

br.II (NMC-C.1981-379); 2 99 and 6 imm.

paratypes (NMC-C.198 1-380). Additional records:

North-central coast: Bousfield stns., 1964; H1

(3 specimens), Vancouver Island, Bousfield stns.,

1955; P6éa (1 specimen). Bousfield stns., 1959;

Voabr (le specimen)... N.A»° Powell ‘stns., 1969:

Wiffen Spit (#319) (1 specimen). Bousfield stns.,

1970; P702, P711 (6 specimens).

Mexico — Gulf of California, San Estaban

Island (28°48’N, 112°34’W): A. Hancock Foun-

dation stns., 562, 729 (200 specimens).

a = ae

Distributional ecology: Geographic range —

Northeastern Pacific, North central coast of

British Columbia (51°39’N,128°09’W) to the Gulf

of California, Mexico. Bathymetric range —

intertidal in B.C. showing a submergence to the

south but rarely deeper than 40 m (Barnard

1967a). Sediment preference — mixed bottom

areas of sand and boulders.

Life cycle: Ovigerous females found in B.C.

waters during July (1), August (2). Pelagic males,

none.

Diagnosis: (Female). Medium-sized Ampelisca

(8 mm) characterized by: head, lower front

margin convex. Antenna 1 reaching end of

peduncle of antenna 2, peduncular segment 2

slightly larger than 1, flagellum setae short.

Antenna 2 about %4 body length, flagellum setae

very long. Mandibular palp, segment 3 subequal

to segment 2 in length. Coxae 1-3, slit at lower

posterior corner, long setae along entire ventral

Figure 6. Ampelisca fageri n.sp. Wiffen Spit, Vancouver Island, B.C. 9 holotype, 8 mm, br II

FAGERI

SCHELLENBERGI

Figure 7. Ampelisca fageri n.sp. Wiffen Spit, Vancouver Island, B.C. 2 holotype, 8 mm, br. II, upper

panel. Ampelisca schellenbergi Shoemaker. Dog Island, Gulf coast, Florida, 9 9 mm, br. II, lower panel.

margin. Coxa 1 weakly expanded distally.

Gnathopod 1, palm shallow and elongate.

Gnathopod 2, segment 5 short, less than twice

as long as segment 6. Peraeopod 3 stout, sub-

equal to peraeopod 4, marginal setae restricted

to distal third of segment 4. Peraeopods 5 and 6,

bases broad, posterior margin of segment 5

armed with 2-3 sets of comb spines. Peraeopod 7,

lower posterior margin of basis oblique nearly

Straight, cut by a series of unevenly spaced

shallow setae bearing notches, segment 4 longer

than 3 and very broad, posterior lobe of seg-

ment 4 bearing 14-16 plumose setae, anterior

margin of segment 5 twice length of posterior

margin and cut by a deep spine bearing notch,

segment 6 long and attached near posterior

margin of segment 5, segment 7 short and stout.

Uropod | reaching end of uropod 2, inner margin

of inner ramus spinulose, outer ramus with basal

spines only. Uropod 2, peduncle longer than

rami, both margins of both rami spinulose.

Uropod 3 short, inner ramus slightly longer than

outer ramus, outer ramus notched at tip, inner

edge of inner ramus microdentate only, no large

teeth present. Telson, lobes notched apically, a

single blunt spine in each notch, dorsal surface of

each lobe with 2 blunt spines in a central row.

Pleon side plate 3, lower posterior corner nearly

quadrate, lacking tooth. Urosomite 1, dorsal

surface weakly produced, posterior margin flush

with urosomite 2. Gills in mature females

flattened, dendritic folds short, narrow, and

curved downward.

(Male). Unknown. Probably similar in form to

that described for A. schellenbergi in Barnard

(1954a).

Remarks: This species is very similar to

Ampelisca schellenbergi Shoemaker 1933, and

Barnard (1954a, 1967a, 1969b) treated it as

merely a geographic variant. The two forms can

be separated on the basis of the shape and seta-

tion of coxa 1, the shape of the basal lobe of

peraeopod 7, and the relative size and shape of

the rami of uropod 3 (see Fig. 7). In addition to

the morphological differences, the geographic

range and differences in ecology confirm that

these two forms are different species.

Etymology: The species 1s named in honour of

the late Dr. E.W. Fager of the Scripps Institute

of Oceanography who was a brilliant and influ-

ential teacher of the use of quantitative methods

in marine ecology.

Ampelisca schellenbergi Shoemaker 1933

Figure 7.

Ampelisca schellenbergi Shoemaker 1933, pp.

2-5), tg.’ 2:

‘Barnard 1954a, in part pp. 14-16, pls. 7b,d,e,

8c.

Material examined: Florida — Gulf coast: Dog

Island, Franklin County, NMNS coll. 1965

(1 specimen); Tampa Bay, NMNS coll. 1976

(1 specimen).

Costa Rica — Pacific coast: Salinas Bay,

A. Hancock Foundation stn. 478 (17 specimens).

Remarks: This species is not found north of

Central America, and is included only as an aid

in defining A. fageri n.sp. (see Fig. 21 and dis-

cussion p. 30). Ampelisca schellenbergi is reported

from the Gulf of Mexico, Caribbean Sea, and in

the= eastern Pacific irom: ‘Costa;/Rica: to)-Peru

(Barnard 19544, 1954b). Barnard’s (1967a, 1971b)

records of this species from Hawaii are suspect,

and should be re-examined more critically

considering the disjunct nature of the Hawaiian

population. However, it should be noted that the

Hawaiian form is much closer to A. schellenbergi

than to A. fageri in the form of its uropod 3,

coxa 1, and peraeopod 7.

Ampelisca hancocki J.L. Barnard

Figure 8.

Ampelisca hancocki Barnard 1954a, pp. 37-38,

pl. 26.

Material examined: British Columbia — North

central coast: C. Levings stns., 1972; Ocean Falls

(52°20'N,127°50’W), (1 specimen). Bousfield

stns., 1964; H1, H37, H55 (12 specimens). Van-

couver Island: Bousfield stns., 1975; Pl4a, P26

(2 specimens). Bousfield stns., 1976; B26, B27

(10 specimens). Bousfield stns., 1977; B1 (1 spe-

cimen). P. O’Rourke stns., 1979; French Creek

(49°20'N, 124°20'W),. FC1, FC2, FCIQ33Rwane

FC12 (15 specimens). Southern mainland: Bous-

field stns., 1977; P2 (1 specimen). Bousfield stns.,

1978; V1, \(1 specimen).

Distributional ecology: Geographic range:

British Columbia (52°20'N,127°50’W) to Costa

Rica (Barnard 1971). Bathymetric range: 9-200 m

(Barnard 1971). 0-73 m (NMNS coll.). Sediment

preference: fine sand.

Life cycle: Ovigerous females found in B.C.

waters in July (1), August (6). Pelagic males in

May (1), July (1), August (1).

Diagnosis: (Female). A small Ampelisca

(4-6 mm), characterized by: head, lower front

margin oblique nearly straight. Antenna | short

not reaching end of peduncle of antenna 2,

peduncular segments | and 2 subequal in length,

flagellum setae medium length. Antenna 2, about

body length, flagellum setae medium length.

Mandibular palp, segment 3 about */, length of

segment 2. Coxae 1-3, lower posterior corner

toothed. Gnathopod 1, stout, palm shallow and

elongate. Gnathopod 2, segment about ’/; length

of segment 5. Peraeopod 3, plumose setae on

distal half of posterior margin of segment 4.

Peraeopods 5-6, only single spine on posterior

margin of segment 5. Peraeopod 7, basal lobe

broad with lower posterior margin obliquely

rounded, segment 3 subequal to 4 in length, seg-

ment 4 produced into a tapering setose posterior

lobe, segment 6 inflated and only slightly longer

than 5, segment 7 very short. Uropod 1, almost

reaching end of uropod 2, peduncle and rami

subequal in length, outer ramus unarmed.

Uropod 2, rami shorter than peduncle, outer

margin of outer ramus spinulose, both margins

of inner ramus spinulose. Uropod 3, rami slender,

lanceolate, and sparsely setose at tips: Telson

lobes, broad rounded apices with two setae and a

small medial notch bearing a blunt spine, dorsal

surface with two pairs of spines along cleft. Pleon

Figure 8. Ampelisca hancocki J.L. Barnard. Diana Island, Vancouver Island, B.C. 9 5 mm, ov.

Departure Bay, Vancouver Island, B.C. & 6 mm.

sideplate 3, posterior margin nearly straight

ending in medium sized acute tooth. Urosomite 1,

weakly produced dorsally into a saddle shaped

carina. Gills flattened, dendritic folds short,

narrow, and curved downward.

(Male). Pelagic males (5.0-6.0 mm) differ from

females in: presence of brush setae on peduncles

of antenna | and 2, flagellum of antenna 1

elongate, extending beyond end of peduncle of

antenna 2, flagellum setae of both antennae very

short. Urosomite 1, dorsal surface notched

anteriorly and produced into a hoodlike carina

posteriorly. Uropod 3, rami bearing plumose

setae on all margins. Gills with long, straight

dendritic folds.

Remarks: Ampelisca hancocki is quite distinct

from other Northeast Pacific Ampelisca and is

easily identified by the straight posterior margin

of pleon sideplate 3, the short dactyl on peraeo-

pod 7, and the rounded apices of the telson lobes

(see key p. 3). A. hancocki is also the only

Ampelisca in the region which has an acute tooth

on pleon sideplate 3 but lacks a subapical spine

on uropod 2.

Ampelisca milleri J.L. Barnard

Figure 9.

Ampelisca milleri Barnard 1954a, pp. 9-11,

pls. 3-4

Material examined: California — Dillon Beach:

D. Markowitz coll. 16 April 1979 (30 specimens).

Distributional ecology: Geographic range —

Dillon Beach, California to Ecuador and the

Galapagos Islands (Barnard 1967a). Bathymetric

range: 0-187 m (Barnard 1971).

Life cycle: Ovigerous females found in Cali-

\) .

Figure 9. Ampelisca milleri J.L. Barnard. Dillon Beach, California. 9 6 mm, ov., & 6 mm

fornia waters in April (5). Pelagic males in

April (15).

Diagnosis: (Female). A small Ampelisca

(5-6 mm) characterized by: head, lower front

margin convex. Antenna | reaching well beyond

end of peduncle of antenna 2, peduncular seg-

ment 2 distinctly longer than segment 1, flagellum

setae long. Antenna 2 short, less than */, body

length, flagellum setae long. Mandibular palp,

segment 2 inflated and setose, segment 3 about

*/; length of segment 2. Coxae 1-3, lower posterior

corner slit. Gnathopod 1, palm elongate, seg-

ments 5 and 6 stout. Gnathopod 2, segment 5

less wthan~ twice *thesdength of. :segment 6.

Peraeopod 3, plumose setae confined to distal

third of posterior margin of segment 4. Peraeopod

5-6, bases broad, posterior margin of segment 5

bearing 2-3 single spines, posterodistal end of

segment 5 produced. Peraeopod 7, lower posterior

corner of basal lobe obliquely rounded, segment 3

longer than 4, segment 4 lacking posterior lobes

but bearing a single long plumose setae at lower

posterior corner, segment 6 broad and slightly

longer than 5, dactyl short and stout. Uropod 1

nearly reaching end of uropod 2, both rami

sparsely spinose. Uropod 2, rami shorter than

peduncle, medial margins of both rami unarmed.

Uropod 3, rami lanceolate and sparsely setose,

inner ramus much broader than outer ramus.

Telson lobes each tapering to a narrow, laterally

notched apex, each notch bearing a single spine,

dorsal surface of each lobe with 2-3 small spines

in a central row. Pleon sideplate 3, posterior

margin convex, ending in a small rounded process

at the lower corner. Urosomite 1 only slightly

elevated dorsally, without strong carina. Gills (in

mature females) flattened and unpleated.

(Male). Pelagic males differ from females in the

presence of brush setae on peduncles of antenna |

and 2, flagellum setae of both antenna very short.

Gnathopod 1, segment 6 narrowed distally so

that upper and lower margins are parallel for a

third of the segment. Urosomite 1, dorsal surface

notched anteriorly and produced posteriorly into

a hood-like carina. Uropod 3, rami more setose.

Gills less flattened and pleated.

Remarks: To date, A. milleri has been collected

only as far north as the San Francisco Bay

region (38°N). The species was treated here

since it represents a morphological type not

otherwise found in the northeast Pacific, and

pelagic males of A. milleri were previously

undescribed. The shape of peraeopod 7 dis-

tinguishes A. milleri from all other Ampelisca in

the area (see key p. 3).

ee aes ace Ae, 3 ‘

Ze Ul Jor i wid * Saal ee

<i ip Mie A

Zag aN

s VW | AN

Ampelisca pugetica Stimpson 1864

Figure 10.

Ampelisca californica Holmes 1908, pp. 513-

515; figs 23

Ampelisca gnathia Barnard 1954a, pp. 46-48,

pls. 33-34

Ampelisca pugetica Barnard 1954a, pp. 49-51,

pls. 35-36

Material examined: Southeastern Alaska:

Bousfield and McAllister stns., 1961; A117,

A150 (2 specimens).

British Columbia — Queen Charlotte Islands:

J.W. Scoggan stns., 1965; JWS 106 (1 specimen).

Bousfield stns., 1957; Ell, El4b, El4c, H2, H4a,

H5 (50 specimens). North central coast: Bousfield

stns., 1964; °H5, H13, 1125;)H50; 1953, S740

specimens). Vancouver Island: Bousfield stns.,

Figure 10. Ampelisca pugetica Stimpson. Alert Bay, Vancouver Island, B.C. 2? 9.0 mm, Los Angeles,

California, A. Hancock stn. 5737. & 9.0 mm

1959; N11, N22, V7, V10 (50 specimens). Bous-

field stns., 1970; P710b (30 specimens) Bousfield

stns., 1975; P17a (25 specimens). Bousfield stns.,

1976; Blla, B11b, B26 (12 specimens). Bousfield

stns., 1977; B1, B21b, E3 (6 specimens). K. Conlan

coll. 1975; Saanich Inlet (3 specimens). D. Ellis

coll; 1979; “Sidney -Channel. .Hero*: Strait

(2 specimens).

Southern California: Allan Hancock Foundation

sti.) 1958;°5737, 33°52 N, 118°3 PW, 34.5 ‘ims;

sand (13 specimens).

Distributional Ecology: Geographic range —

Prince William Sound, Alaska (60°28’N,

146°29’W) to Baja California (Barnard 1971]a).

Bathymetric range — 0-225 m (Barnard 1971a).

0-100 m (NMNS collections). Commonly less

than 40 m. Sediment preference — sand bottoms

(Barnard 1969b and NMNS coll.).

Life cycle: Ovigerous females found in B.C.

waters in February (1), July (1), August (2),

November (1). Mature males: none in NMNS coll.

Diagnosis: (Female). Medium sized Ampelisca

(9-11 mm) characterized by: head, lower front

margin oblique and convex. Antenna | short, not

reaching end of peduncle of antenna 2, pedun-

cular segment 2 twice the length of segment 1,

flagellum setae long. Antenna 2 about body

length, flagellum setae long. Mandibular palp,

segment 3 slightly more than half as long as

segment 2. Coxae 1-2, lower posterior corner

slit. Gnathopod 1, palm elongate and relatively

deep. Gnathopod 2, segment 6 about half as long

as segment 5. Peraeopods 3-4, stout. Peraeopod 3,

segment 4 with plumose setae only on distal third

of its posterior margin. Peraeopods 5-6, posterior

margin of segment 5 with 2-3 sets of comb spines.

Peraeopod 7, lower posterior margin of basal

lobe rounded, segment 3 subequal to 4 in length,

segment 4 with a broad setose posterior lobe

extending the length of segment 5, segment 5 with

a spine bearing notch on anterior margin, seg-

ment 6 inflated and longer than 5 and 7. Uropod |

short, reaching only halfway along rami of

uropod 2, outer ramus armed only with basal

spines. Uropod 2, rami shorter than peduncle,

both margins of both rami spinulose, outer ramus

with long spine at tip. Uropod 3, rami lanceolate,

medial margins of both rami setose. Pleon side-

plate 3, lower posterior corner bearing a medium-

sized tooth. Urosomite 1, dorsal carina saddle-

shaped. Telson, each lobe notched and pointed at

apex, notch with 2-5 setae, dorsal surface of each

lobe with 3-6 long spines originating centrally.

Gills in mature females flattened with a few pleats.

(Male 9.0 mm). A. Hancock stn. 5737. Pelagic

stage males differ from females in: presence of

brush setae on peduncles of antenna | and 2,

flagellum setae short. Gnathopod 1, segment 6

narrowing distally so that dorsal and ventral

margins are parallel for one-third of segment.

Uropods 1-2 more spinulose. Uropod 3 more

foliaceous. Urosomite 1 shallower and more

rounded dorsal carina. Gills less flattened but

more pleated than in females.

Remarks: This species is easily identified from

other northeastern Pacific Ampelisca by its

saddle-shaped urosome, the shape of peraeopod 7,

and the length of uropod | (see key p. 3).

The subspecies A. pugetica mora described by

Barnard (1967b) from bathyal depths is probably

a new species rather than a sub-species and

should be re-evaluated. The ‘‘macrodentata’”’

form of A. pugetica described by Barnard (1954a)

from Mexico and Central America was not

examined so its status cannot be evaluated here.

A. pugetica is represented by two photographs:

a @ (# AAA pug. 5) from the Burrard Inlet

region, and a @ (# AAA pug. 4) from the

Alexander Archipelago. Both specimens have a

yellowish brown ground colour with small areas

of red pigment scattered over the peraeopods and

uropods. The usual concentration of red pigment

is found in the eye region. White pigment

chromatophores are found peripheral to the eyes,

and scattered over the distal portions of the

anterior coxae and peraeopods.

Ampelisca eschrichti Kroyer 1842

Figure 11.

Ampelisca eschrichti Sars 1895, pp. 174-176,

pls'61, fig. 1:

Ampelisca eschrichti eschrichti Gurjanova 1955,

pp. 167-170, fig. 2.

Ampelisca eschrichti Lincoln 1979, p. 118, fig.

50f-}.

Material examined: Alaska — Bering Sea: St.

Lawrence Island (63°N,170°W), July 1980,

30-40 m., J. Oliver coll. (25 specimens).

Distributional ecology: Geographic range:

Circumpolar, through the North Atlantic and

Arctic oceans and into the Bering Sea (Stephen-

sen 1935, Dunbar 1954, Gurjanova 1955). Bathy-

metric range: 0-250 m (Mills 1971). Sediment

preference: sand to silt clay (Stephensen 1935).

Figure 11. Ampelisca eschrichti Kroyer. St. Lawrence Island, Bering Sea, Alaska. 2 25 mm, ov.

Life cycle: Ovigerous females found in the

Bering Sea in July.

Diagnosis: (Female). A very large Ampelisca

(20-30 mm) characterized by: head narrow

anteriorly, lower front margin oblique and

convex. Antenna 1 extending beyond peduncle

of antenna 2, peduncular segment 2 almost three

times length of segment 1, flagellum setae long.

Antenna 2 about half body length, flagellum setae

medium length. Mandibular palp, segment 3

about half the length of segment 2. Maxillipedal

palp, segment 3 produced distally well beyond

attachment of the dactyl. Coxae 1-2, lower

posterior corner toothed. Gnathopod 1, palm

elongate. Gnathopod 2, segment 5 twice length

of 6. Peraeopod 3, segment 4 with plumose setae

along distal */, of posterior margin. Peraeopods

5-6, bases broad, segment 5 with 4 rows of comb

spines along posterior margin. Peraeopod 7,

basal lobe deep with lower posterior margin

oblique nearly straight, segment 3 subequal to 4

in length, segment 4 with a tapering setose

posterior lobe, segment 5 with notch in anterior

margin, segment 6 long, slender and linear,

segment 7 medium length. Uropod | short, not

reaching end of uropod 2, outer ramus with basal

spines only. Uropod 2, outer ramus distinctly

shorter than inner ramus and bearing a large tip

spine, both margins of both rami spinulose.

Uropod 3, rami lanceolate and foliaceous.

Telson lobes tapering to rounded apices with a

single spine set in a small lateral notch, dorsal

surface with 3-4 spines scattered along the

central axis of each lobe. Pereonites 6-7 and

pleonites 1-3 elevated into a low thin keel along

mid-dorsal line. Pleon sideplate 2 slightly

produced at lower posterior corner. Pleon side-

plate 3 weakly convex posteriorly, ending in a

medium-sized acute tooth. Urosomite | produced

dorsally to form a massive carina that ends

subacutely above urosomite 2. Gills in mature

females flattened and unpleated.

(Male). No specimens of pelagic males are

present in NMNS collections. Marjorie Bousfield

of Dalhousie University kindly loaned me a

specimen from Nain Bay, Labrador. The pelagic

male differed from the female in: the presence of

brush setae on peduncles of antennae | and 2,

flagellum setae shorter than in female. Urosomite

1 dorsal surface notched anteriorly and produced

posteriorly into a hood-shaped carina. Uropods 1

and 2 more spinulose. Uropod 3 more setose.

Gills less flattened and strongly pleated.

Remarks: The deep water form of A. eschrichti

found by Barnard (1967a, 1971la) off California

and Oregon would seem to merit species recogni-

tion based on Barnard’s (1971a) illustrations. It

differed from the polar form particularly in the

shape of peraeopod 7. Gurjanova’s (1955) sub-

species A. eschrichti pacifica is also most likely a

separate species although it is difficult to evaluate

its status solely from the illustrations.

Ampelisca eschrichti is easily identified by its

distinctive peraeopod 7, telson, and urosomal

carma (see. key =p. 3):

Ampelisca macrocephala Liljeborg 1852

Figure *F2,

Ampelisca macrocephala G.O. Sars 1895, pp.

12-1735 pl, 60, Ties di:

‘Mills, 1967b, pp. 640-642, fig. 2.

‘Bousfield 1973, pp. 133-134, pl. 36, fig. 1

‘Lincoln 1979, p. 118, fig. 30a; 27

Material examined: Alaska — Bering Sea:

St. Lawrence Island (63°N,169°W), 40 m, July

1980 (30 specimens). Beaufort Sea: (70°43’N,

149°2’W), 50 m, 6 September 1971 (2 specimens).

Diagnosis: (Female). A large Ampelisca (15-24

Figure 12. Ampelisca macrocephala Liljeborg. St. Lawrence Island, Bering Sea, Alaska. 9 22 mm, ov.

mm) characterized by: head, upper front margin

produced slightly forward; lower front margin

oblique, slightly concave under lower eye con-

tinuing convexly to insertion of antenna 2. Eyes

present. Antenna | short just reaching end of

peduncle of antenna 2, peduncular segment 2

variable in length but less than twice segment 1,

flagellum setae short. Antenna 2 about half body

length, flagellum setae short. Mandibular palp,

segment 2 about twice length of segment 3.

Coxae 1-2, lower posterior corner toothed.

Coxa 3, tooth absent in Bering Sea and Beaufort

Sea populations, tooth present in Northwest

Atlantic populations. Gnathopod 1, palm short,

less than one third length of segment 6. Gnatho-

pod 2, segment 6 about half length of segment 5.

Peraeopod 3, plumose-setae confined to distal

half of posterior margin of segment 4. Peraeopods

5-6, posterior margin of segment 5 with 4-5 rows

of comb spines. Peraeopod 7, basal lobe deep,

lower posterior margin obliquely rounded,

segment 3 subequal to 4 in length, segment 4

with tapering, setose, posterior lobe which

extends about halfway down segment 5,

segment 6 much longer than 5, segment 7 sub-

equal to 5 in length. Uropod | reaching tip of

uropod 2, outer ramus with basal spines only.

Uropod 2, rami shorter than peduncle, outer

ramus slightly shorter than inner ramus with

long sub-apical spine, both margins of both rami

spinulose. Uropod 3, rami slender, lanceolate

and foliaceous. Telson lobes tapering to laterally

notched apices, notch bearing blunt spine and

short seta, apices in large specimens (20-25 mm)

bear 1-2 spines on distal lateral margins, dorsal

surface of each lobe with 5-6 short blunt spines in

central row, additional spines occasionally near

lateral margins. Pleon sideplate 3, posterior

margin convex, ending in a large acute tooth.

Urosomite | weakly elevated dorsally to form a

massive saddle-shaped carina. Gills in mature

females flattened and unpleated.

(Male). No specimens in NMNS collections.

Sars (1895) showed the pelagic male to differ in:

the presence of brush setae on the peduncles of

the antenna | and 2; antenna | longer than in

the female and extending beyond peduncle of

antenna 2. Urosomal carina more strongly

developed.

Distributional ecology: Geographic range:

Boreal North Atlantic through Arctic Ocean into

the Bering Sea (Mills 1967b). Bathymetric range:

10-280 m (Mills 1967b). Sediment preference:

sand bottoms (Dickinson et al. 1980).

Life cycle: It takes 18 months to reach breeding

age in fall of the second year. Males then die after

breeding. Females carry embryos (x = 60) 5

months over winter. Juveniles released in spring.

Most females then die (2 years old), but a few live

on and breed again to reach 3 years (Kanneworf

1965).

Remarks: This species is very similar to

Ampelisca careyi n.sp. (see phenogram fig. 21).

However, these two species may be reliably

separated by their armature patterns on the

dorsal surface of the telson and size at maturity.

Ampelisca careyi n.sp.

Figure 13

Ampelisca macrocephala Barnard 1954a, pp.

41-43, Fig. 29.

Material examined: Southeast Alaska

Alexander Archipelago: Bousfield stns., 1980;

S18F2, S18F3 (2 specimens).

British Columbia — Queen Charlotte: Bousfield

stns., 1957; H3 (20 specimens).

North Central Coast: Bousfield stns., 1964; H13,

HL?7, H27, H31,)H34,, N36, H37,.H49 (7S speer

mens). C. Levings, Swanson Bay stns., 1973; 28,

29 (4 specimens). C. Levings, Swanson Bay stns.,

1975; 53°00’N,128°30’W, 18 November 1975,

62 m; holotype ov. 9, 10 mm (NMC-C. 1981-381);

allotype mat. o&, 8 mm (NMC-C.1981-382),

692,4 cco, 8 imm. paratypes (NMC-C.1981-

383). Vancouver Island: Bousfield stns., 1970;

P717. (Inspecimen); (Bousfield., stnsz, 1975, PAO:

P2la, P22 (4 specimens). Bousfield stns., 1976;

B26, B28 (6 specimens). Bousfield stns., 1977;

B16, B18, B2la (4 specimens). Southern Main-

land: Bousfield stns., 1977; P3 (10 specimens).

Bousfield stns., 1978; V4 (20 specimens).

Oregon — OSU coll. 01248, 44°26.1N,124°14.5’W,

21June’ 1976;62. m(Qespecimens);

California — A. Hancock Found. 5270, 34°20’N,

119°53’W, 3 November 1957, 45 m, (7 specimens).

Distributional ecology: Geographic range —

Southeast Alaska (55°N) to Baja California

(Barnard 197la). Bathymetric range: 0-200 m

primarily shallower than 100 m (NMNS coll.).

Sediment preference: sand.

Life cycle: Ovigerous females found in B.C.

waters in the months of March (1), May (1),

June (1), July (30), August (2), November (16),

December (1). Pelagic males: June (2), July (1),

S\N

= N)

“Ng

Figure 13. Ampelisca careyi n.sp. Swanson Bay, B.C. Paratype @ 8 mm. Paratype 0 8 mm

August (1), November (2). This species must have

a different life cycle than that described by

Kanneworf (1965) for A. macrocephala since

ovigerous females are present throughout the

summer months.

Diagnosis: (Female). A medium sized Ampelisca

(7-12 mm) characterized by: head, produced

anteriorly into a dome-shaped process above

antenna 1, lower front margin distinctly concave.

Eyes present. Antenna | extending just beyond

end of peduncle of antenna 2, peduncular seg-

ment 2 more than twice length of segment 1,

flagellum setae short. Antenna 2 about ’/, body

length, flagellum setae short. Mandibular palp,

segment 2 about twice length of segment 3.

Coxae 1-3, lower posterior corner slit to form

small tooth. Gnathopod 1, palm short, extending

less than one-third the length of segment 6.

Gnathopod 2, segment 5 about twice length of

segment 6. Peraeopod 3, plumose setae confined

to distal half of posterior margin of segment 4.

Peraeopod 5-6, segment 5 with 2-3 sets of comb

spines on posterior margin. Peraeopod 7, basal

lobe deep with lower posterior margin obliquely

rounded, segment 3 subequal to 4 in length,

segment 4 with tapering setose posterior lobe

which extends about half the length of segment 5,

segment 6 much longer than 5, segment 7 sub-

equal to 5 in length. Uropod 1 reaching end of

uropod 2, outer ramus with basal spines only.

Uropod 2, rami shorter than peduncle, outer

ramus shorter than inner, with long sub-apical

spine, both margins of both rami spinulose.

Uropod 3, rami slender, lanceolate, and folia-

ceous. Pleon sideplate 3, posterior margin convex,

ending in short acute tooth. Urosomite | elevated

into a saddle-shaped carina ending acutely above

urosomite 2. Telson lobes tapering to laterally

notched apices, notches each armed with a spine

and setule, dorsal surface of each lobe bearing

3-4 scattered long slender spines. Gills flattened

and unpleated in mature females.

(Male). Pelagic males are characterized by:

presence of brush setae on peduncles of antenna |

and 2, flagellum setae shorter than in female;

urosomal carina large and hood-shaped; uropods

1-2 more spinulose; uropod 3 more setose; gills

less flattened and strongly pleated.

Remarks: A. careyi n.sp., 1s very similar to

both A. macrocephala and A. unsocalae (see

phenogram fig. 21). However, it is distinct from

A. macrocephala on the basis of its telson

armature. A. careyi and A. unsocalae differ in

head shape, length of tooth on pleon sideplate 3,

relative length of antenna 1, and the position of

the apical notch in the telson (see key p. 3 and

figs. 13-14).

A. careyi is represented by six photographs:

2 2 2 (HAAA car. 4, HAAA car. 9) from the Alex-

ander Archipelago region and 2 ob o& (HAAA car.

5, HAAA car. 8) and 2 9 9 (#AAA car. 6, fAAA

car. 7) from the Barkley Sound region. The

cuticular ground colour varies from translucent

to off-white to yellow brown among the speci-

mens. This species is relatively heavily pigmented.

In typical pattern, red pigment spots are found in

solid or reticulated masses interrupted along the

dorsal line and on all the appendages and

epimeral plates. The mid portion of the body

above the body-coxae junction line shows the

least pigmentation. The anterior portion of the

head shows the usual concentration of red pig-

ment around the eyes; the number of white

pigment chromatophores varies between speci-

mens, but the pigment is always concentrated in

a dorsoventral band between the frontal and

caudal eyes. Some of the specimens have banding

of white pigment on the distal segments of the

gnathopods and peraeopods 3 and 4, but in other

specimens the white pigment spots are scattered

or lacking on these appendages. The male speci-

mens have larger masses of red pigment associated

with the eyes than found in the females. Other

species of Ampelisca photographed in the Gulf of

Mexico region, A. verrilli (AAA ver. 7, #AAA

ver. 8) and A. agassizi (HAAA aga. 9, HAAA

aga. 10) show this same pattern suggesting that

this male-female difference in pigmentation of the

eyes may be typical in Ampelisca.

Etymology: This species is named in honour

of A.G. Carey of Oregon State University who

is an active researcher in marine benthic ecology

and kindly loaned a great deal of valuable

material from his collections of amphipods from

Oregon coastal waters.

Ampelisca unsocalae J.L. Barnard 1960 new status

Figure 14.

Ampelisca macrocephala unsocalae Barnard

1960, pp. 28-30, fig. 7.

Material examined: British Columbia — Queen

Charlotte Islands: J.W. Scoggan coll. 1965;

Dixon Entrance, stns., 104, 105 (90 specimens).

North Central Coast: NMNS coll. 1967; 51°23’N,

129°08’W, 175 m, 14 August 1967 (2 specimens).

Vancouver Island: NMNS coll., 48°21’,126°08’W,

September 1968 (1 specimen). Southern Main-

land: Bousfield stns., 1977; J2, P3, P4, P6, P7,

P8 (40 specimens). Bousfield stns., 1978; V5,

V6 (100 specimens).

Oregon — Oregon State University Benthic

Invertebrate coll. 01332, 44°34.2’N,124°52.6’W,

16 June 1963, 600 m, identified by J.L. Barnard

(1 specimen).

California — A. Hancock Foundation stn., 6915,

33°59’N,118°48’W, 13 March 1960, 250 m

(13 specimens).

Diagnosis: (Female). A medium-sized Ampelisca

(7-9 mm) characterized by: head, upper front

margin not produced forward, lower front margin

oblique and convex. Antenna | short, not

reaching end of peduncle of antenna 2, peduncular

segment 2 about twice length of segment 1,

flagellum setae short. Antenna 2 about body

length, flagellum setae short. Mandibular palp,

segment 3 about half length of segment 2. Coxae

1-3, small tooth on lower posterior corner.

Gnathopod | palm short, less than '/, of posterior

margin of segment 6. Gnathopod 2, segment 5

about twice length of segment 6. Peraeopod 3,

plumose setae along distal */, of posterior margin

of segment 4. Peraeopod 5-6, posterior margin of

segment 5 with 3 sets of comb spines. Peraeopod 7,

basal lobe deep, segment 3 subequal to 4 in

length, segment 4 with tapering setose posterior

lobe extending half the length of segment 5,

segments 5 and 7 subequal in length, segment 6

much longer than 5. Uropod | reaching end of

uropod 2, outer ramus with basal spines only.

Uropod 2, rami longer than peduncle, outer

ramus shorter than inner and with a long sub-

apical spine, both margins of both rami spinu-

lose. Uropod 3, rami slender, lanceolate and

foliaceous. Telson lobes tapering to a centrally

notched apex bearing a spine and setule, dorsal

surface of each lobe bearing 5 long scattered

ee ee

Figure 14. Ampelisca unsocalae J.L. Barnard. Dixon Entrance, Queen Charlotte Islands, B.C. 9

9 mm, ov.

spines. Pleon sideplate 3, posterior edge convex,

ending in a long slender tooth. Urosomite |

elevated into a carina which rises posteriorly to a

right angle above urosomite 2. Gills flattened

and unpleaied in mature females.

(Male 7.0 mm). The pelagic male is charac-

terized by: the; presence “of “brush setae’ .on

peduncles of antennae, the relative lengths of

antennae are unknown since the flagellae are

broken on available specimens; urosomite 1 is

produced dorsally into a hood-shaped carina;

uropod 3 is more setose.

Distributional ecology: Geographic range:

Queen Charlotte Islands to southern California

(Barnard 1971). Bathymetric range: 50-1700 m

(Barnard 1971). Primarily, 100-400 m (NMNS

coll.). Sediment preference: silt to silt-clay

bottoms.

Life cycle: Ovigerous females found in B.C.

waters during the months of May (1), July (8),

August (13), September (1), November (25).

Pelagic males: July (1), August (2).

Remarks: Barnard (1960) described this species

as a subspecies of A. macrocephala which lost its

eyes in deeper water (400 m). However, the

differences in morphology including head shape,

relative antennal lengths, shape of tooth of pleon

sideplate 3, and shape of apical notch of the

telson warrant full species recognition. The

majority of the specimens in the NMNS were

collected between 100-400 m and have eyes;

specimens collected from greater depths (600 m)

off Oregon lack eyes but seem otherwise identical.

Ampelisca unsocalae is represented by photo-

graphs of three @ specimens (HAAA uns. 3,

H#AAA uns. 4, #AAA uns. 5) from the Burrard

Inlet region. All three specimens show a basically

transparent body with faint traces of red pigment

particularly in the buccal mass and on the bases

of peraeopods 5-7. The anterior portion of the

head has the usual concentration of red pigment

near the eyes with white pigment granules

concentrated between the frontal and caudal

eyes. The distal edges of coxae 1-2 and the distal

segments of the gnathopods and peraeopods 3-4

also have scattered white pigment chromato-

phores.

Ampelisca cristata Holmes

Figure 15.

Ampelisca cristata Holmes 1908, pp. 507-508,

figs. 16-17.

Barnard 1954a, pp. 26-28, pls. 17a-d, 18a, c,

My f;)2.

Material examined: Southeast Alaska —

Alexander Archipelago: Bousfield and McAllister

stns., 1961; A169 (1 specimen).

British Columbia — North Central Coast:

Bousfield stns., 1964; H52 (2 specimens). C.

See Wy

La Ve

AL; ff / |

Levings stns., Swanson Bay, 1973; (51B-031),

1975; (S1B-013) (2 specimens); Vancouver Island:

Bousfield stns., 1977; B13 (1 specimen).

Southern Mainland Coast: Bousfield stns., 1955;

M1 (1 specimen). Bousfield stn., 1959; N22

(1 specimen).

Oregon: Oregon State University Benthic Invert.

coll. 01257, 44°10’N,124°14.7°W, 24 June 1976

(1 specimen).

Distributional ecology: Geographic range —

Alexander Archipelago (56°16’N,135°30’W) to

Costa Rica (Barnard 1954a). Bathymetric range:

0-152 m (Barnard 1971a), 0-70 m (NMNS coll.).

Sediment preference: coarse sand.

Life cycle: No ovigerous females or pelagic

males in NMNS collections.

Diagnosis: (Female). A large Ampelisca (12-17

mm) characterized by: head long, narrowing

anteriorly, lower front margin parallel to upper

Figure 15. Ampelisca cristata Holmes. Swanson Bay, B.C. @ 10 mm, br. I

margin. Antenna | short, not reaching end of

peduncle of antenna 2, peduncular segments 1-2

subequal in length, flagellum setae medium

length. Antenna 2 about % body length, flagellum

setae medium length. Mandibular palp, segment

3 about ’/, length of segment 2. Maxilla 1, seg-

ment 2 of palp with setae along distal portion of

outer margin. Coxa 1, small tooth at lower

posterior corner. Gnathopod 1, palm shallow

and elongate. Gnathopod 2, segment 5 more than

twice length of segment 6. Peraeopod 3, segment

4 with plumose setae along distal */, of posterior

margin. Peraeopods 5-6, segment 5 with 3-4 sets

of comb spines along posterior margin.

Peraeopod 7, lower posterior margin of basal

lobe oblique nearly straight, segment 3 shorter

than 4, segment 4 with small slender setose

posterior lobe, segment 6 inflated and longer

than 5 and 7. Uropod 1, nearly reaching end of

uropod 2, middle portion of upper margin of

peduncle elevated into a lamellar carina, outer

ramus with basal spines only. Uropod 2, rami

shorter than peduncle, outer ramus with long

tip spine, both margins of both rami spinulose.

