Contents

Résumé, vii

Summary, Vii

Introduction, 1

Family Caprellidae, 4

Key to the Caprellidae of the American North Pacific, 6

Cercops, 9

C. compactus, 9

Perotripus, 12

P. brevis, 12

Deutella, 16

D. californica, 16

Mayerella, 19

M. banksia, 19

Tritella, 22

T. laevis, 22

T. pilimana, 25

Metacaprella, 29

M. anomala, 29

M. kennerlyi, 31

Caprella, 35

C. alaskana, 35

C. ferrea, 35

. angusta, 40

. incisa,:42

. verrucosa, 44

. borealis, 47

. californica, 4°

drepanochir, 53

. equilibra, 55

. mendax, 58

C. pilidigita, 60

C. gracilior, 62

C. irregularis, 65

C. laeviuscula, 67

C. rudiuscula, 70

C. pustulata, 72

C. striata, 76

Pseudoliropus, 78

P. vanus, 78

AAaaaanns

ECOLOGICAL AND ZOOGEOGRAPHICAL DISCUSSION, 80

REFERENCES, 87

List of Figures

OMAN MN PWN

Cercops compactus, 11

Perotripus brevis, 14

Deutella californica, 17

Mayerella banksia, 20

Tritella laevis, 23

Tritella pilimana, 27

Metacaprella anomala, 30

Metacaprella kennerlyi, 33

Caprella alaskana, 36

Caprella ferrea, 39

Caprella angusta, 42

Caprella incisa, 45

Caprella verrucosa, 47

Caprella borealis, 50

Caprella californica, 52

Caprella drepanochir, 54

Caprella equilibra, 56

Caprella mendax, 59

Caprella pilidigita, 61

Caprella gracilior, 63

Caprella irregularis, 66

Caprella laeviuscula, 69

Caprella rudiuscula, 71

Caprella pustulata, 75

Caprella striata, 80

Abdomina of 1. Perotripus brevis, 2. Tritella laevis, 3. Deutella californica,

4. Mayerella banksia, 5. Metacaprella anomala, 6. Caprella pustulata, 82

MAPS

Known distribution of (a) Perotripus brevis, (b) Deutella californica, 15

and (c) Mayerella banksia within the American Pacific boreal region,

Known distribution of (a) Tritella laevis, (b) T. pilimana, and (c) Metaca-

prella anomala within the American Pacific boreal! region, 24

Known distribution of (a) Metacaprella kennerlyi, (b) Caprella alaskana,

and (c) C. ferrea within the American Pacific boreal region, 34

Known distribution of (a) Caprelia angusta, (b) C. incisa, and (c) C.

verrucosa Within the American Pacific boreal region, 43

Known distribution of (a) Caprella borealis, (b) C. californica, and

Known distribution of (a) Caprella equilibra, (b) C. mendax, and (c) C.

pilidigita within the American Pacific boreal region, 57

Known distribution of (a) Caprella gracilior, (b) C. irregularis, and (c)

C. laeviuscula within the American Pacific boreal region, 64

Known distribution of (a) Caprella rudiuscula, (b) C. pustulata, and (c)

C. striata within the American Pacific boreal region, 72

Résumé

Le présent ouvrage décrit vingt-six espéces de Caprellidae du littoral nord-

américain du Pacifique, appartenant a huit genres différents. Ce nombre com-

prend un genre nouveau, Pseudoliropus, et six espéces nouvelles: Cercops compac-

tus, Mayerella banksia, Caprella pilidigita, C. rudiuscula, C. pustulata et Pseudo-

liropus vanus. L’auteur traite de la zoogéographie de la région et expose cer-

tains facteurs écologiques qui influent sur la répartition des caprellidées dans

cette région.

Summary

Twenty-six species of Caprellidae, representing eight genera, are described from

the North American Pacific region. Included are one new genus, Pseudoliropus,

and six new species: Cercops compactus, Mayerella banksia, Caprella pilidigita,

C. rudiuscula, C. pustulata, and Pseudoliropus vanus. The zoogeography of the

region is discussed, and certain ecological factors affecting the distribution of the

caprellids within this region have been noted.

Vii

Introduction

Since Mayer’s monographic works (1882, 1890, 1903) on the caprellid amphi-

pods first revealed the diverse nature of the North American Pacific fauna, few

studies have been published on the systematics, distribution, or ecology of

these animals.

Holmes (1904) confined his studies to the Prince William Sound and Aleutian

Island region of Alaska, and (1909) recorded only one species, from California.

La Follette (1914, 1915) and Shaw (1916) published accounts of seven species

from Laguna Beach. Wailes (1931) included two caprellid species in a general

list of intertidal amphipods from British Columbia. Johnson and Snook (1927)

and Ricketts and Calvin (1952) included brief descriptions of a few species of

caprellids from the general North American Pacific area. Dougherty and Stein-

berg (1953) studied the caprellids of California between latitudes 36°N and 38°N

and listed twenty species, including two new species. Saunders (1962, 1966) has

done some interesting work on the ecology and behaviour of a few species, and

has identified eleven caprellids from the San Juan area.

Study of these works shows that, apart from Mayer, no one has investigated

the caprellids of the North American Pacific region as a whole, and that large

parts of this region, e.g., Oregon and British Columbia, have never been inves-

tigated. It is perhaps surprising that more than thirty species of caprellids have

been recorded from this region despite the lack of thorough study.

In contrast, much work has been done on the caprellids of the corresponding

region of the western Pacific. Arimoto (1929, 1930, 1931, 1934) and Utinomi

(1931, 1937, 1943, 1947) in Japan, and Gurjanova (1933), Kudrjaschov and

Vassilenko (1966), Schurin (1935, 1937), Stschapova, Mokyevsky, and Pasternak

(1957) and Vassilenko (1967) in Russia, have published on the caprellids of the

Sea of Japan and the northwestern Pacific, and have listed over sixty species

of caprellids from this region. It is apparent that, as is the case with other ani-

mals (Ekman 1953), the North Pacific caprellid fauna is very rich, and it can be

expected that more intensive investigation will show the North American fauna

to be nearly as diverse as the Japanese fauna.