Uropod 3, rami broadly lanceolate and foliaceous.

Telson lobes, notched laterally and pointed

medially at tip, notches bearing 3-4 spines and

1 setae, dorsal surface with 2-4 pairs of medium

length spines orginating centrally. Pleon side-

plates 2-3, tooth at lower posterior corner, dorsal

surface of pleonite 3 elevated into two low

longitudinal crests. Urosomite 1, dorsal suface

elevated into lamellar carina. Gills flattened in

mature females with a few indistinct pleats.

(Male). Unknown.

Remarks: Ampelisca cristata is easily distin-

guished from other Northeast Pacific ampeliscids

by its head shape and urosomal carina (see key

The status of Barnard’s (1954b) records from

the Caribbean and form “microdentata”’ (Barnard

1954a) could not be evaluted from his limited

illustrations.

A single photograph of A. cristata (HAAA

cri. 4) shows the colour patterns ina 9 specimen

from the Barkley Sound region. The body is

basically translucent, but small amounts of red

pigment dot the dorsal surface of the thorax, the

antennae, peraeopods, and uropods. The anterior

portion of the head is heavily pigmented in red

and white. The red pigment is concentrated

around the eyes in the usual manner. The white

pigment is concentrated in a band between the

frontal eyes, but scattered pigment spots occur

over the entire anterior half of the head and distal

portions of coxae 1 and 2.

Ampelisca brevisimulata J.L. Barnard

Figure 16.

Ampelisca brevisimulata Barnard 1954a,

pp. 33-35, pls. 23-24.

Material examined: Alaska — Bousfield and

McAllister stns., 1961; A42, Lynn Canal,

58°31’N,134°50’W (1 specimen). N.A. Powell

coll. 1969; Izembek Lagoon, 55°20’N,162°55’W

(1 specimen).

British Columbia — Queen Charlotte Islands:

J.W. Scoggan coll. 1965; 104, 105 (3 specimens).

North Central Coast: Bousfield stns., 1964; H34

(1 specimen). C. Levings stns., Swanson Bay,

1973; (1 specimen), 1975; (8 specimens). Van-

couver Island: D. Ellis coll., 1965; Parry Bay,

48°23’N, 123°30’'W (1 specimen). Bousfield stns.,

1975; P13 (1 specimen). Bousfield stns., 1976;

Bl0a (1 specimen). NMNS coll., Numukamis

Bay, 48°55’N,125°00’W (1 specimen). Southern

Mainland: Bousfield stns., 1977; El (1 specimen).

Bousfield stns., 1978; V6 (2 specimens).

Washington — Lopez Island: Upright Head, R.I.

Smith coll. 1955; (1 specimen).

Oregon — Oregon State University Benthic

Invertebrate coll. 43°49’N,124°S50’W, 417 m,

28 June 1976; 46°06’N, 124°13’W, 88 m, 1 October

1974 (3 specimens).

Distributional ecology: Geographic range —

Southeast Alaska (58°30’N,134°50’W) to Panama

(Barnard 1971). Barnard (1954b) also reported it

from the Caribbean Sea, but he did not illustrate

it from that region. Bathymetric range: 20-400 m

(NMNS coll.). Sediment preference: sand to

silt clay.

Life cycle: Ovigerous females found in B.C.

waters during the months of January (1), May (1),

November (1). Pelagic males: November (2).

Diagnosis: (Female). Medium-sized Ampelisca

(7-9 mm) characterized by: head, lower front

margin concave almost parallel to upper margin.

Antenna | short, not reaching end of peduncle of

antenna 2, peduncular segment 2 more than twice

length of segment 1, flagellum setae short.

Antenna 2 about ’/, body length, flagellum

setae short. Mandibular palp, segment 3 about

*/,; length of segment 2. Coxae 1, small tooth at

lower posterior corner. Gnathopod 1, palm

shallow and elongate. Gnathopod 2, segment 5

more than twice length of segment 6. Peraeopod 3,

Figure 16. Ampelisca brevisimulata J.L. Barnard. Cape Beale, Vancouver Island, B.C. 9 7 mm, ov.;

Burrard Inlet, B.C., & 6 mm

plumose setae restricted to distal third of

posterior margin of segment 4. Peraeopods 5-6,

segment 5 with 4-5 sets of comb spines on

posterior margin. Peraeopod 7, lower posterior

margin of basal lobe rounded, segment 3 sub-

equal to 4 in length, segment 4 bearing a slender

setose posterior lobe, segments 5 and 6 inflated,

segment 6 relatively short just slightly longer than

5, segment 7 slender and subequal to 6 in length.

Uropod | long, extending beyond end of uropod

2, both rami spinulose. Uropod 2, rami subequal

to peduncle in length, outer ramus with tip spine,

inner ramus unarmed along medial margin.

Uropod 3, rami broad, lanceolate and foliaceous.

Telson lobes tapering to narrow apices, each

bearing 1-2 setae, dorsal surface bearing 6-12

scattered setae. Pleon sideplate 2, small acute

tooth at lower posterior corner. Pleon sideplate

3, large tooth at lower posterior corner, produced

ps |

into a large, rounded hump above the tooth.

Urosomite 1 slightly produced dorsally, ending

acutely above urosomite 2. Gills flattened and

unpleated in mature females.

(Male). Pelagic males differ from females in:

the presence of brush setae on peduncles of both

antenna; antenna | longer than in female, extend-

ing beyond peduncle of antenna 2; flagellum

setae of both antenna very short. Pleon side-

plate 3, tooth smaller, not as long as process

above it. Urosomite | produced more dorsally

into a hood-shaped carina. Uropods 1-2 slightly

more spinose. Gills moderately pleated and

less flattened.

Remarks: Mature specimens of A. brevisimu-

lata are easily distinguished from other Northeast

Pacific Ampelisca by the shape of the head, pleon

sideplate 3 and the telson (see key p. 3). Juveniles

of A. brevisimulata could be confused with other

species in the A. macrocephala group, but it seems

recognizable at all sizes by its head shape. Based

on its telson, uropod 3, and peraeopod 7,

A. brevisimulata shows a strong similarity to

Atlantic species such as A. verrilli, A. holmesi and

A. brevicornis.

Discussion

The coxal gills of Ampeliscidae are rarely

illustrated and seldom described in taxonomic

treatments of the group. Although Sars (1895)

illustrated a representative gill of one species for

each of his three Norwegian genera, Ampelisca,

Byblis, and Haploops, subsequent major works

on the Ampeliscidae such as Reid (1951),

Barnard (1954a), Karaman (1975) and Bellan-

DS Yj)

“es

Aik onl

CX2

Santini and Kaim Malka (1977) make little or

no mention of gill structure. However, Mills

(1963, 1964, 1967b) described and illustrated

the gills of some species from the western

Atlantic. The present study revealed gill structure

varied among species of Ampelisca, and that

sexual dimorphism typified all species and was

rather striking in most.

The species of Ampelisca from the northeast

Pacific could be divided into two groups on the

basis of gill structure. The first type of gill

resembles an upside-down conifer tree when

viewed in longitudinal section and will be

referred to as the “‘dendritic”’ type of gill in this

paper (see Fig. 17). The surface\ of the veul

lamellae of this type has a series of folds perpen-

dicular to the long axis of the gill. In longitudinal

long-sec

CX6

Figure 17. Coxal gills of Ampelisca agassizi from coxae 2, 4,6 for 2 11 mm, ov., and o 7mm, subadult

section, these folds can be seen to originate oppo-

site each other on either side of the gill. Internally,

the tissue of the gill folds is reticulated with tiny

anastomosing spaces presumably to maximize the

surface area for respiratory exchange. The gills of

mature females are longer and more slender, with

shorter and narrower folds than in males and

juvenile females. The folds of the female gill curve

downward so the folds overlap against the axis of

the gill. Thus, female gills tend to appear much

thinner in longitudinal section than in the male

where the folds are nearly perpendicular to the

gill axis for their entire length. When viewed

laterally (as attached to the coxae), the gills of

the male often appear twisted on their peduncles

so that the longitudinal rather than facial view is

presented. Gill form changes little from coxae 2-6,

but the size varies somewhat with coxal gill 6

usually the smallest. This type of gill structure is

found in A. agassizi, A. fageri, A. hancocki, and

A. lobata (Fig. 18). Within this group, A. Jobata

has more and relatively longer gill folds than the

other three. In A. agassizi, females have relatively

longer and more slender gills than those of the

other three species. The small size of A. hancocki

HANCOCK|

FAGERI

CX2

and the scarcity of specimens of A. fageri rendered

comparisons difficult with the other two species

and were not done here.

The second type of gill is referred to as the

““‘pleated”’ type. Males in this species group have

gills with a series of shallow, relatively broad

folds or pleats which form a venetian blind-like

structure in a facial view (see Fig. 19). The male

gill is broad at the base and tapers distally to a

rounded apex. The pleats are reticulated with tiny

anastomosing spaces. The outer margin of the gill

is translucent. In cross section, the pleats of the

male gill can be seen to orginate in alternate

fashion on the two faces of the gill, and that the

gill tapers distally. Mature females of this group

have gills which are longer, narrower, thinner,

and usually unpleated or occasionally weakly

pleated. As in the male, the gills of females

have a translucent outer margin with a central

area of reticulated respiratory tissue. In form, the

gills show little change from coxae 2-6, but in

size coxal gill 4 is longest and coxal gill 6 is

shortest (Fig. 19). This type of gill structure is

found in A. birulai, A. brevisimulata, A. careyi,

A. cristata, A. eschrichti, A. hessleri, A. macro-

Figure 18. Representative coxal gills of Ampelisca fageri, 9 8 mm, br. II, Ampelisca hancocki, 2 5 mm,

Ov., o& 4 mm, subadult, and Ampelisca lobata, 2 7 mm, ov., & 6 mm, subadult

CX2

CX6

long-sec

Cx6

Figure 19. Coxal gills of Ampelisca macrocephala from coxae 2-6 for 2 20 mm, ov., and o& 20 mm,

subadult.

cephala, A. milleri, A. plumosa, A. pugetica and

A. unsocalae. Within this species group, the gill

differs little in form, but the number of gill pleats

in males varies considerably from 14 pleats in

A. pugetica to 4 pleats in A. eschrichti. The

number of pleats seems to be independent of

species size. Pleat number did vary with maturity.

Thus, the pleat numbers shown in Figs. 19-20

cannot be regarded as diagnostic since terminal

male instar specimens were not always available.

From this study, it was concluded that the

“dendritic” gill is plesiomorphic and the “‘pleated”’

gill is apomorphic. The two sources of evidence

for this conclusion are: 1) both types of gills show

a reduction in the size and number of pleats in

the development of the female, while the pelagic

male stage, which is usually considered conserva-

tive, retains the juvenile gill form; 2) the “‘dendri-

tic’ species group is characterized as plesio-

morphic on the basis of other characters (see

Table 2 and Fig. 21). However, this inference

should be re-evaluated when the gills of a broader

representation of Ampelisca are known. Further

insight will be gained by study of the gills of

related genera (Byblis, Haploops) and related

superfamilies (Dexaminoidea). Since the pattern

of sexual dimorphism in Ampelisca is for a flatter,

less space-consuming gill in mature females, it

suggests that this is an adaptation to allow larger

broods without a corresponding increase in body

size. However, this suggestion should be re-

evaluated when more is known of the activity

patterns and respiratory physiology of brooding

females.

In an effort to elucidate subgeneric groupings

among the species of Ampelisca from our study

area, a cluster analysis was performed using the

24 characters listed in Table 1. Each character

was defined in such a way that every species

could be classified into one of 2 to 4 character

states depending on the character. This coding

process resulted in the data matrix of species

versus characters shown in Table 2. Using the

data matrix, a similarity matrix was generated

using the simple matching coefficient (see Sneath

and Sokal 1973, p. 132). The similarity matrix

gives an index of morphological resemblance

for each possible species pair. From the similarity

CAREY!

hy, ;

CRISTATA

HESSLERI MILLERI

CX2

PUGETICA

Figure 20. Representative coxal gills of Ampelisca brevisimulata, 9 7 mm, ov, & 6 mm, subadult;

Ampelisca careyi 9 8 mm, ov, & 7 mm, subadult; Ampelisca cristata 9 17 mm, ov, & 12 mm, sub-

adult; Ampelisca eschrichti 9 20 mm, ov. & 12 mm, subadult; Ampelisca hessleri 2 7mm ov, & 6mm,

subadult; Ampelisca milleri 9 6 mm, ov, & 6 mm, adult; Ampelisca pugetica 9 10 mm, ov, & 7 mm,

subadult; Ampelisca unsocalae 9 8 mm, ov, & 7 mm, subadult.

matrix, a phenogram was generated using the

method of complete linkage clustering (see Sneath

and Sokal 1973, p. 222). Although these tech-

niques are among the simplest methods of

classification analysis, they seemed sufficient for

the level of discrimination desired and had the

advantage of being easily performed without the

use of a large computer.

In analysing the phenogram (Fig. 21), it should

be kept in mind that the relationships shown in it

reflect only the morphological resemblance of

these species and inference of phyletic relation-

ships from it should be done with caution. The

interpretation of a cluster analysis can be com-

plicated by two factors not considered in the

methodology: convergence and negative matches.

Either of these factors can give two species a

higher phenetic resemblance than appropriate for

their phyletic relationship. However, despite

these weaknesses in the methodology, some useful

insights may be gained from a careful examina-

tion of the phenogram.

PLUMOSA 9.)

BIRULAI 8.5

HESSLERI 7.9

AGASSIZI 7.9

LOBATA 6.9

FAGERI 9.5

HANCOCKI 9.0

MILLERI 9.5

ESCHRICHTI 15.5

PUGETICA 17.0

MACROCEPHALA 14.5

UNSOCALAE 16.5

CAREYI 17.9

CRISTATA 16.9

BREVISIMULATA 15.9

Figure 21. Phenogram showing morphological similarity of species of Ampelisca from Northeast Pacific

region. Number beside each species is index of apomorphy. Scale across the top is the measure of

similarity.

As a further aid in this analysis, each of the

character states was designated as plesiomorphic

(0), intermediate (.5) or apomorphic (1). Using

these rankings, an index of apomorphy was

calculated for each species (see Table 2). A species

which was advanced in all characters would have

an index of 24; whereas a species which was

primitive in all characters would have an index of

zero. Although this technique is rather subjective

and the scoring of certain characters was

questionable, the general trends shown by this

method seem reasonable. The index for each

species is listed on its line in the phenogram (Fig.

PAY,

The phenogram shows 3 distinct clusters of

species: 1) A. plumosa, A. birulai, A. hessleri, and

A. agassizi; 2) A. lobata, A. fageri, A. hancocki and

A. milleri; 3) A. eschrichti, A. pugetica, A. macro-

cephala, A. unsocalae, A. careyi, A. cristata and

A. brevisimulata. The first of these clusters

contains 4 species which have very similar

urosomal carinas and telsons. Three of the species

(A. plumosa, A. birulai, and A. hessleri) are so

similar in their morphologies that any suggestion

of convergence can be dismissed reasonably. The

relationship of A. agassizi to the other three

species must be rather distant since they differ in

so many characters (see Table 2) including the

structure of their gills, presence of coxal teeth,

shape of basal lobe of peraeopod 7, presence of

comb spines on peraeopods 5-6, relative lengths

of mandibular palp segments, and structure of

uropod 3. All four of the species in the A. plumosa

cluster are relatively plesiomorphic with indices

between 7 and 9.

The second cluster contains a_ rather

anomalous group of plesiomorphic species

which probably have little phyletic relation-

ship. A. fageri and A. lJobata share enough

Smaracters in the structure of their gills,

mandibular palp, urosomal carina, uropod 3,

and telson to indicate a fairly close phyletic

relationship despite the large differences in

their peraeopod 7. The addition of A. hancocki

and A. milleri to this pair seems. artificial

despite the near equivalence of their apo-

morphic indices. Their similarity is mainly due

to their generalized structure which results in

negative matches (see Table 2) and perhaps

some convergence due to their small size. In

any case, the differences in the mouthparts,

peraeopod 7, and telson seem sufficient to

rule out any close relationship. A. milleri

seems distinct from other North Pacific

species, and its closest relatives are North

Atlantic species such as A. abdita Mills 1963,

and A. vadorum Mills 1964, which are

extremely similar in their morphology (Mills

1965). A. hancocki is difficult to group. It is

clearly closely related to A. shoemakeri des-

cribed by Barnard (1954a) from the Pacific

coast of central America. The presence of a

tooth on pleon sideplate 3 and the shape of

the urosomal carina suggest that it may be

distantly related to the more apomorphic

species in cluster 3, but it clearly has no close

relatives in the British Columbia region.

The third cluster contains a group of

apomorphic species all of which have a tooth on

pleon sideplate 3, an apical spine on uropod 2,

elongated segment 5 on gnathopod 2, comb

spines on peraeopods 5-6, and lanceolate third

uropod. The three subgroupings within this

cluster also seem to reflect probable phyletic

relationships. A. pugetica and A. eschrichti both

have an elongated uropod 2, and an anterior

notch in segment 5 of peraeopod 7. This combina-

tion of characters is so distinctive as to suggest

a close phyletic relationship. A. coeca Holmes

1908, a bathyal species off California, is also

clearly related to this species pair. A. macro-

cephala, A. careyi and A. unsocalae were previously

considered one species and are clearly closely

related. A. cristata and A. brevisimulata have

similarities in their head shapes, coxal teeth, and

peraeopod 7 which suggest some degree of

phyletic relationship. However, A. brevisimulata

is much more similar to the North Atlantic

species A. verilli Mills 1967 and A. holmesi Pearce

1908, and A. cristata is a closer relative of the

tropical Pacific species A. cristoides Barnard 1954a.

Although the cluster analysis has been useful in

pointing out possible phyletic relationships

among the Northeast Pacific Ampelisca, it has

also underlined the difficulty in trying to sub-

divide this large genus. Almost any of the twenty-

four characters used in the analysis would give a

unique set of clusters of the 15 species if used

alone. This mosaic pattern of morphologies found

in Ampelisca suggests a high degree of conver-

gence in the group. It seems inadvisable to

attempt any formal subdivisions of the genus

until the morphologies of the world fauna are

better documented, and even more importantly, a

better understanding of their functional morpho-

logy is achieved.

The zoogeography of the 14 species of

Ampelisca found in the Northeast Pacific region

is summarized in Table 3. Three of the species

(A. birulai, A. eschrichti and A. macrocephala) are

arctic forms not found south of the Bering Sea.

Nine species (A. agassizi, A. brevisimulata, A.

careyi, A. cristata, A. fageri, A. hancocki,

A. lobata, A. pugetica and A. unsocalae have

broad geographic ranges extending from the

Queen Charlotte Islands to Mexico. This pattern

would suggest that their habitat requirements

and ecological tolerances must be relatively

broad, but it also reflects the continuity of the

cold temperate subtidal sand habitat along the

Pacific coast of North America. The new species

A. hessleri has been found only in northern

British Columbia waters to date. Two species

(A. milleri and A. plumosa are not known north

of central California.

Acknowledgements

This study has been made possible by a post-

doctoral fellowship from the Natural Sciences

and Engineering Research Council of Canada

which allowed the author to work in the National

Museum of Natural Sciences in Ottawa. I would

like to thank the museum staff who provided me

with all the necessary facilities to carry out this

work. I would especially like to thank Dr. E.L.

Bousfield who gave generously of his time in

offering continuous advice, stimulating discus-

sion and critical insight throughout the duration

of this study. Dr. Eric Mills improved the manu-

script by his helpful review. I would also like to

thank Kathleen Conlan, Norma Jarrett, Rama

Chengalath, Diana Laubitz, Craig Staude,

Brenda Hann and Marjorie Bousfield for helpful

discussions on particular aspects of this work.

The line illustrations were prepared with the

assistance of artists Floy E. Zittin and Susan

Bourque. The manuscript was typed by Joyce

Tynan-Byrd.

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West Indies. Am. Mus. Novit. 598: 1-24

1955. Amphipoda collected at the Arctic Laboratory, Office

of Naval Research, Point Barrow, Alaska, by G.E. Mac-

Ginitie. Smithson. Misc. Collct. 128: 1-78

Sneath, P.H.A. and R.R. Sokal, 1973. Numerical taxonomy.

W.H. Freeman, San Francisco, 573 pp.

Stephensen, K. 1935. The Amphipoda of north Norway and

Spitsbergen with adjacent waters. Tromsé Mus. Skr. Vol. III.,

140 pp.

*Stimpson, W. 1864. Descriptions of new species of marine

invertebrates from Puget Sound, collected by the naturalist of

the Northwest Boundary Commission, A.H. Campbell, Esq.,

Commissioner. Proc. Acad. Nat. Sci., Philadelphia. 16: 153-165

*Stout, V.R. 1913. Studies in Laguna Amphipoda. Zool. Abbreviations For Figures

Jahrb. Abt. Syst. 34: 633-659

Legend for all figures

Hd head UL upper lip

Gn gnathopod EE lower lip

P peraeopod Md mandible

Cx coxal plate Lft left

U uropod Rt right

r telson Mx maxilla

Mxpd maxilliped

Table 1. List of Characters used in Cluster Analysis

Relative length of antenna | to peduncle of

antenna 2

shorter (+), equal (0), longer (-)

Shape of lower head margin

concave (+), straight (0), convex (-)

Number of coxal teeth 0, 1, 2, 3

Relative length of mandibular palp segment 3 to

segment 2

mete {-),.< .1> (+)

Mandibular palp segment 2 inflated (+), not

inflated (-)

Length of palm of gnathopod |

elongate (0), simple (-), short (+)

Relative length of segment 5 to segment 6 on

gnathopod 2 ;

=> .50 (+), < .50 (-)

. Setation on posterior margin of segment 4 of

peraeopod 3

entire (-), '/, (0), '4% (+)

Comb spines on posterior margin of segment of

peraeopods 5-6

well developed (+), reduced (0), absent (-)

Notch on anterior surface of segment 5 of

peraeopod 7

present (+), absent (-)

. Shape of basal lobe of peraeopod 7

D shaped (-), “triangular” (+)

a7]

24.

. Urosomal carina

Length of segment 6 of peraeopod 7

elongated (+), normal (-)

Relative length of segment 3 to segment 4 on

peraeopod 7

longer (+), subequal (-)

. Pleon sideplate 2

with tooth (+), without tooth (-)

Pleon sideplate 3

with acute tooth (+), without tooth (-)

Gills pleated (+), dendritic (-)

lamellar (+), massive (0),

absent or reduced (-)

. Uropod | with outer ramus

spinulose (+), unarmed (-)

. Uropod 1 reaching end of uropod 2 (-),

only reaching halfway along rami (+)

. Outer ramus of uropod 2 with apical spine (+),

without apical spine (-)

. Rami of uropod 3

long and lanceolate (-), short and flattened (+)

Apex of telson lobes

notched (+), narrowed (0), blunt (-)

Relative length of peduncular segment 2 to seg-

ment | of antenna |

SDA es 2m. f=)

Dorsal spines of telson

scattered (+), central rows (0), along cleft (-)

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Table 3. Geographical distribution of Northeast Pacific Ampelisca

Cross Sd.

to Dixon’ Northern Central

Arctic Entrance B.C. & B.C. &

Ocean & Northern Prince & Queen’ Northern Southern Washington California

Beaufort Bering William Charlotte Vancouver Vancouver & &

Species Sea Sea Sound Islands Island Island Oregon Mexico

Ampelisca

birulai xX X

Ampelisca

macrocephala »4

Ampelisca

eschrichti xX X

Ampelisca

pugetica Xx

Ampelisca

agassiZzi

Ampelisca

careyi

Ampelisca

cristata

Ampelisca

brevisimulata

Ampelisca

lobata

Ampelisca

hessleri

Ampelisca

unsocalae

Ampelisca

hancocki

Ampelisca

fageri x ?

Ampelisca

milleri

Ampelisca

plumosa

oo oo

EE 9 og

~ «~ KM Mm MO

~ ~*~ KM KM

et oo

~~ Ke Kh KM OM

3°

S - a

all il = _

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. ,

The Amphipod Superfamily Corophioidea in the

Northeastern Pacific Region. Family Ampithoidae:

Systematics and Distributional Ecology

K.E. Conlan and E.L. Bousfield

National Museum of Natural Sciences

National Museums of Canada

Ottawa Canada

ABSTRACT

The gammaridean family Ampithoidae of the boreal eastern Pacific coastal region (Alaska to Pt.

Conception, California) is examined on a taxonomic, biogeographical and ecological basis. Keyed and

described are fifteen species within the. genera Cymadusa Savigny, Ampithoe (Leach, sens. str.) and

Peramphithoe n.gen. This latter genus is erected for all species of Ampithoe bearing a transverse first

gnathopod. Ampithoe dalli Shoemaker is reinstated as a species distinct from A. simulans Alderman.

Ampithoe sectimanus n. sp., earlier confused as a variant of A. po/lex Kunkel and A. simulans Alderman,

is recognized as a distinct new species. The taxonomic status of Peramphithoe mea (Gurjanova), P. tea

(Barnard), P. plea (Barnard) and P. annenkovae (Gurjanova) is clarified. For comparison, two anti-boreal

eastern Pacific species, Peramphithoe lessoniophila n.sp. and P. femorata (Kréyer) are described. Thirteen

of the fifteen North Pacific species are endemic to the boreal region and eight of these are restricted to the

North American coast. Greatest diversity occurs on the central and Vancouver Island coasts (12 species)

of British Columbia, pointing to this region as being the centre of distribution in the North-eastern

Pacific.

RESUME

La famille de gammares Ampithoidae de la partie septentrionale de la céte orientale du Pacifique (de

P Alaska a la Californie — Point Conception) a fait l’objet d’une étude taxinomique, biogéographique et

écologique. L’auteur fournit des clés et des descriptions de quinze espéces des genres Cymadusa Savigny,

Ampithoe (Leach, sens. str.) et Peramphithoe n. gen. Ce dernier regroupe toutes les espéces d’Ampithoe qui

ont un premier gnathopode transversal. Ampithoe dalli Shoemaker reprend le rang d’espéce, distincte

d’A. simulans Alderman. Ampithoe sectimanus n. sp. antérieurement considérée comme une variante

d’A. pollex Kunkel et d’A. simulans Alderman est maintenant reconnue comme une nouvelle espéce

distincte. On fournit des éclaircissements sur le statut taxinomique de Peramphithoe mea (Gurjanova),

P. tea (Barnard), P. plea (Barnard) et P. annenkovae (Gurjanova). A des fins de comparaison, la descrip-

tion de deux espéces anti-boréales, Peramphithoe lessoniophila n. sp. et P. femorata (Kréyer), apparait

dans ce travail. Parmi les espéces du Pacifique-nord, treize sur un total de quinze sont endémiques a la

région boréale, tandis que huit d’entre elles sont restreintes a la cote de l’Amérique du Nord. La diversité

la plus grande se rencontre en Colombie-britannique au centre de la céte et sur la céte de Vile de

Vancouver, ce qui indique que cette région est le centre de la distribution dans le nord-est du Pacifique.

Introduction

Members of the corophioidean family Ampi- The Ampithoidae is a primarily warm water

thoidae inhabit self-constructed tubes in coastal family but species extend into the boreal regions

marine algae. The family was first described by of both the Atlantic and Pacific. This paper

Stebbing (1888), individuals being distinctive in examines the Ampithoidae of the boreal and anti-

the incision of the outer lobes of the lower lip and boreal eastern Pacific. To date, the boreal Pacific

the possession of shortened pad-like rami on the Ampithoidae has been studied in California and

third uropod, the outer ramus of which bears one Oregon by Barnard (1954, 1964, 1965, 1969b,

or two large reverted uncini. 1970), on the Pacific coast of the USSR by

Gurjanova (1938, 1955) and Kudryashov (1979),

and in Japan by Nagata (1960). Only 2 of the

11 known genera have boreal representatives

(Cymadusa and Ampithoe). The intent of this

study is to extend our knowledge of the boreal

northeastern Ampithoidae from the southern-

most limits of the region (Point Conception,

California) to the northernmost limits in Alaska.

In addition, information on a new anti-boreal

species from South America is given.

The specimens from which information was

compiled were collected mainly by National

Museum survey expeditions. Station data for the

expeditions of 1955 (Vancouver Island and

Georgia Strait, B.C.), 1957 (Queen Charlotte

Islands, B.C.), 1959 (northern Vancouver Island),

1961 (southern Alaska coast) and 1964 (northern

and central British Columbia coast) are published

in Bousfield (1957 and 1963), Bousfield and

McAllister (1962), and Bousfield (1968), respec-

tively. Station data for 1966 (Washington

and Oregon), 1970, 1975 and 1977 (Vancouver

Island and southern British Columbia mainland

coast) and 1980 (southern Alaska coast) are

presented in Bousfield and Jarrett (1981).

In order to facilitate identification, the keys

and descriptions are constructed with sex and

age independent characters, except where stated.

Mature males are recognizable by their relatively

large size for the species, enlarged and usually

differentiated second gnathopod, and by the

presence of paired penial papillae on the sternum

of peraeon 7. Mature females are recognizable by

their relatively large size and the presence of

setose brood plates attached to the inner margin

of coxae 2-5, below the sternum. Immature

specimens are recognizable by their relatively

small size, lack of brood plates and undifferen-

tiated second gnathopod. Terminology of the

appendages is by vertical orientation in relation

to the body.

SYSTEMATIC SECTION

Family Ampithoidae Stebbing 1888!

Ampithoidae Stebbing, 1906, p. 631; J.L.

Barnard, 1969a,, p.,/141-143, fig... 61;

Bousfield, 1982, p. 285.

Amphithoidae Stebbing, 1899, p. 211

Diagnosis: Body smooth, little compressed.

Pair of short setae on the dorsum of urosomites

1 and 2. Head, rostrum lacking, anterior lobe

short and blunt, inferior antennal sinus shallow;

eyes lateral, rounded, medium to small. Antennae

medium to large. Antenna | peduncular segment

3 short, accessory flagellum short, vestigial or

lacking. Antenna 2 peduncle strong.

Buccal mass directed below the head. Upper

lip rounded below. Mandible, molar strong,

palp slender or lacking. Left lacinia mobilis with

5 or more cusps. Lower lip, outer lobes with

characteristic medial notch or emargination.

Maxilla 1, inner plate small, outer plate with 10

(rarely less or more) apical spine-teeth. Maxilla 2,

plates apically and medially setose, outer plate

somewhat broadened. Maxilliped plates large,

palp slender and dactylate.

Coxae 1-4 overlapping, deep, smooth or lightly

setose below. Gnathopods usually strongly

subchelate, 2 larger and sexually dimorphic.

Peraeopods 3 and 4 glandular, segment 2

expanded, distal segments short, dactyls with

gland duct. Peraeopods 5-7 dissimilar, distal

segments may be reversed, segment 6 may

expand and form a weak subchela with the

dactyl; coxae 5, 6 and sometimes 7 strongly

anterolobate, coxa 5 often as deep as coxa 4.

Pleopods normal, retinacula more than 2.

Urosome segments separate, not shortened.

Uropods | and 2 normally biramous. Uropod 3

biramous, rami very short, quadrate, inner

setose, outer with 2 (occasionally 1) strong apical

uncini. Telson short, apex usually with cusps.

Brood plates with hook-tipped marginal setae (on

peraeopods 2-5). Gills laminar, plate-like, short

pedunculate on peraeopods 2-6.

Genera: Amphithoides Kossman 1880, Amphi-

tholina Ruffo 1953, Ampithoe (Leach 1814),

Cymadusa Savigny 1816, Exampithoe K.H.

Barnard 1925, Macropisthopous K.H.

Barnard 1916, Paradusa Ruffo 1969, Peram-

phithoe new genus, Paragrubia Chevreux

1901, Pleonexes Bate 1857, Sunamphitoe

Bate 1857, Pseudoamphithoides Ortiz 1976

Animals are medium to large (5-35 mm) and

1. Some doubt exists concerning the orthography of the

family name. Leach (1814) dedicated his genus to the

Nereid ‘Amphithoe’ of Homer but erroneously translit-

terated the greek ‘phi’. This has led to a confusion of

spelling in subsequent publications. Although the authors

agree with Ruffo (1969) and Myers (personal communica-

tion) that the philologically better form ‘Amphithoe’ is

preferable, it is felt that formal alteration by the Inter-

national Commission on Zoological Nomenclature is first

necessary.

spin isolated tubes (with the cement glands of

peraeopods 3 and 4) on algae, eelgrass, stones

and detritus, or in the case of Amphitholina,

burrow into kelp stipes and holdfasts. All are

littoral and shallow-water marine or epibiontic,

tropical, warm temperate to arctic-boreal in

both northern and southern hemispheres.

Key to Genera of the Ampithoidae of the Northeastern

Pacific (Alaska to Northern California)

1. Antenna 1, accessory flagellum multi-segmented; palm of gnathopod | oblique; peduncular

spinous process projecting distally below the rami of uropods | and 2. .............00e0 eee

Cymadusa Savigny 1816 (p. 43)

Antenna |, accessory flagellum vestigial or absent; palm of gnathopod | transverse or

oblique; peduncular spinous process well developed if palm transverse but absent if palm

oblique

2. | Gnathopod 1, palm oblique, coxa 1 produced forward; peraeopods 3 and 4, segment 2

slender, less than */, the width of its coxa; uropod | peduncular spinous process absent

beele B1e SB e@ @eoeeseseeviecocoeostoanuveeeseoeevnueeesvesevoseeseveeeseesesee

Ampithoe Leach 1813-14 (sens. str.) (p. 45)

Gnathopod I, palm transverse, coxa | not produced forward; peraeopods 3 and 4, segment 2

strongly inflated, more than */, the width of its coxa; uropod 1 peduncular spinous process

present

Genus Cymadusa Savigny 1816, Barnard 1969a

Cymadusa Savigny, 1816

Grubia Czerniavski, 1868

Acanthogrubia Stout, 1912

Type species: Cymadusa filosa Savigny 1816

(Monotypy). See Chevreux and Fage, 1925 (as

Grubia hirsuta), Shoemaker, 1935 (as Grubia

filosa).

Diagnosis: Head lobe produced, inferior

antennal sinus moderate. Antenna 1, accessory

flagellum with 1'/, to 6'/, segments. Mandibular

palp moderately strong. Maxilla 1 palp broad.

Gnathopod 1, palm oblique, coxa | produced

forward; gnathopod 2 subchelate, equal to or

larger than 1. Peraeopods 3 and 4, segment 2

moderately inflated. Peraeopods 5-7, segment 6

not strongly widened apically, spines not restricted

to the antero-distal region. Uropod 1, peduncle

extended postero-distally into a long spinous

process between the rami. Uropod 3, outer ramus

with two hooked uncini. Telson with two small

apical cusps. About 10 species, generally tropical

to subtropical to cool temperate; littoral.

Northeastern Pacific species: Cymadusa

uncinata (Stout 1912)

Cymadusa uncinata (Stout 1912, Barnard 1969b)

igure |.

Acanthogrubia uncinata Stout, 1912, p. 146,

figs. 81-83

Paragrubia uncinata Shoemaker, 1941, p. 188;

Hewatt, 1946, p. 199

Peramphithoe n. gen. (p. 60)

Cymadusa uncinata J.L. Barnard 1965, p. 40,

figs. 26-28

J.L. Barnard, 1969b, p. 86

Material examined: Alaska — Puffin Bay,

Baranof Island: 12 immature specimens from

Bousfield and McAllister 1961, stn. A171. British

Columbia — Queen Charlotte Islands: 4 imma-

ture specimens from Bousfield 1957 stn. W8.

Northern mainland: 48 specimens from Bousfield

1964 stns. HS53, H65 and the collection of D.E.

McAllister, 1965. Vancouver Island and southern

mainland: 78 specimens from collections of

Bousfield 1977 (stn. B6a), 1959 (stns. O5, O11,

V4b, cat. 2606), 1955 (stns. P7, F2) and the collec-

tions of J.F.L. Hart 1934, 1939, 1941. Washing-

ton and Oregon — 54 specimens from Bousfield

1966 stns. W40, W42 and the collection of R.I.

Smith, 1955. Smithsonian collections (USNM):

Bousfield 1966 Stn. W40, 1 o&, 1 2 subadult,

6 immatures.

Distribution: Baranof Island, Alaska (56°16’N,

134°48’W) to Laguna Beach, California (33°S’N,

117°8’W).

Ecology: Occurs amongst kelp and Phyllospadix

in high salinity waters of exposed coasts at low

tide, in the shallow subtidal or in tide pools.

Females brood young May-August. Large over-

wintered females which have not developed setae

on the brood plates can be found in the spring.

Diagnosis: Antenna | longer than 2, peduncular

segments weakly setose, peduncle 1 bearing spines

on postero-distal edge. Antenna 2 peduncle not

_—

‘e'

Figure 1. Cymadusa uncinata Stout 0 20.0 mm, Trial Island Point, Victoria, B.C. 18 May 1977; 2 23.0

mm, Shipwreck Point, Clallam County, Washington. | Aug. 1966

greatly stouter, flagellum equal in length to

peduncle segments 4 and 5 together. Mandible,

incisor with 9 teeth, 11 spines; lacinia mobilis

with 6 teeth; palp segment 1 bare of setae,

segment 2 setose on distal and inner margins,

segment 3 longer than 2, distally rounded and

strongly setose on distal and ventral margins.

Lower lip outer lobes, apical somewhat longer

than medial. Maxilla 1, inner plate with 3 setae;

palp more than half the width of the outer plate,

tipped with 10 spines and 8 setae. Maxilliped

inner plate apically spinous; outer plate inner

edge smooth, teeth smooth. Coxae 1-4 setose on

lower margin. Coxa | produced forward, anterior

edge straight; coxae | and 2 not shallower than

3-5. Gnathopod 1, both sexes, segment 2 with a

small antero-distal lobe; segment 3 not lobed,

segment 5 narrowed below into a shallow forward

produced lobe; palm oblique, well defined, dactyl

serrated. Gnathopod 2 (co) much larger than 1,

segment 2 bearing long plumose setae on the

anterior, medial and posterior margins; neither

segment 2 nor 3 lobed; segment 5 bearing

feathery setae on the upper margin; at about

15 mm body length, hand splitting into two large

teeth, the ventral tooth much the Jomger,

approaching half the length of the hand; dactyl

slightly sinuous, projecting beyond the hand.

Gnathopod 2 ( 2) somewhat larger than gnatho-

pod 1, lacking plumose setae or teeth as in the

male. Peraeopod 3 similar to, though slightly

longer than 4, segment 2 expanded, but less than

the width of its body segment, segment 4 slightly

less than half the width of segment 2; male

peraeopod 3, anterior edge of segment 2 bearing

plumose setae. Peraeopods 5-7, anterior edge of

segment 2 spinous, segment 4 longer than 5,

segment 6 bearing 5-7 groups of strong spines.

Epimeral plates, lower hind corners acute but not

notched and lacking a lateral ridge. Pleopod

outer rami slightly shorter than inner, bearing

5-6 anchor-shaped coupling hooks. Urosome |

sternum setose. Uropods | and 2 bearing lateral

tufts of setae on the peduncle and outer ramus,

peduncular process well developed, outer ramus

shorter than inner, spines small and abundant,

2-4 at tip. Uropod 2 reaching somewhat beyond 1.

Uropod 3 long, peduncle bearing 2 central spines

and 12 distal ‘“‘crown” spines, inner ramus with 2

central and 5 distal spines, outer ramus with |

central and two well developed hooked uncini.

Telson with a row of 10-15 apical setae. Female

brood plates relatively small, reaching not more

than two-thirds the length of peraeopod

segment 2. Body length at maturity: & 15-35 mm,

2 20-35 mm. This is the largest species of

Cymadusa and one of the largest amphipods on

this coast.

Remarks: Cymadusa uncinata can be distin-

guished from the North Atlantic species C. filosa

and C. compta by its much greater body size (up

to 35 mm as opposed to 15 mm), the unequal

sizes of gnathopods | and 2 and the deeply incised

palm of gnathopod 2(o). Cymadusa most closely

resembles Ampithoe (Sens. str.) but differs mainly

in the presence of a distinct accessory flagellum,

long peduncular process of uropod | and more

setose maxilla | plate (3-6 setae as opposed to

0-2 setae). These characteristics are considered

plesiomorphic, and suggest that Cymadusa 1s

ancestral to Ampithoe.

Genus Ampithoe (Leach 1813-14) emend. (sens.

str.)

Ampithoe Leach, 1813-14, p. 403, 432; Stebbing,

1906, p. 631-632;

J.L. Barnard, 1969a, p. 143, 144, fig. 61

Type species: Ampithoe rubricata (Montagu

1808) (type by original designation).

Species contained in the genus Ampithoe have

recently been found by the authors to be of three

basic types. These differ on a significant basis in

several prime morphological characters, hence

justifying subdivision into the genera Ampithoe

(sens. str.), Peramphithoe and Pleonexes. Details

of this analysis are to be published separately.

(Conlan, 1982).

Diagnosis: Head lobe produced, antennal sinus

present. Antenna | accessory flagellum minute,

one-segmented. Mandibular palp moderately

weak to strong. Maxilla 1 palp relatively broad.

Gnathopod 1, palm oblique, coxa | produced

forward. Gnathopod 2 subchelate, equal to or

larger than 1. Peraeopods 3 and 4, segment 2

moderately inflated. Peraeopods 5-7, segment 6

little distally expanded, spines usually not

restricted to the antero-distal region. Uropod 1,

peduncular process vestigial or absent. Uropod 3,

outer ramus with two apical hooked spines or

uncini. Telson with two small apical cusps. About

37 species, arctic-boreal to tropical, littoral.

Northeastern Pacific species: Ampithoe valida

Smith 1873, A. lacertosa Bate 1858, A. simulans

Alderman 1836, A. volki Gurjanova 1938,

A. dalli Shoemaker 1938, A. plumulosa Shoe-

maker 1938, A. rubricatoides Shoemaker 1938,

A. kussakini Gurjanova 1955, A. sectimanus n.sp.

Key to Species of Ampithoe of the Northeastern Pacific

Gnathopod 1, lower lobe of segment 5 broad, more than half the length of the full segment.

Peraeopods 3 and 4 slender, segment 4 less than half the width of segment 2. Gnathopod

Zee ), both segments 2 and 3 produced into a large anterodistal lobe ......6..0...0 0055 000% 2

Gnathopod 1, lower lobe of segment 5 less than half the length of the full segment. Peraeopods

3 and 4 strong, segment 4 more than half the width of segment 2. Gnathopod 2 (), only

segment 2 produced into an anterodistal lobe

Antenna 2 with dense plumose setae on peduncle 5 and flagellum. Epimeron 3, hind margin

evenly rounded. Gnathopod | (o’), segment 5 shorter than segment 6. Gnathopod 2(<),

Pam sinuous, slightly oblique ..............

etches Ampithoe plumulosa Shoemaker 1938 (p. 50)

Antenna 2 lacking dense plumose setae. Epimeron 3, hind margin slightly to strongly notched.

Gnathopod | (o’), segment 5 longer than segment 6. Gnathopod 2(o), palm

EMS x, LE en Re EOE 0 we OS, « «

Antenna | peduncle | setose but lacking ventral spines. Coxae 1-4 with a group of long setae

on the lower hind margin. Epimeron 3, hind margin only faintly notched and lacking a lateral

ridge. Gnathopod | (0°), front margins of segments 5 and 6 bearing long overhanging simple

setae. Gnathopod 2 (<o), palm with a median quadrate tubercle, hind margin not

rea eet SE Gee eet Cate ieee cree paca s sw. oe ree FUR Ampithoe valida Smith 1873 (p. 49)

Antenna | peduncle | setose and bearing one to several spines on the ventral margin. Coxae

1-4 lacking a group of long setae on the lower hind margin. Gnathopod | (*), front margins

of segments 5 and 6 bearing upright plumose setae. Epimeron 3, hind margin with a strong

notch, from which radiates.a lateral ridge. Gnathopod 2 (o’), palm sinuous, hind corner

produced downwards with age, lacking median tubercle ...... Ampithoe lacertosa Bate 1858 (p. 47)

Antenna | subequal to antenna 2. Antenna 2, peduncle moderately to strongly setose;

flagellum shorter than segments 4 and 5 together. Uropods 1 and 2, rami tipped by several

spinessCoxae | and.2(o¢)same depthas coxae 3-5 2... FSi ws eden ss dave el ode 5

Antenna | shorter than antenna 2. Antenna 2, peduncle weakly setose; flagellum as long as

segments 4 and 5 together. Uropods | and 2, rami tipped by a single heavy spine. Coxae 1 and

D his) Seaweed lage 3-5) hoi Vale aise sacees w slg le we, Sele ge ale oes 4 aa eee bald pote a 8

Antennae | and 2, peduncles strongly setose; antenna | peduncle | without a posterodistal

spine. Anterior margin of segment 2 of peraeopods 3-5 strongly setose. Maxilliped outer plate

teeth serrated. Gnathopod 2 (0), palm strongly incised to form a pointed tooth ............

SAG tp a es Sead. hyo Rieter eres. sos \ Waa omen ota Ampithoe sectimanus n. sp. (p. 54)

Antennae | and 2, peduncles not strongly setose; antenna | peduncle | with a spine on the

posterodistal angle. Anterior margin of segment 2 of peraeopods 3-5 bare. Maxilliped outer

plate teeth smooth. Gnathopod 2 (0), palm not strongly incised, defining tooth small and

Seer e Or IRIS, washings aeahiele ohare. « « «calles wed nase oe ge mee ee we 6

Mandibular palp, segment 3 distally pointed and oblique, setae apical; segment 2 with | setae.

Gnathopod 2 (o’'), segment 5 as long as deep; palm incised to form a square tooth at posterior

CORUCT Bremyey Giant aaions: Sa tial. +« » enter aits Ampithoe volki Gurjanova 1938 (p. 53)

Mandibular palp segment 3 rounded, apically and laterally setose; segment 2 with more than

1 seta. Gnathopod 2 (o’), segment 5 longer than deep; palm oblique, small rounded tooth at

POSEMONCOMC tien onladid bon aenkis sutewadte eda eae Ampithoe kussakini Gurjanova 1955 (p. 52)

Antenna 2, peduncle 5 and flagellum with conspicuous groups of setae on hind margin.

Uropod | strongly spinose, with about 15 spines on the outer margin of the peduncle and

about 25 on the outer ramus. Uropod 3 peduncle with mid-dorsal spines in addition to the

usual distal crown spines. Gnathopod 2 (0), hand not greatly enlarged, less than twice the

Size Ob the hand of enathopod 1, palmeritecth absent . 2.0... 2 .)en5..0 4s os eel

nent ome. eteonitee Sul, ENR: swiefater.. 3 dai Ampithoe rubricatoides Shoemaker 1938 (p. 56)

Antenna 2 weakly setose. Uropod 1 with less than half as few spines on peduncle and rami.

Uropod 3 with distal crown spines only. Gnathopod 2 (0), hand greatly enlarged, more than

twice the size of the hand of gnathopod 1, with a strong tooth at the hind corner of the palm

anise tne dacty) linge it 2 Ralallieaeels 2 CaS Bs bs 2 ee 8

Mandibular palp, terminal segment acutely oblique, distal margin demarcated by a definite

angle from the inner margin and setose for about half the length. Gnathopod 1 (o&),

developing (at about 8 mm size) plumose setae on segments 2-5. Gnathopod 2 (oc), palm

barely concave, hind margin produced into a small tooth which in the full adult does not meet

PS AC ee eee ees. Pea eas Nes Gees oS Ampithoe dalli Shoemaker 1938 (p. 56)

Mandibular palp, distal margin of segment 3 very oblique, rounding smoothly into the inner

margin and setose for nearly the full length. Gnathopod | (0), developing (at about 12 mm

size) plumose setae on segment 2 only. Gnathopod 2 (’), palm strongly concave, hind margin

produced intoalong.tooth which meets ‘the: dactyl tassiciew. &. 0a bade eel

The northeastern Pacific species of Ampithoe

may be clustered into three subgroups which, on

further study, may warrant formal taxonomic

recognition:

1. A. lacertosa, A. valida, A. plumulosa

2. A. kussakini, A. volki, A. sectimanus

3. A. rubricatoides, A. dalli, A. simulans.

Characteristics which order the species are:

reduction in size of the antennal sinus, shortening

of antenna | with concurrent strengthening

of antenna 2, increasing obliqueness of mandi-

bular palp segment 3, increasing roundness of

the coxae, narrowing of the segment’5 lobe of

gnathopod 1, increasing obliqueness and con-

cavity of male gnathopod 2, increasing glandu-

larity of peraeopods 3 and 4, strengthening of

the uropod 3 uncini, loss of the lateral ridge on

the epimera, an overall deepening of the body

and shallowing of the coxae. The functional

significance of these morphological changes

suggest a trend towards greater domicoly.

Group of Ampithoe lacertosa, A. valida,

A. plumulosa

Head, lateral lobe and antennal sinus prominent,

eye medium. Antenna | equal to or longer than

antenna 2, peduncle» poorly setose. Antenna 2

very little stronger than antenna 1, flagellum

equal to or shorter than peduncular segments 4

and 5 combined. Mandibular palp segment 3

apically blunt, distally setose. Coxa | anterior

margin straight or slightly upcurved; coxae | and

2 (oc), shallower than 3-5; coxa 5 in both sexes,

lower corners acutely rounded. Gnathopod 1,

posterior lobe of segment 5 broad, more than

half the length of the full segment and in the

male, produced distally under segment 6.

Gnathopod 2 (co), both segments 2 and 3

produced into a prominent antero-distal lobe,

palm transverse or slightly oblique. Peraeopods

3 and 4 slender, segment 4 less than half as wide

as segment 2. Peraeopods 5-7 slender, spines

weak. Uropods | and 2, rami ending in a group

of 2-4 spines. Epimera 1-3 with a lateral ridge.

Pleopods with 8-11 coupling hooks. Uropod 3

long, apical uncini of outer ramus weak.

The species trend phyletically in the increased

shortening of antenna | relative to antenna 2,

increasing obliqueness of mandibular palp

segment 2 and increasing obliqueness of the male

palm of gnathopod 2. Ampithoe japonica Stebbing

1888 and A. cavimana Sivaprakasam 1970, seem

referable to this group.

Ampithoe lacertosa Bate 1858, Barnard 1965

Figure 2.

Amphithoe lacertosa Bate 1858, p. 362; 1862,

p. 236-237, pl. 41, fig. 5; Gurjanova, 1951,

p. 895-897, fig. 622.

Ampithoe lacertosa Stebbing, 1906, p. 633-634;

J.L. Barnard, 1954, p. 31-33, pls. 29-30;

J.L. Barnard, 1965, p. 9-12, figs. 4, 5;

Nagata, 1960, p. 175-176, pl. 16, figs. 95-96;

Heller, 1968, p. 1-132; J.L. Barnard, 1969b,

p. 83

Amphithoe macrurus Stephensen, 1944, p. 80-83,

figs. 30-31

Dexamine scitulus Harford, 1877, p. 116

Amphithoe scitulus Holmes, 1904, p. 314-315,

pl. 36, figs. 21-24

2? Amphithoe stimpsoni Boeck, 1871, p. 14-15,

fig. 5; Stebbing, 1906

Material examined: Alaska — Aleutian Islands:

1 immature specimen from Izembek Lagoon,

Unimak Is., 1969, N.A. Powell collector. South-

eastern Alaska: 123 specimens from Bousfield

and McAllister 1961 stations, A3, A7, Al8, A27,

A48, A87, A91, A92, A96, A107, A115, Ai31,

A139, A147, A151, A174, A175; 5 specimens

from Bousfield 1980 stns., S8B1 and S23F1.

British Columbia — Queen Charlotte Islands: 60

specimens from Bousfield 1957 stns., W3a, W4a,

W14, H2, H2a, Sb; H9, H10; HES, BS RI

El4a, El4c, E24, E25. Northern mainland: 169

specimens from Bousfield 1964 stns., H1, H3, H8,

H10, H23, H25, H26,.H29, H30; 1135, H47, 4s.

HS50, H52, H56, H57, H65. Vancouver Island

and southern mainland: 1336 specimens from the

collections of Bousfield 1977, 1976, 1975, 1970,

1964, 1959, and 1955, and from the collections

of Cy Carl, K.E.*Conlan, DVV. Ells, J.Psl: Fiari;

D. Kittle, C.D. Levings, C. Lobban, R.J. Long,

N.A. Powell.

Washington and Oregon — 77 specimens from

Bousfield 1966 stns., W8, W10, W13, W30, W33,

W42, W44, W47, W64 and 10 specimens from

the collections of S. Helen, 1967 and N.

McDaniel, 1977.

Smithsonian Collections (USNM): Bousfield

1975: stn Pili6a, doitmleos

Distribution: Aleutian Islands, Alaska (SO°N,

163°W) to Magdalena Bay in Baja California

(25°N,112°W); Japan, south toyShizuoka

prefecture (35°N,138°E).

Ecology: Found amongst algae, eelgrass or

woody debris, on mud, sand and gravel beaches,

in tidepools or on wharf pilings at low water or

Figure 2. Ampithoe lacertosa Bate 0 21.5 mm; 2 30.0 mm, Friday Harbor, San Juan Is., Washington.

April 1977

subtidally to about 10 m depth. It is equally

abundant in high salinity exposed coastal waters

and mesohaline shallow coastal waters but is

rarely found in brackish water. Females are

ovigerous from May to August.

Diagnosis: Antenna | longer than antenna 2,

peduncle 1 with 1-4 posterior marginal spines

(although absent in immatures). Antenna 2

peduncle not spinose, flagellum equal in length

to peduncular segments 4 and 5 together. Mandi-

bular palp segment 3 blunt, apically setose, 1

small seta on the distal corner of segment 2.

Lower lip outer lobe, apical and medial lobes

well separated, apical longer than medial.

Maxilla 1 inner plate with 1 seta, palp strong,

with about 12 marginal spines. Maxilliped palp

segment | lacking setae. Coxae 1-4 lacking long

setae on lower margin. Gnathopod | both sexes,

segment 6 slender, width less than half the length.

Gnathopod 1 (0), segment 5 longer than 6,

margins of segments 5 and 6 developing a growth

of plumose setae when body length is 12 mm or

more. Gnathopod 2 (o'), segment 2 poorly

setose, palm transverse, sinuous, lacking a

median tubercle, hind edge produced downwards

progressively with age to form a wide thumb.

Male sternum of peraeon 7 lacking a median keel

between the penes. Epimera 1-3 with strong

Sona

ee ee ee

notch in hind corner (in immatures as well)

from which radiates anteriorly a lateral ridge.

Uropod 3 peduncle bearing dorsal spines along

its length in addition to a crown of spines on

the distal margin.

Body colour in life is orange to brown, heavily

speckled. Distinctive white spot on the dorsum of

each segment; some white spotting also on the

coxae. Body length at maturity: Male 12-24 mm,

female 10-23 mm. The male gnathopod 2 becomes

transverse at as small a size as 7 mm but

gnathopod | does not become plumose until the

body is about 12 mm in length. A plot of the

body length of 81 specimens against geographic

distribution indicates a northward trend towards

increased body length at maturity. The average

size of maturity in Vancouver Island, southern

B.C., Washington and Oregon was 16 mm gd,

14 mm Q, while in Alaska and northern B.C. the

average was 17 mm oc, 17 mm @.

WZx

Remarks: Heller (1968) described extensively

the biology and development of this species.

Ampithoe valida Smith 1873

Figure 3.

Amphithoe valida Smith, 1873, p. 563; Paulmier,

1905, p. 164-165, fig. 34

Ampithoe valida, Stebbing, 1906, p. 635;

Alderman, 1936, p. 68; J.L. Barnard, 1954,

p. 34-35, pl. 31; Nagata, 1960, p. 176, pl. 16,

figs. 97-98; J.L. Barnard, 1965, p. 34-36, figs.

22, 23; Bousfield, 1973, p. 180-181, pl. LV.1

Ampithoe shimizuensis Stephensen, 1944, p.

77-80, figs. 27-28

Material examined: British Columbia

Vancouver Island and southern mainland: 67

specimens from the collections of Bousfield

1976 (stns. B3, Blla, B13); 1975 (stns. P6a, P6b,

Péc, P18a),; 19°70 (stns. P706, P709), 1959 (sta:

N17))-1955 fstns, Fl; F2a, F110; Gil Gispars

Figure 3. Ampithoe valida Smith & 12.0 mm, Sarita Bay, Barkley Sound, Vancouver Is., B.C. 2 July 1977;

2 12.0 mm Brady’s Beach, Barkley Sound, Vancouver Is., B.C. 31 July 1975

Mla, M2, M4, MS, M8, M10, M11). Four speci-

mens also examined from the collections of

M.A. Bousfield, K.E. Conlan, D.V. Ellis and

R.J. Long.

Washington and Oregon — 12 specimens from

Bousfield 1966 stns. W28, W47 and W64.

Smithsonian collections (USNM): Bousfield 1975

Stiwroa, 27d OAc e ee

Distribution: Pacific Ocean: British Columbia

and Vancouver Island at 51° latitude south to

Newport Bay, California (45°N,124°W), ? Japan

at Shizuoka Prefecture (35°N,138°E). Atlantic

Ocean: Piscataqua estuary, New Hampshire

(43°N,70°W) south to Chesapeake Bay, Virginia

(37°N,76°W).

Ecology: A warm temperate species occurring

along sheltered coasts and estuaries, mainly in

mesohaline to brackish waters. It builds tubes on

algae and eelgrass on muddy, gravelly beaches in

saltmarshes, tidepools and log fouling communi-

ties, at low water level to 30 m depth. Females

brood from May to August. Immatures are

brooded for about 2-5 weeks and grow about

1 mm/week, reaching maturity in about 6 weeks

(Nicotri, 1980).

Diagnosis: Antenna | slightly longer than (c)

or equal to (2) antenna 2; peduncle | setose but

lacking posterior marginal spines. Antenna 2

peduncle not spinose, flagellum shorter than

segments 4 and 5 together. Mandibular palp

segment 3 obliquely truncate, apically setose,

1 small seta on the distal corner of segment 2.

Lower lip, outer lobe, apical and medial lobes

closely appressed, apical only slightly longer than

medial. Maxilla | inner plate lacking setae, palp

rather weak, with about 6 marginal spines.

Maxilliped palp segment 1 setose. Coxae 1-4

bearing a group of long setae on the lower

margin. Gnathopod 1, both sexes, segment 6

subcircular, width more than half the length.

Gnathopod 1 (oc), segment 5 longer than 6,

upper margins of segments 5 and 6 fringed with

dense overhanging setae. Gnathopod 2 (o), hind

margin of segment 2 strongle setose, palm

transverse, developing a median quadrate tubercle

at about 6 mm body length, but not produced

antero-distally with age. Male sternum 7 lacking

a median keel between the penes. Epimera 1-3

with lateral ridge, hind margins rounded, with

one short seta at hind corner. Uropod 3 lacking

dorsal spines other than the usual marginal

crown spines.

In life, body olive green to brown, and heavily

speckled. Body length at maturity: Male 6-12 mm,

female 5-12.5 mm.

Ampithoe plumulosa Shoemaker 1938

Figure 4. (after Shoemaker 1938)

Ampithoe plumulosa Shoemaker 1938, p. 16-19,

fig. 1; 1942, p. 39; J.L. Barnard, 1959, pa37:

1964, p. 111; 1965, p. 20, figs. 11, 12; 1969b,

p. 84.

Material examined: No material available. |

Information compiled from Shoemaker (1938),

Barnard (1964, 1965 and 1969b).

Distribution: Patos Island, British Columbia

(48°48.7’N) to Salinas, Ecuador; Galapagos

Island (O°N,90°W).

Ecology: A warm water species found amongst

algae and Phyllospadix on pilings, floating docks,

mud beaches and in tidepools, intertidal to a

depth of about 15 m.

Diagnosis: Antenna | slightly longer than 2,

peduncle | with up to 6 ventral spines. Antenna 2

peduncle may bear a few dorsal spines on

segment 3 (but see remarks below); peduncular

segment 5 and flagellum of antenna 2 clothed in

abundant plumose setae. Mandibular palp seg-

ment 3 obliquely truncate, apically and some-

what laterally setose, 3 long setae on the margin

of segment 2. Lower lip outer lobe, apical and

medial lobes well separated, apical longer than

medial. Maxilla 1 inner plate with 4 setae, palp

strong, with about 8 marginal spines. Maxilliped

palp segment | setose. Coxae 1-4 lacking long

setae on hind margin. Gnathopod 1, both sexes,

segment 6 slender, width less than half the length.

Gnathopod 1, male, segment 5 shorter than 6,

upper margins of segments 5 and 6 not strongly

setose. Gnathopod 2, male, hind margin of

segment 2 not strongly setose, palm slightly

oblique, sinuous, forming a low flat tooth with

age. Male sternum 7 bearing between the penes a

median, lamellar, oval keel armed with marginal

teeth which extends about 7/, the depth of the

penes (see Shoemaker, 1938). Epimera 1-3 hind

margins evenly rounded, lateral ridge on each

(but see remarks below). Uropod 3 lacking dorsal

spines other than the usual marginal crown

spines. Body length at maturity: Male 13-16 mm,

female 12 mm.

Remarks: The illustrations of Ampithoe

plumulosa collected in Bahia de San Quintin by

Barnard (1965) differ in some respects from

Shoemaker’s type collected on Catalina Island,

California. Antenna 2: type male (16 mm) lacks

Ea es

BES

(ee SE

MX|

PENIAL KEEL

Figure 4. Ampithoe plumulosa Shoemaker & 16 mm, La Jolla, California. 20 Sept. 1918. Reconstructed

from Shoemaker, 1938

spines on peduncle, while the male (13 mm)

figured by Barnard bears 4 anterior marginal

spines on peduncle segment 3. Epimera | and 2:

lateral ridges are illustrated on Shoemaker’s

type, yet are stated by Barnard to be absent. Only

three specimens have been documented (by Shoe-

maker, 1938) for the Pacific coast north of

lower California.

Group of Ampithoe kussakini, A. volki,

A. sectimanus

Head, lateral lobe and antennal sinus prominent;

eye small. Antennae 1 and 2 subequal, peduncle

moderately to strongly setose. Antenna 2 some-

what heavier than antenna 1, flagellum shorter

than peduncular segments 4 and 5 together.

Mandibular palp segment 3 oblique, laterally

setose. Coxa 1 produced forward, anteriorly

upturned; coxae | and 2 (co) as deep as 3-5;

coxa 5 in both sexes, lower corners evenly

rounded. Gnathopod 1, lower lobe of segment 5

narrow, less than half the length of the full

segment and not produced under segment 6.

Gnathopod 2 (oc), segment 3 not produced into

an anterior lobe, palm oblique and weakly

concave, developing a small to long projection at

the hind corner. Peraeopods 3 and 4 normal,

segment 4 more than half as wide as segment 2.

Peraeopods 5-7 normal, spines strong. Epimera

1-3 with or without a lateral ridge. Pleopods

with 6-9 coupling hooks. Uropods | and 2, rami

tipped by a group of 2-4 spines. Uropod 3 long,

uncini of outer ramus moderately strong.

The species sequentially trend to reduction of

the antennal sinus, increasing strength of setation

on antennae, peraeopods and uropod 1, the

increasing obliqueness in mandibular palp

segment 1, increasing degree of incision of the

male gnathopod 2 palm and a loss of the lateral

ridge of epimeron 3.

Other species exhibiting affinities with this

group: Ampithoe djakonovi Gurjanova 1938,

Ampithoe longimana Smith 1873, Ampithoe

platycera Sivaprakasam 1970, Ampithoe ramondi

Audouin 1826, Ampithoe zachsi Gurjanova 1938.

Ampithoe kussakini Gurjanova 1955

Figure: S,

Ampithoe kussakini Gurjanova 1955, p. 215-

Pie. Migs. 22). 23

Material examined: Alaska — Aleutian Islands:

63 specimens from the collections of C.E. O’Clair,

1970 and 1972 and N.A. Powell 1969. South-

eastern Alaska: 144 specimens from Bousfield

and McAllister 1961 stns. A7, Al9, A20, A25,

A27, A30, A43, A68, A70, A71, A75, A86, A87,

A91, A92, A98, A99, A105, A106, A114, A115,

A121, A131, A136, A139, Al47, A151, A153,

A164, A175; 40 specimens from Bousfield 1980

stns. S4B1-4, S8B1, SI13B1, S18B1, S19B1.

British Columbia — Queen Charlotte Islands:

131 specimens from Bousfield 1957 stns., ES,

E9, El4a, El4b, El4c, E21, E24, E25, H4a, HS,

H8a, H8b, H9, H11, W5, W15b. Northern main-

land: 35 specimens from Bousfield 1964 stns.,

H8, Hi2, Hi3,-H17, H32, B33, Hotesnoe

Northern Vancouver Island: 7 specimens from

Bousfield 1959 stn. 04. Smithsonian collections

(USNM): Bousfield 1957 stn. H4a,2 0 oh,2 2 9,

immature. Zool. Inst. (Leningrad): Bousfield

1961 stn. A27, 1 o&; Bousfield 1961 stn. A25,

2 2, 1 immature; Bousfield 1957 stn. El4a,

2 WO Grn Gee

Distribution: Aleutian Islands, Alaska, (55°N,

163°W) to Quatsino Sound, Vancouver Island,

B.C. (50°30'’N,128°06’W). Otradnaya Bay,

Shikotan Island, Tatar Strait, Kurile Islands,

USSR.

Ecology: A northern cold-temperate species

found in the mid and low intertidal to a depth of

about 15 m amongst algae and eelgrass, primarily

on protected and less often exposed coasts,

where summer temperatures are 10-17°C, and

salinities are high, seldom brackish. Females

are ovigerous June to August.

Diagnosis: Antenna 1 shorter than 2 (co),

Figure 5. Ampithoe kussakini Gurjanova & 17.5 mm; 9 13.0 mm, Yakoun Bay, Masset Inlet, Queen

Charlotte Is., B.C. 27 August 1957

longer than 2 (9); antenna | peduncle | with

a distal ventral spine (present also in immatures);

antennae | and 2 moderately setose. Mandibular

palp segment 3 evenly rounded, setae apical

and lateral; segment 2 with about 8 setae. Maxilla

| inner plate with | seta, palp slender. Maxilliped

outer plate teeth smooth, palp segment 2 lacking

setae. Gnathopod 2 (0°), segments 4-6 elongate,

palm oblique, sinuous, thumb at hind corner

small or absent, groups of dense setae developing

on inner side of hand at about 9 mm body length.

Anterior margin of peraeopods segment 2 not

strongly setose. Epimera 1-3 with a lateral ridge.

Pleopods moderately setose. Uropod 3 moderately

long, inner ramus with | medial and 4 apical

spines. Colour pattern: body orange to green or

brown; antennae brown and white banded. Body

length at maturity: Male 9-18 mm, female

9-20 mm.

Remarks: There are some discrepancies between

this and Gurjanova’s description (types not

re-examined). Her specimens are much larger

than those examined here, the male being 31 mm,

the female 29 mm, yet the flagella of antennae |

and 2 are much shorter, comprised of 15 and 6

segments respectively, as opposed to 30 and

13-14 in our specimens half their size. Gurjanova

stated that the palm of the female gnathopod 2

lacks an obturator spine, an unusual feature, yet

one is present in the North American specimens.

The number of setae on the telson of Gurjanova’s

specimens is only about half the North American

number. Perhaps, however, these characters

reduce with age. Otherwise the eastern and

western Pacific specimens show close resemblance.

Ampithoe volki Gurjanova 1938 ?

Figure 6.

Amphithoe volki Gurjanova 1938, p. 359, fig. 52;

1951, p. 899-901, fig. 624.

Material examined: Alaska — St. Makarius

Bay, Amchitka Island (51°N,179°W), collection

of CLE’ O' Clair,

2 immatures.

1968. 1 male, 1 female,

LZ,

——— SS See

Figure 6. Ampithoe volki Gurjanova? o& 8.0mm; @ 8.0 mm, St.

Is., Alaska. 20 July 1968

Makarius Bay, Amchitka, Is., Aleutian

Distribution: Amchitka Island, Alaska; Sea of

Japan, Prebrazheu’e Bay region, USSR (45°N,

130°E), Tatar Strait and northern Kurile Islands.

Ecology: A cold temperate species inhabiting

exposed coasts in tidepools overgrown with algae

and in beds of Laurencia, intertidal and subtidal

to 3 m depth.

Diagnosis: Antenna | shorter than 2 (ov), or

subequal (¢); antenna | peduncle 1 with a

postero-distal spine (absent in immatures);

antennae | and 2 moderately setose. Mandibular

palp segment 3 acutely oblique, tip pointed, setae

apical; segment 2 with 1-2 setae. Maxilla 1 inner

plate with 1 seta, palp moderately widened.

Maxilliped outer plate teeth smooth, palp

segment 2 setose. Gnathopod 2, male, segments

4-6 not greatly elongate, hand laterally incised

to form a short truncate thumb which is not

long enough to reach to the end of the hand.

Anterior margin of peraeopods segment 2 not

strongly setose. Epimera 1-3 with a slight lateral

ridge. Pleopods moderately setose. Uropod 3 not

greatly elongate, inner ramus with 3 apical and

no medial spines. Body colour in life translucent

yellowish-grayish-brownish (Gurjanova 1951).

Body length at maturity: Male 8 mm, female 8 mm.

Remarks: Table 1 illustrates that there are

several differences between these specimens and

Gurjanova’s types which may warrant designa-

tion of a new species name.

Ampithoe sectimanus n. sp.

Figure 7.

Ampithoe pollex: J.L. Barnard 1954, p. 29-31,

pls. 27-28 (not Kunkel 1910)

? Ampithoe simulans: J.L. Barnard 1965,

p. 27-30, fig. 18 (not Alderman, 1936)

Material examined: Point east of Point Marsh,

Figure 7. Ampithoe sectimanus n.sp. & 11.0 mm; 9 11.5 mm, Point Marsh, Prince of Wales Is., Alaska.

1 June 1961

Prince of Wales Island, Alaska (54°43’N,

132°17’W). Bousfield stn. A6, 1 June 1961. Tide-

pools and under stones. LW to MW. 9.5°C,

30.0°/.0. Holotype & (NMC-C-1981-961);

allotype @ (NMC-C-1981-962); paratypes

(NMC-C-198 1-963).

Additional material: Alaska — Southeastern

coast: About 40 specimens from Bousfield &

McAllister 1961 stns. A6 (holotype, allotype,

paratypes), A98, A17I-2; 5 specimens from

Bousfield 1980 stns. S4B5 and SI1BI.

British Columbia — Queen Charlotte Islands:

About 55 specimens from Bousfield 1957 stns.,

W4a, W8, W12. Northern mainland: 66 specimens

from Bousfield 1964 stns. H12, HS50, HS3.

Vancouver Island and southern mainland: 9

specimens from Bousfield 1977 stns., B6a, B11b,

B19a, B19b; about 30 specimens from Bousfield

1976 stns., B7, B28; 3 specimens from Bousfield

"975 stns., PSc, Pl7d; 174 specimens from

Bousfield 1970 stns., P702, P710, P711, P712,

P719; 18 specimens from Bousfield 1959 stns.,

01, 03, 05, 07b, V4B, V5, N6 and cat. #2606;

6 specimens from Bousfield 1955 stn., P7; also 3

specimens from the collections of C. Haylock

meee RK: Lee 1971 and G.C. Carl 1934.

Oregon — | specimen from Bousfield 1966 stn.,

W60, Otter Rock at Marine Gardens, Lincoln Co.

Smithsonian collections (USNM): Bousfield 1961

stn. Al71-2, 1 &, 1 9; Bousfield 1970 stn. P712,

3 o6od,3 2 2, Zool. Inst. (Leningrad); Bousfield

70’ stn. P702, 1 3, 2 2 2; Bousfield 1957 stn:

m2. 2 6d juveniles, 2 9 QO.

Distribution: Prince William Sound, Alaska,

(60°2’N,146°47’W), south to Cape Arago,

Oregon (43°30'N,124°26’W).

Ecology: A cold-temperate species restricted

to high salinity, exposed coasts, amongst algae

on rocky headlands at low water level, in low

summer temperatures (9.5°-15°C) and high

salinities (23.3-33.7°/..). Females ovigerous

May to August.

Description of male holotype, 11.0 mm:

Antennae | and 2 subequal, peduncles strongly

setose; antenna | peduncle | lacking a ventral

spine. Mandibular palp segment 3 acutely

oblique, tip pointed, setae apical; segment 2

with 3 setae. Maxilla | inner plate bare, palp

broad. Maxilliped outer plate teeth serrated,

palp segment 2 setose. Gnathopod 2, segments 4

and 5 not greatly elongate, segment 6 enlarged,

palm distally incised (beginning at 7 mm body

length), to form a pointed thumb which splits

increasingly with age to more than half the

length of the hand. Anterior margin of

peraeopods 3-7 segment 2 strongly setose (degree

and extent varies with age). Epimeron 3 with a

faint indication of a lateral ridge. Pleopods

strongly setose. Uropod 3 short, inner ramus with

4 apical and no medial spines. Body colour in

life: mottled orange to chestnut, legs and

antennae orange and white banded. Body length:

male 7-10 mm, female 8.5-12.5 mm.

Description of female allotype, 11.5 mm: Body

and appendages as in the male, except gnathopod

2 which is similar in form and somewhat larger

than gnathopod 1.

Etymology: (L.: secti = split; manus = hand),

referring to the strongly cleft palm of the male

second gnathopod.

Remarks: Ampithoe sectimanus was figured by

Barnard (1954) as Ampithoe pollex Kunkel 1910.

The latter species differs in many features —

smaller body size, weaker setation of antennae

and peraeopods (yet stronger setation of peraeo-

pods 3 and 4), different shape of gnathopods and

near occlusion of the inner lobes of the lower

lip. Barnard (1965) also figured gnathopod 2 of

two male specimens from station 40a, Coal Oil

Point, Goleta, California, that he called aberrant

forms of Ampithoe simulans. The illustration of a

longer palmar tooth on the hand combined with

the statement that the lobes of the lower lip are

well separated, indicates that these are more

likely specimens of Ampithoe sectimanus. If so,

these specimens mark the southernmost distribu-

tional limit of this species.

Group of Ampithoe rubricatoides, A. dalli,

A. simulans

Head, lateral lobe and antennal sinus shallow;

eye small. Antenna 1 shorter than 2, peduncle

poorly setose. Antenna 2 much heavier than

antenna 1, flagellum equal in length to peduncular

segments 4 and 5 together. Mandibular palp

segment 3 oblique, laterally setose. Coxa l,

anterior margin straight or slightly upcurved,

coxae | and 2 of the male shallower than 3-5;

coxa 5 in both sexes, lower corners evenly

rounded. Gnathopod 1, lower lobe of segment 5

narrow, less than half the length of the full

segment and not produced forward under seg-

ment 6. Gnathopod 2 (co), segment 3 not

produced into an anterior lobe, palm strongly

concave, with or without a small “thumb” on the

hind corner. Peraeopods 3 and 4 normal, segment

4 more than half as wide as segment 2. Peraeopods

5-7 strong, slightly expanded distally, spines

strong. Epimera 1-3 without a lateral ridge.

Pleopods with 6-11 coupling hooks. Uropods 1

and 2, rami tipped by a single strong spine.

Uropod 3 short, uncini of outer ramus strong.

Ampithoe corallina Stout 1913, which might also

be found in the northeastern Pacific, seems

referable to this group.

Ampithoe rubricatoides Shoemaker 1938

Figure 8. (reconstructed from Shoemaker, 1938)

Ampithoe rubricatoides Shoemaker 1938, p. 22,

figs. 3, 4

Amphithoe rubricatoides Gurjanova 1951, p. 878-

880, fig. 613

Material examined: No material available.

Information extracted from Shoemaker (1938).

Distribution: Alaska — Aleutian Islands:

Pribilof Islands (57°N,172°W).

Ecology: A cold water species found subtidally

at 10-18 m depth in sand and mud sediments.

Diagnosis: Head, antennal sinus absent, eye

small. Antenna 2 moderately setose. Mandibular

palp segment 2 short, less than twice the length

of segment 1; one corner and a group of 4 lateral

setae; segment 3 oblique, apically setose for about

Y/, its length. Gnathopod | palm very oblique,

passing by a scarcely perceptible angle into the

hind margin; male segment 2 lacking plumose

setae. Gnathopod 2 (co), segment 5 lacking a

dorsal tubercle; hand not greatly enlarged, less

than twice the size of the hand of gnathopod 1;

palm concave, lacking teeth, other than the

normal obturator spine. Peraeopod 7 segment 6

with 5 marginal spine groups. Epimeron 3 hind

margin evenly rounded. Uropods very spinose,

uropod | peduncle with about 15 outer spines,

outer ramus with about 25 spines. Uropod 3

peduncle with about 15 crown spines and 2

dorsal spines. Body length: Male 24 mm, female

length not stated.

Ampithoe dalli Shoemaker 1938

Figure 9.

Ampithoe dalli Shoemaker 1938, p. 19-22, fig.