New material has become available that further extends our knowledge of

the range and numbers of caprellid species in the North American Pacific.

During the period 1955-1966 inclusive, Dr. E. L. Bousfield (1956, 1961, 1963,

etc.) of the National Museums of Canada made summer collections of crustacea

and other invertebrate animals from the Pacific coast, from Oregon, at approx-

imately latitude 44°N, north to Prince William Sound and Seward, Alaska.

Caprellid amphipods from these intertidal invertebrate collections were studied

as well as those from general collections made by other National Museums of

Canada investigators, and from collections in the United States National Mu-

seum, from the same geographical region. Although no special effort had been

made to collect caprellids during these surveys, 171 of the 502 localities did con-

tain caprellid material, which yielded twenty-six species in eight genera.

The purpose of this paper is, therefore, to describe all the species recorded

in this new material; to construct a key for the identification of these species;

and to reach a tentative conclusion on the effect of temperature and salinity on

the distribution of these animals, with a view to predicting which areas should

be particularly rewarding in future study.

It has been found that in the American Pacific between latitudes 40°N and

60°N, approximately, general intertidal collections contained twenty-six species

of caprellid amphipods, six of which were new to science. Two of the eight genera,

Cercops and Mayerella, are new to this region, and one, Pseudol’ropus, is new

to science. Knowledge of the ranges of twenty species has been extended.

Although most of the species found had extensive ranges within this region, due

to the general uniformity of temperature, more specific indications of the effects

of local temperature and salinity values have been noted for the commoner

species.

It has also been noted that of the forty caprellid species recorded from the

boreal region of the American Pacific, eleven are found also in the northwestern

Pacific, two are found also in the Atlantic, and twenty-seven of the species are

apparently endemic.

The author thanks the National Museums of Canada for making this study

possible, particularly Dr. E. L. Bousfield for the use of collections and for his

invaluable advice and criticism; Dr. John C. McCain, of Oregon State Univer-

versity, for his advice and for the loan of material in the United States National

Museum; Dr. Marvin P. Jessen, of Grand View College, Iowa, for the use of his

collection of Cercops compactus n. sp.; and Dr. Tetsuo Matsumura, of the

National Research Council, for his help in the translation of the Japanese

literature.

Illustrations

All illustrations were made to scale, using a projector. Scale is shown only

for the whole mount, where it equals 1 mm unless otherwise indicated.

Key to symbols used:

A = antenna MDP = mandibular palp

ABD = abdomen MX = maxilla

BR = brood plate MXPD = maxilliped

GN = gnathopod P = pereopod

HD = head PEN) = penis

LEY = leit PER = pereonite

LL, . = lower lip RT = right

MD = mandible UR = uropod

Maps

These show the known distribution of the species within the region under

consideration. The closed circles indicate localities from which specimens were

examined. The closed triangles indicate references in the literature.

Collecting Stations

Detailed locality data has not been given for most of the species here de-

scribed. NMC Expedition Station Lists have been published for nearly all the

stations cited (Bousfield 1956, 1963, 1968, in prep., .. . and McAllister 1962).

The five unpublished stations are as follows:

3532, 1935. July 1, 1935. Skidegate Channel, Queen Charlotte Islands.

JWS 85 August 6, 1965. Lawn Point, B.C., latitude 53° 32’ N, longitude

131° 09’ W. 30 fm.

JWS 92 August 8, 1965. Queen Charlotte Islands, latitude 51° 05’ N, longi-

tude 132° 22’ W. Trawl, 1550 fm.

NAP66-201 July 16, 1966. Whiffin Spit, Sooke Harbour, Vancouver Island.

WVV 1966 August 30, 1966. Dixon Entrance, B.C., latitude 54° 16’ N, longi-

tude 132° 40’ W. Dredge, 80 fm.

Measurements

All measurements were made by recording the distance from the anterior of

the head, between the points of attachment of the two pairs of antennae, to the

posterior tip of the abdomen, through the longitudinal mid-line of each pereo-

nite. The lengths recorded for each species are of the largest male found and

of an average large ovigerous female.

Family CAPRELLIDAE

According to Stebbing (1906):

Head fused with Ist segment of peraeon. Palp of maxilliped 1- to 4-jointed. Peraeon

often with fewer than 7 pairs of legs; 2, rarely 3, segments of peraeon with branchial

vesicles; 2 segments of 9 with marsupial plates; Ist joint of gnathopods and peraeopods

wanting. Pleon and its legs rudimentary. Eyes small, 1 pair. Hepatopancreatic tubes 2;

rectal glands none. Heart with 3 pairs of ostia. Posterior ganglia of nerve-chain very

small, none situated in pleon.

All specimens examined by the author showed at least a part of the suture

between the head and pereonite I, indicating that even in the most advanced

forms (e.g., Caprella) the fusion between these parts is not complete. All speci-

mens have coxal plates on the gnathopods, and usually also on pereopods 5, 6,

and 7. Stebbing’s family definition is therefore not satisfactory and must be

changed to read: “‘Head often partially fused with first segment of pereon.. .

first joint of gnathopods and pereopods reduced.”

The Caprogammaridae (Kudrjaschov and Vassilenko 1966), which was origi-

nally included in the Gammaridea, was assigned to the Caprellidae by McCain

(1968). This move is fully justified by the morphology of this genus. Capro-

gammarus and Cercops should probably be placed in a sub-family separate

from the ‘typical’ caprellids, on the basis of their well-developed abdomen.

However, our knowledge of how these two genera compare with other primitive

caprellid genera, e.g., the Proto group, is at present insufficient to support such

a separation.

The characters used to distinguish the genera within the family Caprellidae

are those originally formulated by Mayer (1890, 1903). Some of these characters

appear to be unsatisfactory, and the number of monotypic genera that have

resulted suggests that too much emphasis has been placed on what may be

unstable features of caprellid organization.