2; Gurjanova, 1951, p. 887-890, fig. 618

Ampithoe simulans Barnard, 1965, p. 27-30,

fig. 17 (not Alderman 1936)

Material examined: Alaska — Aleutian Islands:

9 specimens from the collections of C.E. O’Clair,

1972-74. Southeastern Alaska; 207 specimens

Figure 8. Ampithoe rubricatoides Shoemaker & 24.0 mm, Kyska Is., Aleutian Is., Alaska. 1873. Recon-

structed from Shoemaker, 1938

Figure 9. Ampithoe dalli Shoemaker o& 18.0 mm; o subadult 11.0 mm, 2 20.0 mm, Kakul Narrows,

Baranof Is., Alaska. 9 June 1961

from Bousfield and McAllister 1961 stns., A2,

AS, A7, Al8, A23, A25, A27, A33, A37, A43,

A48, A68, A71, A98, A105, A115, A121; SO

specimens from Bousfield 1980 stns., SILI,

784. 5/B5, S8B1, $11B1, S13B1, $18B1, S20B2,

S20B4, S20B5, S20B6.

British Columbia — Queen Charlotte Islands: 40

specimens from Bousfield 1957 stns., E25, H&b,

H14, H15, W1, W5a, W9b, W12, W15. Northern

mainland: 191 specimens from Bousfield 1964

eed). (H113, H29;- H33, H35, H40, -H44.

Vancouver Island and southern mainland: 1

specimen from Bousfield 1977 stn. B12c; 33 speci-

mens from Bousfield 1976 stns., B2, B3, B12a,

B13, Bl4b; 5 specimens from Bousfield 1975

stns., P5d, P18; 25 specimens from Bousfield 1970

stns.,.P 703, P704,-P705; P7144P7 lS 4AP7 fp Zane

cimens from Bousfield 1959 stns., 03, 04; 115

specimens from Bousfield 1955 stns., P2, P4, P7,

F2a, F5, Gl, Gl1, G13, M2, M5, M11; in addi-

tion, 464 specimens from the collections of

Jj.G. Carl, Sx Cross. Voces, De Kittle; UKs.

Lee, C. Levings, C. Lobban, D.E. McAllister and

R.I. Smith.

Washington and Oregon — 3 specimens from

Bousfield 1966 stns., W5, W34, W64. Smithsonian

collections (USNM): Bousfield 1961 stn. A27,

lus vhost gguwenile,. 1) 492

Distribution: Known authentically from

Aleutian Islands, Alaska (51°N,179°W) south

to Cape Arago, Oregon (60°30’N,124°26’W).

Pacific coast of the USSR.

Ecology: A cold-temperate species occuring

amongst algae and eelgrass on exposed and

protected beaches, in tidepools, under stones

and amongst log fouling organisms. Occurs in

the low intertidal to 10 m depth in a wide

range of salinities (10-33°/..), and in summer

temperatures (8-18°C). Females ovigerous March-

August.

Diagnosis: Head, antennal sinus slight,

eye small. Antenna 2 poorly setose. Mandi-

bular palp segment 2 more than twice the

length of segment 1 and bearing a single

corner ‘seta; seoemrent 3° acutely oblique, a

definite angle between apical and lateral

margins, apically setose for about '/, its length.

Gnathopod 1 palm less oblique than in

A. rubricata, definite angle from the palm to

hind margin; male segment 2 with dense,

long plumose setae on the front, inner and

hind margins at about 6-7 mm body length,

the plumosity extending by 8-12 mm _ body

size to the lower margins of segments 3, 4

and 5. Gnathopod 2 (oc), segment 5 with a

low dorsal tubercle; hand enlarging with age,

sides diverging and palm lengthening, a

medial tooth forming at the dactyl hinge and

another at the hind corner of the palm; dactyl

increasingly curved with age, failing to meet

the hind palmar tooth in old specimens.

Peraeopod 7 segment 6 with 5-7 marginal

spine groups (depending on age). Epimeron 3

rounded, hind corner with slight indentation

holding a small seta. Uropod 1 with not

more than 6 spines on the outer peduncle and

10 spines on the outer ramus. Uropod 3 with

6 crown spines and no central spines. Body

colour in life: uniformly green to brown

speckled. Body length at maturity: male 8-18

mm, female 7-20 mm.

A plot of the body length of 272 specimens

against geographic distribution showed a

general increase in body size with latitude. In

latitude 45°-52°, mean length was 10.8 mm

(of) (range 8-17mm), 10.5 mm (@) (range

7-15 mm). In latitudes 53°-65°, mean length

was 12.2 mm (o) (range 7-19 mm), 14.1 mm

(2). (range 7-20"mm).: Ino the female; the

presence of brooding immatures was found to

occur at a size at least 1.5 mm greater than

in the condition where the brood pouch was

empty but brood lamellae were setose, indi-

cating that a moult occurred between these

two stages. Some seasonal differences in size

of maturity were also observed, individuals in

the spring (presumably having overwintered)

being about 1-2 mm larger than later indi-

viduals which presumably were early offspring

able to mature more quickly in warmer

summer water. This indicates that gonadal

growth requires a higher temperature than

does somatic growth.

Remarks: Barnard (1954) synonymized

Ampithoe dalli with Ampithoe simulans, in

the belief that A. dalli represented the mature

form of A. simulans, the transformation

occurring at about 15 mm. However, his con-

clusions were based on the evidence or

subadult male specimens which do resemble

each other closely but diverge with age. The

characters which distinguish these species

are body size at maturity, eye size, shape of

the mandibular palp, degree of plumosity of

male gnathopod 1, shape of male gnathopod 2,

spination of peraeopod 7, shape of epimeron 3

and body colour.

Ampithoe simulans Alderman 1936

Figure 10.

Ampithoe simulans Alderman 1936, p. 68-

79, figs. 44-47; J:-L. Barnard, igsa

p. 33-34, 1 fig.; not J.L. Barnarayy ies

p. 27-30, fig. 17;\? J.L. Barnard; ean:

p. 85

Material examined: Alaska — Aleutian

Islands: 1 specimen from the collection of

C.E. O’Clair, 1972 (St. Makarius Bay, Amchitka

Island). Southeastern Alaska: 23 specimens

from Bousfield and McAllister 1961 stns.,

A99, All4, A147, A168; 2 specimens from

Bousfield 1980 stn. S18B1.

British Columbia — Queen Charlotte Islands:

3 specimens from Bousfield 1957 stns., WI,

W9a. | specimen from collection of M. Frazer,

1935. Northern mainland: 9 specimens from

Bousfield 1964 stns., H12, H16. Vancouver

Island: 2 specimens from Bousfield 1975

stns., P5a; 25 specimens from Bousfield 1970

stns., P702; P710; P712, P715; 2 Spegemens

from Bousfield 1959 stn., N1; 4 specimens

from the collections of R.K. Lee, 1973 and

N.A. Powell, 1966. Washington and Oregon

— 10 specimens from Bousfield 1966 stns.,

4 OS il

ISS

eee ea eaee

Figure 10. Ampithoe simulans Alderman & 18.0 mm; & subadult 13.0 mm; 2 15.5 mm, Gonzales Bay,

Victoria, B.C. 29 July 1970

W34, W35, WS50; 1 specimen from the collec-

tion of N. McDaniel.

Smithsonian collections (USNM): R.K. Lee stn.

oe. io, 1 Y.

Distribution: Known authentically from St.

Makarius Bay, Amchitka Island, Alaska (51°N,

179°W) south to Cannon Beach, Clatsop Co.,

Oregon (45°54.5’N, 123°58’W).

Ecology: A cold temperate species occurring

from low water level to 4 m depth, amongst

algae and eelgrass on semi-protected and

exposed coasts in summer __ temperatures

10-16°C, salinities 29-34°/,,, occasionally in

brackish water.

Diagnosis: Head, antennal sinus slight, eye

larger than in A. dalli, Antenna 2 poorly setose.

Mandibular palp segment 2 more than twice the

length of segment 1 and bearing | or 2 setae;

segment 3 very oblique, passing by a barely

perceptible angle into the lower margin and

setose for nearly the full length. Gnathopod 1

palm less oblique than in A. rubricata, palm

verging into the hind margin at a definite angle;

male segment 2 with a few plumose setae on the

hind margin only (beginning at 11-15 mm body

length), but plumosity not extending to other

segments. Gnathopod 2 (o'), segment 5 with a

low dorsal tubercle; hand enlarging with age and

sides diverging, palm becoming increasingly

concave, hind corner lengthening to form a

broad tooth, a smaller medial tooth developing

at the dactyl hinge; dactyl increasingly curved

with age but always meeting the long hind

tooth. Peraeopod 7 segment 6 with 6-8 marginal

spine groups (depending on age). Epimeron 3

rectangular, hind corner indented more strongly

than in A. dalli. Uropod 1 with not more than

6 spines on the outer peduncle and 10 spines on

the outer ramus. Uropod 3 with 6 crown spines

and mo central spines. Body colour in life:

uniformly green to brown speckled. Body length

at maturity: Male 11-23 mm, female 12-30 mm.

Remarks: Alderman’s description of the type

male is based on the characteristics of a subadult:

viz. the absence of plumose setae on gnathopod 1,

a hardly enlarged gnathopod 2 that lacks a hinge

tooth but retains a serrated dactyl. His speci-

mens were 12-24 mm in length, a size at which

plumosity and enlargement of the gnathopod

would have occurred if the specimen had been

A. dalli.

The two palmar teeth of the male gnathopod 2

of A. dalli and A. simulans grow at the location of

and eventually replace the obturator spine and

slender dactyl hinge spine (see Fig. 10). The

large teeth probably prevent the enlarged dactyl

from crimping the female during amplexus.

Barnard (1965) suggested that Ampithoe

simulans is probably synonymous with Ampithoe

corallina Stout (1913), in the belief that Stout

might have confused the male with a female

because of the undifferentiated second gnathopod

of the male. However, Stout mentioned that

females of 6-8 mm were carrying eggs, indicating

that this species matures at a much smaller size

than A. simulans. A. corallina differs also in the

following: head very deep and broad, relatively

strongly setose antennae, large number of seg-

ments in antennal flagella, lower lip outer lobes

subequal in size, mandibular palp obliquely

acute with only 6 setae.

The two aberrant specimens of A. simulans

shown by Barnard to have a deeply incised hand

of gnathopod 2 are indeed not this species but

males of Ampithoe sectimanus n.sp.

Genus Peramphithoe new genus

Type species: Ampithoe femorata Kréyer 1845

Component species: A. orientalis Dana 1853,

A. humeralis Stimpson 1864, A. eoa Bruggen

1907, A. falsa K.H. Barnard 1932, A. annen-

kovae Gurjanova 1938, <A. lindbergi |

Gurjanova 1938, A. mea Gurjanova 1938, ¢

A. plea Barnard 1965, A. tea Barnard 1965,

A. spuria Krapp-Schickel 1978, A. aorangi

Barnard 1979, P. humeralis (not Stimpson

1864) of Griffiths, 1979, P. lessoniophila

n.sp.

Diagnosis: Head lobe produced, antennal sinus

present. Antenna | accessory flagellum absent.

Mandibular palp moderately weak. Maxilla 1

palp slender. Gnathopod 1, palm transverse,

coxa 1 not produced forward. Gnathopod 2

subchelate, equal to or larger than 1. Peraeopods

3 and 4, segment 2 strongly inflated. Peraeopods

5-7, segment 6 not distally expanded, spines

usually not restricted to the antero-distal

region. Uropod 1 peduncular process well

developed. Uropod 3, outer ramus with two

hooked uncini. Telson with two small apical

cusps.

About 14 species, arctic-boreal to tropical,

littoral.

Etymology: From the Greek, pera meaning

beyond or across, referring both to the trans-

verse form of the first gnathopod and the

advanced form in relation to Amphithoe. The

correct spelling of ‘Amphithoe’ is herein applied

as derived from the Greek amphi = around, both,

and thoé = quick.

Northeastern Pacific species: Peramphithoe

humeralis (Stimpson 1864), P. mea (Gurjanova

1938), P. lindbergi (Gurjanova 1938), P. tea

(Barnard 1965), P. plea (Barnard 1965).

Key to Species of Peramphithoe

of the Northeastern Pacific

1. Body large, 19-35 mm at maturity. Antenna 2 flagellum slender, proximal segments not

fused. Gnathopod 1, segment 5 shorter than segment 6. Peraeopod 5, segment 4 longer than 5.

Male gnathopod 2 hardly enlarged, dactly less than twice the length of the dactyl of

gnathopod |

Body small, 6-12 mm at maturity. Antenna 2 flagellum heavy, proximal segments fused.

Gnathopod 1, segment 5 equal to or longer than segment 6. Peraeopod 5, segments 4 and 5

subequal. Male gnathopod 2 enlarged, dactyl more than twice the length of the dactyl of

gnathopod 1

2. | Gnathopod 1, segment 5 longer than 6. Gnathopod 2 (both sexes) subequal to 1, palm trans-

verse; segment 5 longer than deep and as long as segment 6. Peraeopod 7 larger and about one

third longer than peraeopod 6

Peramphithoe humeralis (Stimpson 1864) (p. 61)

Gnathopod |, segment 5 about equal in length to 6. Gnathopod 2 (both sexes) larger than 1,

palm oblique; segment 5 not longer than deep and shorter than segment 6; peraeopod 7

hardly larger than peraeopod 6

Peramphithoe mea (Gurjanova 1938) (p. 63)

3. Antenna | peduncle | lacking a postero-distal spine. Antenna 2 slender and nearly as long as

antenna |. Gnathopod | dactyl overlapping the palm by hardly more than the length of the

nail. Gnathopod 2 (o), hand rectangular, dactyl sinuous, palm with a low rounded palmar

process at dactyl hinge

oeeeeeeeee eee ee ee ee © we we we ee wee He ww

Peramphithoe plea (Barnard 1965) (p. 67)

Antenna | peduncle with a posterodistal spine in mature individuals. Antenna 2 heavy, less

than */, length of antenna |. Gnathopod | dactyl overlapping the palm by considerably more

than the length of the nail. Gnathopod 2 () proximally broader, dactyl evenly curved, palm

with or without process at hinge

Ce |

4. Antenna 2 about */, the length of antenna 1; flagellum 1 times the length of peduncle 5,

proximal segments of flagellum fused in pairs. Gnathopod 2 (’), palm with tubercle at

dactyl hinge, dactyl reaching with age to the length of the hand

eoeeeveveer eee eeeeeeee eee ee ee ew ee ee we wee we ee ee ew ww ee we wo

ee )

Peramphithoe tea (Barnard 1965) (p. 65)

Antenna 2 about ” length of antenna 1; flagellum about equal to the length of peduncle 5,

proximal 5-6 segments of flagellum fused together. Gnathopod 2 (o’), palm lacking tubercle

at hinge, dactyl never more than half the length of the hand

eeceoveveveevee eee eee eee ee ee we ew wee em hh Oh em hm ho hm hm hm Om hh Ow

he Bastern North Pacific species of

Peramphithoe are divisible into two groups, which

may later warrant formal taxonomic recognition —

1) P. humeralis, F. mea

2) P. lindbergi, P. tea, P. plea

Characteristics which order the species are,

progressive decrease in body size, compression

of the antenna 2 flagellum and increase in sexual

dimorphism. This may represent an evolutionary

gradient.

Group of Peramphithoe humeralis and P. mea

Body large, 19-35 mm at maturity. Eye medium

to small. Antenna 2 flagellum slender, proximal

segments not fused. Lower lip outer lobes, apical

longer than medial. Gnathopod 1, segment 5

longer than or equal to 6. Gnathopod 2 not

greatly enlarged, scarcely sexually dimorphic.

Peraeopods 3 and 4, segment 4 barely extending

over segment 5. Peraeopod 5, segment 4 longer

than 5. Peraeopods 6 and 7 anterior and posterior

setae short, about equal in length. Pleopods

with 8-9 coupling hooks.

Peramphithoe humeralis (Stimpson 1864)

Figure 11.

Amphithoe humeralis Stimpson 1864, p. 156;

@aaman, 1898, p:° 271-273, pl. 33; fig. 4;

Holmes, 1904, p. 241; Hewatt, 1946, p. 199,

204.

Ampithoe humeralis: Stebbing, 1906, p. 636;

oeeeeeeeeee eee eee ee ee ewe ee ew ee

Peramphithoe lindbergi (Gurjanova 1938) (p. 64)

J.L. Barnard, 1954, p. 29; J.L. Barnard, 1965,

p. 7, figs. 2, 3; J.L. Barnard 1969b, p. 83,

not Griffiths, 1979, p. 131-138, figs. 1-3.

Material examined: Alaska — southeastern

coast: 36 specimens from Bousfield and McAllis-

ter 1961 stns., A3, A6, A59, A75, A80, A811,

A83, A131, A151, A168, A171-2, A174, A175;

1 specimen from Bousfield 1980 stn. S23F1.

British Columbia — Queen Charlotte Islands: 11

specimens from Bousfield 1957 stns., ES, H2a,

H3, H14, W1; 5 specimens from the collection of

W. Spreadborough. Northern mainland: 6 speci-

mens from Bousfield 1964 stns., H3, H7, H26,

H30, H47, H65. Vancouver Island and southern

mainland: 2 specimens from Bousfield 1977

stns., B6b, B8; 1 specimen from Bousfield 1975

stn., P5c; 18 specimens from Bousfield 1970 stns.,

P710, “P71 P7127 P7135; 3 specimens irom

Bousfield 1959 stns., 015, V4b; 3 specimens from

Bousfield 1955 stns., F8, P7; 23 specimens from

the“ collections of \J.F:L. “Hart, D: “Kittle, N-A:

Powell, D.B. Quayle, D. Zittin. Washington — |

specimen from Bousfield 1966 stn., W35; 4 speci-

mens from the collection of R.M. O’Clair, 1974.

Smithsonian collections (USNM): Bousfield 1961

sta ATS, 2°e¢ of} ) 9 -subadult, ‘1 O?paveniic:

3 immatures; stn. A6, 1 9, 1 immature.

Distribution: Prince William Sound, Alaska

(60°40’N,145°36’W), south to Guadalupe Island,

Baja California.

Ecology: Occurs amongst eelgrass and kelp,

Figure 11. Peramphithoe humeralis (Stimpson) & 15.0mm; ? 19.0 mm, Trevor Channel, Barkely Sound,

Vancouver Is., B.C. 25 May 1977

curling a frond around itself to form a tube.

Several animals, such as a brood of young

accompanying the parent may use the same tube

(Jones, 1971). Mainly intertidal in the north,

subtidal in California to a depth of 70 m. Usually

in high salinity, exposed or semi-protected shores

where summer water temperature is 9.5-14.5°C,

salinity 14.3-32.7°/o.. Females ovigerous June

to August.

Diagnosis: Eye medium. Gnathopod 1, segment

5 longer than segment 6. Gnathopod 2, both

sexes, not enlarged, palm transverse, segment 5

equal to or longer than 6, hind lobe broad and

rounded. Peraeopod 5 segment 2 nearly as broad

as long, lower hind edge concave. Peraeopod 7

much stronger than 6 and about a third longer,

segment 6 with about 12 spine groups. Uropod 3

peduncle long, more than twice the length of the

rami. Body colour in life: uniformly orange to

brown. Body length at maturity: male up to

34.5 mm, penial papillae developing at 5 mm;

female 19-35 mm.

Remarks: Superficially, an immature specimen

may resemble the female or immature of another

species. If the specimen is 8 mm or greater it

can be recognized within the genus by the lack

of brood plates and unaltered gnathopod 2.

Other distinguishing features are the long

segment 5 of gnathopods 1 and 2, slender

antenna 2 and long peraeopod 7. The South

African Ampithoe humeralis of Griffiths (1979)

differs from the North American type in the form

of the antennae, coxae, gnathopods, peraeopods

5-7, uropod 3, mandibular palp, lower lip and

size of maturity. These differences are sufficient

to designate the South African specimens as a

new species. Its habits are similar to those of the

North American P. humeralis in rolling a kelp

blade into a tube and living in a colony. The

upper walls of the tube are consumed by the

occupants, necessitating progressive extension

of the tube downwards (Jones, 1971).

Peramphithoe mea (Gurjanova 1938)

Figure 12.

Amphithoe mea Gurjanova 1938, p. 361-364,

fig. 53; 1951, p. 882-885, fig. 616; ? J.L.

Barnard, 1966, p. 60

Material examined: Alaska — Aleutian

Islands: Stag Point, Deer Island, 1 male from

the collection of P. McRoy and P.A. Lebednik,

1970; Cold Bay, Amchitka Island, 1 male,

2 immature females from the collection of

een, O’Clair 1970. |

Distribution: Aleutian Islands, Alaska (51°N,

179°W); Japan Sea (45°N,130°E).

Ecology: A cold water species found amongst

eelgrass and algae at 5-60 m depth, rarely

intertidal.

Diagnosis: Eye small. Gnathopod 1, segment 5

about equal in length to segment 6. Gnathopod 2,

Sages ot

Pa. GN2

both sexes, larger than 1, palm oblique; segment 5

shorter than 6, narrowed into an acute hind lobe.

Peraeopod 5 segment 2 slender, longer than wide,

lower hind edge evenly rounded. Peraeopod 7

little stronger than 6, segment 6 with about 9

spine groups. Uropod 3 peduncle normal, twice

the length of the rami. Body length at maturity:

male 18-22 mm, female probably about the same.

Remarks: The specimens examined closely

resemble Gurjanova’s holotype but lack a spine

on the postero-distal margin of antenna 1,

peduncle |. In view of the large size reached by

this species (18-22 mm), the probability, as

suggested by Barnard (1965), that P. mea is

simply an early growth stage of the much smaller

(maximum 8 mm) P. annenkovae is highly unlikely.

Barnard’s (1965) inclusion of his Ampithoe

sp., within P. mea is unjustified. This species is

only 8 mm yet has a more strongly developed

second gnathopod resembling that of P. lindbergi,

a very large eye, shorter segment 5 on the first

Jv'Q°6en2

‘

Figure 12. Peramphithoe mea (Gurjanova) o& 18.0 mm, Deer Is., Aleutian Is., Alaska. 17 Sept. 1970

gnathopod, much more expanded segment 2 and

shortened segment 4 on peraeopod 5 and

generally more setose appendages.

Barnard’s (1969b) inclusion of his Oregon

Ampithoe eoa (Barnard 1954, not P. eoa

(Bruggen 1970)) is similarly not justified. At

10 mm the male gnathopod 1s greatly enlarged,

similar in shape to that of P. tea with two palmar

tubercles, segment 5 of the first gnathopod is

shorter than segment 6 and more strongly lobed,

segment 4 of peraeopods 3 and 4 are more

strongly expanded, segment 2 of peraeopod 5 is

broader than deep and segment 4 is subequal to 5,

uropod 2 peduncular process is very small,

uropod 3 peduncle bears a central spine, the

appendages are more strongly setose, and the

outer lobes of the lower lip are equal in length.

Peramphithoe eoa (Bruggen 1907) is probably

also a separate species, rather than a later growth

stage as Barnard (1965) suggested. In this species

the apical lobes of the lower lip are much longer

than the medial, the palp of maxilla | is broader

and more spinose, the peduncle of uropod 3 is

much longer, the gnathopod 2 of the female is

less oblique and the palm crenulate, and in the

male the hand of the second gnathopod is

rectangular and the dactyl much longer, reaching

back to segment 4.

Group of Peramphithoe lindbergi, P. tea and

P. plea

Body small, 6-12 mm at maturity. Eye moderately

large. Antenna 2 flagellum heavy, 5-6 proximal

segments fused. Lower lip outer lobes, apical

and medial subequal. Gnathopod 1, segment 5

shorter than 6. Gnathopod 2 enlarged, strongly

sexually dimorphic. Peraeopods 3 and 4, segment

4 extending downwards over segment 5S.

Peraeopod 5, segments 4 and 5 subequal.

Peraeopods 6 and 7, posterior setae longer than

anterior setae. Pleopods with 5-6 coupling hooks.

Peramphithoe lindbergi (Gurjanova 1938)

Figure 13.

Amphithoe lindbergi Gurjanova 1938, p. 351-

354, fig. 49; 1951, p. 892-895, fig. 620

Figure 13. Peramphithoe lindbergi (Gurjanova) & 11.0 mm; ¢? 10.0 mm, Haines Is., Barkley Sound,

Vancouver Is., B.C. 8 August 1975

Ampithoe femorata Kréyer 1845: J.L. Barnard,

1952, p. 24-28, pls. 6, 7 (not Krdyer, 1845)

Ampithoe lindbergi: J.L. Barnard, 1965,

p. 12-15, figs. 6, 7; 1969b, p. 83, 84

Material examined: Alaska — southeastern

coast: 37 specimens from Bousfield and

McAllister 1961 stns., A68, A70, A80, A86, A90,

A92, A115, A131, A151, A174, A175.

British Columbia — Queen Charlotte Islands:

13 specimens from Bousfield 1957 stns., E25,

H4a, H9, H11, W2. Northern mainland: 20 spe-

cimens from Bousfield 1964 stns., H4, HS, H17,

H29, H33, H47, H53, H65. Vancouver Island and

southern mainland: 4 specimens from Bousfield

1977 stn., E3; 1 specimen from Bousfield 1975

stn., P25; 2 specimens from Bousfield 1970 stns.,

P715, P721; 1 specimen from Bousfield 1964 stn..,

H43; 15 specimens from Bousfield 1959 stns.,

meet, V5, V10, V17, V19, V20; 1 specimen

from Bousfield 1957 stn. P2; 3 specimens from

Bousfield 1955 stns., F2, F6; 15 specimens from

the collection of L. Daniels 1975. Smithsonian

collections (USNM): Daniels 1975 collection

moo, ! 2,1 9 ‘subadult.

Distribution: Prince William Sound, Alaska

(60°43’N,146°7’W) south to Corona del Mar,

California (35°5’N,118°W); Bering Sea, Okhotsk

Sea, Japan Sea (45°N,130°E).

Ecology: Found amongst eelgrass and algal

holdfasts at low water level to 18 m depth on

exposed and protected coasts. Summer tempera-

tures: 9.8-17.5°C, salinity 14.8-33+°/,.. Females

ovigerous June to September.

Diagnosis: Body stout. Antenna | peduncle |

with a postero-distal spine in mature individuals.

Antenna 2 short and stout, '/, length of antenna 1;

flagellum strongly setose, proximal 4-7 segments

of flagellum fused. Gnathopod 1 dactyl over-

lapping the palm by considerably more than the

length of the nail. Gnathopod 2 (@), lower lobe

of segment 5 pointed. Gnathopod 2 (0°), hand

broadened, palm oblique, somewhat concave

but without a tubercle at the dactyl hinge, dactyl

evenly curved, never extending more than half

the length of the hand. Peraeopod 5, lower hind

edge of segment 2 straight or concave. Peraeopods

6 and 7 moderately to strongly setose, segment 2

proximally broadened. Uropods 1 and 2, rami

broad. Body colour in life: uniformly yellow to

olive green. Antennae yellow, red and white

banded. Body length at maturity: Male

6-10.5 mm, female 6.5-12.5 mm.

Remarks: The length of the flagellum of

antenna 2 is quite variable and the loss of

segments in the male as described by Barnard

(1965) seems to represent a difference in the

degree of segment fusion. A count of the number

of setal bundles on flagellum 2 of 22 individuals

gave a value of 8 in 7 mm specimens, to 12-14 in

10-12 mm specimens, consistently about 2 bundles

less than in Peramphithoe tea. Immatures and

females which lack antennae 2 are very difficult

to distinguish from P. tea.

P. lindbergi cannot be a younger stage of

P. annenkovae or P. plea, as Barnard (1965)

Suggested, in view of the differences stated

herein, the nearly identical range in size at

maturity of P. plea (7-10 mm), and the smaller

size at maturity of P. annenkovae (max. 8 mm).

Peramphithoe tea (Barnard 1965)

Figure 14.

Ampithoe tea Barnard 1965, p. 30-34, figs.

19—21; ? 1969b, p. 85

Material examined: Alaska — southeastern

coast: 10 specimens from Bousfield and McAllis-

ter 1961 stns., A3, A8, Al8, A91, A92, A96,

A136, A164, A171-2; 8 specimens from Bousfield

[980 stns., SILI, SI8P3, S23F 1.

British Columbia — Queen Charlotte Islands: 7

specimens from Bousfield 1957 stns., H4a, El4a,

W14. Northern mainland: 46 specimens from

Bousfield 1964 stns., H8, H12, H22, H25, H26,

H30. Vancouver Island and southern mainland:

3 specimens from Bousfield 1977 stns., B7a, B21b;

3 specimens from Bousfield 1975 stns., P5b, P25;

2 specimens from Bousfield 1970 stns., P706,

P712; 11 specimens from Bousfield 1955 stns.,

Fl, P6a. In addition, 9 specimens from the

collections of D.V. Ellis, J.F.L. Hart, OR. Vee

N.A. Powell, J.W. Scoggan and C.H. Young and

W. Spreadborough.

Smithsonian collections (USNM): Bousfield

1961 stn. A91, 1 o&; Bousfield 1961 stn. A92, 1 9.

Distribution: Prince William Sound, Alaska

(60°46’N,146°31’W), south to Baja California

QPN, LIS? WwW).

Ecology: Intertidal to a depth of 67 m, amongst

algae on exposed and semi-protected high salinity

coasts, in summer temperatures of 9.7-14.5°C

and salinities 17.0-33+°/... Females ovigerous

May to August.

Diagnosis: Body relatively stout. Antenna 1,

peduncle segment | with a postero-distal spine

in mature individuals. Antenna 2 heavy, about */,

the length of antenna 1, flagellum moderately

Figure 14. Peramphithoe tea (Barnard) 0 8.0mm; 2 7.5 mm, Whiffen Spit, Sooke, Vancouver Is., B.C.

11 August 1969. Intersex, Yakoun Bay, Queen Charlotte Islands, B.C.

setose, proximal segments of flagellum fused in

pairs. Gnathopod 1 dactyl overlapping the

palm by considerably more than the length of

the nail. Gnathopod 2 ( 9 ), lower lobe of segment

5 pointed. Gnathopod 2 (oc), hand broadened,

palm crenulate; at about 7.5 mm body length,

palm developing a square or crenulate tubercle

near the dactyl hinge. Dactyl evenly curved or

slightly sinuous, increasing with age to the full

length of the hand. Peraeopod 5, lower hind edge

of segment 2 straight or concave. Peraeopods 6

and 7 moderately setose, segment 2 proximally

broadened. Uropods | and 2, rami broad. Body

length at maturity: male 7.5-12 mm, female

6-10 mm.

Remarks: The male illustrated herein, the same

size as Barnard’s holotype, has a much shorter

dactyl on the second gnathopod and is less

strongly setose. In no specimens in this collection

is the dactyl more than */, the length of the palm,

although body lengths are comparable.

The tubercle in the palm of the male second

gnathopod of this species and P. plea probably

arises from the obturator spine and may act to

prevent the dactyl from closing too tightly. The

first gnathopod sometimes appears to be para-

chelate, a tendency that Barnard (1970) noted in

a new Hawaiian species.

Peramphithoe plea (Barnard 1965)

Figure 15.

Ampithoe plea Barnard 1965, p. 15-20, figs. 9, 10

Material examined: British Columbia — Queen

Charlotte Islands: 2 immatures from Image Point,

Skidegate Inlet (Bousfield, 1957 collection, stn.

E5). Vancouver Island: Dodger Channel, Barkley

Sound; mature male and female from the collec-

tion of D. Zittin, 1976.

Distribution: Queen Charlotte Islands, B.C.

(53°15’N,132°00’W), south to Santa Barbara,

California (34°N,120°W).

Ecology: Occurs amongst kelp holdfasts on

high salinity exposed coasts, intertidally in the

north and subtidally to 17 m in the south.

Diagnosis: Body slender. Antenna | peduncle |

lacking a ventral distal spine. Antenna 2 long

and slender, '/, to */, length of antenna 1;

flagellum weakly setose, proximal 2-4 segments

fused. Gnathopod | dactyl overlapping the palm

by only about the length of the nail. Gnathopod 2

(2), lower lobe of segment 5 rounded. Gnatho-

pod 2 (co), hand rectangular, dactyl sinuous,

increasing with age to the full length of the

hand; palm smooth, with a low, rounded process

near the dactyl hinge. Peraeopod 5, lower hind

edge of segment 2 convex, evenly rounded.

Peraeopods 6 and 7 weakly setose, segment 2

slender, margins parallel. Uropods | and 2, rami

slender. Body length at maturity: Male 7-10 mm,

female 7.5-12.5 mm.

Remarks: The individuals described here differ

Figure 15. Peramphithoe plea (Barnard) & 10.0 mm; ? 12.5 mm, Dodger Channel, Barkley Sound,

Vancouver Is., B.C. 28 June 1976. Queen Charlotte Is., B.C. 19 August 1957

slightly from Barnard’s holotype in that antenna 2

flagellum is less setose, the dactyl of gnathopod 1

is somewhat longer and the dactyl of the male

gnathopod 2, although of a specimen 1.5 mm

longer than the holotype, does not reach the

full length of the hand.

The two species from the boreal coast of South

America warrant inclusion in the monograph

because their morphology verifies conclusions

justifying the transfer of the transverse-handed

Ampithoe to the new genus Peramphithoe. They

meet all diagnostic criteria for the genus and

most closely resemble members of the P. lindbergi

group. The first appears to be, from the descrip-

tions of Stebbing (1906) and Kreibohm de

Paternoster & Escofet (1976), Ampithoe femorata

Kréyer 1845 (herein re-described and assigned

the new generic title). The second is a new species

found in colonies amongst the fronds of Lessonia

in self-constructed chambers (H.K. Schminke,

pers. comm.). It is consequently assigned the

name Peramphithoe lessoniophila.

Peramphithoe femorata (Kréyer 1845)

Figure 16.

Ampithoe femorata Kréyer 1845, p. 335,

figs. 4a-1

Amphithoe femorata: Stebbing, 1906, p. 636-

637; Chilton, 1921, p. 88; Schellenberg, 1931,

p. 245; Kreibohm de Paternoster & Escofet,

Figure 16. Peramphithoe femorata (Kréyer) & 18.0 mm; o& subadult 12.5 mm; 2 subadult 13.0 mm.

Cape Horn Is., South America. 1970.

1976, p. 77-91, figs. 1-3; (not J.L. Barnard,

1952, p. 24-28, pls. 6-7); Alonso, 1980,

p, 4-5, fig. 1...

? Amphitoe gaudichaudii H. Milne Edwards 1840,

p, 31

Amphithoe brevipes Dana 1852, p. 216; 1853-

55, p. 941, pl. 64, figs. 5a-n.

Ampithoe brevipes: Stebbing, 1906, p. 637;

1914, p. 371, ? K.H. Barnard, 1916, p. 255-

256, fig. 34.

? Amphithoe peregrina Dana 1853-55, p. 940,

pl. 64, figs. 4a-b; 1862, p. 247, pl. 43, fig. 1

? Amphithoe falklandi Bate 1862, pp. 237-248,

ies; 1, 2, 6. 7

2 Amphithoe rubricata: Della Valle,

p. 456, 459 (not Montagu 1808)

? Grubia crassicornis: Della Valle, 1893, p. 456

Material examined: Cape Horn Island, South

America (55°S,77°W), 1970 stn. 27896, J. Mark-

ham collector, 1 adult o’, 1 subadult o, 1 sub-

adult 9, 1 immature; Banco de los Tacas, Isla

Navarino, South Chile (55°05’S,67°04’W),

5 February 1970, stn. 27924, J. Markham collec-

tor, 9.5°C, 2 subadult dc.

Distribution: New Zealand, South Africa, Chile,

Argentina, Brazil?.

Ecology: No sampling data available. Kreibohm

de Paternoster and Escofet (1976) found the

species in tubes rolled in the fronds of Macro-

cystis pyrifera.

Diagnosis: Body relatively stout, eye medium.

Antenna | long, about % body length; peduncle

segment 1 without a posterodistal spine.

Antenna 2 stout, about % length of antenna 1,

flagellum strongly setose, proximal segments of

flagellum fused. Lower lip outer lobes, apical

subequal to medial. Gnathopod 1, segment 5

somewhat shorter than 6, dactyl overlapping the

palm by considerably more than the length of

the nail. Gnathopod 2 (o'), hand enlarged, palm

crenulated, nearly straight, bearing a tubercle;

dactyl sinuous, reaching to 7/, the length of the

hand; in the juvenile (10.5 mm and 12.5 mm),

palm slightly concave, not crenulated, tubercle

absent; dactyl evenly curved, '/, - '/, length of

the hand. Gnathopod 2 (92) segment 5 shorter

than 6 and narrowed into a posterior lobe;

segment 6 similar in form to, but broader than

segment 6 of gnathopod 1. Peraeopods 3 and 4,

segment 4 strongly overhanging segment 5.

Peraeopod 5, lower hind edge of segment 2 evenly

curved or slightly concave; segment 4 slightly

longer than 5. Peraeopods 6 and 7 moderately

1893,

setose, segment 2 proximally broadened. Uropods

1 and 2 slender. Body length reaching 22 mm.

Peramphithoe lessoniophila n.sp.

Figure 17.

Material examined: Near Coquimbo, Chile

(30°S,71°W). A. Viviani, collector. Holotype o

(NMC-C-1981-964); allotype 9 (NMC-C-1981-

965); paratypes (NMC-C-198 1-966).

Smithsonian collections (USNM): 1 #, 1 9.

Ecology: Found living in little chambers self-

constructed in the fronds of Lessonia (H.K.

Schminke, pers. comm.)

Description of male holotype, 9.5 mm: Body

stout, eye medium. Antenna | short, less than

'/, the body length; peduncle segment | without

a posterodistal spine. Antenna 2 about */, the

length of antenna 1, flagellum strongly setose,

proximal segments of flagellum fused. Lower lip

outer lobes, apical longer than medial. Gnatho-

pod 1, segment 5 subequal to 6, dactyl over-

lapping the palm by more than the length of the

nail. Gnathopod 2, hand enlarged, palm smoothly

concave; dactyl evenly curved, reaching to '/, the

length of the hand. Peraeopods 3 and 4, segment 4

strongly overhanging segment 5. Peraeopod 5,

lower hind edge of segment 2 concave; segment 4

slightly longer than 5. Peraeopods 6 and 7

moderately setose, segment 2 proximally

broadened. Uropods 1 and 2 slender. Body

length: male 9.5 mm, female 7-8.5 mm.

Description of female allotype, 7 mm: Gnatho-

pod 1 as in the male. Gnathopod 2, segment 5

shorter than 6 and narrowed into a posterior

lobe; segment 6 similar in form to, but broader

than segment 6 of gnathopod 1. Appendages

otherwise as in the male.

Etymology: “‘Lessonia - loving”, referring to

the construction of tubes in the fronds of the

kelp Lessonia.

Discussion and Conclusions

Biogeography and Ecology

Tables 2 and 3 summarize the geographic and

ecological distributions of the northern Pacific

Ampithoidae. The species are divisible into the

following groups:

1. Pan-Pacific, cold water species further sub-

divisible into

a) subarctic species Ampithoe volki, Peramphi-

thoe mea, and

b)subarctic and boreal species Ampithoe

ly é

A] f

\ ;

} 4

Figure 17. Peramphithoe lessoniophila n.sp. & 9.5 mm; 2 7.0 mm. Near Coquimbo, Chile

kussakini, A. lacertosa and P. lindbergi,

these occurring in a broader range from

Alaska to California. The species generally

occur in high salinity exposed and semi-

protected coasts although A. Jacertosa,

having the broadest geographical and eco-

logical range, occurs also in lower salinity

protected embayments and estuaries.

2. Aleutian-endemic, subarctic species Ampithoe

rubricatoides occuring in high salinity cold

waters.

3. American-endemic, boreal species Cymadusa

uncinata, Ampithoe sectimanus, A. dalli,

A. simulans, P. humeralis, P. tea and P. plea

occurring variously from southern Alaska to

California in exposed and semi-protected

meso- and polyhaline waters.

4. American-endemic, primarily warm water

species Ampithoe valida and A. plumulosa.

These occur south of central British Columbia.

A. valida occurs also in the Atlantic and is

restricted to low salinity protected coasts.

All species occur amongst algae and debris in

tide-pools, the intertidal and subtidal coast, to

the limits of the photic zone. Greatest diversity

in the northeastern Pacific is achieved in the

environs of Vancouver Island (12 species). Seven

species occur as far north as the Aleutian Islands

of Alaska and probably beyond.

Taxonomic Considerations

Table 4 shows that there are many differences

between the northeastern Pacific Cymadusa,

Ampithoe and Peramphithoe which, on further

investigation by Conlan (in press) prove to be

universal. All three genera are wide ranging in

both hemispheres, although Cymadusa and

Ampithoe concentrate in the tropics while

Peramphithoe predominates in boreal waters. The

more apomorphic Peramphithoe is less diverse,

and morphologically less variable than the more

plesiomorphic Cymadusa and Ampithoe. This is

demonstrated by the two South American species

of Peramphithoe which little differ from the

North American species. The alteration in palm

configuration in Peramphithoe, coincident with

strong development of the peraeopod spinning

glands and uropod spinous processes must incur

a measurable change in living and tube-building

habits. An examination of the evolutionary

relationships of these genera with other Amphi-

thoidae is presented separately (Conlan, in press).

Acknowledgements

The authors wish to thank the many agencies

and individual collectors who donated specimens.

In particular, C. Staude (University of Washing-

ton), C.D. Levings (Department of Fisheries and

Oceans, Canada), P. Lambert (B.C. Provincial

Museum) and S. Cross (University of Victoria)

provided special assistance. The preparation of

the illustrations by Mrs. F.E. Zittin, Van-

couver, B.C. is gratefully acknowledged.

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fauna No. 5. Pleonexes lessioniae, a new species of the family

Amphithoidae. Trans. R. Soc. N.Z. 81(4): 619-626

Jones, L.G. 1971. Studies on selected smal! herbivorous

invertebrates inhabiting Macrocystis canopies and holdfasts

in southern California kelp beds. Jn: North, W.J., ed. The

biology of giant kelp beds (Macrocystis) in California. Nova

Hedwigia, 32: 343-367

Kudryashov, V.A. 1979. The fauna and ecology of amphipod

crustaceans from the intertidal zone of the northern Tatar

Strait (in Russian). Trans. Acad. Sci. USSR Far East Sci. Cen.

No. 15: 123-136

Krapp-Schickel, G. 1978. Die gattung Amphithoe (Crustacea,

Amphipoda) in Mittelmeer. Bijdr. Dierkd. 48(1): 1-15

Kreibohm de Paternoster, I. and Escofet, A. 1976. La fauna de

anfipodes asociada a los bosques de Macrocystis pyrifera en el

chubut: Amphithoe femorata (Kréyer) (Ampithoidae) y

Bircenna fulva Chilton (Eophliantidae) Physis. Secc. A.

Buenos Aires 35(90): 77-91

Kunkel, B.W. 1910. The Amphipoda of Bermuda. Trans.

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Nicotri, M.E. 1980. Factors involved in herbivore food

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Ortiz, M. 1976. Un nuevo genero y una nueva especie de

anfipodo de aguas Cubanas (Amphipoda, Gammaridea,

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Paulmeier, F.P. 1905. Higher Crustacea of New York City.

Bull. 91, N.Y. St. Mus.

Rabindranath, P. 1972. Marine Gammaridea (Crustacea:

Amphipoda) from the Indian region. Family Ampithoidae.

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1969. Studi sui crostacei anfipodi LXVII. Terzo contributo

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1938. Three new species of the amphipod genus Ampithoe

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15-25

Sivaprakasam, T.E. 1970. Amphipods of the family Ampi-

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34(5-6): 633-659

Table 1. Comparative characters of the North American and type (USSR) specimen of Ampithoe volki

Character

Size at maturity

Antenna | — length

flagellum

Lower lip

Maxilla | inner plate

Maxilliped palp

Gnathopod 2 &

Epimeron 3 o&

Uropod 3 — peduncle

outer ramus

Type specimen (as published)

Gurjanova 1938

< 6 mm

1 slightly > 2

17-22 segments

apical= medial

lacking setae

segment 3 = segment 2

thumb triangular

segment 2 not lobate

posterodistal angle forms an

extended denticle

stout, nearly as wide as long

bearing a central spine

North American Specimens

> 8 mm (@ not ovigerous at

this size)

1 slightly to markedly < 2

14 segments

apical > medial

bearing | seta

segment 3 < 2

thumb square (but not appearing

to have been broken)

segment 2 lobate

posterodistal margin smoothly

angled

slender, length 1-'/, times

the width

lacking a central spine

Table 2. Distribution of Northeastern Pacific Ampithoidae listed in geographic order

Aleutian William to Dixon

Es.

Northern

Prince Cross Sd. B.C. and Central

Queen __—iB..C.. and

Sound Entrance Charlotte Vancouver

Species Alaska Alaska Alaska _ Is. B.C. Is., B.C. Washington Oregon California Other Records

Peramphithoe mea x a — — — — — _ USSR: Japan Sea

Ampithoe

rubricatoides xX — — — — — — —

Ampithoe volki xX _ — — — —- a — USSR: Japan Sea

Preobrazhen’e Bay

Ampithoe kussakini xX xX xX x O — — ao USSR: Shikotan Is.

(Otradnaya Bay)

Ampithoe dalli xX xX xX Xx x X xX —

Ampithoe simulans x x x Xx xX xX xX _

Ampithoe lacertosa O xX x 4 xX xX xX xX Japan: Schizueka

Prefecture

Ampithoe sectimanus _ x x x x — 4 —

Peramphithoe

humeralis — x Xx x xX 4 xX xX

Peramphithoe

lindbergi — 4 xX x xX x xX xX USSR: Bering Sea,

Okhotsk Sea,

Japan Sea

Peramphithoe tea — x xX xX xX x xX xX

Cymadusa uncinata — _— O O 4 O — x

Peramphithoe plea _— — — x xX x xX x

Ampithoe valida — —_ — — X X xX 4 Japan, US mid

Atlantic coast

Ampithoe plumulosa — — —_ — O — — xX Mexico, Ecuador,

Galapagos Is.

Number of species i! 8 9 10 12 9 9 8

X abundant, O occasional, — absent

Table 3. Habitats of Northeastern Pacific Ampithoidae

Coastal Exposure

Open and

semi-

Species

Peramphithoe lindbergi

Peramphithoe tea

Cymadusa uncinata

Peramphithoe plea

Ampithoe valida

Ampithoe plumulosa

X abundant,

Peramphithoe mea Xx

Ampithoe rubricatoides X

Ampithoe volki Xx

Ampithoe kussakini X

Ampithoe dalli X

Ampithoe simulans X

Ampithoe lacertosa X

Ampithoe sectimanus xX

Peramphithoe

humeralis X

wats’

O occasional,

Table 4. Character differences in the Northeastern Pacific Cymadusa, Ampithoe

Character

Antenna |, accessory flagellum

Mandible,

incisor, no. teeth

lacinia mobilis, no. teeth

no. spines

palp setosity

Maxilla 1,

inner plate, no. setae

palp, no. spines

no. setae

Maxilliped,

outer plate, inner margin

teeth

Gnathopod 1,

sexual dimorphism

obturator spine

coxa produced forward

Coxae 1-5,

lower margin setose

Peraeopods 3 and 4,

segment 2

segment 4

Pleopods,

no. coupling hooks

Uropod 1,

spinous peduncular process

Uropod 3,

no. peduncular marginal spines

outer ramus serrations

uncini

Telson,

no. setae at cusp

No. species

Percentage of total

— absent

Cymadusa Savigny

multisegmented

5-6

6-10

strong

smooth

absent

strong

yes

weakly inflated

moderately inflated

5-6

well developed

weak

medium

many

Salinity Range

Ampithoe (Leach)

vestigial

moderate to strong

0-1

5-12

3-7

fringed

usually smooth

present

strong

yes

rarely

weakly inflated

moderately inflated

6-11

vestigial or absent

0-13

weak

medium

l-many

Depth Range

Marine Meso- Oligo- _— Fresh-

polyhaline haline haline —- water LW- MW-

protected Protected (2 28°/0) (10-27°/o0) (1-9°/00) (< 1°Zo0) Subtidal MW HW

_ X — _— — to60m 0 ?

o- X _— — — tol&’m — ?

— Xx _ — to15m X in tidepools

O xX x O — to1l5m X_ in tidepools

» 4 Xx x O — tol0m X _ in tidepools

— x O O — to4m_ X in tidepools

xX x Xx O O to 10 m_ X_ in tidepools

— xX O _ — to3m _ X in tidepools

O xX O _ — to70m X_ in tidepools

O x O _ — to1l8m X_ in tidepools

O x O _ — to67m X in tidepools

O x — =_ — to7m = X in tidepools

— x — — — tol7m X in tidepools

x O x Xx — to 32m X in tidepools

NO DATA to 75 m_ X_ in tidepools

and Peramphithoe

Peramphithoe n.gen

absent

6-17

6-14

weak

—

-4

smooth

serrated

absent

weak

always

strongly inflated

4-8

well developed

1-6

strong

]

The Superfamily Corophioidea in the North Pacific Region.

Family Aoridae:

Systematics and Distributional Ecology

K.E. Conlan and E.L. Bousfield

National Museum of Natural Sciences

National Museums of Canada

Ottawa Canada

ABSTRACT

Amphipod crustaceans of the corophioidean family Aoridae were studied in extensive collections from

the northeastern Pacific coastal marine region. Redefined herewith is the genus Aoroides Walker and the

species A. columbiae Walker, Lembos (Lembos) concavus Stout and Lembos (Arctolembos) arcticus

(Hansen). Newly described are Columbaora new genus and C. cyclocoxa new species, Aoroides inermis

new species, A. intermedius new species, A. exilis new species, and A. spinosus new species. The geographic

range of Lembos (Lembos) concavus Stout is extended north from California to British Columbia and of

Lembos (Arctolembos) arcticus east to the Bering Sea. A key to recorded and probable species of Aoridae

of the northeastern Pacific region is provided and phyletic relationships are examined. Of the boreal

merochelate Aoridae, Columbaora is hypothesized to be closest to the ancestral form, from which

separately evolved Aora and Aoroides.

RESUME

Les auteurs ont étudié les Crustacés amphipodes corophioidés de la famille des Aoridae qui se trouvent

réunis dans d’importantes collections provenant des eaux cétiéres du nord-est du Pacifique. Le genre

Aoroides Walker et les espéces A. columbiae Walker, Lembos (Lembos) concavus Stout et Lembos (Arcto-

lembos) arcticus (Hansen) sont redéfinis. Le nouveau genre Columbaora et les nouvelles espéces C. cyclo-

coxa, Aoroides inermis, A. intermedius, A. exilis et A. spinosus sont décrits pour la premiére fois. Les

auteurs ont étendu l’aire de répartition de Lembos (Lembos) concavus Stout vers le nord depuis la

Californie jusqu’a la Colombie-Britannique, et celle de Lembos (Arctolembos) arcticus vers \’est jusqu’a

la mer de Béring. Une clé des espéces signalées et probables d’Aoridés du nord-est du Pacifique apparait

et les auteurs examinent les relations phylétiques. Ils formulent l’hypothése que, parmi les Aoridés

mérochéliformes de l’hémisphére boréal, Columbaora serait la forme la plus rapprochée de l’ancétre dont

auraient évolué séparément le genre endémique du Pacifique, Aoroides, et le genre endémique de

PAtlantique, Aora.

Introduction

The family Aoridae encompasses a diverse group Isaeidae and Photidae into the Corophiidae,

of tube-building corophioidean amphipods that producing a large family of diverse origins. Bous-

is diagnosed by a combination of morphological field (1973) and (1982) and Myers (1981) have

features, rather than by any single character. retained Aoridae as a separate family and

Myers (1969 p. 95) described the family as those expanded the. concept «to amcludes several

members of the Corophioidea ‘‘which exhibit convergently depressed-bodied genera such as

greater development of gnathopod 1 than Unciola, Pseudunciola and Rildardanus. The

gnathopod 2, coupled with a tendency towards separate family concept is maintained in this

retention of such primitive characters as an study.

accessory flagellum on the antennule, uropod 3 Of the 38 genera and more than 100 species

with two rami, contiguous coxae, and no marked currently included in the family Aoridae, only

depression of the pleon.”’ Barnard (1973) pro- one species, Aoroides columbiae Walker 1898, had

posed to combine the Aoridae, along with the previously been recorded from boreal eastern

Pacific waters. A second species, A. californica

Alderman 1936, had been synonymised with

A. columbiae by Barnard (1954). Apart from the

records of Thorsteinson (1941), no other illus-

trated documentation has been made of the

Aoridae from the east Pacific north of 48°

latitude. This report attempts to fill the distri-

butional hiatus and clarify the attendant

taxonomic problems through examination of

more than 6,000 specimens of Aoridae acquired

by the National Museum of Natural Sciences and

many individual collectors from Oregon to the

Aleutian Islands of Alaska. Station data are

recorded in Bousfield (1957, 1963) Bousfield and

McAllister (1962) and Bousfield (1968) for

expeditions to Vancouver Island and Georgia

Strait, B.C., Queen Charlotte Islands, the

southern Alaska coast and the north and central

British Columbia coast, respectively. Station data

for 1966 (Washington and Oregon), for 1970,

1975 to 1977 (Vancouver Island and southern

British Columbia mainland coast) and 1980

(southern Alaska coast) are provided in Bousfield

& Jarrett (1981).

In order to facilitate identification the keys

and descriptions are constructed with sex and

age independent characters, except where stated.

In addition, information on pigmentation pat-

terns is provided. These pigments should remain

in alcohol-preserved specimens for up to 10 years.

Table 5 lists the locality of specimens deposited

in the Smithsonian Institution (USNM) and the

Zoological Museum, Leningrad, USSR.

SYSTEMATIC SECTION

Family Aoridae Stebbing 1899

Barnard, J.L. (1969), p. 147

Myers, A.A. (1981), p. 9, 10

Bousfield (1973, p. 165 1979, 1982, p. 284)

Body smooth, slender, often depressed or

broadened. Colour pattern characteristically

disruptive. Mid-sternal spines or bulges occa-

sionally present; pleon and urosome occasionally

sparsely setose dorsally, segments rarely

coalesced. Head shallow, rostrum generally

absent or short. Eyes small, rounded, or lacking.

Interantennal head lobe not pronounced.

Antennae slender to stout, | usually the longer;

inferior antennal sinus shallow; peduncles long,

moderately spinose or setose, that of 2 often stout

or incrassate in o'; accessory flagellum multi-

articulate, short or vestigial, rarely lacking.

Upper lip rounded below, epistome not produced.

Mandible, molar strong; palp slender, segment 3

normally setose. Lower lip, mandibular lobes

often elongate. Maxilla 1, inner plates setose

or bare; outer plate usually with 10 apical spine-

teeth; palp large. Maxilla 2, normal, inner plate

bearing facial setae. Maxilliped, plates strong;

palp medium.

Coxae 1-4 medium deep to very shallow,

weakly overlapping or not contiguous; coxa |

often produced forward. Gnathopod 1 larger

than gnathopod Z, subchelate or, in the male,

often carpochelate or merochelate. Gnathopod 2

subchelate, segments 5 and 6 slender. Peraeopods

3 and 4 weakly glandular, basis not expanded.

Peraeopods 5-7 unequal, 7 longest, bases usually

differently expanded; coxa 5 shallower than

coxa 4. Pleopods normal, peduncle slender.

Abdominal sideplates shallow, weakly overlap-

ping or separated distally. Uropods 1 and 2

occasionally unequally biramous or uniramous,

postero-distal peduncular spinous process usually

present. Uropod 3 biramous or uniramous, outer

ramus occasionally minutely 2-segmented; apices

with simple spines lacking uncini. Telson entire,

fleshy with small dorsal subapical cusps. Coxal

gills sac-like, present on peraeopods 2-6. Brood

plates large, laminar; marginal setae simple.

The family now contains nearly 180 species in

38 genera. Size is generally small (2-12 mm) but

the deep water Neohela and the northern species

Lembos (Arctolembos) arcticus (Hansen) reach

30 mm. Most species are shallow marine and

estuarine and occur occasionally in tidal (rarely

supratidal) freshwaters; a few are bathyal and

abyssal. Diversity is greatest in tropical and

warm-temperate regions of the northern hemi-

sphere. Most members of the Aoridae are tube

builders but some species crawl freely on the sea

bottom and among sponges or occupy scaphopod

shells or tubes of other animals.

On the northeastern Pacific coast (northern

Alaska to northern California) the following

species are herewith recorded: Aoroides columbiae

Walker 1898, A. exilis n.sp., A. inermis n.sp.,

A. intermedius n.sp., A. spinosus n.sp., Colum-

baora cyclocoxa n. gen., n.sp., Lembos (Lembos)

concavus Stout 1913 and Lembos (Arctolembos)

articus (Hansen 1887). Aoroides secunda

Gurjanova 1938 could conceivably occur in the

northern regions of the northeastern Pacific;

species in the southern limits might include

Acuminodeutopus heteruropus Barnard 1959, Shoemaker 1942, Neohela pacifica Gurjanova

Amphideutopus oculatus Barnard 1959, Lembos 1953, Neomegamphopus roosevelti Shoemaker

(Lembos) audbettius Barnard 1962, L.(L.) macro- 1942, Rildardanus tros Barnard, and Rudilemboides

manus (Shoemaker 1925), Microdeutopus schmitti stenopropodus Barnard 1959.

Key to Genera of the Aoridae of the Northeastern Pacific

(Alaska to California)

rors) ANC 2-INWer FAN VESUIPIAl fo. ee eee eee eee essen yet oat Rildardanus Barnard

ees WAT) 2 MISE TEM WEIPORVCIODER 65 3 OU oe i ae Ss oe ew wad vies ve baw liale Oe i Z

IIS, ATIICE TALIMIS AOSD. fy cect a oh eceie dine vein cee bees tnd s eleva owelews Neohela Smith

Meonod 3, inner ramus short and scale-like or fully developed ......0c%...000ceevevereues 3

IMCS RIC TING VOI CAUIOEU Scie a Wore ate ws divest s Wa Rten es see eS eae Ne wt See a BIN 4

ISRO WLODER at bl iid ale PSs Vale Toe Onkol RS ye eee iy ie ee ele lee ee 6

4. _Uropod 3, inner ramus less than half the length of the outer .......... Acuminodeutopus Barnard

frenows, inner ramus more than half the length of the outer 2.45... 0.00. 6. ies sees dees 5

5. Antenna | segment 3 as long as segment 1. Mandibular palp segment 3 subclavate.

SIO? | (ues. ) CarOeiiel ate Faia. % ee OS 6 Fe dea Ses 2 ee Neomegamphopus Shoemaker

Antenna | segment 3 shorter than segment 1. Mandibular palp segment 3 slender, elongate.

memernonod le) usually carpochelate ey... incase. bane eee Rudilemboides Barnard

6. Antenna 1, accessory flagellum minute, vestigial. Mandibular palp slender, segment 3

cylindrical and weakly setose. Gnathopod 2 (c) merochelate, coxa rectangularly elongate

a bie ea brsies ena ore Os cbs vesbctten barn cacdaess a AOROIQES Wi aimct one

Antenna | accessory flagellum multisegmented. Mandibular palp well developed, segment 3

more or less falcate and strongly setose. Gnathopod 2 () usually carpochelate or subchelate

Be eLOCHMIALG, CORA CIFGUIDELY CRIAROEd 5. wwe eek een ea ve cn ea ee nde eee Nas eaten 7

7. | Gnathopod | (¢) carpochelate. Gnathopod | ( 2) segment 5 wider than 6, posterodistal

emermuev em omtt: Cty DANG a Ns. RENE « oe SOD. SSO Ee et Microdeutopus Costa

Gnathopod 1 (’) not carpochelate. Gnathopod | ( 2) segment 5 not wider than 6, postero-

EMer MORUOOLIE the Ei Sehe. . .. wa BRI RR CS 4 2s PN es 8

8. Pleon lacking a pair of setae on the dorsum. Gnathopod 1 (o’) merochelate, segment 5 much

longer than 6; coxa | (o) circularly enlarged. Gnathopod | ( 2), segment 5 subequal to 6;

meemronr slender distally-tapered)s icine. ...6 sets See Columbaora n. gen. (page 83)

Pleon with a pair of setae on the dorsum of at least one segment. Gnathopod | (c’) subchelate,

segment 5 shorter than 6; coxa | (o’) not circularly enlarged. Gnathopod | ( 2), segment 5

muemerinan.6;sepment6 distally broadened .. oi ..004.5 5 ee a Lembos Bate (page 80)

Key to North Pacific Species of Aoroides

meee ropod 2, peduncle lacking antero-distal spinous process .........0.000c0000 ve eeneeeees 2

/eeped 2, peduncle with antero-distal process well developed ...4.. 0.0. cc0c cans ete alee le 3

2. | Gnathopod 1 (c¢), anterior margin of segments 2 and 5 bearing dense plumose setae;

eaacnepod | ( 2); segment 2 lacking anterodistal group of setae «2.0.8... ede slew ewes’

heehee lst hel) cise ae ree Seen. 10's ee barh REE WS Aoroides secundus Gurjanova 1938

Gnathopod | (o’), anterior margin of segments 2 and 5 bearing a few simple setae only;

gnathopod 1 ( 9 ), segment 2 with well developed anterodistal group of setae ...............

Ut ar) ores REN ere wel P a isis. sada Wide Ghee be SBR Aoroides nahili J.L. Barnard 1970

3. | Uropod 3 outer ramus usually bare, rarely with 1-2 small spines. Mandibular palp, segment 2

setose. Gnathopod 2, both sexes, palm slightly oblique, dactyl overlapping by more than the

length of the nail; segment 2 of the female with a group of long setae on the distal anterior

eer ein. Male gnathopod 1,hind margin of segment 2 bare... 66. cee ee eee ce cue waee 4

Uropod 3 outer ramus with 1-3 strong spines. Mandibular palp, segment 2 bare. Gnathopod 2,

both sexes, palm transverse, dactyl overlapping by only the length of the nail; segment 2 of the

female without group of long setae. Male gnathopod 1, hind margin of segment 2 setose

4. Body pigmented in broad bands, parts of the head, segments 6 and 7 bare, giving a “‘saddle-

back”’ appearance. Maxilliped outer plate, teeth strongly serrated, lower teeth with at least 1-4

cusps each. Gnathopod 1 (o’), segment 5 broader than segment 2, dorsal margin of 5 bare

except for one distal group of short setae

A. columbiae Walker 1898 (page 89)

Body pigmented in discrete speckles throughout. Maxilliped outer plate, lower teeth with 0-2

cusps each. Gnathopod 1, (oo), segment 5 not broader than segment 2, dorsal margin of

5 setose

5. | Body pigment (speckles) never concentrated in the lower hind corners of body plates 1-5.

Maxilliped outer plate, teeth cusped. Gnathopod | (o’), dorsal margin of segment 5 with 5-7

bundles of setae

oeeee er eee eee eee ee ee we we we ewe hl Ohl Ohl Ohl Ohl Oh Ohh HO OO Hh OO OO Oh wo ow

A. intermedius n.sp. (page 87)

Body pigment (speckles) usually concentrated in the lower hind corners of body plates 1-5.

Maxilliped outer plate, all mid to lower teeth smooth. Gnathopod | (o’), dorsal margin of

segment 5 with 8-15 bundles of setae

oeee eee ee ee ee eee eh Om HH oO oO hm he wm em eh ewe we

A. inermis n.sp. (page 86)

6. | Body pigmented in broad bands, parts of the head, segments 6 and 7 bare, giving a “‘saddle-

back” appearance. Peraeopod 7 segment 2 slender (width/length = 39%). Gnathopod 1 (<),

segments 2 and 3 densely setose, setae as long as the width of the segment

A. exilis n.sp. (page 92)

Body diffusely speckled throughout. Peraeopod 7, segment 2 broad (width/length = 56%).

Gnathopod | (o’), segments 2 and 3 weakly setose, setae shorter than the width of the

segments

Genus Lembos Bate 1857 — Barnard 1969, Myers

1981

Subgenus Arctolembos Myers 1981

Diagnosis: Anterior lateral cephalic lobes bilobed;

maxilliped palp article 2 extending beyond the

distal end of the outer plate for '/, its length;

maxilla 1 outer plate with more than 10 spines;

lower lip, outer plate, anterior margin broadly

truncate.

Type Species: Microdeutopus arcticus Hansen

1887

Lembos (Arctolembos) arcticus (Hansen 1887)

Figure. 1.

Microdeutopus arcticus Hansen 1887, p. 231,

pl 22, ig. 3-Stebbing, 1894, p. 43.

Autonoe arctica Della Valle 1893, p. 406, pl. 56,

figs. 35-36.

Lembos arcticus, Stebbing, 1895, p. 207; 1906,

p. 595; Bruggen, 1909, p. 39, pl. 3, figs. 22-28;

Gurjanova, 1951, p. 836, fig. 586 (part-

figures on extreme right apply to Proto-

medeia grandimana Bruggen); Myers, 1981,

pp. 269-273, figs. 200-202.

Material examined: Alaska — Bering Sea; |

juvenile o&, 1 subadult 9 from St. Lawrence

Island (64°N,169°W). Slattery stn. 23, 33-38 m,

1 and 7 July 1980.

Distribution: Kara Sea (70°N,63°E) (type

eoeeeeeeeeeeeeeee ee ee ee ee ee eee eee em eo He em em ew oe we wm oe He wm we hw ww ow ow

A. spinosus n. sp. (page 93)

locality), Bering Sea (64°N,169°W).

Ecology: An arctic species occurring subtidally

on a sandy substratum.

Diagnosis: Male — Body with a pair of short

setae on the dorsum of pleon segment 3 and

urosome segment 1. Sternal processes absent.

Head lobe shallow, bilobed. Coxal plates shallow,

coxa 5 nearly as deep as coxa 4. Cuticle not

especially shiny. Antenna | peduncle 1 with 4

ventral spines, accessory flagellum well developed,

with up to 8 segments. Antenna 2, flagellum with

up to 7 segments, first two the longest, terminal

two segments distally spinose. Mandible with

maximum 5 teeth and 8 raker spines; lacinia

mobilis with maximum 4 teeth; palp strongly

falcate, segment 2 with numerous inner marginal

setae and two groups of outer setae; segment 3

subequal, inner margin strongly concave with a

marginal row of long setae and in addition 4

groups each of long lateral and outer marginal

setae; no seta at base of palp. Maxilla 1 inner

plate with several minute setae; outer plate

with up to 13 teeth; palp without setae at the

base or on segment 1, 11 terminal spines and

numerous lateral and marginal setae on seg-

ment 2. Maxilla 2 inner plate smaller than the

outer. Maxilliped inner and outer plate teeth

smooth and minute; palp segment 2 very long,

exending beyond the distal end of the outer

LFT MD

Figure 1. Lembos (Arctolembos) arcticus (Hansen) juvenile o& (16.0 mm), subadult 2 (25.0 mm), St.

Lawrence Is., Bering Sea. 1 and 7 July 1980

plate by about '/, its length; segment 3 without

a distal protruberance, dactyl moderately large.

Gnathopod 1, coxa | smaller and shallower than

coxa 2, anterodistally produced; segment 2,

anterior and lateral margins nearly bare, posterior

margin with a group of long setae; segment 4 not

prolonged into a long distal tooth; segment 5,

posterior margin more than half the length of the

anterior margin; segment 6 longer than 5, dis-

tally broadened, palm transverse and with a

small defining tooth at the obturator spine

(spine lacking in adult), dactyl not elongate,

serrated in the juvenile, less so in the adult.

Gnathopod 2, segment 2, anterior and lateral

margins nearly bare, posterior margin with a

group of long setae; segment 5 broadened dis-

tally, segment 6 slightly longer than 5, palm

transverse; anterior margins of segments 5 and 6

not densely setose. Peraeopods 3 and 4, hind

margin of segment 2 bearing a group of long

setae; segment 5 not tapered proximally, slightly

shorter than segment 4. Peraeopods 5-7, hind

margin of segment 2, densely setose; no spine at

posterodistal corner; dactyls progressively elon-

gated, from half the length of segment 6 in

peraeopod 5 to four-fifths the length in peraeo-

pod 7. Peraeopods 5 and 6, segment 5 lacking

groups of comb spines. Peraeopod 7, segment 2

slender. Epimera 1-3, hind corner slightly

notched, lateral ridge absent; lower margin of

epimeron 2 not setose. Uropod spines strong;

peduncular process of uropods | and 2 nearly

half the length of the longest ramus. Uropod 2,

inner ramus sinuous. Uropod 3, peduncle bearing

lateral spines, lateral setae and terminal setae;

inner ramus bearing lateral spines and terminal

setae. Telson, each dorso-lateral crest with a

group of 7-9 setae and a small apical cusp;

posteroventral portion acutely produced. Gills

short, plate-like.

Female. Coxa | not shallower than coxa 2;

gnathopod 1, palm lacking a small tooth at the

defining corner. Features otherwise as in the

male.

Size range: male to 25 mm, female to 30 mm

(not mature at 23 mm).

Remarks: This is a primitive aorid, as evidenced

by the multi-segmented accessory flagellum of

antenna 1, well developed falcate mandibular

palp, larger number of mandibular raker spines,

well developed maxilla | palp, unaltered segments

4 and 5 of peraeopods 3 and 4, well developed

peduncular spinous process of uropods | and 2

and small, plate-like gills. It appears to have a

number of features in common with Leptocheirus

and Protomedeia (Table 1) and in addition, many

features not common to its genus. As such, full

generic status may be justified, in recognition of

its apparent transitional importance in linkage

with the Isaeidae (Photidae).

Subgenus Lembos Myers 1981

Diagnosis: Anterior lateral cephalic lobes not

bilobed; maxilliped palp segment 2 not extending

beyond the distal end of the outer plate; maxilla 1,

outer plate with 10 spines; lower lip, outer plate,

anterior margin rounded.

Type species: Lembos (Lembos) websteri Bate

1857

Additional Species: A series of recent papers

by Myers (summarized in Myers, 1981) has

revealed numerous new species of Lembos in the

Mediterranean and Caribbean regions, at least

one group of which will probably require new

subgeneric status (Myers, personal communication)

Lembos (Lembos) concavus Stout 1913?

Figure 2.

Lembos concavus, Stout 1913, p. 36-41;

Barnard, 1962, p. 7-9, fig. 2.

Material examined: British Columbia —

northern mainland: Townsend Pt., St. John

Harbour; 7 specimens from Bousfield 1964 stn., —

HS53, (52°12’N,128°29’W). Surf exposed bedrock,

boulders, Phyllospadix, kelp, Corallina: LW-MW;

temperature 14.2°C, salinity 30.3°/... Vancouver

Island: Sea Otter Cove; 3 specimens from Bous-

field 1959 stn. O2B (50°40’N,128°21’W). Dredge

haul on stony bottom. Temperature 12.8°C,

salinity 33:0°7 se.

Distribution: There is some uncertainty that

these specimens are indeed Lembos concavus (see

remarks below) and certainly this record is far

north of its southern California habitat.

Figure 2. Lembos (Lembos) concavus Stout ¢& 4.0mm, 9 approx. 5mm, mouth of San Josef Bay, Barkley

Sound, Vancouver Is., B.C. 18 July 1959

Ecology: Temperate species living subtidally

amongst algae.

Diagnosis: Male — Body with a pair of short

setae on the dorsum of pleon segments 1-3 and

urosome |. Sternal processes absent. Head lobe

medium, not bilobed; coxal plates shallow;

coxa 5 as deep as coxa 4. Cuticle not especially

shiny. Antenna | peduncle | with posterodistal

spine; rest of antenna lost. Mandible with

maximum 6 teeth and 6 raker spines; lacinia

mobilis with maximum 4 teeth; palp falcate,

segment 2 with 5 inner setae, and no outer setae,

segment 3 the longer, inner margin convave with

a marginal row of short comb setae, another of

long setae plus several groups of lateral setae;

no seta at base of palp. Maxilla 1 inner plate

with | seta; outer plate’ with 10 teeth; palp with

2 setae at base, | seta on segment 1, 7 teeth and

4 setae on segment 2. Maxilla 2 inner plate

nearly as broad as outer. Maxilliped inner and

outer plate teeth smooth and well developed;

palp segment 2 not extending beyond the distal

end of the outer plate; segment 3 without a distal

protruberance, dactyl moderately long. Gnatho-

pod 1 subchelate, not elongate; coxa | neither

enlarged nor shallower than 2 nor produced

anteriorly; segment’ 2, margins nearly bare;

segment 4 not prolonged into a long distal tooth;

segment 5, posterior margin half the length of the

anterior margin, not broadened distally; segment

6 longer than 5, palm transverse and with a small

tooth at the obturator spine in the subadult, a

larger square tooth proximal to the defining

Sommer in the adult; dactyl not elongate,

serrated. Gnathopod 2, segment 2, margins nearly

bare; segment 5 not broadened distally; segment 6

as long as 5, anterior margins of each densely

setose; palm transversely rounded. Peraeopods

3 and 4, hind margin of segment 2 bare; segment 5

tapered proximally, slightly shorter than seg-

ment 4. Peraeopods 5-7, hind margin of segment 2

moderately setose, lacking a spine at the postero-

distal corner; dactyls barely increasing in length

posteriorly. Peraeopods 5 and 6, segment 5

bearing two groups of comb spines. Peraeopod 7,

segment 2 slender. Epimera 1-3, hind corner

notched, lateral ridge present; lower margin of

epimeron 2 setose. Uropod spines strong;

peduncular process of uropods | and 2 nearly

half the length of the longest ramus. Uropod 3,

Outer ramus with lateral setae, inner ramus

laterally spinose and terminally setose. Telson

with a few setae at the apical cusps. Gills long,

sac-like.

Female — Ganthopods barely sexually

dimorphic. Gnathopod 1, palm apparently more

oblique (see Barnard, 1962). Features otherwise

as in the male.

Size range — male to 6 mm, female to 6 mm.

Remarks: There are some differences between

these specimens and those of Barnard (1962)

from southern California which may only be

due to differences in age. In B.C. material, coxa |

is not so acutely produced forward; the palm of

the subadult male gnathopod | is oblique rather

than transverse; segment 2 of gnathopod 2 is

not antero-distally produced, margins are bare,

the obturator spine is present, peraeopods 3 and 4

are much less densely setose and the lower margin

of epimeron 2 is setose. The California male has a

pair of setae on the dorsum of urosome | but in

the B.C. specimens, pairs of dorsal setae are

borne also on pleon segments 1, 2 and 3.

Genus Columbaora new genus

Diagnosis: Body with a pair of short setae on

the dorsum of urosome |. Segment | with a

sternal hump or process. Head lobe strongly

produced in the male, not bilobed. Coxal plates

moderately deep, coxa 5 shallower than coxa 4.

Antenna | accessory flagellum multi-segmented.

Antenna 2 flagellum 6-7 segments, lower 3

segments distally spinose. Mandibular palp

strong, falcate. Maxilla 1 inner plate with

1 setae; outer plate with 10 teeth. Maxilla 2

inner plate nearly as broad as outer. Maxilliped

palp segment 2 not extending beyond the distal

end of the outer plate; segment 3 with a distal

protruberance, dactyl short. Gnathopods strongly

sexually dimorphic. Male gnathopod | elongate,

merochelate; segment 4 extending the full length

of segments 5 and 6 together. Gnathopod 2, both

sexes, subchelate, palm oblique, obturator spine

small; segment 5 longer than 6, in the male

becoming greatly elongate and densely setose.

Peraeopods 3 and 4 slender, segment 5 slightly

shorter than segment 4 and tapered proximally.

Peraeopods 5-7, hind margin of segment 2 with-

out long setae, lower hind corner not spinose,

dactyls barely differing in length. Peraeopods 5

and 6, segment 5 with 3 groups of spines.

Peraeopod 7, segment 2 with a broad hind lobe.

Epimera 1-3, hind corner notched; lateral ridge

present; lower margin of epimeron 2 setose.

Uropod spines moderately strong; peduncular

process of uropods 1 and 2 long, more than

half the length of the longest ramus. Uropod 3

both rami spinose and terminally setose. Telson

with a few setae at the apical cusps. Gills long,

sac-like.

Type species:

species.

Etymology: A compound word derived from

British Columbia, the coastal region of most

frequent occurrence and from Aora and Aoroides

(in which the form of the first gnathopod (<) is

similarly merochelate).

Relationship: The first gnathopod of the male

resembles that of Aoroides in the young, but

diverges with age. Unlike Aoroides the second

gnathopod is also strongly sexually dimorphic,

Columbaora cyclocoxa new

resembling that of the male Lembos macromanus

(Shoemaker). Columbaora shows closer relation-

ships with Lembos also in the form of the

mandibular palp and in the possession of setae

on the lower margin of epimeron 2.

Columbaora cyclocoxa n.sp.

Figure’ 3.

Type locality: Klokachef Is., Chichagof Is.,

Alaska (57°25’N,135°52’W). Bousfield and

McAllister 1961 station A168, 24 July 1961.

Under boulders and amongst Laminaria at low

water. Temperature 12.0°C, salinity 30.9°/o.

Holotype &0, NMC-C-1981-975; Allotype 9,

NMC-C-1981-958; Paratypes NMC-C-1981-

959-960.