The characters that must be particularly suspect are the number of segments

in pereopods 3, 4, and 5, and the organization of the abdomen. Their diagnostic

worth cannot be fully assessed until more is known of their developmental

stability and function. If the abdomen is undergoing reduction, as seems likely

from its vestigial appearance in most caprellids, then its organization must not

be relied on too heavily for generic diagnosis. Thus the separation of Metaca-

prella from Caprella, solely on the basis of the female abdomen, is highly

unsatisfactory. Similarly, pereopods 3, 4, and 5 appear to be vestigial structures

in many genera; for example, in Tritella pereopods 3 and 4 are minute and

unsegmented, and in Capre/la they are absent.

Characters that deserve further investigation are the brood plates, the second

antennae, and particularly the mouthparts.

The nature and arrangement of the brood plate setae appears to be stable

within each genus. It has not been possible to determine whether it is also

characteristic for each genus.

The presence or absence of ‘swimming’ setae on the antenna 2 must be a

generic character; but the organization of these setae, particularly on the fla-

gellum, appears to be more specific (see Trite/la laevis and T. pilimana).

The most useful generic characters will probably be found among the mouth-

parts, but there has not been sufficient study to show which are the stable char-

acters. The presence or absence of the mandibular molar process must be at

least a generic character, reflecting feeding habits. The proportions of the lobes,

and probably other features, of the maxilliped, and the number of spines on

the outer lobe of maxilla I, also seem reliable. However, even casual observation

shows that variation does occur. Thus, while Perotripus normally has six spines

on the maxilla I outer lobe, the specimen figured here had only four spines on

one of the maxillae; Mayer (1903) recorded similar variations for other genera.

Similarly, the mandibular setal row of Capre/la appears to be stable, with two

setae on the right mandible and three setae on the left; yet in C. alaskana, all

the investigated specimens had a greater number of setae on at least one man-

dible.

McCain (1968) has urged a thorough examination of caprellid mouthparts.

Such an investigation must include as many genera and as many individuals as

possible, thereby to determine whether such variations as have been cited are

truly random.

Pending such an investigation, the present system of diagnosis, derived from

Mayer, will continue to be used.

The terminology used in this paper is as follows:

The cephelon is the partly fused head and pereonite I complex.

The swimming setae are the long hairs present along the lower edge of the anten-

na 2 peduncle, and, frequently, flagellum, in some genera. Although they are

generally called swimming setae, they play a more important part in feeding

than in swimming (Wetzel 1932).

The pereopods are numbered according to the pereonite to which they are

attached. The first and second pereopods are the gnathopods. Despite the

presence of coxal plates, the pereopods are treated as six-segmented limbs,

with the propodus and dactylus being segments 5 and 6, respectively. The palm

of the propodus of the pereopods, and particularly of gnathopod 2, has its own

special nomenclature. The large, usually paired, spines between which the tip of

the dactylus rests when closed, are the grasping spines. In Caprella there is

usually one pair of these spines on pereopods and gnathopod 1. The configura-

tion of the palm of the gnathopod 2 is important in the diagnosis of some

species. The grasping spine, which is frequently single, may have one or more

accessory spines at its base. The poison spine is so-called for historic reasons.

It is the spine-like projection which is not demarcated at its base from the palm

of gnathopod 2. As it is a secondary sexual character in many species (e.g.,

Tritella laevis, Deutella californica, Caprella laeviuscula), being very large in

the male and minute in the female, it is doubtful that it always has a poisonous

nature. However, Wetzel (1932) has shown that in some species it does appear

to secrete a venom.

For the abdomen, McCain’s (1968) system of discriminating between the

obvious, projecting appendages, and the less developed and usually setose lobes

has been used.

iS)

KEY TO THE CAPRELLIDAE OF THE AMERICAN NORTH

PACIFIC

. Gills on pereonites II, III, and IV; mandibular molar

POLS Sed SING il creases PEER, Sse clon cdhtss piondeve Ora tael Ne a ee

Gills on pereonites HI and IV; mandibular molar pro-

SCSSUDI CRC M5 te rere need 8 tert suet at ne eet Gia

Pereopods 3 and 4 one-segmented, pereopod 5 six-seg-

mented; abdomen five-segmented.............occeececeeeeteeeeeee

Pereopods 3 and 5 three-segmented, pereopod 4 one-

SESIMEMLER | DOGOMICT MUIMUGS. oo. scas.csercsvessecmenencadeesecnensce

. Rudimentary pereopods present on pereonites III and

W= mandibular palp presemt, a... he .wecece 0.3 shoes cadens -vessane

No pereopods present on pereonites III and IV; mandi-

bular palp absent (Caprella and Metacaprella)................

Pereopods 3 and 4 two-segmented; antenna 2 with no

SOTTO cae gaa a REN RR I a Rees a ir

Pereopods 3 and 4 one-segmented; antenna 2 with swim-

PIU ISCHAE. CRT ERCIIG ) en oe vile od snae craniccis ceiuniguoatetotoss a tiex

Pereopod 5 six-segmented; head with spine or tubercle;

male gnathopod 2 attached anteriorly on pereonite II...

Pereopod 5 three-segmented ; head smooth; male gnatho-

pod 2 attached posteriorly on pereonite II...............000....

Body spines anteriorly pointing; antenna 2 flagellum

Si@Uits WEA SHOrE SCLAE S48 ic. ah ccc Bath he Incest Tos hae Re ewdaanee

Body spines laterally pointing; antenna 2 flagellum

SleM@er: With Ome Selae Uh Nee thee ans soRcionceess

Single ventral spine between the insertions of gnatho-

No ventral spine between the insertions of gnathopods 2

Long, slender, anteriorly pointing head spine; male

gnathopod 2 propodus four times as long as broad........

No head spine; male gnathopod 2 propodus twice as

homes EOE) A): cate ies: ar Ww Celene ene aoa ce tah.

Pereonite V with lateral projections anteriorly; large

antero-laterally directed lateral spines at base of gnatho-

Pereonite V without lateral projections anteriorly;

lateral spines at base of gnathopods 2 small or absent..

Cercops compactus

(p. 9)

Perotripus brevis

(p. 12)

Deutella californica

(p. 16)

Mayerella banksia

(p. 19)

Tritella laevis (p. 22)

Tritella pilimana

(p. 25)

Caprella californica

(p. 49)

Caprella equilibra

(p. 55)

10.

12.

13.

14.

15;

16.

17.

Gnathopod 2 dactylus setose; no lateral spine at base of

LOS i Sa OSE A 6 Se rer

Gnathopod 2 dactylus not setose; small lateral spine at

LE 0) oT 0) 0 A Rar A Re A 0 or

Head without: Spine OF tubercle nic ise... ersss eee cceecsv ee cccceveas

Fiead With: one Spite OF tubercles. 5... kc sccee

Head with paired spines or tubercles..............0......00000...

Propodus of pereopods slender, with median grasping

spines and scarcely delineated palm; male gnathopod 2

with basis much longer than propodus, and dactylus

NS eee sco vac Te Re Tee AMS OF ATE. Fon:

Propodus of pereopods stout, with proximal grasping

spines and concave or well delineated palm; male

gnathopod 2 with basis shorter than propodus, dactylus

RON ee NSS hin: sbiye Saiog a os cote naa terep x ond aa ee eee

Propodus of gnathopod 2 with two accessory spines at

base of grasping spine. In male, gnathopod 2 propodus

with antero-dorsal projections; pereonite I three times

DS USCE Le 10 0(o7 10 anaes Re te tent RR Bs STS ae Cee

Propodus of gnathopod 2 with less than two accessory

spines at base of grasping spine. In male, gnathopod 2

propodus without antero-dorsa! projections; pereonite

isnot more than twice.as long as head............0..:.00..0000.00

Dorsal spines or tubercles present, at least on posterior

pereonites; antenna | more than half body length in

female, more than two-thirds in male.......0.............cccceee.

No dorsal spines or tubercles present; antenna 1 less

earn body length ns. DA en eae eee.

Sub-littoral; flagellum of antenna 1 longer than pe-

duncle; in male, antenna 2 equal in length to peduncle

ONAN Wis ccna veg Meta AH ee ae hd

Intertidal; flagellum of antenna 1 shorter than peduncle;

in male, antenna 2 shorter than peduncle segments 1

2 ZSC UIST 1016 a ee ee en oe

Gills round. In male, gnathopod 2 setose, poison spine

normal. In female, gnathopod 2 attached near middle

“1 SHIGSOrDTI/ jal i IC ae ale alt Mean oi oe Sa Mt AL alee aE bebe Pret

Gills long, oval. In male, gnathopod 2 not setose, poison

spine enormous. In female, gnathopod 2 attached at

Mammo CNG Of PETCOnite VE) Scie. fc. b) ce oye ndicin. abhi sb beads

Headspine an anteriorly pointing triangular projection..

Headspine an upward pointing rounded knob................

Caprella pilidigita

(p. 60)

Caprella mendax

(p. 58)

Caprella gracilior

(p. 62)

Caprella irregularis

(p. 65)

Caprella striata

(p. 76)

Caprella alaskana

(p. 35)

Caprella drepanochir

(."33)

Caprella laeviuscula

(p. 67)

18.

19.

20.

21.

22,

Visy

Dorsal body tuberculations minute or absent, never on

pereonite I; in male, gnathopod 2 attached anteriorly on

percomite WIA ies. 6k Ack teees See oy

Dorsal body tuberculations obvious, present on all per-

eonites; in male, gnathopod 2 attached medianly on

PCTCOMIPC Ie oF shi asdsicansiigeea tata ea eee

Body tuberculations large; in male, propodus of gnatho-

poe 2 shorter than, pereonite.[L..... 2 ee eee

Body tuberculations small; in male, propodus of gnatho-

pod 2.asvione as perconite Ws ot eek ee

Head and body covered with large and small tubercula-

tions; antenna 2 with many long swimming setae. In

male, gnathopod 2 and much of body setose..................

Body with few dorsal tuberculations; antenna 2 with

few short swimming setae. In male, gnathopod 2 and

DOM VINO Sebo see ied a6, sy dicenes Re re seen kee hee

Paired tubercles on head; female abdomen with one

PATNI OWS reise ce ec icin ee oh ones akc

Paired spines on head; female abdomen with one pair

of one-segmented limbs (Metacaprella)..............0.00cc000000044

Large dorsal spines on posterior pereonites. In male,

gnathopod 2 poison spine normal; pereonite I shorter

bit wOn. SMe TO CA Nee te dese esac cath oocesaaentirs

Small dorsal tuberculations on posterior pereonites. In

male, gnathopod 2 poison spine enormous; pereonite I

NGAGE MAM IIS AG ec sous scnis cap scsi aw ehovubedaeede bata atts. lees

Flagellum of antenna 1 longer than peduncle. In male,

pereonites I and II not greatly elongated; antenna 1

peduncle slender, NOt. setose22 .. jee... ee

Flagellum of antenna 1 shorter than peduncle. In male,

pereonites I and II much elongated; antenna | peduncle

Stout, \ensely SeIOse.. 7s i trek Sean eh sek ee Pee es

Caprella angusta

(p. 40)

Caprella verrucosa

(p. 44)

Caprella incisa (p. 42)

Caprella pustulata

(p. 72)

Caprella borealis

(p. 47)

Pips

Caprella ferrea (p. 35)

Caprella rudiuscula

(p. 70)

Metacaprella anomala

(p. 29)

Metacaprella kennerlyi

(p. 31)

Genus CERCOPS Krgyer 1842-43

Antenna 2 without swimming setae, flagellum bi- or tri-articulate; mandibular

palp triarticulate, with three terminal bristles; molar absent; maxilliped lobes

very small, outer larger than inner; gills on pereonites II, III, and IV; pereopods

3 and 4 one-segmented, pereopod 5 six-segmented; abdomen five-segmented,

male with two pairs of well-developed biarticulate limbs on segments 4 and 5,

plus two pairs of very rudimentary limbs on segments | and 2; female with two

pairs of biarticulate limbs on abdominal segments 4 and 5.