Material examined: Alaska — southern tip: 10

Figure 3. Columbaora cyclocoxa n. gen., n.sp. holotype & 6.5 mm, allotype 9 7.0 mm, Klokachef Is.,

Chichagof Is., Alaska, 24 July 1961; paratype juvenile & 4.0 mm, Codfish Pass, Miles Is., B.C. 5 August

1964

specimens from Bousfield and McAllister 1961

stns. A168, A171-2. British Columbia — northern

mainland: 50 specimens from Bousfield 1964

stns., HS, H8, H30, H47 (paratype juvenile),

H50, H65. Vancouver Island: 2 specimens from

Bousfield 1977 stn. B6a; 1 specimen from Bous-

field 1975 stn. P16b; 8 specimens from Bousfield

1970 stns. P709, P710, P715; 4 specimens from

Bousfield 1959 stns. O26, V11. Washington: 3

specimens from Mukkaw Bay, Bousfield 1966

stn. W40.

Distribution: Chichagof Is., Alaska (57°25’N,

135°52’W) south to Mukkaw Bay, Washington

(48°19’N,124°40’W). |

Ecology: Occurs amongst algae in the low

intertidal and subtidally to 20 m depth on surf-

exposed high salinity coasts. Summer tempera-

tures 10-14°C, salinity 30.5-33.0°/... Female

ovigerous in July and August.

Diagnosis: Male holotype (6.5 mm). Head lobe

strongly produced with age. Cuticle very shiny.

Antenna | peduncle with a posterodistal spine;

accessory flagellum 1!/, segments. Mandible with

maximum 6 teeth, 4 raker spines; lacinia mobilis

with maximum 4 teeth; | seta at base of palp;

segments 2 and 3, inner margins setose. Maxilla 1

palp with 3 setae at base, no setae on segment 1,

maximum 9 teeth and 8 setae on segment 2.

Maxilliped inner and outer plate teeth serrated

and well developed. Coxa | circular, enlarging

with age to enclose the head; gnathopod 1,

segment 2 bare, segment 4 prolonged into a long

distal tooth which extends with age beyond

segment 6 to meet the dactyl; segment 5 elongate,

in oldest specimens produced into a ventrodistal

tooth; segment 6 shorter than 5, in old specimens

produced at the location of the obturator spine

into a ventrodistal tooth; dactyl not elongate,

serrated in young, smooth in old; segments 2, 3

and dactyl without setae, segments 4, 5 and 6

strongly setose. Gnathopod 2, segment 2, anterior

margin setose; segment 5 longer than 6, not

broadened distally, but elongating with age and

margins densely setose; segment 6, margins

densely setose, palm oblique, obturator spine

small; dactyl not overlapping the palm.

Female allotype (7.0 mm) — Head lobe

moderately produced. Coxa 1 smaller and

shallower than coxa 2; gnathopod 1 somewhat

larger than gnathopod 2, subchelate; segment 5

subequal to segment 6, not posteriorly lobed;

palm transverse, dactyl strongly overlapping.

Gnathopod 2, segment 2, nearly bare; segment 5

slightly longer than segment 6, anterior margin

bare; segment 6, anterior margin bare, palm

oblique; dactyl not overlapping the palm. Fea-

tures otherwise as in the male.

Size range: Male 4-6.5 mm, female 4-7 mm.

Pigmentation pattern: Brown bands on head

and segments 1, 2, 4, 5 and 7.

Etymology: The species designation is descrip-

tive of the large circular first coxa, distinctive

in the male.

Remarks: In common with Lembos (Lembos)

concavus and Lembos (Arctolembos) arcticus, the

gnathopods of this species do not differ greatly

in size. However, they exhibit considerable

alteration from the normally subchelate form of

the female. Primitive features are the presence of

(albeit shorter) accessory flagellum, well

developed mandibular palp and maxilla | palp.

The shorter accessory flagellum, fewer mandi-

bular raker spines, less setose maxilla 1 inner

plate, altered segments 4 and 5 of peraeopods 3

and 4 and long, plate-like gills suggest a more

apomorphic form and a closer relationship to

Lembos (Lembos) concavus.

By comparison with the merochelate genus

Aora which occurs in the Indo-Pacific and

Atlantic, Columbaora appears to be somewhat

more primitive in the possession of a more falcate

mandibular palp and well developed peduncular

spinous process on both uropods | and 2. The

greatly enlarged circular first coxa, more elongate

fourth segment of the first gnathopod and greatly

elongate fifth segment of the second gnathopod

of the male is distinctive of the species.

Genus Aoroides Walker (revised)

Type species: A. columbiae Walker 1898

Additional northeastern Pacific species:

A. inermis new species

A. intermedius new species

A. exilis new species

A. spinosus new species

Diagnosis: Body with a pair of short setae on

the dorsum of pleon 3 and urosome |. Sternal

bump or process on segment 1. Head lobe

medium, not bilobed. Coxal plates shallow,

coxa 5 as deep as coxa 4. Cuticle not especially

shiny. Antenna 1, peduncle | lacking a ventro-

distal spine; accessory flagellum minute, vesti-

gial. Antenna 2, flagellum of 3'/, segments, first

much longer, each segment distally spinose.

Mandibular palp slender, cylindrical, segment 2

with 0-2 setae, segment 3 slightly longer, inner

margin with 2-7 setae, no setae at base of palp.

Maxilla 1 inner plate with 1 seta; outer plate

with 10 teeth. Maxilla 2 inner plate much more

slender than outer. Maxilliped inner plate teeth

serrated, outer plate teeth serrated or smooth;

palp segment 2 not extending beyond the distal

end of the outer plate; palp segment 3 with a

distal protruberance, dactyl short. Gnathopods

strongly sexually dimorphic. Male gnathopod 1

elongate, merochelate; coxa elongating with age

to mask laterally the buccal mass; segment 2,

anterior, lateral and in some species, posterior

margin setose; segment 4 prolonged into a long

distal tooth extending to the junction of segments

5 and 6; segment 5 shorter than 6, as broad as 5 in

young but slendering with age; never a distal

tooth; dactyl serrated in young, losing cusps,

elongating and bearing dense marginal setae at

maturity; all segments moderately to densely

setose. Gnathopod 2, both sexes, subchelate;

segment 2, anterior margin setose, posterior

margin bare; segment 5 subequal to 6, not

broadened distally, margin not densely setose;

segment 6, margins not densely setose, palm

transverse, obturator spine normal, dactyl over-

lapping the palm. Peraeopods 3 and 4 slender,

segment 5 longer than segment 4 and barely

tapered proximally. Peraeopods 5-7, hind margin

of segment 2 with or without long setae, lower

hind corner marked by a long slender spine;

dactyls barely differing in length. Peraeopods

5 and 6, segment 5 with 2 groups of spines.

Peraeopod 7, segment 2 slender or moderately

expanded. Epimera 1-3, hind corner notched,

lateral ridge lacking; lower margin of epimeron 2

bare. Uropod spines slender; peduncular process

of uropods 1 and 2 about '/, the length of the

longer ramus. Uropod 3 rami with or without

lateral spines, both rami terminally setose, outer

more so than inner. Outer ramus 2-segmented,

distal segment minute. Telson with a few setae

at the apical cusps. Gills long, sac-like.

Type species: Aoroides columbiae Walker 1898.

Coloration: Basic body colour is white or

orange, in some species alternating in bands on

the antennae with red. Superimposed 1s a darker

pigmentation in distinct speckles, concentrated

spots or broad bands, the pattern which, except

in recently moulted individuals, remains distinc-

tive in alcohol for up to 10 years storage. Females

brood up to 45 eggs. Young are released soon

after hatching.

Distribution: Endemic to the north Pacific

region (Alaska to California); previous records

from Hawaiian Islands, Pacific coast of USSR,

and Japan await confirmation.

Remarks: The reduction of the accessory

flagellum of antenna | to a microscopic button,

reduction in size and setosity of the mandibular

palp and strong dimorphism of the gnathopods

imply a more advanced relationship of Aoroides

to Lembos and Columbaora.

Aoroides inermis n.sp.

Figure 4.

Type Locality: Departure Bay, Nanaimo, Van-

couver Island, B.C. Bousfield stn. B2, 14 May

1977. Sand, eelgrass, 1-3 m. Holotype o&’, NMC-

C-1981-946, paratypes NMC-C-1981-948-950.

Mouth of Trevor Channel, Vancouver Island,

B.C. Bousfield stn. B18, 30 May 1977. Sand and

fine shell, 32-36 m. Allotype 2 NMC-C-1981-949.

Material examined: British Columbia

northern mainland: 2 specimens from Bousfield

1964 stns. H49, H57. Vancouver Island: Approxi-

mately 1000 specimens from the collections of

Bousfield 1977, 1976, 1975, 1970, 1959, 1955, and

the collections of K.E. Conlan, D.V.) Ell

L. Daniels, J.F.L. Hart, G.C. Carl, J.W. Scoggan;

southern mainland: 53 specimens from the collec-

tions of Bousfield 1978, 1977, 1959 and the

collections of C.D. Levings and P. O’Rourke.

Washington — approximately 75 specimens from

Bousfield 1966 stns. W7, W8, W13, W42.

Geographic Distribution: Goose Island, B.C.

(51°56’N,128°26’W) south to Shipwreck Pt.,

Clallam Co., Washington (48°19’N,124°27.5’W).

Ecology: A boreal species occurring primarily

on sand bottoms in the low intertidal and sub-

tidally to 75 m depth on high salinity exposed

and protected coasts. Summer temperature:

8.0-18.3°C, salinity 25.6-33+°/... Females ovi-

gerous May-November. This species apparently

prefers warmer waters and sandier sediments

than other northeastern Pacific species of

Aoroides and has not been recorded from the

Queen Charlotte Islands or Alaska.

Diagnosis: Male holotype (5.0 mm). Antenna 2

flagellum weakly setose. Maxilla 1 palp teeth

weakly serrated. Maxilliped outer plate teeth

smooth below the upper three. Gnathopod 1,

coxa | slender and elongate, 1 spine and a few

setae at the anterodistal corner; segment 2 front

and lateral margins densely setose, hind margin

bare; segment 3 front margin densely setose;

segment 4 tip abruptly narrowed; segment 5 not

Figure 4. Aoroides inermis n.sp. holotype o& 5.0 mm, Departure Bay, Nanaimo, Vancouver Is., B.C.

14 May 1977; allotype 9 5.5 mm, Trevor Channel mouth, Barkley Sound, Vancouver Is., B.C. 30 May

1977

broader than segment 2, upper margin lined by

8-15 bundles of long setae. Gnathopod 2, palm

slightly oblique, dactyl overlapping by more than

the length of the nail; coxa with only a few short

setae. Peraeopods 3 and 4, segment 5 not spinose.

Peraeopod 7, segment 2 slender, marginal setae

short. Uropod 3, outer ramus usually lacking

lateral spines, but occasionally with 1 or 2 small

spines; inner ramus with 0-2 lateral spines.

Pigmentation: Antenna 2 flagellum segment |

red banded in life. Pigment speckles scattered

throughout, concentrating in the lower hind

corners of body segments 1-5. Size range: male

3.5-6 mm, female 3-6.5 mm.

Female allotype (5.5 mm). Gnathopod 1,

somewhat larger than 2, coxa not enlarged,

anterodistal margin of segment 2 bearing a group

of long setae, segment 4 not produced, segments

5-7 similar in form to those of gnathopod 2.

Otherwise as in the male.

Etymology: Refers to the smooth mid and lower

inner marginal teeth of the outer plate of the

maxilliped.

Aoroides intermedius n.sp.

Figure 5.

Type locality: Haines Island, Barkley Sound,

Vancouver Island, B.C. L. Daniels, collector,

8 August 1975. In kelp holdfasts at 3-6 m depth.

Holotype o&, NMC-C-1981-951: Allotype 9,

NMC-C-1981-952; Paratypes NMC-C-1981-953.

Material examined: Alaska — southern tip: 87

specimens from Bousfield 1961 stns. A3, A6, A7,

Al15, A171-2, A174, A175; 50 specimens from

Bousfield 1980 stns. S18F3 and S23F1. British

Columbia — Queen Charlotte Islands: 52 speci-

Figure 5. Aoroides intermedius n.sp. holotype & 5

Vancouver Is., B.C. 8 Aug. 1975

mens from Bousfield 1957 stns. E25, H3, H14,

W3a, W4a, W8, W12, W14. Northern mainland:

approximately 200 specimens from Bousfield

19G64estns) Hi, 113,95; 7, Hs, 10, 7; 21,

H23, H25, H26, H30, H31, H33, H37, H47, H48,

H49, H50, H51, H53, H57, H58, H65. Vancouver

Island: approximately 1,000 specimens from the

collections of Bousfield 1977, 1976, 1975, 1970,

1964, 1959, 1955, K. Conlan 1976, L. Daniels

1975; D.V.. Ellis 1979, J-F.L: Hart and:G:C., Carl

1938, and D. Kittle 1972. Southern mainland:

8 specimens from Bousfield 1977 stn. E4, and

Bousfield 1955 stn. G4. Washington — approxi-

mately 75 specimens from Bousfield 1966 stns.

W40, W42.

Geographic distribution: Puffin Bay, Baranof

Island, Alaska (56°16’N,134°48’W) south to

Mukkaw Bay at Sooes Pt., Clallam Co.,

Washington (48°19’N,124°40’W).

MXPD PL

.0 mm, allotype 2 5.5mm, Haines Is., Barkley Sound,

Ecology: A boreal species occurring amongst

algae and eelgrass on sand and gravel bottoms in

the low intertidal and subtidally to 63 m on

exposed and semi-protected coasts. Summer

temperatures from 9.8-15.0°C, salinity 12.8-

33+°/o0. Females ovigerous March to August.

Diagnosis: Male holotype (5.0 mm). Antenna 2

flagellum strongly setose. Maxilla 1 palp teeth

moderately to strongly serrated. Maxilliped outer

plate teeth weakly serrated, lower teeth with 0-2

cusps each. Gnathopod | adult coxa slender;

segment 2 front and lateral margins densely

setose, hind margin bare; segment 3 front margin

densely setose; segment 4 tip abruptly narrowed;

segment 5 not broader than segment 2, upper

margin lined by 5-7 bundles of long setae.

Gnathopod 2, palm slightly oblique, dactyl

overlapping by more than the length of the nail;

coxa with a few short setae only. Peraeopods 3

and 4, segment 5 not spinose. Peraeopod 7,

segment 2 broad, marginal setae short. Uropod 3,

outer ramus usually lacking lateral spines, but

occasionally with | or 2 small spines; inner

ramus with 0-2 strong lateral spines.

Pigmentation: Antenna 2 flagellum segment 1

red banded in life. Pigment speckles scattered

throughout the body, with no concentration in

segment corners.

Size range: male 3-5.5 mm, female 3-6 mm.

Female allotype (5.5 mm). Gnathopod | some-

what larger than 2, coxa not enlarged, antero-

distal margin of segment 2 bearing a group of

long setae, segment 4 not produced, segments 5-7

similar in form to those of gnathopod 2. Other-

wise as in the male.

Etymology: Refers to the serration of the

marginal teeth of the maxilliped outer plate,

intermediate in degree between A. inermis (with-

out) and A. columbiae (with).

Remarks: The intensity of serration of the

maxilliped outer plate teeth is constant with age

(although the number of teeth will vary). Table 2

gives the average amount expected. The upper-

most 2-3 teeth are always densely serrated as they

blend in size into the marginal plumose setae

above. The teeth in the lower two-thirds of the

plate are those of diagnostic value.

Aoroides columbiae Walker 1898

Figures 6-8.

Aoroides columbiae Walker (1898), p. 285,

pl. 16, figs. 7-10: Thorsteinson, 1941,

pp. 83-84, pl. 6, figs. 65-66; not Barnard,

1954, pp. 24-26, pl. 22. ? Barnard, 1959,

p. 33; ? Barnard, 1961, p. 180; ? Barnard,

1964, pp. 217-218; ? Barnard, 1969, pp.

89-90; ? Barnard, 1970, pp. 68-70, figs.

31-32; ? Nagata, 1960, p. 175, pl. XVI,

fig. 94.

Aoroides californica Alderman, 1936, pp. 63-66,

figs. 33-38.

Material examined: Alaska — Amchitka Island:

1 specimen from St. Makarius Bay, P.A. Lebed-

nik, Coll. 1976; southern coast: 83 specimens

from Bousfield and McAllister 1961 stns. A6, A7,

A48, A75, A&1, A87, A91, A96, A147, A151,

A153, A165, A168, A171-2, A175; 20 specimens

from Bousfield 1980 stns. S4B1-4, S5B2, S11B1,

. S11B2; S18B1, S21L1, S22F2. British Columbia

Figure 6. Aoroides columbiae Walker, lectotype 2 5.0 mm, Puget Sound, Washington, 1897. Illustration

of appendages mounted on British Museum slide #1908.3.10.15. The dotted line demarcates an air bubble.

Figure 7. Aoroides columbiae Walker, plesiotype 9 6.0 mm, Willis Beach, Victoria, Vancouver Is., B.C.

17 May 1977

— Queen Charlotte Islands: 28 specimens from

Bousfield 1957 stns. E5, El4a, H2a, H8b, H11,

H14, W8, W11, W14, W15b; northern mainland:

approximately 250 specimens from Bousfield

1964 sins -Ei?y HS SHIO7 Hi2. H21, W22,

123 ,cH25, F261 29; T3502 8131,.H33,,H43, H47,

H50, H53, H56a, H58, H65; Vancouver Island:

approximately 1500 specimens from the collec-

tions of Bousfield in 1977, 1976, 1975, 1970,

1959 and 1955 and J.M. Green 1975; southern

mainland: 2 specimens from Vancouver Harbour

stn. 14,. F. ‘Ratt and 'C.. Levings, coll. 1976.

Washington and Oregon — approximately 80

specimens from Bousfield 1966 stns. W13, W34,

W35, W39, W40, W58, W63, W64.

Geographic distribution: Amchitka Island,

Alaska (51°N,179°W), south to Oregon

(45°26.5’N,123°57’W); Bahia de San Quintin,

Baja California (?); Hawaii (?); Japan (?).

Ecology: A boreal species occurring on high

salinity exposed and protected coasts amongst

algae and sponges or under stones on mixed sand

and stony beaches in the low intertidal, to a

depth of 85 m (or to 180 m?). Summer tempera-

tures 8.0-14.5°C, salinity 14.8-33+°/,.. Females

ovigerous April-September.

Diagnosis: Eyes small, subcircular. Antenna 2

flagellum strongly setose. Maxilla 1 palp teeth

weakly serrated. Maxilliped outer plate teeth

strongly serrated, lower teeth with 1-4 cusps each.

Gnathopod | (0), adult coxa | broad, all setae

simple; 1 spine and a few setae at the anterodistal

corner; segment 2 front and lateral margins

densely setose, hind margin bare; segment 3 front

margin densely setose; segment 4 tip abruptly

narrowed; segment 5 broader than segment 2,

upper margin bare except for a small distal bundle

of short setae. Gnathopod | (¢@), anterodistal

margin bearing a group of long setae. Gnathopod

2, both sexes, palm slightly oblique, dactyl over-

lapping by more than the length of the nail; coxa

of older males with several long marginal setae.

Peraeopods 3 and 4, segment 5 sometimes spinose

in the male. Peraeopod 7, segment 2 broad,

marginal setae short. Uropod 3, outer ramus

usually lacking lateral spines, but occasionally

MXPD PL

Figure 8. Aoroides columbiae Walker plesiotype ¢& 6.0 mm, Willis Beach, Victoria, Vancouver Is., B.C.

18 May 1977. Juvenile plesiotype & 3.5 mm, Witty’s Lagoon, Victoria, Vancouver Is., B.C. 17 May 1977

with | or 2 small spines; inner ramus with 0-2

strong lateral spines.

Pigmentation: Antenna 2, flagellum not red

banded in life. Body segments dark banded; head

and parts of segments 6 and 7 white; thereby

giving a “‘saddle-back”’ pattern.

Size range: male 3.5-6 mm, female 3.5-6.5 mm.

Remarks: Only two of Walker’s 9 female

syntypes remain, these being preserved in slide

mounts. British Museum (Natural History) slide

#1908.3.10.15 is designated as the holotype

because it contains the first gnathopod and

maxilliped, the appendages which in combination

are diagnostic in the female (see Figs. 5 and 6).

The long setae on the anterior margin of gnatho-

pod 1 segment 2 distinguish this species from

A. exilis n.sp. and A. spinosus n.sp., and the

serration of the maxilliped outer plate teeth

further eliminate A. inermis n.sp. A. intermedius

closely resembles A. columbiae but in A. inter-

medius the lower teeth of the maxilliped outer

plate are rarely cusped.

The present male specimens (Fig. 8) resemble

Thorsteinson’s (1941) male plesiotype in the

form of the gnathopods: in gnathopod 1, pre-

sence of a distal spine on the coxa, absence of

setae on the hind margin of segment 2, segment 5

wider than 2 and lacking dorsal setae; in

gnathopod 2, palm oblique, dactyl longer than

the palm.

Specimens from Oregon described by Barnard

(1954) are almost certainly A. spinosus n.sp., as

evidenced by the lack of a spine on coxa | of

the male gnathopod 1, presence of setae on the

hind margin of segment 2, reduction of length

and number of setae on the anterior margin of

segment 2, absence of setae on the anterior

margin of segment 3 and gradual, rather than

abrupt, distal narrowing of the older male

segment 4.

Aoroides californica Alderman (1936), could

be synonymous with A. columbiae but not for the

reasons given by Barnard (1954). The species

have in common a lack of lateral spines on the

outer ramus of uropod 3, a group of long setae

at the anterodistal corner of segment 2 of the

female gnathopod 2, absence of dorsal setae on

the male gnathopod 1 segment 5 and greater

breadth of segment 5 relative to segment 2.

However, the body pigmentation (evenly scattered

spots) is more indicative of Aoroides intermedius.

Furthermore, since Alderman failed to illustrate

the marginal setae of segment 2, which are pre-

sent in all male species of the genus, it is conceiv-

able that he could also have overlooked the setae

of segment 5 diagnostic of A. intermedius.

Re-erection of Alderman’s name would therefore

necessitate examination of his holotype.

In view of the species diversification within

the genus, previous references to Aoroides

columbiae from southern California and Hawaii

by Barnard (1959, 1964, 1969 and 1970) and from

Japan by Nagata (1960) require verification,

using the diagnostic characters described herein.

Similarly, re-examination of Aoroides nahili

Barnard (1970) of Hawaii is warranted.

Aoroides exilis n. sp.

Figure 9.

Type locality: Trial Island Point, Vancouver

Island, B.C. Bousfield 1977 stn. B6a, 18 May

1977. Phyllospadix, bedrock, Egregia and algae,

LW-subtidal. Holotype o&', NMC-C-1981-942;

Allotype 9, NMC-C-1981-943; Paratypes, NMC-

Figure 9. Aoroides exilis n.sp. holotype o& 4.0 mm, allotype ? 5.0 mm, Trial Island Point, Vancouver Is.,

B.C. 18 May 1977; paratype intersex, Rennison Is., B.C. 20 July 1964

C-1981-944; Rennison Island, B.C. Bousfield

1964 stn. H30, 20 July 1964. Sand and kelp, 7-22

m. Intersex, NMC-C-1981-945. Departure Bay,

V.I., Bousfield 1977, stn. Bl, May 14, 1977.

Paratype immature, NMC-C-1981-947.

Material examined: Alaska — southern tip: 79

specimens from Bousfield 1961 stns. A3, A6, A7,

A8, A168, A171-2, A174, A175; 30 specimens

from Bousfield 1980 stns. S4B1, S20B5, S23F1.

British Columbia — Queen Charlotte Islands:

156 specimens from Bousfield 1957 stns. E24,

E25, H2, H2a, H3, H5, H8b, H9, W2, W4a,

W4b, W8, WII. Northern mainland: approxi-

mately 450 specimens from Bousfield 1964 stns.

fees. 1/7, H8, H10, H12, H23, H30, H33,

H39, H47, H48, H49, H50, H53, H56a, H57,

H65. Vancouver Island: approximately 500

specimens from the collections of Bousfield

fer? 1976, 1975, 1970, 1959, 1955. Southern

mainland: 14 specimens from Bousfield 1959

stns. N16, N18, N22. Washington and Oregon —

approximately 80 specimens from Bousfield 1966

stns. W8, W18, W30, W34, W36, W40, W64.

Geographic distribution: Klokachef Is.,

Chichagof Is., Alaska (57°25’N,135°52’W) south

to Netart’s Bay at Wilson Beach, Tillamook Co.,

Oregon (45°25.5’N, 123°56’W).

Ecology: Occurs amongst algae, eelgrass and

sponges under stones and in tide pools on sand

and gravel beaches in the low intertidal and

subtidally to 50 m depth. Abundant on high

salinity exposed coasts, semi- and well protected

waters. Summer temperatures 8-15+°C, salinity

12.8-33+°/... Females ovigerous May - August.

Diagnosis: Male holotype (4.0 mm). Appen-

dages slender. Antenna 2 flagellum poorly setose.

Maxilla | palp teeth strongly serrated. Maxilliped

outer plate teeth strongly serrated, lower teeth

with 2-4 cusps each. Gnathopod | coxa slender;

segment 2 front and lateral margins densely

setose at all ages, hind margin weakly setose;

segment 3 front margin densely setose; segment 4

tip abruptly narrowed; segment 5 broader than 2,

upper margin bare except for a small distal

bundle of short setae. Gnathopod 2, palm trans-

verse, dactyl overlapping only by the length of

the nail; coxa of male with several long marginal

setae. Peraeopods 3 and 4, segment 5 spinose

on hind margin. Peraeopod 7, segment 2 slender,

marginal setae short. Uropod 3, both rami with

1-3 strong lateral spines (lateral spines lacking

in immatures of less than 3 mm).

Pigmentation: Antenna 2 not red banded in

life. Body segments dark banded; head and parts

of segments 6 and 7 white, thereby giving a

*“‘saddle-back”’ pattern.

Size range: male 3.5-5.5 mm, female 2.5-6 mm.

Female allotype (5.0 mm). Gnathopod | some-

what larger than 2, coxa not enlarged, antero-

distal margin of segment 2 nearly bare, with

only a few very short setae, segment 4 not

produced, segments 5-7 similar in form to those

of gnathopod 2. Otherwise as in the male.

Remarks: Figure 8 illustrates the gnathopods

of an intersex. The peraeopods and first gnatho-

pod bear small, poorly setose brood plates. The

first gnathopod resembles that of an immature

male: coxa slightly acute but not elongate, the

anterior margin of segment 2 with several long

setae and segment 4 somewhat extended below

segment 5.

Aoroides spinosus n.sp.

Figure 10.

Aoroides columbiae: Barnard, 1954, plate 22

(not Walker, 1898)

Type locality: Willis Beach, Victoria, Vancouver

Island, B.C. Bousfield 1977 stn. B7a, 19 May

1977. Mud, rock and algae, LW-subtidal.

Holotype o&, NMC-C-1981-954; Allotype 9,

NMC-C-1981-955: Paratypes, NMC-C-1981-956.

Material examined: Alaska — southern coast:

110 specimens from Bousfield 1961 stns. A7, A8,

Al8, A20, A21, A86, A91, A92, A117, A136,

A147, A151, A175; 3 specimens from Bousfield

1980 stns. SSB1, S19B1, S19B3. British Columbia

— Queen Charlotte Islands: 224 specimens from

Bousfield 1957 stns. E5, El4a, E25, H2a, H8b,

H9, H11, W1. Northern mainland; 30 specimens

from Bousfield 1964 stns. H25, H29, H64.

Vancouver Island: 25 specimens from Bousfield

1977 stns. B4a, B6b, B7a, B7b; 10 specimens from

Bousfield 1976 stns. B7, B10d, B14; 9 specimens

from Bousfield 1975 stns. P17a, P17d; 30 speci-

mens from Bousfield 1970 stns. P709, P715,

P717, P718; 31 specimens from Bousfield 1959

stns. O4, O11, V3, V5, V17, V18; 78 specimens

from Bousfield 1955 stns. P2, F2, F3, F4, F6, F9,

G2, G4; 47 specimens from the collections of

KE. Conlan 1975, J.F.Lo Hart and eG:C. Cari

1955. Southern mainland: 61 specimens from

Bousfield 1959 stns. N11, N17, N18. Washington

and Oregon — 11 specimens from Bousfield 1966

stns. W7, W64.

Geographic distribution: Prince William Sound,

Alaska (60°44’N, 146°10’W) south to Netart’s Bay

RT MD

Figure 10. Aoroides spinosus n.sp. holotype & 5.5 mm, allotype 2 6.0 mm, Willis Beach, Victoria,

Vancouver Island, B.C. 19 May 1977

at Wilson Beach, Tillamook Co. Oregon

(45°2525'N2123°56 W). Coos. Bay. (Barnard,

1954)?

Ecology: Occurs amongst algae and debris on

mixed sand, gravel sediments and bedrock in the

low intertidal and subtidally to 45 m depth.

Abundant in meso- to polyhaline, semi- and

well-protected waters. Summer temperatures

8-16°C, salinity 12.8-32+°/,.. Females ovigerous

April-August.

Diagnosis: Male holotype (5.5 mm). Append-

ages slender. Antenna 2 flagellum poorly setose.

Mandibular palp segment 2 lacking setae, seg-

ment 3 with 2-6 setae. Maxilla 1 palp teeth

strongly serrated. Maxilliped outer plate teeth

strongly serrated, lower teeth with 2-3 cusps

each. Gnathopod | coxa slender; segment 2 front

and lateral margins poorly setose, setae reducing

in number and length with age, hind margin

weakly setose; segment 3 front margin nearly

bare; segment 4 tip gradually narrowed; segment 5

broader than 2, upper margin bare except for a

small distal bundle of short setae. Gnathopod 2,

palm transverse, dactyl overlapping only by the

length of the nail; coxa of male with several

long marginal setae. Peraeopods 3 and 4, segment

5 not spinose. Peraeopod 7, segment 2 broad,

marginal setae long in male. Uropod 3, both

rami with 1-3 strong lateral spines (lateral spines

lacking in immatures of less than 3 mm).

Pigmentation: Antenna 2 not red banded in

life. Body segments light brown, pigment diffusely

speckled.

Size range: male 3.5-7 mm, female 3.5-6.5 mm.

Female allotype (6.0 mm). Gnathopod | some-

what larger than 2, coxa not enlarged, antero-

distal margin of segment 2 nearly bare, with only

a few very short setae, segment 4 not produced,

A. A. inermis. B. A. intermedius. C. A. columbiae. D. A. exilis. E. A. spinosus.

Figure 11. Body pigmentation patterns in Aoroides.

segments 5-7 similar in form to those of gnatho-

pod 2. Otherwise as in the male.

Remarks: A. spinosus also shows a tendency

to intersex. Six intersexes were noted in material

from Vancouver Island near Victoria and another

from the Queen Charlotte Islands.

Juvenile males are distinguishable from other

juveniles by the slender second gnathopod, the

setose hind margin of segment 2 of the first

gnathopod and paucity of setae on the front

margin (Fig. 10.)

Barnard’s (1954) Aoroides columbiae is clearly

not this species but Aoroides spinosus. The shape

of the first gnathopod 1s distinctive in the setation

pattern of coxa and segments 2, 3 and 5, the

shape of segment 4 and the larger size of seg-

ment 5 relative to segment 2. His comments about

growth variability in Aoroides columbiae must be

viewed with the consideration that more than one

species was being examined.

Discussion and Conclusions

Biogeography and Ecology

Tables 3 and 4 summarize the geographic and

ecological distributions of the northeastern

Pacific Aoridae. All species occur amongst algae

and debris on mixed sediments in the intertidal

and subtidally to the limits of the photic zone.

The species are restricted to meso- and polyhaline

waters, generally on open and semi-protected

coasts. The genus Aoroides ranges through the

entire boreal coast. A. inermis is the most res-

tricted of the species, failing to penetrate into

Alaska and occurring primarily on sandy sub-

strates. Columbaora and Lembos (Lembos) occur

in more southerly waters. The present record for

Lembos concavus is its most northerly extension

in the northeastern Pacific coastal region. Lembos

(Arctolembos) is a rare arctic genus, this present

record extending its range eastward to the

Bering Sea.

Taxonomic considerations

Within the superfamily Corophioidea, the

Aoridae appear to be most closely related to the

Isaeidae (Photidae), and the new family Neo-

megamphopidae (Myers, 1981) which also show

strong development of the first gnathopod.

Lembos (Arctolembos), the most primitive of the

northeastern Pacific Aoridae, possesses many

characteristics in common with the primitive

photid Protomedeia, Subsequently, in increasing

order of advancement, are Lembos (Lembos),

Columbaora and Aoroides. These genera trend

phylogenetically towards reduction of the

accessory flagellum, weakening of the mandibular

palp, reduced setation of the maxilla 1 inner

plate, increasing dimorphism of the gnathopods,

loss of the hind setae on segment 2 of the append-

ages and loss of the marginal setae on epimeron 2.

Comparison of the merochelate genus Aora,

which occurs in the Indo-Pacific and Atlantic,

indicates .a close resemblance with Lembos

(Lembos) and Columbaora in the form of the

female gnathopods, semi-falcate to falcate

mandibular palp and prominent accessory

flagellum.

Phyletic relationships therefore seem to link

these Aoridae in the order, Protomedeia to

Lembos to Columbaora to Aora, with Aoroides

being an outlier, possibly a derivative of Colum-

baora, rather than of Aora. This linkage may be

bidirectional, as Barnard (1973) indicates, with

the Columbaora-Aora-Aoroides \ine originating

separately from Lembos. The latter would sub-

sequently have led to Lembopsis and Lemboides.

Within the genus Aoroides, close similarity of

the component species suggests a relatively recent

evolution. The northeastern Pacific species are

divisible into two groups: the stouter bodied,

more setose group comprising A. columbiae, A.

intermedius and A. inermis, and a more slender

bodied, less setose group comprising A. exilis and

A. spinosus. The individual species relate to each

other in the presented order: A. columbiae with

strongly cusped maxilliped teeth and dorsally

bare segment 5, leading through A. intermedius

as its name implies, to A. inermis with smooth

maxilliped outer plate teeth and dorsally setose

5th segment of the male gnathopod 1. The

second group has in common with A. columbiae

the cuspate maxilliped teeth, the dorsally bare

segment 5, enlarged segment 5 in relation to

segment 2, setose male coxa 2, and stronger

body pigmentation. A. exilis appears to be more

closely related to A. columbiae than does

A. spinosus by having in common a similar

pigmentation pattern and strong setation on the

male gnathopod | segments 2 and 3. The first

gnathopod of the male A. spinosus, which shows

stronger setation and similar form in the juvenile,

with age becomes poorly setose and greatly

elongated to hang way below the body.

The two groups appear to be sufficiently

dissimilar to suggest that the gradient of evolu-

tion was not linearly from A. inermis to A.

spinosus, but bidirectional; thus, from a hypo-

thetical ancestral type evolved in one direction

A. columbiae and thence A. intermedius and

A. inermis, and in the other direction A. exilis

and A. spinosus. The ecological data of Tables 3

and 6 give some support to this hypothesis.

A. columbiae is the most primitive and most

widely distributed of the five species, having

spread north into the Alaskan Aleutian Islands,

possibly westward into Japan, and possibly also

southward into Hawaii and Baja California

(assuming that species identifications outside of

this study area are accurate). A. intermedius and

A. exilis have a narrower but similar distribution

with a focal point in British Columbia, while A.

spinosus spans a more northerly range and

A. inermis is restricted to warmer waters and a

narrower habitat. The two species of Aoroides

not found in the northeastern Pacific, A. secundus

Gurjanova* and A. nahili Barnard, were

probably derived from the Asian Pacific

and Hawaiin populations of A. columbiae

respectively.

*Preliminary examination of material of Aoroides secundus

Gurjanova 1938 from the Okhotsk Sea, kindly supplied very

recently by Dr. Nina Tzvetkova, Zoological Museum, Lenin-

grad, indicates that this species is intermediate between the

A. exilis-spinosus group, and the A. columbiae-inermis-

intermedius group, with somewhat closer affinities (overall) to

the former group.

Off

Acknowledgements

We wish to thank the many donors of valuable

material, too numerous to list, and the British

Museum of Natural History for the loan of

A.O. Walker’s type specimens of Aoroides

columbiae. We are greatly indebted to Mrs. F.

Zittin of Vancouver, B.C. for her capable

execution of the illustrations.

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4(2): 50-96

Walker, A.O. 1898. Crustacea collected by W.A. Herdman,

F.R.S., in Puget Sound, Pacific coast of North America,

Trans. Liverpool. Biol. Soc. 12: 268-287

Table 1. Uncommon characteristics of Lembos (Arctolembos) arcticus

Character Differences from Lembos (Lembos)

Head lobe shallower

Eye smaller

Lower lip mandibular lobes not acutely attenuated

Maxilla | inner plate

outer plate

Maxilliped outer plate

palp segment 2

Coxae

Gills

Gnathopod | segment 6

Gnathopods | and 2, peraepods

3 and 4, segment 2

Peraeoods 5-7, segment 2

dactyls

Uropods | and 2, rami

Uropod 3, outer ramus

ISimilar to Protomedeia

Similar to Leptocheirus

lacking a long apical seta, bearing many fine setae!

bearing more than 10 spines

shorter

more elongate

deeper, especially the anterior lobe of coxa 5

smaller, plate-like

subchelate, relatively small and unaltered?

posterior margin bearing a group of long setae

hind margin densely setose!

lengthen posteriorly

sinuous!

setose as well as spinose?

Table 2. Range in number of cusps on the maxilliped outer plate teeth of Aoroides inermis, A. intermedius

and A. columbiae.

Range in number of cusps per tooth*

Tooth No.

uppermost to A. inermis A. intermedius A. columbiae

] 0-15 4-9 5-15

2 0-15 4-8 2-13

3 0-4 1-4 3-8

‘| 0 0-3 2-9

3 0 0-1 1-7

6 0 0-1 2-6

f 0 0-1 2-8

8 0 0-1 0-4

9 0 0-1 1-4

10 0 0 1-3

11 0 0 1-3

*Values are given for 3.5-7 mm individuals. Specimens with fewer teeth carry the same range in values.

Table 3. Distribution of Boreal Northeastern Pacific Aoridae Listed in Geographic Order

Northern

B.C. and Central

Prince Cross Sd. Queen B.C.-and

Aleutian Is., William Sd., To Dixon Charlotte Vancouver Washing-

Species Alaska Alaska Ent., Alaska Is.; B.C. Is., B.C. ton Oregon Other Records

arcticus

Aoroides columbiae X x Me ».& xX ».& X Baya California ?

Hawaii?

Japan ?

Aoroides spinosus 0 >. b.€ se Xx K x

Columbaora cyclocoxa 1S) ‘S) Xx x x Xx 3)

Aoroides intermedius 'o) (3) Xx Xe x .& 16)

Aoroides exilis 2) oO X x Xe Xe x:

Aoroides inermis 0 (2) 3) x x x 1S)

Number of species Z 2 5 6 fi 6 3

Table 4. Habitats of Boreal Northeastern Pacific Aoridae

Coastal Exposure Salinity Range Depth Range

Open and Protected Marine Oligo- —_- Fresh-

semi- and Polyhaline Mesohaline _haline water

Species protected Estuarine (> 28°/o0) (10-27°/o0) (1-9°/00) (<< 1°/o0) > 25 m 1-25 m LW-MW Substrate

Lembos (Arctolembos)

arcticus x (o) >. 0 0 0 ».« (2) 12) Sandy sediment

xX x b. & ».€ (2) 0 X ».& xX Amongst algae, eelgrass and

columbiae wood debris on mixed sediments

Aoroides x x >. ».& 0 io) X x X Amongst algae, eelgrass, wood

spinosus debris on mixed sediments

Columbaora xX 16) x 12) 12) 0 12) i xX Amongst algae and eelgrass on

cyclocoxa coarse sediments

Aoroides >. 0 oh x, 0 0 ps x X Amongst algae, eelgrass,

intermedius sponges and tunicates on coarse

sediments

Aoroides >. & >. & xe x 0 0 (2) x xX Amongst algae, eelgrass, wood

exilis debris, sponges and tunicates on

mixed sediments

Aoroides ».& >. >. X 0 0 x > xX Amongst algae on sandy

inermis sediment

Lembos x 0 x 16) 6) 16) 6) xX 6) Amongst algae on stony

concavus substrate

100

Table 5. Specimens of Aoridae Designated for Deposition in the Smithsonian Institution (USNM) and

Zoological Museum (Leningrad, USSR).

Smithsonian Institution (USNM)

Species

Aoroides inermis*

Aoroides intermedius*

Aoroides columbiae*

Aoroides exilis*

Aoroides spinosus*

Aoroides spinosus

Columbaora

cyclocoxa*

National

Museums

Accession No.

1977-181

1975-205

1976-157

1977-181

1959-112

1970-152

Station

No.

Haines Is.

P710

See ee

Zoological Museum (Leningrad USSR)

National

Museums

Accession No.

1977-181

1975-205

1976-157

1977-181

1959-112

1977-181

1964-198

*Material also deposited in the British Museum (Natural History) London.

101

Station

No.

Haines Is.

Studies on the amphipod family Lysianassidae

in the Northeastern Pacific region. Hippomedon: and related genera

Systematics and distributional ecology

N.E. Jarrett & E.L. Bousfield

National Museum of Natural Sciences

National Museums of Canada

Ottawa Canada

ABSTRACT

This study analyzes the systematics and distributional ecology of the genus Hippomedon (sens. lat.), a

group of medium large, mainly infaunal lysianassid amphipod crustaceans, in shallow wters of the

northeastern Pacific coastal marine region. Utilizing numerical taxonomic methodology, the 17 regional

species are grouped in four genera, viz. Hippomedon (sens. str.), the most apomorphic group, containing

about 40 species world-wide of which 10 occur in the North Pacific and of which H. columbianus n.sp. is

newly described; the monotypic, intermediate Paratryphosites Boeck of Arctic and North Atlantic

regions; the relatively primitive Wecomedon new genus, consisting of five species endemic to the North

Pacific region and of which W. similis is newly described, and the most primitive genus Psammonyx

Bousfield, consisting of four species, two of which are endemic to the northwestern Atlantic region, and

two, P. kurilicus (Gurjanova) and P. longimerus new species are endemic to the North Pacific. Keys,

figures and diagnoses are provided for the regional species and genera.

RESUME

Il s’agit d’une étude taxinomique et de distribution écologique du genre Hippomedon (sensu lato), un

groupe de crustacés amphipodes lysianassidés de taille moyenne et se rencontrant surtout sur les hauts-

fonds des eaux cdtiéres dans le nord-est du Pacifique. Suivant une méthodologie propre a la taxinomie

numérique, l’auteur groupe les dix-sept espéces de la région en quatre genres : 1° Hippomedon (sensu

stricto), le groupe le plus évolué (quarante espéces dans le monde, dont dix dans le Pacifique nord), dont

H. columbianus n.sp. décrit ici pour la premiére fois; 2° Paratryphosites Boeck de |’Arctique et de

’Atlantique nord, genre monotypique et intermédiaire ; 3° Wecomedon, genre relativement primitif

nouvellement décrit, constitué de cinq espéces indigénes dans le Pacifique nord, y compris W. similis

décrit ici pour la premiére fois; 4° Psammonyx Bousfield, genre le plus primitif, constitué de quatre

espéces dont deux sont indigénes au nord-ouest de |’Atlantique et deux, de nouvelles espéces P. kurilicus

(Gurjanova) et P. longimerus, sont indigénes au Pacifique nord. Des illustrations, des données techniques

d’identification et des clés des genres et des espéces de la région étudiée apparaissent dans ce travail.

Introduction

The lysianassid genus Hippomedon (sens. lat) including the Sea of Okhotsk and Japan Sea.

comprises about 75 species of benthic fossorial Holmes (1908), Shoemaker (1955), Barnard

amphipod crustaceans that occur mainly in sub- (1954, 1964b, 1966, 1971), and Hurley (1963)

arctic and boreal coastal marine regions of the have collectively treated an additional 6 species

northern hemisphere. Several species are known and 3 subspecies or forms from the northeastern

from littoral and sublittoral regions of the (North American) Pacific coast, from Point

southern hemisphere and a few have been Barrow, Alaska to California. However, much

recorded offshore and from the abyss. In the of the Pacific rim coastal marine region reamins

North Pacific region (Pacific rim) a total of virtually unsampled with respect to material of

13 species and 4 subspecies have previously been Hippomedon. Furthermore, the present basis for

recorded. Gurjanova (1962) included 6 speices recognition of the genus and species composition

and 2 subspecies from the northwestern Pacific, encompasses major taxonomic problems. The

103

present study was undertaken in an attempt to

fill this distributional hiatus and to resolve these

taxonomic problems.

This study utilizes extensive lysianassid

material from the northeastern Pacific region

obtained by field expeditions of the National

Museum of Natural Sciences since 1955 (see

Bousfield (1958, 1963, 1968), Bousfield and

McAllister (1962) and Bousfield and Jarrett (in

press) for pertinent station lists). Other source

material in the NMNS and in other museums

have been utilized, as noted in the text. Recently

material from Vancouver Island and southeastern

Alaska has been collected and photographed live

by E.L. Bousfield and Mr. Ron Long, Simon

Fraser University, and the resulting slide trans-

parencies have been utilized in some of the

species descriptions.

SYSTEMATIC SECTION

Family Lysianassidae Dana, 1849 (emend.

Bousfield, 1979)

Subfamily Uristidinae Hurley, 1963

In an attempt to subdivide the large and

unwieldly family Lysianassidae into natural

groupings of allied genera, Hurley (1963) tenta-

tively proposed the subfamilies Uristidinae and

Lysianassinae. Although these two subfamilies

as defined would encompass less than half the

known lysianassid genera, Hurley (1963) did

include the genus Hippomedon (and allied genera)

in the subfamily Uristidinae. This subfamily is

characterized primarily by a subchelate or

imperfectly subchelate first gnathopod. Addi-

tional features include: antenna 1, peduncle

generally not carinate; antenna 2, peduncular

segments not noticeably dilated; coxal plates |

and 2 not appreciably smaller than 3 and 4;

coxal plate 1, lower front angle not hidden by

coxal plate 2; uropod 3 biramous. This subfamily

delineation is very incomplete and does not

clearly demonstrate a natural or phyletic basis for

its recognition. However, component genera

appear somewhat more plesiomorphic than that

of Lysianassinae in the following morphological

tendencies: mouthparts basic (e.g., mandibular

molar present and palp inserted opposite or

distal to molar); coxal gills simple; uropod 3

usually natatory; telson lobes usually separate.

The subfamily Uristidinae also includes the

genera Uristes, Tryphosa, Orchomenella, Anonyx

and Allogaussia.

The genus Hippomedon was first established

by Boeck (1870) with the Arctic species Anonyx

holbolli Krdéyer, 1846, as the type, and for many

years encompassed species mainly from the

North Atlantic and Arctic oceans. The genus

was distinguished from other lysianassid genera

by such characters as a short first antenna with

a large first flagellar segment and comparatively

small accessory flagellum, a slender first gnatho-

pod with an oblique palm and a long dactyl, a

tooth on the third epimeral plate, a gill on

peraeopod 7, rami of uropod 3 not setose, and an

oblong, deeply cleft telson.

Gurjanova (1962) fused Paratryphosites to

Hippomedon and Barnard (1969) provisionally

synonymized with it also Paracentromedon

Chevreux and Fage, 1925, and Elimedon J.L.

Barnard, 1962. The fusion of genera and the sub-

sequent submerging and assignation of many new

species within Hippomedon has resulted in a

morphologically diverse genus which is excessively

large, poorly defined, and taxonomically

unwiedly. In this study, endemic North Pacific

species, type North Atlantic and other selected

species have been re-examined on the basis of

23 taxonomic characters of presumed or probable

generic value. (see Tables 1 & 2). The above

species are grouped into distinct genera as

diagnosed in the text and in the key below.

Key to Hippomedon and Related Genera

Mandible, palp seement 3 very short, length ’/, or less sepment2 .........5.....5.-. eee ps

Mandible, palp sepment 3 normal, length nearly equal to'sepmient 2 .......5..5....: «oe 3

Trae ASG NOBCSMse 17 pO PReANel eos cn ak Ch neeiae Bm he or POR thee on eee ae Elinedon

WReISCIO Ges MUSEO ee iets Se prc ae Seca: cad ate er oe eee ase eee treed eee ate Paracentromedon

Telson short, lobes fused '/, or more, each lobe broad, with 7 to 9 pical spines; antenna 2

more than twice as long as antenna |’; eill peraecopod 7 lacking .........6....7.++4+ eae

oeeeeeree ee eee ee ee ewe wee we wwe www eee wee ewe eee

Paratryphosites Stebbing, restored status (p. 120)

Telson long, lobes fused basally less than '/, length, each lobe broad, with 1-5 apical spines;

if antenna 2 more than twice as long as antenna |, gill peraeopod 7 present

Antenna 1, basal flagellar segments fused (conjoint per Barnard 1967); telson lobes well

tapered, apices with single spine (rarely 2-3); coxal gill on peraeopod 7 present, small

Hippomedon Boeck, sens. str. (p. 105)

Antenna |, basal flagellar segments not conjoint; telson lobes weakly or not tapered,

apices truncate, with multiple spines; coxal gill on peraeood 7 lacking

5. Antenna 1, peduncular segments 2 and 3 each longer than flaggellar segment 1, not tele-

scoped, peraeopod 5 much (25%) shorter than peraeopods 6 and 7; peraeopod 7 longest

Psammonyx Bousfield emend. (p. 118)

Antenna |, peduncular segments 2 and 3 each shorter than flagellar segment 1; peraeopod 5

slightly shorter than peraeood 6, peraeopod 6 longest

Genus Hippomedon Boeck, 1870 (sens. str.,

revised)

Hippomedon

— Sars, G.O., 1895, p. 55

Stebbing, T.R.R., 1906, p. 58

Gurjanova, E.F. 1962, p. 93 (part,

excluding Paratryphosites)

Purley, D.E., 1963,:p: 135) (part)

Barnard, J.L., 1964a, p. 5 (part)

Batnard, J.L., 1969, p. 345 (part,

excluding Paracentromedon, Elimedon

and Paratryphosites)

— Bellan-Santini, D. 1965, p. 161 (part)

Diagnosis: Body slender, elongate. Head small,

partially hidden by coxa 1. Eyes imperfectly

developed, often not visible in preserved speci-

mens. Coxal plates 1-4 moderately deep, narrow.

Epimeral sideplate 3 produced posterodistally

into an upturned tooth. Antenna | short, pedun-

cular segment | inflated, more or less produced

anterodistally into a beak-shaped process;

segments 2 and 3 short, deep, telescoping into 1;

flagellum, proximal segments fused (conjoint);

accessory flagellum relatively short. Antenna 2

slender, much longer than antenna 1; flagellum

of antenna | and antenna 2 calceolate in male

only. Mandibular molar strong, ridged; palp

slender, level with or slightly distal to molar;

segment 2 with a distal setal row; segment 3 more

than '/, the length of segment 2, medial margin

with comb setae. Maxilla 1, palp with broad

segment 2, bearing a row of apical spine-teeth;

outer plate with eleven apical pectinate spine-

teeth; inner plate with 2-5 apical plumose setae.

Maxilla 2, inner plate slightly shorter and broader

than outer. Maxilliped, outer plate reaches well

beyond segment 2 of palp, inner plate bearing

three strong teeth and two spines apically, a row

of plumose setae medially. Gnathopod 1, seg-

ment 6, palm long, oblique, usually weakly

105

Wecomedon new genus (p. 113)

defined; dactyl long, slender. Peraeopods 3 and

4, dactyls long, slender. Peraeopod 5 not

shortened, segment 2, hind margin tapering

distally. Peraeopods 5, 6 and 7, distal segments

slender; peraeopod 6 slightly longer than 5 and 7.

Coxal gills simple on peraeoods 2, 3 and 4, lobed

on 5 and 6, simple, reduced on 7. Uropod 3,

rami spinose, not setose. Telson oblong, lobes

fused basally, each tapering distally, apex with

single apical spine (rarely 2).

Type species: Hippomedon holbolli (Kréyer,

1846)

Remarks: To date, 42 species of Hippomedon

(sens. str.) are known in the world; most are

recorded from the North Atlantic and Arctic

oceans, but 6 species are known from the

southern hemisphere, especially the Tasman Sea.

H. geelongi Stebbing, 1888, from Australia,

H. tracatrix Barnard, 1971, from Oregon,

H. macrocephalus Bellan-Santini, 1972, from

Antarctica, and H. incisus K.H. Barnard, 1930,

from New Zealand, are questionably included in

this genus. Although their original limited

descriptions (in the literature) do not conform

with the present generic diagnosis, these three

species are retained in Hippomedon (sens. Str.)

until specimens can be examined and all

characters analyzed.

Barnard (1964) provisionally synonymized

Paracentromedon Chevreux and Fage, and

Elimedon Barnard with Hippomedon. However,

the unexpanded coxa 1, the short second antenna,

the short third segment of the mandibular palp

and the narrow segment 2 of peraeopod 5 and 6

exclude both of these genera from Hippomedon

(sens. str.) Barnard (1969) includes Paratrypho-

sites in his different brief diagnosis of Hippo-

medon, so that his diagnosis of the telson differs

from that given above for Hippomedon (sens. Str.).

Key to North Pacific and Related Species of Hippomedon (sens. str.)

1. Epimeral sideplate 3 with basal notch dorsal to the posterodistal tooth (rudimentary on

coecus)

Epimeral sideplate 2 without notch dorsal to the posterodistal tooth

2. Gnathopod 2, palm long, concave, dactyl much shorter than palm

Gnathopod 2, palm short, dactyl equal to palm

3. Antenna 1, peduncular segment | strongly produced anterodistally into an acute process,

reaching beyond segment 2 (North Atlantic)

oeeeeeeeeeeeeee eevee

denticulatus (Bate) (p. 108)

Antenna 2 peduncular segment | not strongly produced anterodistally, not reaching beyond

segment 2 (North Pacific)

4. Uropod 3, rami broad, margins parallel, tapering only at the tips, segment 2 of outer ramus

short

Cr |

orientalis Gurjanova (p. 112)

Uropod 3, rami with margins gradually tapering distally, segment 2 of outer ramus relatively

elongate

oeeeoveeeseeeeeeeeeeeee ee ee ee we we ewe em eh Oh Ohh Ohh OC HO HO HOO Om Om Ohm OO OOO oO oO OO hm Om Om Ol hh hl

Crystalline eye lens not present; urosome segment | dorsally smooth

6. Gnathopod 2, dactyl distinctly shorter than palm

Gnathopod 2, dactyl equal to palm

oeeeveeeeeeeeeeeeeeeeeeeeeeeeeee eee eee ewe eee ewe we eww wee wwe ewe wm em me wm em em hm Oh hh hh Oe

Peraeopod 7, posterior margin of basis weakly or not tapered distally

5. Crystalline eye lens present; urosome segment | with a prominent dorsal keel

7. Peraeopod 7, posterior margin of segment 2 (basis) strongly tapered distally

coecus (Holmes)

holbolli (Kroyer) (p. 106)

8. Gnathopod 1, segment 6 (propod) broadened distally; peraeopods 5, 6 and 7, posterior

marginal serrations of basis without setae

eous Gurjanova

Gnathopod 1, segment 6 not broadened distally; peraeopods 5, 6 and 7, posterior proximal

serrations of basis each with inserted seta

9. Gnathopod 2, segment 5 twice as long as segment 6; telson elongate, twice as long as broad ..

eeeeeeeeeeeeeeeeeeeee ee eee eeee eee ee eee eee eee eee eee eee eee ee

granulosus Gurjanova (p. 111)

Gnathopod 2, segment 5 less than twice as long as segment 6; telson of medium length, less than

PEE) | RnR rs Ei. as RS ats. SS A tenax Barnard

10. Uropod 3, rami broad, margins parallel, tapered only at tip ............... punctatus Gurjanova

Uropod 3, cami slender, margins pradually tapered distally +s... 2.0. 0.0. 0290 0.09 7 ee 11

11. Peraeopods 5, 6 and 7, serrations of posterior margins of basis without setae ..... propinquus Sars

Peraeopods 5, 6, and 7, posterior marginal serrations of basis with inserted setae ........... 12

12. Peraeopod 4, segment 5, posterior margin with 3 strong spines; uropod 2, peduncle and rami

spinose

Ce er

subrobustus Hurley

Peraeopod 4, segment 5, posterior margin lacking spines; uropod 2, peduncle and rami

smooth

Hippomedon holbolli (Kréyer, 1846)

Figure |

Anonyx holbolli Kréyer, 1846, t. 15, fig. 1 a-s

Hippomedon holbolli

— Sars, 1895, p. 58, pl. 21 no. 2

— Stebbing, 1906, p. 58

— Gurjanova, 1951, p. 229

— Gurjanova, 1962, p. 104, fig. 93

Material examined: 10 samples, 40 specimens.

Greenland, various coll., 1900 to 1933; Peary-

land, North Greenland, 4 June 1966, 105 m;

Murchison Sound (77°31.6’N,70°40’W), 24

August 1968, 50 m; northwestern Greenland

oeeveeeveeeeeeeeeeeeee eee eee eee wee ewe ere ee ewe ee ee ee ee em eH eH he eh ee em eH eH ee ee oo

106

zetesimus Hurley

(75°53.5’N,66°29.7°W), 8 August 1980, 72 m,

J. Just coll.; Beaufort Sea (70°33’N,145°40’W),

9 August 1972, 50 m, R/V Glacier, Department

of Oceanography, Oregon State University coll.;

70°13’N, 140°50’W, 4 April 1914, stomach

contents of seal, F. Johansen coll.

Diagnosis: Female ovig. 18.0 mm, Beaufort

Sea. Body integument with distinct surface

striations. Eyelobe not developed. Eyes with a

crystalline lens; crimson pigment in the eye region

which is visible in live specimens, disappearing

quickly in preservative. Antenna 1, flagellum

12-segmented, basal segments fused, segment 1

Figure 1. Hippomedon holbolli (Kréyer) Beaufort Sea 2 ov., 18.0 mm.

and 2 each bearing a strong posterodistal spine;

accessory flagellum 5-segmented. Antenna 2,

peduncular segment 5 slender; flagellum approxi-

mately 45-segmented. Mandibular palp, segment

3 slightly shorter than segment 2, strongly tapered

distally, inner margin with a dense setal row.

Maxilla 1, inner plate with 2 distal setae. Gnatho-

pod 1, segment 6, palmar margin oblique, convex,

subequal to posterior margin. Gnathopod 2,

segment 5 not distally bulbous; dactyl shorter

than palm. Peraeopods 3 and 4, segments 4, 5

and 6, posterior margin lined with many strong

spine groups. Epimeral sideplate 3 posterodistal

tooth moderately stout, upturned. Urosome

segment | dorsally with a depression followed by

107

a rounded crest. Uropod 1, peduncle and rami

with numerous short outer marginal spines.

Uropod 2, margins with few short spines.

Uropod 3, margins of rami non-plumose, with

short spines only. Telson lobes fused in basal

'/;, apex with notch and single small spine.

Male. 15.0 mm, Beaufort Sea. Antenna |

slightly longer than in female, 16-segmented.

Antenna 2 much longer than in female, approxi-

mately 70-segmented.

Remarks: Specimens from the Beaufort Sea are

very similar to those from Greenland. In large

males (15 mm) from Greenland, peduncular

segments 4 and 5 are armed anteriorly with

numerous brush setae. In seal stomach contents

from the Beaufort Sea, brush setae could not be

seen on smaller males although calceoli were

visible. Uropod 3 exhibits slight variation in the

rami; in Greenland material the rami are slightly

more broadly lanceolate than in specimens

from elsewhere.

Distributional ecology: A widely distributed

circumpolar species; in the Pacific Ocean,

extending possibly (but not authentically) into

the northern part of the Bering Sea.

Hippomedon denticulatus (Bate)

Figure 2

Anonyx denticulata Bate, 1857, p. 139

Hippomedon denticulatus

— Sars, 1895, p. 56, pl. 20

— Stebbing, 1906, p. 59

— Chevreux & Fage, 1925, p. 53, fig. 37

— Gurjanova, 1955, p. 233, fig. 96

— Gurjanova, 1962, p. 106, fig. 23

Lincoln, 1979; p. 75, fig. 28

Non Hippomedon denticulatus

Non

Barnard, 1954, p. 4, pl. 2-3

Barnard, 1964b, p. 80

Barnard, 1971 (forma — subrobustus

Hurley (from Oregon) p. 28, figs. 17-18

Barnard, 1971 (propinquus shelf form of

California) p. 30, fig. 193-m

Barnard, 1971 (form with gaped gnatho-

ned 2)ep. slits. 21

Barnard, 1971 (propinquus of north-

western Atlantic Ocean) p. 31, fig. 20

Hippomedon denticulatus orientalis

Gurjanova, 1962, p. 104, fig. 22.

Non

Hippomedon denticulatus orientalis

Gurjanova (off Alaska) = Barnard, 1971

p. 30, fig. 19 a-d.

Non Hippomedon denticulatus — Hurley, 1963,

p. 137, fig. 45

Material examined: 2 samples, approximately

Figure 2. Hippomedon denticulatus (Bate) Thor. I., Skaggerak & 11.0 mm, 92 9.5 mm.

108

50 specimens, Skagerrak, 6 March 1903, 140 m.

S.C. Johansen coll.; 28 October 1904, 44 m.

Description: Male 11.0 mm. Eyelobe small,

acute. Eyes narrow, linear, slightly widened

below, pigment (in life) light red with a few

opaque white stripes horizontally. Antenna 1,

peduncular segments 1 and 2 each produced

strongly anterodistally into an acute process.

Flagellum | 1-segmented, first fused segments and

article | with a sharp spine posteriorly; accessory

flagellum 3-segmented. Antenna 2, peduncular

segments 3-5 with brush setae anteriorly and a

few long setae posteriorly; flagellum 41-

segmented. Mandibular palp segments slender,

segment 3 distinctly shorter than 2, inner

marginal setae sparse. Maxilla 1, inner plate

with 2 apical setae. Gnathopod 1, coxal plate

broadened distally, distal margin nearly straight;

segment 6 slender, untapered; palm oblique,

about equal to posterior margin, defined by one

palmar spine. Gnathopod 2, segment 6, palm

short, transverse, dactyl equal to palm. Peraeo-

pods 3 and 4, segments 4, 5 and 6, posterior

margins with slender spine tufts; segment 4

produced anterodistally, dactyl long and thin.

Epimeral plate 3, hind margin produced into a

long, slender, well-tapered tooth with a distinct

dorsal notch. Uropod 3, rami long, slender,

sparsely spinose. Telson lobes fused basally,

inner margins proximally parallel, distally

diverging, apices with one spine.

Female (ov.) 8 mm. Antenna 1, flagellum 9-

segmented; antenna 2, flagellum 33-segmented.

Remarks: Sars (1895) figured males in which

antenna | was both long and short. All male

specimens from the Skagerrak examined here

had short antenna 1.

Barnard (1964b) opined that H. propinquus

Sars and H. denticulatus (Bate) are not specifi-

cally distinct; rather that H. propinquus is a

juvenile stage of the latter in which the dorsal

notch of the third epimeral plate and the antero-

distal process of the first two segments of

antenna 1 have not yet reached the sexually

mature form. Barnard (1971), further elaborated

this view, with particular attention to the epimeral

notch. He interpreted some small (5-6 mm)

sexually mature specimens from California and

Oregon that lacked an epimeral 3 notch to be a

dwarf southern race of H. denticulatus. He

identified mature specimens (larger than 5 mm)

with a notch from California and Oregon as

various forms of H. denticulatus, whereas those

109

specimens lacking a notch he designated as a

“propinquus”” phase. Barnard concluded that

Hippomedon populations from Oregon and Cali-

fornia form a complex of phenotypes of a single

species which could ultimately be linked to

H. denticulatus orientalis Gurjanova, 1962 of

the far eastern seas of the USSR.

In addition to AH. denticulatus, a notch on

epimeron 3 has been described in five other

species, viz., H. coecus Holmes, H. reticulatus

Stephensen, H. massilensis Bellan-Santini,

H. mercatoris Pirlot, and H. denticulatus

orientalis Gurjanova. H. denticulatus orientalis

was designated a subspecies by Gurjanova

(1962); however, this form is morphologically

distinct from the type species in many species-

significant characters of the overall body, eyes,

antenna 1, gnathopod 1, uropod 3, and telson

and merits full species status.

In juvenile and adult specimens (6.0 mm to

11.0 mm) of both H. denticulatus and H. propin-

quus from the Skagerrak, characters which

separate these species as adults were equally

applicable in the juvenile stage. Thus, in smallest

specimens of H. denticulatus (6.0 mm) the pro-

nounced first antennal anterodistal process is

distinct; moreover, the third epimeral tooth is

long and slender, with a distinct dorsal notch,

and quite unlike that of HA. propinquus of

comparable size. Such well defined ‘“‘mature’’

characters of specimens 6 mm in body length

cannot be growth-dependent, thereby confirm-

ing their value in species identification. These

two species are separated by characters given in

the key and in the text, notably by the first coxa

which, in H. denticulatus, is maximally expanded

at the flattened distal margin.

Other species from the northwest Pacific that

bear an epimeral notch (e.g., H. columbianus

n.sp.) are also found to be morphologically

distinct from Atlantic material of H. denticulatus

and H. propinquus (see also p. 111).

Hippomedon columbianus n.sp.

Pieure 3:

Hippomedon denticulatus Barnard, 1954, p. 4,

peed

Barnard, 1971, p. 31-34, fig. 21 (form with

gaped gnathopod 2)

Hurley, 1963, p. 137-140, fig. 45

Material examined: 36 specimens from 18

samples. British Columbia: Swanson Bay stns.,

J (53°0.58’N, 128°30.6’W) 67 m, 4 April 1973,

Figure 3. Hippomedon columbianus n.sp. & 7 mm 51B-029 Sta. J. Swanson Bay, B.C.

holotype male (7 mm figd. NMC-C-1981-1051),

allotype female ov. (8 mm figd. NMC-C-1981-

1052), NMNS slide mounts, paratype male

imm., 3 juvs. NMC-C-1981-1053; (53°0.76'N,

128°30.05’W) 45-47 m, 18 November 1975,

C.. kevirigs -coll.;, Verdier, Pt., Saanich Inlet

(48°22.5’N,123°29.W) 4-7 m, 19 March, 1 May

1975; Kx Conlan; cotl.;, French -Creek, north of

Nanaimo (49°20'N,124°25’W), 23 August 1977,

P. O’Rourke coll.; Bl4 Trevor Channel (48°

48.9'N,125°11°W) 40-50 m, 25 May 1977,

E.L. Bousfield coll.; W5-B, McCauley Pt.,

(33°507,130°20°W)~ 353°.m; 26.Aupust 1976,

G. O’Connell coll.; Vancouver Harbour (49°

18’N,123°12’W), 17 June 1976, C. Levings coll.

Diagnosis: Male holotype (7 mm). Eyelobe

moderately deep, subacute. Eye not visible in

preserved specimens. Antenna 1, peduncular

segment 2 with | large sensory seta distally; main

flagellum 10-segmented, segment 1 equal to first

110

two segments of 3-segmented accessory flagellum.

Antenna 2, peduncular segment 4 with 10, 5 with

14 short setal tufts anteriorly, long seta

posteriorly; flagellum 38-segmented. Antennae

calceolate. Mandible palp segment 3, setae on

medial margin not dense. Maxilla 1, inner plate

with 4 apical plumose setae. Maxilla 2, inner

plate broadened basally. Gnathopod 1, coxa

expanded distally; segment 6 slightly expanded

distally, setal groups on posterior margin and

medio-distally; palm oblique, distinctly shorter

than posterior margin of hand, dactyl hooded.

Gnathopod 2 segment 6 slightly expanded

distally, dactyl much shorter than concave

palm. Peraeopods 3 and 4, segments 4, 5 and 6

with long, slender spine groups posteriorly.

Peraeopods 5, 6 and 7, segment 2 weakly serrate

posteriorly; segments 4, 5 and 6, anterior margins

3-5 spine groups; posterior margins with a few

short spines. Epimeral sideplate 3 with a short

notch dorsally over a medium-sized tooth.

Urosome | with dorsal depression. Uropod 1,

peduncle lateral margin bearing 6-7 spines,

medial margin 9-10, margins of rami sparsely

spinose proximally. Uropod 2 peduncle with 4-5

spines, rami | or 2 each. Uropod 3 each ramus

bearing 2 or 3 short marginal spines. Telson with

3 to 4 pairs of dorsal spines, apices notched,

each with one spine.

Female allotype 8.0 mm ovigerous. Antenna 2,

flagellum shorter than male, 22-segmented.

Distributional ecology: H. columbianus has

been collected from the middle and southern

mainland coasts of British Columbia, and the

inner and outer coasts of southern Vancouver

Island in depths of 4 m to 67 m. Ovigerous

females were present in samples taken in March,

April, May and August.

Remarks: In H. columbianus specimens, the

epimeral notch is not visible at 2.0 mm, barely

visible at 3.0 mm to 4.0 mm, and easily visible

at 5.0 mm.

H. columbianus differs from Atlantic H. denti-

culatus in lacking a pronounced anterodistal

process on the peduncle of antenna 1, and the

third epimeral tooth is shorter and stouter, with

a smaller notch. It differs from both H. denticu-

latus and H. propinquus in the smaller relative

length of segments 5 and 6, the shorter and less

oblique palm of gnathopod 1, and in the presence

of 4, not 2, apical setae on the inner plate of

maxilla 1.

From the North Pacific species, H. subrobustus

Hurley and H. zetesimus Hurley, H. columbianus

is separable primarily by its third epimeral notch;

moreover, its telson is shorter, with broader

lobes than that in H. subrobustus, and segment 6

of gnathopod 1, the coxal plate of peraeopod 4,

and the spination of uropod 2 differ from

Hurley’s figures of H. zetesimus. The rather

slender tapering rami of the third uropods, and

the long palm of gnathopod 2 distinguish

H. columbianus from H. orientalis Gurjanova.

The third epimeral notch, visible in specimens

5 mm and larger, as well as the long palm of

gnathopod 2, separate H. columbianus from

H. denticulatus (f. subrobustus), (Barnard,

1971), and from H. denticulatus ( propinquus shelf

form) (Barnard, 1971). H. columbianus is possibly

synonymous with H. denticulatus (Barnard, 1954),

H. denticulatus (form with gaped gnathopod 2)

(Barnard, 1971), and H. denticulatus (Hurley,

111

1963). Although corresponding specimens have

not been examined, the figures and descriptions

agree reasonably closely with H. columbianus.

Hippomedon granulosus Bulycheva, 1955

Figure 4.

Hippomedon granulosus Bulycheva 1955, p. 195,

fig. 2.

Hippomedon granulosus Gurjanova 1962, p. 123,

fig. 31.

Material examined: Two samples, 2 specimens.

St. Lawrence Is., Bering Sea (64°N,169°W),

1 July 1980, 40 m; 10 July 1980, 30 m, coll.

J. Oliver.

Diagnosis: Female ov. (13 mm). Eyelobe

moderate, subacute. Eyes not visible in preserved

material. Antenna 1, anterodistal processes of

peduncular articles 1 and 2 moderate; main

flagellum 10-segmented; conjoint segments equal

to first 2 segments of 3-segmented accessory

flagellum. Antenna 2, peduncular segment 5

distinctly longer than segment 4, flagellum 25-

segmented. Mandibular palp, segment 3 strongly

tapered distally, not markedly shorter than

segment 2. Maxilla 1, inner plate with 3 apical

setae. Maxilliped outer plate densely toothed on

the inner margin. Gnathopod |, article 6, palm

oblique, distinctly shorter than posterior margin

of hand, defined by two palmar spines; dactyl

slightly longer than palm. Gnathopod 2 segment

6 short, broad; palmar margin long, concave;

dactyl considerably shorter than palm. Peraeopods

3 and 4, posterior margins of segments 4, 5 and 6

lined with long slender spines. Peraeopods 6 and

7, segment 2 barely tapered distally, posterior

margins serrate, notches with short setae.

Posterodistal tooth of epimeral plate 3 large,

acute. Uropod 3, rami elongate, strongly tapered

distally, 2 to 3 short spines on the outer margin.

Telson lobes fused basally, long, well tapered,

blunt distally, apices each with 3 small, unequal

spines.

Distributional ecology: Gurjanova (1962)

recorded this species from the Sea of Japan and

the Sea of Okhotsk at depths of 60 m to 134 m.

The samples from gray whale feeding pits,

northeast of St. Lawrence Island at a depth of

25 m are the first records of H. granulosus from

the north Bering Sea.

Remarks: Present specimens differ from those

of Gurjanova (1962) in being larger (to 13 mm

vs. 10 mm), possessing a few small spines on the

Figure 4. Hippomedon granulosus Bulycheva. St. Lawrence I. Bering Sea. 92 ov., 13.0 mm., o& penult.

9.0 mm.

rami of uropod 3 versus none in the type, and

having three apical spines on the telson lobes

versus two in Gurjanova’s specimens.

Hippomedon species indeterminate

The following regional taxonomic entities

recorded in the literature could not be critically

evaluated because specimens were unavailable for

examination, and the original descriptive com-

mentary and illustrations were too limited. These

include three forms of H. denticulatus from

Pacific coastal regions described by Barnard

(1977), viz. H. denticulatus (f. subrobustus

Hurley) from Oregon; H. d. orientalis Gurjanova

112

from Alaska; and H. d. (propinquus shelf form)

from California. As outlined previously (p. 109)

all Pacific denticulatus-like forms are _here-

with considered distinct from the Atlantic species,

H. denticulatus. Barnard’s figures of H. denticu-

latus f. subrobustus differ from H. subrobustus

Hurley in the following respects: antenna 2, ped.

segment 5 is distinctly longer than 4 rather than

subequal, peraeopod 5, segment 6 is longer with

more spines anteriorly; uropod 3, terminal

segment of outer ramus is much shorter; and the

telson is shorter and broader with 2 apical spines

per lobe.

H. denticulatus orientalis Gurjanova from

Alaska as described by Barnard (1971) differs

from H. denticulatus orientalis Gurjanova (1962)

from the U.S.S.R. in the following respects;

segment 6 of gnathopod | has parallel sides;

segment 6 of gnathopod 2 is barely expanded; the

distal lobes of coxa 5 are almost equal; segment 2

of peraeopod 6 is slightly tapered distally; and

segment 6 of peraeopods 6 and 7 have | or 2

spines posteriorly. And finally, figures of H.

denticulatus ( propinquus shelf form) differ signi-

ficantly from Atlantic specimens in the form of

gnathopod 1, especially the expanded coxa, sub-

equal segments 5 and 6, shorter palm, and

presence of two spines, and the posterior tooth

of epimeral plate 3 has a small dorsal notch.

These forms are all distinct from North Pacific

species treated herewith and in other papers.

Precise taxonomic placement of these three forms

awaits re-examination of the original material,

and more extensive series of specimens.

Wecomedon new genus

Diagnosis: Body elongate, moderately deep. Head

small, partially hidden by coxa 1; anterior head

lobe small to moderate, variable. Eyes whitish

in preserved specimens, moderately large, oval,

slightly widened below. Coxal plates 1-4 very

deep; coxa 4 weakly serrate on the posterodistal

margin. Epimeral plate 3 produced postero-

distally into an upturned tooth. Antenna 1

moderately short; segment 1, moderate to long,

weakly inflated, slightly produced anterodis-

tally; segments 2 and 3 not strongly telescoped

into 1; flagellum proximal segments normal, not

fused; non-calceolate. Antenna 2 slightly longer

than antenna | in the female, elongate and

occasionally calceolate in the male. Mandible,

palp, segment 1, distal setae few or lacking;

segment 3 more than '/, segment 2, medial

margin comb-setose, remaining surfaces sparsely

setose. Maxilla 1, palp densely toothed distally;

outer plate with eleven pectinate spines; inner

plate with 2 to 5 apical setae. Maxilliped, inner

margin of inner plate setose; outer plate extends

distad of palp segment 2, inner margin closely

spinose. Gnathopod | segment 5 longer than 6;

segment 6 long, narrow, palm slightly oblique,

distinctly shorter than posterior margin; dactyl

strong. Gnathopod 2 minutely subchelate.

Peraeopods 3 and 4, segments 4, 5 and 6, posterior

margins with slender spine tufts. Peraeopods 5,

6 and 7 unequal, 5 distinctly shortest, 6 longest;

coxal plate of 5 as deep as anterior margin of

segment 2; segment 4 moderately expanded.

Paraeopod 6 segment 2 broad, untapered;

peraeopod 7 very broad, weakly or not tapered

distally, posterodistal margin straight. Peraeo-

pods 2, 3 and 4 with simple, non-lobate gills,

peraeopod 5 coxal gill with one long, slender

accessory lobe, peraeopod 6 gill with 3 short

accessory lobes, peraeopod 7, gill lacking.

Uropod 3 rami subequal, lanceolate, margins

spinose, setose in male, sometimes setose in

female. Telson lobes broad, weakly tapered,

truncate apically, each bearing 2-5 spines

(rarely 1).

Etymology: A combining form of Hippomedon

and wecomus the type species.