Cercops compactus n. sp. (Figure 1)

Material examined:

Female holotype, NMC 10776, Puffin Bay, Baranof Island, latitude 56°16’N,

longitude 134°40’W;; intertidal. Station A171-172, 1961. Male allotype, NMC

11201, Middle Bay of Cape Arago, Oregon; intertidal, on Plocamium pacificum.

July 8, 1960. 2 female paratypes, Middle Bay of Cape Arago, Oregon, July 8,

1960. 1 male paratype, Cape Blanco, Oregon, July 10, 1960.

Description:

Female: Head with small median dorsal projection behind eye, concave over

eye, and with small median spine above antenna 1 attachment. Body without

spines. Length 3.8mm. Cephalon and pereonite II subequal, pereonites III

and IV slightly longer, pereonite V equal to cephalon, pereonites VI and VII

decreasing in size.

Eye with very few facets.

Antenna 1 equal in length to cephalon plus pereonite II, flagellum with five

articles. Antenna 2 approximately equal to peduncle of antenna 1; flagellum

triarticulate.

Mandible with triarticulate palp, having two bristles on median article, and

two long bristles plus one shorter one on distal article; right incisor five-

toothed, lacinia mobilis five-toothed, setal row of four long, slender plus five

short, stout plumose setae. Maxilla I outer lobe with six serrate spines; palp

with few heavy setae. Maxilla II lobes small and slender. Maxilliped inner

lobe minute, convex distally, with two apical setae; outer lobe small, with

several marginal and surface setae, median margin not serrate; dactylus of palp

heavy. Upper and lower lip not known.

Propodus of gnathopod 1 almost square, with palmar surface beginning approx-

imately halfway along ventral surface; palmar surface serrate, armed with one

pair of proximal grasping spines, and with spines and setae; dactylus heavy.

Propodus of gnathopod 2 almost circular, shorter than basis, with large proxi-

mal protrusion carrying a large spine, and small distal protrusion; palmar sur-

face with spines and setae. Dactylus thickened proximally and tapering distally.

Basis has a small lateral projection distally and is attached anteriorly on pere-

onite II; ischium square, slightly produced anteriorly, and with a small lateral

projection; merus square, and smaller than ischium; carpus very small, rec-

tangular.

97949—2

Gills small, oval, the posterior one being the largest.

The brood plates have plumose setae around their margins, the anterior plate

being the more setose.

Pereopods 3 and 4 are alike, being very small, one-segmented, with two

setae distally, and are attached at the base of the gills. Pereopods 5, 6, and

7 are six-segmented, increasing in size posteriorly; propodus with up to three

pairs of grasping spines on a proximal projection, palmar surface concave, with

spines and setae; dactylus heavy, tapered distally.

The abdomen is approximately one-tenth of the length of the body, five-seg-

mented, with two pairs of biarticulate limbs with a single longitudinal row of

comb setae on each article. The proximal article has a distal spine-like projection

which also carries comb setae.

Male: Very like female in general body proportions. Length of largest male

3.8 mm.

Mouthparts as in female; left mandible with five-toothed incisor, five-toothed

lacinia mobilis, one accessory plate, and setal row of seven or eight setae.

Gnathopod 2 basis very heavy; palmar surface of propodus setose, and having

distal projections more obvious than in female. Abdomen with very rudimen-

tary pleopods on segments 1 and 2; these consist of a minute unsegmented

appendage terminating in a long seta. The biarticulate uropods on segments 4

and 5 are similar to those of the female, except for the anteriorly projecting

basal knob on the uropod 1 of the male; this knob is possibly a copulatory

organ. Penes lateral.

Variation

Both female paratypes had only two articles to the flagellum of antenna 2.

Remarks

The manuscript description of the single female type specimen had been

completed when four more specimens of this species were received from Dr.

Marvin P. Jessen. The male allotype was selected from this material and has

been deposited in National Museums of Canada collections. The holotype, a

mature female with three larvae in the brood pouch, was dissected and mounted

as well as its poor condition would allow. The specific name refers to the very

solid appearance of this specimen. Unfortunately, the other specimens had

been whole mounted, and it was considered inadvisable to try to remove them

from the slides. The species description has therefore been based on all five

available specimens.

Distribution

Type locality: Puffin Bay, Baranof Island, Alaska.

Other localities: Cape Arago and Cape Blanco, Oregon.

Discussion

The only other species in this genus is Cercops holbolli Krgyer, which has

been reported from S. Greenland and from Japan (Tsugaru Strait). The new

species differs from this in the absence of body spines, in having a triarticulate

flagellum in antenna 2, in the setation of the inner and outer lobes of the maxil-

liped, and in the much less slender conformation, with different form of palmar

surface, of the appendages. Because of the segmentation of antenna 2 flagellum,

it has been necessary to change the genus diagnosis.

FIGURE 1— Cercops compactus. Male allotype, lateral view; scale equals 0.5 mm. Female

holotype, mouthparts and appendages. Male allotype and paratype, gnathopod

2 propodus, abdomen, uropods and left mandibular palp and incisor region.

97949—23

Genus PEROTRIPUS Dougherty and Steinberg 1953

Antenna 2 without swimming setae, flagellum biarticulate; mandibular palp

triarticulate with one terminal seta, molar absent; outer lobe of maxilliped

equal to or larger than inner lobe, both minute; gills on pereonites II, III, and

IV ; pereopod 3 three-segmented, 4 one-segmented, 5 three-segmented; abdomen

of male with one pair one-segmented appendages plus one pair setose lobes,

of female with one pair of setose lobes.

Perotripus brevis (La Follette 1915) (Figures 2, 26, Map 1)

Synonymy

Paedaridium breve La Follette 1915

Perotripus brevis—Dougherty and Steinberg 1953; 1954

Material Examined

Prince William Sound: two males, stations A87, A92, 1961; NMC 10820,

10821.

Dall Island: two males, station A3, 1961; NMC 10819.

Queen Charlotte Islands: six males, ten females, station W11, 1967; NMC

10839.

Hecate Strait: over thirty males, over fifty females, stations HS, H25, H26,

H29, H30, H33, 1964; NMC 10822-10827.

Queen Charlotte Sound: five males, six females, stations H50, H53, H58,

1964; NMC 10829-10831.

Vancouver Island: one male, one female, station H44, 1964; NMC 10828.

Description

Male: Body smooth, but pereonites IJ and III sculptured with lateral pro-

jections, as shown in Figure 2. Length 4.9 mm. Cephalon and pereonites II and

III equal in length; IV is longer; V plus VI approximately equal to half the body

length; VII and the abdomen are minute. The anterior pereonites are stout, the

posterior one slender.

Antenna 1 shorter than cephalon plus pereonite II, stout, basal article distally

produced to form a frill around base of second article; articles 2, 3, and flagel-

lum sparsely setose; flagellum with two articles, the basal one being about as

long as the third peduncular article. Antenna 2 slightly shorter than antenna 1

peduncle; articles 3 and 4 sparsely setose; flagellum minute with spindle-shaped

first article.

Mandible with triarticulate palp, middle and terminal articles each with a

single seta; left mandible with five-toothed incisor, five-toothed lacinia mo-

bilis, one accessory plate, one seta; right mandible with five-toothed incisor,

lacinia mobilis not denticulate, one accessory plate, one seta. Maxilla I with six

serrate spines on outer lobe; palp with few stout setae. Maxilla II with very small

slender lobes with few setae. Inner lobe of maxilliped convex distally, with two

median setae and outer tooth-like projection; outer lobe with slightly serrate

apical margin, and marginal setae; dactylus of palp heavy and barely tapered,

with serrate proximal and setose distal margin. Lower lip with very weakly

developed inner lobes.

Propodus of gnathopod 1 triangular, ventral surface with proximal projection

with serrate edge, bearing four grasping spines, palmar suface convex bearing

spines and a few setae. Propodus of gnathopod 2 twice as long as it is broad,

with proximal ventral projection bearing two pairs of grasping spines and a

small proximal spine; palmar surface slightly convex and with spines; digit

thickened proximally, tapered distally. Basis just shorter than propodus, at-

tached anteriorly on pereonite II; ischium square, merus round, carpus trian-

gular.

Gills very small, oval.

Pereopod 3 attached at base of gill; first segment with short distal seta, second

segment very small with long seta, terminal segment long with subterminal

seta and terminal plumose seta. Pereopod 4 attached at base of gill, with two

terminal setae, one plain and one plumose. Pereopod 5 with first and second

segments of same length, first segment with one large seta, second segment

shaped rather like a propodus; third segment a short claw, bearing a large plu-

mose seta. Pereopods 6 and 7 six-segmented, with segments 4 and 5 elongated

and bearing spines on their anterior surface; propodus with no well-defined pal-

mar surface; dactylus slender.

Abdomen with lateral penes; one pair of unsegmented appendages terminating

in a long bristle, and one pair of lobes with two to four bristles.

Female: Length 4.8 mm. In general the female is similar to the male. Pereoni-

tes III and IV are very much more swollen laterally and dorsoventrally; the

anterior brood plate has plain setae all round its margin; the posterior brood

plate has plain setae on the posterior margin only. The abdomen has one pair of

lobes with one or two bristles.

Remarks

This species when present in a preserved collection can easily be detected

because of the characteristic bent shape of its body (see Figure 2). La Follette

records the color as “‘light pink to white.”

Distribution

Type locality: Laguna Beach, California.

Other localities: San Juan area, Washington.

New records: Pacific coasts of Alexander Archipelago, Alaska, and Vancouver

Island, B.C.; southern Prince William Sound, Alaska; British Columbia main-

land coast from Banks Island to Rivers Inlet.

Discussion

Although this species differs from La Follette’s type description in having

three pairs of gills, there is little doubt that it is the same animal as his Paedari-

dium breve. In all other characters mentioned by him there is no difference,

FIGURE 2 — Perotripus brevis. Male, lateral view and appendages. Dorsal view of anterior

pereonites of male, and of pereonites III and IV of female, drawn to scale.

except those which would be expected between his, presumably, juvenile

specimens and the more mature specimen described above. As the gills are

very small, the first pair, which is normally between the bases of the second

gnathopods, could easily be overlooked.

In his description, La Follette did not give his reasons for assigning this

species to genus Paedaridium. According to Mayer’s diagnostic characters, this

species does not belong in genus Paedaridium, and on the basis of the segmenta-

tion of pereopods 3 to 5 it was rightly given new generic status by Dougherty

and Steinberg (1953). However, on the basis of such characters as were used

by Mayer and of our limited knowledge of the mouthparts of the various genera,

Perotripus appears to be at least as close to Paedaridium as it is to the six genera

listed by Dougherty and Steinberg.

ALASKA

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Map fig. 1 — Known distribution of (a) Perotripus brevis, (b) Deutella californica, and (c)

Mayerella banksia within the American Pacific boreal region.

Genus DEUTELLA Mayer 1890

Antenna 2 without swimming setae, flagellum biarticulate; mandibular palp

triarticulate, setal formula for terminal article 1 + x + 1; molar present; outer

lobe of maxilliped larger than inner lobe; gills on pereonites III and IV; pereo-

pods 3 and 4 two-segmented, pereopod 5 six-segmented, inserted at posterior

end of pereonite V; abdomen of male with one pair of non-segmented appen-

dages and one pair of setose lobes, of female with one pair setose lobes.

Deutella californica Mayer 1890 (Figures 3, 26, Map 1)

Synonymy

Deutella californica Mayer 1890; 1903 — Dougherty and Steinberg 1953;

1954 — Steinberg and Dougherty 1957 — Gardella 1962 — McCain 1968

Material Examined

Vancouver Island: two males, two females, stations 07b, 1959, NAP66-201;

NMC 10777, 10781; USNM 172360.

Washington Coast: one male, station W22, 1966; NMC 10779.

Oregon Coast: over thirty individuals, station W63, 1966; NMC 10780.