Type species: Hippomedon wecomus Barnard

197M.

Additional species:

Wecomedon similis new species

W. wirketis (Gurjanova) 1962

W. boreopacificus (Gurjanova) 1962

W. minusculus (Gurjanova) 1938

Key to Species of Wecomedon

1. Uropod 3, outer ramus, segment 2 longer than '/, segment 1; telson lobes, apices blunt, each

mera laree thick spine: : 4...) . 16... Di eee

ee ee es ee ear W. minusculus (Gurjanova)

Uropod 3, outer ramus, segment 2 less than '/, segment 1; telson lobes apices truncate, each

with more than one medium spine ...........

2 Anterior head-lobe long, narrow, acute; uropod 3, outer ramus, outer margin with 4 or fewer

ET ha 8 ia cia hpi Mew catia ini Map e-g Sie b een

Anterior head-lobe short, stout, subacute; uropod 3, outer ramus, outer margin with more

Samer nelc Spies, proximal spines Offen IN STOUPS 2 1. on. cdyed sw OU ee Soe dis dco ates we ee 4

3. | Gnathopod 1, segment 6, palm defined by two large locking spines; telson lobes relatively

emt. Otoad, fused baSally /, ...........06.6%

CA iar < aakors W. wirketis (Gurjanova) (p. 116)

Gnathopod 1, segment 6, palm indistinct, locking spines lacking; telson lobes elongate, length

nearly 3 times width, fused basally Y, ........

atthe Se ee W. boreopacificus (Gurjanova) (p. 118)

4. | Gnathopod 1, segment 2, anterior margin strongly setose; antenna | and 2( 9 ) subequal;

epimeral plate 3 posterodistal tooth, long, slender

W. similis n.sp. (p. 115)

oeoe eee eee eee eee eee eee

Gnathopod 1, segment 2, anterior margin sparsely setose; antenna 2 ( @ ); distinctly (25%)

longer than antenna 1; epimeral plate 3, posterodistal tooth, short, stout

Wecomedon wecomus (Barnard, 1971)

Figure 5.

Hippomedon wecomus Barnard, 1971,

p. 37, figs. 24-25

Material examined: 53 specimens from 14

samples. Alaska: Bousfield stn. 1980, S11

Lisianski Strait, (58°06.4’N,136°27’W), 3 to

10 m., S18 Kamenoi Pt., Kruzof Is., (55°08’N,

135°34’W), 2 to 7 m, 2 August 1980. British

Columbia, Vancouver Island: Bousfield stn.,

1959, V22 Oyster Bay (49°55’N,125°11’W),

intertidal; 1975, P22 Trevor Channel (48°

49’N,125°12.5’°W) 27.5 m; 1976, B22 Dodger

Channel (48°50.4'N,125°12.1°W), 18 m;

1977, B18 Trevor Channel (48°48.8’,125°

oeeeeeee eee eeeeeee

W. wecomus Barnard (p. 114)

13.5°W) 36 m, B21 off Brady’s Beach

(48°50.2’N,125°08.5’W) 18-28 m.

Washington, USA: Bousfield stn. 1966, W235

Agate Beach Clallam Co., (48°15’N,124°16’W)

intertidal.

Description: Female (10.0 mm). Eyes verti-

cally ovate, bright red in life, indistinct in

preserved specimens. Antenna 1, peduncular

segment 1 long, weakly inflated; flagellum

17-segmented; accessory flagellum 4-segmented.

Antenna 2 slightly longer than antenna 1;

peduncular segment 3 with long setae medio-

distally, segment 4 setose posterodistally;

segment 5 slightly longer than 4 with long setal

tufts posteriorly; flagellum about 20-segmented.

aA AA

Figure 5. Wecomedon wecomus (Barnard) 9 br. II, 10 mm, Stn. B27, Dodger Channel, V.I., B.C.

114

Mandible, palp segment | with 2 short distal

setae. Maxilla 1, inner plate with 2 apical setae.

Gnathopod 1, anterior margin of segment 2

sparsely setose; segment 6 typical, weak setal tufts

on margins. Peraeopods 5 and 7, segments 4 and

5 subequal in length, segment 4 of peraeopod 5 as

broad as long; peraeopod 6 segment 4 slightly

longer than 5. Posterodistal tooth of epimeral

plate 3 short, stout. Urosome segment | with a

weak saddle-shaped, mid-dorsal depression.

Uropods | and 2, peduncles spinose, rami weakly

spinose. Uropod 3 outer margins of rami strongly

spinose, inner margins weakly setose. Telson

lobes relatively long, dorsolaterally with 2-3 pairs

of spines, apices each with 3 strong spines.

Male (penultimate. instar) 10.0 mm.

Antenna 2 flagellum 40-segmented, non-

calceolate.

Distributional ecology: Present material

NY zi

4 SOP We

8 oa a \ \

MS RTL

Figure 6.

bi5

ranges from southeastern Alaska, where its

distribution overlaps with W. similis, to

Oregon in sandy shallows, lower intertidal to

depths of 28 m (52-100 m Barnard, 1971). No

ovigerous females were present in_ the

material studied, (June-August).

Remarks: The subadult female (10 mm)

figured here agrees well with small specimens

(6.8 mm) figured by Barnard (1971). Larger

specimens (13 mm) possess longer antennae,

a more setose mandibular palp, more strongly

toothed maxilliped outer plate, more heavily

setose rami of uropod 3, and more dorsally

spinose telson lobes.

Wecomedon similis n.sp.

Figure 6.

Material examined: 21 specimens from 10

samples. Alaska stns.; St. Lawrence Island

Wecomedon similis n.sp. Stn. H13, Lelu I., B.C. 9 11 mm. o& 10 mm.

1965, whale stomach contents, G. Pike, coll.

Bering Sea, St. Lawrence Is., (64°N169°W)

13 m and 29 m, 10 July 1980, J. Oliver coll.

North central coastal British Columbia:

Bousfield ‘stn; 1 31° Lelu: Island. (54°21’N,

130°18’W), intertidal, fine, dark, sand flats,

14 July 1964, holotype female (11.0 mm figd.

NMC-C-1981-1054), allotype male, penulti-

mate (10.0 mm), NMC-C-1981-1055 NMNS

slide mounts; 2 paratype imm. females,

NMC-C- 1981-1056.

Diagnosis: Female holotype 11.0 mm. Eyelobe

short, rather broad. Eyes oval, indistinct in

preserved specimens. Antennae about equal in

length. Antenna 1, peduncular segment 1 long,

cylindrical; flagellum 14-segmented; accessory

flagellum 4.5 segmented. Antenna 2, peduncular

segment 3 setose mediodistally; segment 4 setose

posterodistally; segment 5 slightly shorter than 4,

bearing tufts of long setae posteriorly; flagellum

18-segmented, each segment with long postero-

distal setal tuft. Mandible, segment | of palp with

a long setal tuft distally. Maxilla 1, inner plate

with 2 apical setae. Maxilliped, outer plate

almost reaching segment 3 of palp. Gnathopod 1

anterior margin of segment 2 densely setose,

segment 6 typical, setal tufts on the anterior and

posterior margins. Gnathopod 2, segment 5 well

expanded distally with a prominent postero-

distal “‘cushion’’; segment 6 untapered distally.

Peraeopods 5 and 6, segment 4 slightly longer

than 5; peraeopod 7 segments 4 and 5 subequal.

Epimeral plate 3 posterodistal tooth moderately

long and slender. Uropod 1, peduncle outer

margin with 6-8 long spines, inner margin with

6 to 7 shorter spines; rami with 2 and 3 spines

proximally. Uropod 2 peduncle outer margin

with 4 to 5 spines; rami with 3 and | marginal

spines. Uropod 3, peduncle with 6-10 strong

spines distally; rami broadly lanceolate, outer

margins with 5 to 6 spines, inner margins 0 to

few spines. Telson lobes relatively short, broad,

each bearing 2 to 4 strong apical spines, and 3 to

4 dorsal spines.

Male allotype. (penultimate instar 10.0 mm).

Antenna 1, flagellum 21-segmented. Antenna 2

flagellum 46-segmented. Uropod 3, rami spinose,

inner margins setose.

Distributional ecology: W. similis was present in

samples from the North Bering Sea, south to the

north-central coast of British Columbia, bathy-

metrically from intertidal sand flats to depths of

29 m. No ovigerous females present.

116

Remarks: Material from the Bering Sea

(received after types had been designated and

figured) contained larger, more mature specimens.

The largest female (19.0 mm) agrees with the

figured specimen in most respects. The body

parts are more strongly developed, the armature

of spines and setae more prominent, but the

number of apical spines on the telson are fewer

(3-4). Of two mature males, one (16.0 mm) had

three calceoli on the second antenna, while the

other (14.0 mm) had none. Antennal calceoli may

be in the process of reduction and/or loss within

members of the genus.

Differences noted between the sibling species

W. similis and W. wecomus (Barnard) are slight.

W. similis 1s somewhat larger, more setose, and

more northerly in distribution. It is distinguished

by the moderately long, slender epimeral tooth,

by the subequal antenna (females), and by the

lack of a dorsal depression of urosome | (present

in W. wecomus). Both species are larger than

W. minusculus Gurjanova. This western Pacific

species also has a shortened segment 5 of

gnathopod 2, and a longer terminal segment of

the outer ramus of the third uropod.

Wecomedon wirketis (Gurjanova, 1962)

Figure 7.

Hippomedon wirketis Gurjanova, 1962, p. 115,

figs. 28a, 28b.

Material examined: 2 samples, 11 specimens.

Bering Sea, northeast of St. Lawrence Is.,

(64°N,169°W), July 1980, J. Oliver coll.

Diagnosis: Female ov. (18.0 mm). Anterior

head lobe narrow, sharp; inferior antennal sinus

distinct, bounded behind by cusp. Eyes moderate,

indistinct in preserved specimens. Antenna 1,

peduncular segment | moderately long; flagellum

15 segmented; accessory flagellum 4'/, segmented.

Antenna 2 slightly longer than antenna 1;

peduncular segment 3 with mediodistal setal tuft;

segment 4 shorter, broader than 5, posterior

margin weakly setose and posterodistally with

tuft of long setae; segment 5, posterior margin

with several setal tufts; flagellum 27 segmented,

proximal segments with posterodistal setal tufts.

Mandible, palp, segment 1 lacking setae.

Maxilla 1, inner plate with 4 or 5 apical setae.

Maxilliped, outer plate large, almost reaching

distal end palp segment 3; inner marginal teeth

present distally, reduced in size proximally.

Gnathopod 1, segment 2 setose anteriorly;

segment 6 palm oblique, strongly defined by

Figure 7. Wecomedon wirketis (Gurjanova) St. Lawrence I., Bering Sea. 2 ov. 18.0 mm., o& penult.

13.0 mm.

two locking spines, strongly setose anterior and

posterior margins. Gnathopod 2, segment 5

moderately expanded distally; segment 6 elongate,

slender, tapering distally, palm short. Peraeopods

5 and 6, segments 4 and 5 subequal in length;

peraeopod 7 segment 4 slightly shorter than 5.

Coxal gills typical for genus. Epimeral plate 3,

posterodistal tooth moderately stout. Uropod 1,

peduncle with 4 longish, marginal spines.

Uropod 2, peduncle and outer ramus, each with

about 7 short spines. Uropod 3, inner ramus

distinctly shorter than outer, both margins

weakly spinose; outer ramus, outer margin with

4 or 5 spines. Telson lobes fused in basal '/,, inner

margins rounded apically and bearing | or 2

apical spines.

117

Male. (penultimate instar 13.5 mm). Antenna 1,

flagellum 21-segmented. Antenna 2, flagellum

36-segmented. Uropod 3 rami spinose and setose

marginally.

Distributional Ecology: North Bering Sea,

25 m depth, to the Sea of Okhotsk and the Sea

of Japan at depths of 45 m to 115 m. Ovigerous

females in July.

Remarks: Material from the Bering Sea differs

slightly from Gurjanova’s description of

W. wirketis. The rami of uropod 2 are spinose

rather than smooth, and the rami of uropod 3 are

unequal, not subequal.

The presence of a small lower antennal lobe,

the well defined palm of gnathopod 1, and the

4 or 5 apical setae on the inner plate of maxilla 1

distinguish W. wirketis from the closely related

W. boreopacificus. The narrow, acute eyelobe,

and the elongate sixth segment of gnathopod 2

separate W. wirketis from W. similis n.sp., and

W. wecomus (Barnard).

Genus Psammonyx Bousfield 1973

Tmetonyx Stebbing, 1906 (in part)

Diagnosis: Body slender, elongate. Eyes when

present, small, subovate. Antenna | elongate;

peduncular segment 1 long, cylindrical, slightly

produced anterodistally, peduncular segments 2

and 3 not shortened; flagellum proximal seg-

ments not fused, accessory flagellum with

several segments. Antenna 2 less than twice

length of antenna 1. Antennae 1 and 2 calceolate

(male); 1 and/or 2 not always calceolate (female).

Mandible, molar strong; palp segment | usually

with distal setae; segment 2 with mediodistal

setae; segment 3 both margins setose. Maxilla 1,

inner plate with | or 2 apical setae. Maxilla 2,

inner plate slightly smaller than outer. Maxilliped,

outer plate short, not extending beyond palp

segment 2; palp stout. Gnathopod | subchelate,

palm well defined, dactyl strong. Gnathopod 2

minutely chelate or subchelate. Peraeopods 3 and

4, segment 4 strongly produced anterodistally;

segment 6 with 2 short spines posterodistally.

Coxa 4, posterodistal margin notched. Peraeo-

pod 5 much shorter than 6 and 7, coxa equal to

or deeper than segment 2; segments 4 and 5

expanded. Peraeopod 7 longest, lacking coxal

gill. Epimeral plate 3 with or without a postero-

distal tooth. Uropod 3 large, rami extending

well beyond uropods | and 2, rami marginally

spinose, and plumose. Telson lobes elongate,

fused basally, weakly tapered, apices subtrun-

cate, each with 2-5 spines.

Type species: Anonyx nobilis Stimpson, 1853

Additional species:

P. terranovae Steele, 1978

P. kurilicus (Gurjanova) 1962

P. longimerus new species

Key to Species of Psammonyx

1. Epimeral plate 3 with well developed posterodistal tooth; antenna 2 not calceolate in female;

gnathopod 1, segments 5 and 6 moderately long, narrow

Epimeral plate 3, posterodistal tooth minute or lacking; antenna 2 calceolate in female;

gnathopod 1, segments 5 and 6 short, broad

2. Epimeral plate 3, posterodistal tooth with a dorsal notch; gnathopod 2, segment 5 elongate,

slender, margins subparallel; uropod 3, segment 2 outer ramus minute

oeoeeveeeee ee eeeeeeeeeeeeee eee eee ee ewe wee ewww ww ewe wee wwe eee eee wee ee

P. longimerus n.sp. (p. 118)

Epimeral plate 3, posterodistal tooth lacking a dorsal notch; gnathopod 2, segment 5

relatively short, broadening distally, uropod 3, segment 2 outer ramus stout

oseeeoererweer ee ee eo @

P. kurilicus (Gurjanova)

3. Coxae | to 4, lower margins lined with setae; telson lobes each with 2 short apical spines;

anterior head lobe moderately produced, rounded

P. nobilis (Stimpson)

Coxae | to 4, lower margins bare; telson lobes each with 4 medium apical spines; anterior

head lobe slightly produced, subacute

Psammonyx longimerus n.sp.

Figure 8.

Material examined: 48 specimens from 15 samples.

Northern mainland coastal British Columbia:

Bousfield stn. H59, off Bolivar Is. (15°35’N,

128°08’W), 14.5 m, 9 August 1964, holotype male

(14.0 mm fig’d) NMC-1981-1057; allotype female

ov. (14.0 mm) NMC-C-1981-1058, NMNS slide

mounts, paratype male, 4 paratype female and

4imm. paratypes, NMC-C-1981-1059; H62, head

of River’s Inlet (51°41’N,127°1S’W) 18-27 m,

10 August 1964.

Southern mainland coastal British Columbia

118

P. terranovae Steele

and Vancouver Island: Bousfield stns. 1959;

N6 Raynor Pt. (51°08’N,127°41’W); V22 Oyster

Bay (49°55’N,125°11’W); 012 Ahous B., Vargas

Is. (49°11’N,126°01’W); 1964; H41 Jordan R.

(48°25’N,124°02’W); 1970; P703 McKenzie Beach

(48°45’N,125°15’W); 1975; P14 Keena Bay

(48°47’N,125°11’W); all collections intertidal.

Washington, USA: Bousfield stns. 1966: W22 Pt.

Grenville, Gray’s Harbor Co. (47°18'N,

124°16’W); W36 Clallam Bay, Clallam Co.,

(48°15’N,124°16’W); collections intertidal.

Oregon: Coos Bay, (43°20'N,124°30°W) 50-200 m,

April 1979 to May 1980, coll. J. Trautman.

ise

PARA

7 ett

’ Ssoaset

Figure 8. Psammonyx longimerus n.sp. Stn. H59, Bolivar I., B.C. 0 14 mm. 2 14 mm.

Diagnosis: Holotype male 14.0 mm. Anterior

head lobe small. Eye small, indistinct in pre-

served specimens. Antenna | elongate, flagellar

segments widest distally, distal margins fringed

with short setae, and bearing setal tufts anteriorly

and posteriorly; flagellum 23-segmented, calceo-

late; accessory flagellum 6-7 segmented, proximal

segments distally with long setal tufts. Antenna 2,

peduncular segments 4 and 5 with long posterior

marginal setae; flagellum 37-segmented, calceo-

late. Mandible, palp segment 3 slightly shorter

and more slender than segment 2, with 3 to 4 setal

tufts on the outer and inner margins; segment 2

inner margin with 3-4 setal groups. Maxilla 1,

palp broad with apical spine teeth in dense row,

inner plate with 2 setae. Maxilliped, inner plate,

apex transverse with 3 spine teeth. Gnathopod 1,

segment 5 longer than 6; segment 6 narrow, palm

slightly convex, distinctly shorter than posterior

margin, dactyl barely exceeding palm. Gnatho-

Ob)

pod 2, segment 5 elongate, margins subparallel;

segment 6 short, broad, minutely subchelate.

Peraeopod 3 anterodistal lobe of segment 4 over-

hangs one-third of segment 5; peraeopod 4,

anterodistal lobe of segment 4 overhangs two-

thirds the length of segment 5; segments 4, 5

and 6, posterior margin densely lined with long

setal groups. Coxa 4, posterodistal margin with

about 11 setal notches. Peraeopod 5, coxa as

deep as segment 2. Peraeopods 6 and 7, segment 2

deep, posterior margin with strong notches and

longish setae; peraeopod 7 distal margin of seg-

ment 2 straight. Peraeopods 5 to 7, anterior

margin of segments 4 to 6 each with 4-5 groups

of longish spines. Coxal gills on peraeopods 2,

3 and 4 simple, on 5 with 1, and on 6 with 3

accessory lobes; gill lacking on peraeopod 7.

Epimeral plate 3 with a slender, tapering,

posterodistal tooth, dorsal margin convex with a

distinct notch bearing | short seta. Uropod 1,

peduncle and rami strongly spinose, spines longer

and more numerous (about 8) on inner ramus.

Uropod 2 peduncle with long, slender spines;

lateral margin of outer ramus with short spines,

inner ramus smooth. Uropod 3, rami elongate,

narrowly tapering; inner ramus with both

margins bearing plumose setae; outer ramus with

inner margin bearing plumose setae, outer margin

with 8 pairs of spines. Telson lobes fused only

near base, each lobe with 3-4 dorsal spine groups,

2-4 single outer marginal spines, and 5 apical

spines.

Female ov. (14.0 mm). Both antennae lack

calceoli. Antenna 2 slightly longer than

antenna 1; peduncle 5 with long anterodistal

setae; flagellum 26-segmented; proximal segments

with tufts of long setae posterodistally; all

segments with short setal rows anterodistally.

Distributional ecology: From northern coastal

British Columbia south to Coos Bay, Oregon,

mainly on sandy bottoms from shoreline and

shallows to 200 m in depth. Ovigerous females

in July and August.

Remarks: The addition of P. longimerus n.sp.

and the transfer of H. kurilicus Gurjanova to

Psammonyx extends this previously Atlantic

genus into the North Pacific region. Both

Pacific species differ in having well developed

posterodistal teeth on epimeral plate 3, longer,

more slender segments 5 and 6 of gnathopod 1,

and a shorter segment 6 of peraeopod 7. These

and other differences might justify separate sub-

generic status for the Pacific species complex,

but formal proposal awaits further study.

Psammonyx longimerus is distinguished from

P. kurilicus by the elongate article 5 of gnatho-

pod 2, the pronounced anterodistal process of

segment 4, peraeopod 4, and the well developed

dorsal notch on the third epimeral tooth.

Genus Paratryphosites Stebbing, 1899

Paratryphosites Stebbing, 1906, p. 42

Diagnosis: Body elongate. Head relatively broad;

anterior head lobe small. Eyes (in alcohol)

medium large, oval. Antenna | peduncular

segment 1 slightly produced anterodistally;

segments 2 and 3 telescoped; flagellum short in

the female and elongate in the male; accessory

flagellum multiarticulate. Antenna 2 flagellum

very elongate in the male, terminal segments very

long and thin. Antennae | and 2 calceolate in

male. Mandible, palp segments 2 and 3 subequal;

segment 3 inner margin with comb setae; molar

120

massive. Maxilla 1, inner plate with 5 apical setae.

Maxilliped, inner plate short, not extending

beyond segment 2 of palp, margins of apices

concave. Gnathopod 1, segment 5 longer than 6;

segment 6 palm oblique, well defined. Gnatho-

pod 2, subchelate. Peraeopod 5 not shortened;

peraeopods 5, 6 and 7, segment 2 with posterior

margins coarsely serrate; peraeopod 6 longest.

Epimeral plate 3 produced in an upturned

posterodistal tooth. Peraeopods 2, 3 and 4 with

simple coxal gills; PS gill with one, P6 with 2

accessory lobes, peraeopod 7 coxal gill lacking.

Uropod 3 rami foliaceous (c'). Telson short,

broad, lobes fused '/,; apices broadly truncate,

each with 7-9 spines.

Type species: Paratryphosites abyssi (Goes,

1866),

Paratryphosites abyssi (Goes, 1866)

Lysianassa abyssi Goes, 1866, p. 519, plate

372 fig:

Paratryphosites abyssi — Shoemaker, 1930,

p. 18; fig: 10 (¢ - 7)

Hippomedon abyssi — Gurjanova (1962)

p.. 126-132). figs: 32a;°b, 33a, b

Material examined: 100+ specimens (d'o’,

13 mm - 16 mm.), Gulf of St. Lawrence,

Laurentian Trench, 300 m, July 1969. Coll:

Marine Sciences Centre, McGill University,

Montreal. 1 2 ovigerous (13 mm.) Entrance to

Baie des Chaleurs, stn. HP112M (48°18’07’N,

64°21722”W), 119 m., 26 June 1969.%ea

P. Brunel.

Remarks: In Paratryphosites abyssi the flagel-

lum of antenna | in the female is approximately

12 segmented, whereas it is approximately 30

segmented in the male. This sexually dimorphic

character of the first antenna is unique among

all the genera included in this study. Based on

the relative lengths of the antennae Shoemaker

(1930) has figured a female rather than a male.

Unfortunately, the size of the specimen figured

was not given and the single specimen has not

been re-examined. Coxal gills were lacking on

peraeopod 7 in all specimens.

Through the courtesty of Dr. Pierre Brunel we

reproduce herewith a description of the colour

present in a mature male taken from a specimen

about a week after capture, the specimen having

been preserved in 4% formaldehyde. “The eyes,

which have never been observed or figured so far,

were orange red, and so were three oblique

lateral stripes on the metasome, which may be

muscle insertions. Whole animals were other-

wise whitish.”

Addendum

Material of the following species, from

Gurjanova’s original localities in the far eastern

seas of the USSR, has very recently been made

available to us through the courtesy of Dr. Nina

Tzvetkova, Zoological Museum, Leningrad:

Hippomedon wirketis (Gurjanova 1962) — 9

br. II Kurile Straits

Hippomedon kurilicus (Gurjanova 1962 — ¢&

and Q specimens, N. Kuriles.

Hippomedon pacificus Gurjanova 1962 — ¢&

and 9 — Okhotsk Sea

Hippomedon minusculus Gurjanova 1938 — 4

29 — S. Kuriles.

Hippomedon propinquus eous Gurjanova 1962

— oo and 2 — Okhotsk Sea.

Hippomedon denticulatus orientalis Gurjanova

1962 — o& & @ — Shikotan I.

Hippomedon punctatus Gurjanova 1962 — o&

and 9 —N. Kurile Islands.

The specimens of H. pacificus, propinquus

eous, and punctatus fairly closely match the

descriptions and illustrations of Gurjanova

(1962). However, some discrepancies were noted

when comparing the specimens of H. wirketis,

kurilicus, minusculus, and d, orientalis with the

original descriptions and illustrations. In these

instances, the specimens were apparently not

from the type localities but from distributional

localities of the original text. The possibility that

Dr. Gurjanova’s original material concealed

greater taxonomic diversity than described in her

very fine and comprehensive monograph (1962)

would merit further investigation.

Discussion

A character analysis was carried out on 21 species

of Hippomedon (sens. lat) and 2 species of

Psammonyx using numerical taxonomical methods

(Sneath & Sokal, 1973). Specimens were unavail-

able for the following eleven species (character

data from litt. only): H. minusculus Gur). (Sea of

Japan), H. punctatus Gurj. (Sea of Okhotsk),

H. kurilicus Gur}. (Sea of Okhotsk, Bering Sea),

H. pacificus Gurj. (Kuriles), H. denticulatus

orientalis Gurj. (Chuckchi and Bering Seas),

H. coecus (Holmes) (California), H. subrobustus

Hurley (California), H. zetesimus Hurley

(California) and H. tenax Barnard (California).

Owing to limitations imposed by the literature,

the analyses were restricted to twenty-three

characters considered to be of generic value.

Character states were assigned values of 0 =

primitive or plesiomorphic, .5 = intermediate,

1 = advanced or apomorphic (Table 1), and a

raw date matrix compiled therefrom (Table 2).

In this analysis, advanced species would have

high total scores, primitive species would have

low total scores. The similarity coefficient used

to calculate the similarities between all pairs

of species was that of Rogers and Tanimoto.

(Sneath & Sokal 1973, p. 132) and formed the

basis for the similarity matrix of Table 3. The

clustering technique used to define clusters of

mutually similar taxa was that of Complete

Linkage Clustering (Sneath & Sokal 1973, p. 222)

from which the phenogram (fig. 9) was constructed.

This phenogram reveals four species groupings

having an overall dissimilarity of 50% or greater

in 23 selected characters. Differences of such mag-

nitude between these groupings confirms the

validity of the existing generic concepts of

Hippomedon Boeck (Sens. Str.), and Psammonyx

Bousfield, the re-establishment of Paratryphosites

Stebbing as a separate genus and creation of the

new genus Wecomedon herewith.

These four generic groups are quite unequal in

number of species and in relative plesiomorphy-

apomorphy. Thus, in the north Pacific region,

Hippomedon (Sens. Str.) is by far the largest

group with 13 species, all relatively apomorphic

(values of .57 - .85); Wecomedon is next largest

with 5 known species, of medium apomorphy

(values of .45 - .50); Psammonyx is yet smaller,

with two Pacific species, of greatest plesiomorphy

(values of .30 - .35), whereas the monotypic

Paratryphosites of holarctic and North Atlantic

affinities (not yet actually recorded from the

North Pacific) contains only one species, of inter-

mediate apomorphy (value of .57).

The general correlation between the range of

plesiomorphy-apomorphy values and the

pertinent species groupings tends to confirm

these genera as “‘natural” rather than “‘artificial”’

or convergent.

The temptation is strong to carry this analysis

to subgeneric levels (at 50-75% similarity levels).

Thus, within Psammonyx, some justification

might be found for formal subgeneric recogni-

tion of the Pacific /ongimerus and kurilicus from

the Atlantic nobilis and terranovae on the basis

punctatus

granulosus

col umbi anus

subrobus tus

orientalis

denticulatus

tenax

coecus

7/4 pacificus

eous

Lo)

ie)

ie}

i=)

lee)

zetesimus

foo)

propinquus

ine)

holbolli

abyssi

-50 wirketis

minisculus

boreopaci ficus

similis

we comus

kurilicus

longimerus

terranovae

nobilis

—~s

[<> }

Figure 9. Phenogram of species and genera of Hippomedon sens. lat. complete linkage cluster analysis.

Number beside each species is index of apomorphy.

of their prominent epimeral 2 tooth, less fossorial

peraeopod 5, lack of antennal calceoli in the

o&, etc. Such temptation must be resisted in the

much more diverse genus Hippomedon since only

a small fraction of known world species are

represented in the Pacific regional fauna. Despite

their relative overall apomorphy, all members of

Hippomedon retain some plesiomorphic charac-

ters such as calceolate antennae (co'), and the

presence of coxal gills on peraeopod 7. A litera-

ture review of additional species showed that the

Atlantic species exhibit the most advanced form

of the first gnathopod, the palm becoming so

122

indistinct that it is almost simple (H. serratus),

and also of the anterodistal process of antenna |

which is more pronounced (H. denticulatus,

H. nasutus, H. serratus) than in any of the

Pacific species. Many species of Hippomedon

previously thought to be lacking eyes have

proven to have eyes of a pigmentary type which

disappear completely after preservation. Descrip-

tions of eyes within this genus should be based

on animals which have been studied in the living

condition (Vader 1968). A more primitive condi-

tion exists in H. holbolli previously known to have

a crystalline eye lens in preserved specimens and

now known to have an additional large pigmen-

tary component which is visible in live specimens

(J. Just pers. comm). This eye type was also

described by Bellan-Santini (1965) for H. massi-

liensis from the Mediterranean. Preserved

specimens of H. gorburnovi and H. rylovi from

the Arctic have crystalline eye lenses. However,

formal recognition of other supraspecific groups

awaits a more comprehensive study of the world

fauna.

The five species presently comprising the new

genus Wecomedon are less advanced than those of

Hippomedon in most respects, except for the loss

of the coxal gill on peraeopod 7 and apparent

incipent loss of the calceoli on the male antenna.

Analysis is hampered by a scarcity of mature

males in the collections, but in the single calceo-

late male specimen available, the number of

calceoli was reduced to 3. The relatively small

spread of character indices (.46 - .50) suggests a

relatively tightly knit complex of species of

relatively low diversity as compared to the genus

Hippomedon. The peduncle of antenna | has a less

expanded first segment, and segments 2 and 3 are

less telescoped. The proximal flagellar segments

are not fused, anc although segment | is slightly

longer than the following segments, it is always

much shorter than the accessory flagellum.

Antenna 2 ( 9 ) is not much longer than antenna 1,

and the weakly tapered telson lobes with multiple

apical spines are also primitive. Wecomedon

appears intermediate in almost every respect

between the apomorphic genus Hippomedon and

the plesiomorphic genus Psammonyx.

Psammonyx as the most primitive of the genera

h23

analyzed. The antennae, uropods and telson are

especially plesiomorphic (see Table 1). On the

other hand, the fifth peraeopod is highly special-

ized and fossorial in function.

The posterodistal tooth of epimeral plate 3,

rudimentary in the Atlantic species, is developed

to a much greater extent in their Pacific

counterparts.

In gnathopod | of the Pacific species, the coxa

is not reduced and segments 5 and 6 are longer

and more slender, and segment 6 of peraeopod 7

is not as long as in Atlantic species. Both groups

possess setose mandibular palps, the setose third

uropods, and the long, deeply cleft, multi-spined

telson, all primitive characters.

The monotypic Paratryphosites, having an

intermediate character index of .57, exhibits

characters present to some extent in each of the

other genera but in a combination that is unique

to this genus. In addition, the short broad telson,

cleft less than half-way, the massive flat mandi-

bular molar, the shortened article 5 of gnathopod

2 and the sexually dimorphic antennae are not

found in any other genera.

Future analysis of the remaining lysianassid

material from the Canadian Pacific coastal areas

will hopefully provide insights into taxonomic

relationships which exist between these genera

and others. Hopefully also, the true nature and

significance of geographical differences between

populations and between species and genera will

be revealed through more extensive collections

both geographically and bathymetrically and

more extensive application of analytical

methodology.

Acknowledgements

The writers are greatly indebted to Mrs. Floy

E. Zittin, Vancouver, B.C., for her most capable

assistance with the illustrations. They are grateful

to Dr. Jean Just, University Museum, Copen-

hagen, for the supply of regional material of

H. holbolli, H. denticulatus and H. propinquus;

to officers of the Department of Oceanography,

Oregon State University for providing vital

material of Hippomedon from the North Bering

Sea; and to D.H. Steele, Memorial University,

St. John’s, Newfoundland, for specimens of

Psammonyx terranovae. Dr. Wim Vader, Tromso,

Norway, and Dr. Pierre Brunel, Université de

Montréal, kindly reviewed the manuscript and

Dr. John J. Dickinson supplied frequent helpful

advice, and assistance with the material.

124

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afrikas 3: 111-204

125

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Table 1. Data matrix showing character state as defined in Table 2 of North Pacific and selected North

Atlantic species of Hippomedon (sens. lat.)

P.A.

CARN ee Ae ere ee are ee 1S De eee

Ps. nobilis 0050.0 OO). Bie Ws TT Oe DO Oct Oy rr

Ps. kurilicus OO. O00. ONE SD TO LT O-8- 3 3 PF Resi eee

Ps. longimerus OO O2070; 0 pr O2 AL res 1 0.1.06 1. 0.0 4.1 Af) 0) ee

Ps. terranovae 0-0 O'0O'O 96 8 eo 1 - fr B® 2 O01 1 2e0 at. On ieee

W. minusculus O00 1 0 ) Ne NE NC’ 10.5 0 0 5:0 0 LL. fd 2a

W. similis Ue oS Ooo eo. Oe |) kd, Or uees: Ameo al 103 tee

W. wirketis 0:0 OO. Net a? 0 OT OO Fe! 3 Oo Uy ey 1S

PaorpercoracincHs oc) Oy On O00 kh NEING NC «1 OP for OR Ss S25 rd | ne ae ee 50

W. wecomus OO sO INC Oe dee TO Od Oeud Bien det Oiled hy) cde yd

Pa. abyssi aStLO1Ltt?t oO 1 °1.0 OO £45 21-0 Oy dae eee

H. punctatus Sieben tone wis Oo. O 1°00 F OM 2D 2-150 eee 57

H. columbianus oy ae ee A Ed Ogee Oe OS. Oe lee). (Oirte ale ote 0 .63

H. eous Se OAT SOAP ON Re OB Pe a ce .64

H. granulosus Seer hee: (dnt Ok el mS Oot Ss OT OT 0?) 3° eee

H. coecus ieleal. Med toe at. b -O) d OOS OCP INE” ee .69

H. orientalis 20 ott ft. ONC UCP ok Oe 6 3 FY PO 01250 Sa .70

H. propinquus og halite oo ke olpy Lo Orel bo, Wh 0 te

H. pacificus 2 1 te tty fONCANG yh 3S Of 25 0 1. Sopdet 62) bea .74

H. tenax PAL bay fi NCINCH NC fo OS. .5 OT Se. NC. ih, oe ae 74

H. holbolli Sted sinh his eh es 1. aR 2 OO a 0a i ise ee .76

H. subrobustus NOD ie > ee Perak £ ob) bob OV Ne Sew 719

H. denticulatus PS a Oe Bh SB Bie “ok. oh hg Ow. ung geen .80

H. zetesimus oot) be ok INR Td NCO Pb Dd INC. oes .85

P.A. Index is the index of plesiomorphy vs. apomorphy. The higher the number the more advanced the character

states.

State 0 = Character primitive (plesiomorphic)

.5 = Character intermediate

1 = Character advanced (apomorphic)

NC = No comparison, missing data

126

Table 2. Qualitative Multistate Characters

A, ped. seg.1 anterodistal process well

developed

absent

A, ped. seg. | short, well inflated, segs.

2 & 3 short

A, ped. seg. | long, weakly inflated, segs.

2 & 3 long

. A, main flag. proximal segs. conjoint

segs. distinct not conjoint

A, mainflag.short ,

elongate,

A, more than twice length A,

less than twice length A,

A, flag. segs. simple

well developed

Antennae not calceolate

calceolate

Mandibular palp seg. | without distal setae

with distal setae

Mandibular palp seg. 3 comb setose medially

not comb setose (other setation)

Mx, inner plate 2 setae apically

more than 2 setae apically

. Coxa | reduced

not reduced

Gn, seg. 5 greater than 6

seg. 6 greater than 5

Table 3. Similarity Matrix

nobilis | a4

terranovae By. 2) Xx.

kurilicus 1459 52:63 XX

abyssi | Nees 2 ales yam oe 6

wirketis B20 2028 3k. 39) x

minusculus Gr 40.240 56° 034. .26..40 -X

wecomus HF 236.2238. .38* 230 242) 32) 56

similis bE 39) 39.439 <52.A4: 48) 356

Wereopaciicus J .29 .29 .38 .38 .29 .60 .73

holbolli mK 09 212. 5 19° ST) 24 Bl

Seaeeatus L .15 .15 .21 .18 .35..24 .35

propinquus Me 15) 12 2026 239 124 26

Cammpianus N .12 .12 .18 .26 .35 .35 .35

pacificus Ph) es 938 725

eous O10) 10. 1322) ".29 29 33

orientalis Bets 2.16. 167.38 22.29

punctatus Si toi 1922: 38 nF eek

coecus LT. 24: 2 33. .29' 3 31 Al

zetesimus 8 3 NS) AS AS 23.586

supbropustus V .16 .16 .24 .21 .45 .20 .36

tenax W 09° 09.13. 23: 36-19. 33

o-_

—

Or Or Or Or Or Or Or Or OF OF OO

—

$42

23).

Gn, palm oblique, not well defined

palm transverse, defined

Gn, palm short

palm long

P, and 4dactyls long

dactyls moderate

. P,shortened, coxa as deep as seg. 2

not shortened, coxa not as deep as seg. 2

. P, seg. 2 tapered

not tapered

. Gillabsent, P,

present, P,

. Epimeral sideplate 3 with a well-

developed tooth

sideplate 3 without a well-

developed tooth

U, and U,, ped. and rami mainly smooth

mainly spinose

. U,rami spinose

rami setose

U, rami margins tapered throughout length

rami margins parallel, tapered only at tips

Telson lobes well tapered, | spine apically

lobes weakly tapered, multiple

spines apically

.66 X

54 .80 X

Or Or Or Orroroo;

—

J. Kod SME ONS OU oP RS i ie Se any

32.58 64. X