Prince William Sound: one male, station A153, 1961; NMC 10778.

Description

Male: Head bears a small anteriorly pointing median spine; pereonite II

with antero-lateral anteriorly pointing projections; other body tuberculations

minute or absent. Length 5.5 mm.

Antenna | equals approximately half the body length, flagellum with up to

twelve articles. Antenna 2 shorter than antenna 1 peduncular articles 1 plus 2.

Mandible with triarticulate palp, with one seta on penultimate article, and

setal formula 1 + x + 1 for the terminal article; left and right mandible with

incisor five-toothed, lacinia mobilis five-toothed, setal row of three plumose

setae. Maxilla I outer lobe with six serrate spines, palp with few setae. Maxil-

liped inner lobe flat distally, with few apical setae; outer lobe with one apical

and few marginal and surface setae, median margin not serrate; palp with

distal projection on subterminal article, heavy dactylus with apical seta.

Propodus of gnathopod 2 nearly rectangular; palmar surface with proxima]

grasping spine, and median poison spine separated from the distal portion by

a deep cleft; proximal margin finely denticulate, distal margin coarsely denticu-

late; palmar and distal portion of propodus bear long hairs. Basis attached an-

teriorly on pereonite II.

Pereopods 3 and 4 small, similar, with a ring of setae at the distal end of the

first segment, second segment minute with three terminal bristles. Pereopods

5, 6, and 7 six-segmented; propodus with concave palmar surface, having a

proximal series of knobs bearing grasping spines.

Abdomen with lateral penes; one pair of one-segmented appendages armed

distally with small teeth, and one pair of setose lobes.

Female: Length 3.8 mm. Differs from male in having a small median knob

on head; second gnathopod propodus palmar surface without cleft or large

poison spine and with short hairs; anterior brood plate with plumose setae

around margin, posterior plate with plumose setae on posterior margin only.

Abdomen with one pair setose lobes.

g FD

Figure 3 — Deutella californica. Male, lateral view and appendages. Female brood plates

and gnathopod 2.

Distribution

Type locality: Cape Mendocino, California.

Other localities: Monterey Bay and Mussel Point, California; “‘probably

Port Aransas, Texas” (Steinberg and Dougherty 1957); San Juan area, Washing-

ton.

New records: Pacific coasts of Oregon, Washington, and Vancouver Island;

Juan de Fuca Strait, B.C.; southern Prince William Sound, Alaska.

Discussion

As has been pointed out by McCain (1968), the Texas locality record for this

species is very doubtful; it is probable that Deutella californica is endemic to

the Pacific.

Genus MA YERELLA Huntsman 1915

Antenna 2 without swimming setae, flagellum biarticulate; mandibular palp

triarticulate with one seta on terminal article; molar present; outer lobe of

maxilliped larger than inner lobe; gills on pereonites [II and IV; pereopods 3

and 4 two-segmented, pereopod 5 three-segmented; abdomen of male with

one pair of appendages and one pair of lobes, of female with one pair of lobes.

Mayerella banksia n. sp. (Figures 4, 26, Map 1)

Material Examined

Male holotype, NMC 10832, Rennison Island, B.C., 4-12 fm, station H30,

1964.

Female allotype, NMC 10833, Cox Point Inlet, Trutch Island, B.C., station

H26, 1964.

Forty male paratypes, twelve female paratypes, NMC 10834-10836, Hecate

Strait, B.C., stations H25, H26, H30, 1964.

One male, one female, NMC 10837, Queen Charlotte Sound, B.C., station

H37, 1964.

One male, NMC 10838, Chicagof Island, Alaska, station A165, 1961.

Three females, Department of Fish and Game, California, latitude 122°

24’ N, longitude 37° 5’ W.

Description

Male: Body smooth except for small antero-lateral projections on pereonites

II and III. Length 5.9 mm. Cephalon and pereonite II subequal, longer than

III which equals IV; pereonite V is the longest, VI shorter, VII minute; the

abdomen protrudes ventrally.

Antenna | shorter than cephalon plus pereonite II, flagellum with five articles,

peduncular articles 2 and 3 with sparse setae; antenna 2 equal in length to

peduncle of antenna 1, flagellum very small.

Mandible with triarticulate palp with one seta on median article, and one

apical seta on terminal article; incisor five-toothed, lacinia mobilis five-toothed

on left and indistinctly toothed on right, setal row three on left and two on

right. Maxilla I outer lobe with seven serrate spines, palp with three spines plus

setae. Maxilla II lobes with few setae. Inner lobe of maxilliped flat distally, with

few plumose setae and one tooth; outer lobe with one plumose and one plain

apical seta, median margin strongly denticulate, few surface setae; palp with

finely tapered dactylus.

Propodus of gnathopod 1 slender and triangular, palm with one pair proximal

grasping spines, margin serrate and armed with spines and plumose setae;

‘comb setae’ present midway between dorsal and ventral margins. Dactylus

stout, inner margin scalloped and minutely serrate. Propodus of gnathopod 2

more than three times as long as broad; palm with proximal protrusion bearing

a grasping spine, distal poison spine separated by cleft from rectangular pro-

jection in the angle of the dactylus; margin of palm finely serrate and with a

few setae. Dactylus heavy, distal third strongly curved inwards. Basis attached

posteriorly on pereonite II; longer than propodus, and with slight antero-

lateral projection distally. Ischium and merus approximately equal, carpus very

small and triangular.

FIGURE 4— Mayerella banksia. Male holotype, lateral view. Male paratype, appendages.

Female paratype, brood plates and gnathopod 2.

Gills small and oval, the posterior ones being smaller.

Pereopods 3 and 4 small, attached to base of gills; first segment with two setae,

terminal segment with one plumose and one plain seta. Pereopod 5 with one

seta at base of proximal segment, distal seta on second segment, terminal

segment with three plain and one plumose setae. Pereopods 6 and 7 six-seg-

mented; palmar region of propodus only slightly developed; most of anterior

margin of pereopod armed with spines.

Abdomen with one pair of uniarticulate limbs, separated by median chitinous

hump bearing very small teeth posteriorly; one pair of setose lobes, not dis-

tinctly separated.

Female: Length 3.9 mm. Differs from male in that anterior segments do not

elongate with maturity, and pereonites III and IV are swollen. Gnathopod 2

arises anteriorly on pereonite II, and its propodus lacks both poison spine and

palmar cleft and is armed with a proximal grasping spine and many palmar

spines; margin of palm serrate. Anterior brood plate has plumose setae around

the margin, posterior plate has setae on posterior margin only. Abdomen with

one pair of lobes with setae.

Distribution

Type locality: Rennison Island, B.C.

Other localities: Rennison Island, Rivers Inlet, and Trutch Island, B.C.;

Chichagof Island, Alaska; south of Golden Gate, California.

Discussion

This is the first species of this genus to be reported from the Pacific and is

named after Banks Island, one of the larger islands in the type region. M.

limicola Huntsman and M. redunca McCain both occur in the northwest

Atlantic and differ from M. banksia in the structure of both gnathopods, and

of pereopods 6 and 7. The most striking differences are in the abdomen: in M.

limicola the penes are medial, and in M. redunca the appendages are long,

slender, and recurved at the tip; neither species appears to have the median

chitinous hump found in M. banksia. Mayerella is probably Mayer’s genus

Incertum (1903) from California. The only real difference between these genera

is the absence of the mandibular palp in Mayer’s single specimen. However,

the mandibular palp is frequently not observable in Mayerella unless the

mandible is dissected; the palp is either held very close to the head or is easily

lost.

Genus TRITELLA Mayer 1890

Antenna 2 with swimming setae, flagellum -biarticulate; mandibular palp

triarticulate, setal formula for terminal article 1 + x + 1; molar present; outer

lobe of maxilliped larger than inner lobe; gills on pereonites III and IV; pereo-

pods 3 and 4 one-segmented, pereopod 5 six-segmented; abdomen of male and

female with 1 pair of setose lobes.

Tritella laevis Mayer 1903 (Figures 5, 26, Map 2)

Synonymy

Tritella laevis Mayer 1903 — Dougherty and Steinberg 1953; 1954 — McCain

1968.

Material Examined

Queen Charlotte Islands: fifteen males, six females, stations H2a, W8, W11,

W12, 1957; NMC 10784-10787.

Queen Charlotte Sound: eight males, eleven females, stations H1, H50, H53,

1964; NMC 10793, 10795, 10796.

Vancouver Island: eight males, nine females, stations Fl, P7, 1955; N6, Ol,

O7b, Oll, V4b, 1959; H44, 1964; NMC 10782, 10783, 10788-10792, 10794;

two males, two females, NMC 10801; one female, USNM No. 172360.

Juan de Fuca Strait: two males, stations W34, W42, 1966; NMC 10797,

10798.

Oregon Coast: five males, seven females, stations W57, W60, 1966; NMC

10799, 10800.

Description

Male: Body smooth dorsally, anteriorly directed anterolateral spines on

pereonites II, III, and IV; no spines over gills. Length 9.4 mm. Cephalon

shorter than pereonite II, pereonite III is the longest, [V and V approximately

equal, VI plus VII shorter than V.

Antenna 1 longer than cephalon plus pereonite II, flagellum with eleven

articles. Antenna 2 just longer than peduncle of antenna 1, flagellum two-thirds

previous article, stout and with heavy setae.

Left mandible with five-toothed incisor, lacinia mobilis five-toothed, three

setae; right mandible with toothed lacinia mobilis and two setae. Maxilla I

outer lobe with seven serrate spines. Maxilliped inner lobe distally flat with

plumose setae and one tooth; outer lobe with one or two apical plumose

setae, one tooth and denticulate median margin; palp with distal projection

on subterminal article, dactyl heavy, with apical seta.

Propodus of gnathopod 2 curved, twice as long as broad, palm concave and

densely setose, with proximal grasping spine and poison spine. Dactylus with

proximal and distal thickenings, tapering abruptly distally, densely setose inner

edge. Basis attached anteriorly on pereonite II.

Papi

FIGURE 5 — Tritella laevis. Male, lateral view and appendages. Female brood plates and

gnathopod 2. Dorsal view of male pereonite II, showing orientation of lateral

spines, and lateral view of anterior pereonites of female, not to scale.

60°N

150° w =f va

Prince William a , Prince William @& ee a Prince William © ——

Sound ie Sound a Sound

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PACIFIC OCEAN

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135°W °

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<, COLUMBIA

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OLUMBIA

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Map fig. 2 — Known distribution of (a) Tritella laevis, (b) T. pilimana, and (c) Metacaprella

anomala within the American Pacific boreal region.

Pereopods 3 and 4 similar, having three apical setae, one of which is plumose.

Pereopods 5, 6, and 7 normal, propodus thickened proximally with palmar

knobs supporting the grasping spines, the knobs being most developed on

pereopod 7.

Abdomen with lateral penes, and one pair of lobes with three setal clumps on

each.

Female: Length 6.2 mm. Differs from male in showing no lengthening of

pereonite I with maturity; gnathopod 2 has heavier propodus, median and

minute poison spine, few hairs on palm or dactylus. The anterior brood plate

has plumose setae around the margin, the posterior plate has setae on the

posterior margin only. Abdomen has one pair of setose lobes.

Remarks

The fifty-seven specimens of this species in the NMC collections were ex-

amined in greater detail individually than were many other species, and among

them were found two ‘female’ intersexes. One (station H53 /64) has the male

gnathopod 2 and abdomen with penes, and the female brood plates and some

evidence of a female genital opening on pereonite V. The other (station

W60 /66) has the male gnathopod 2 and abdomen with penes, and developing

brood plates but no sign of the female opening on pereonite V. There is no

evidence of intersexes among the specimens of 7. pilimana examined. Apart

from Mayer’s (1903) record of two “Hermaphrodite”’ specimens of Caprella

bispinosa, no reference to caprellid intersexes has been found in the literature.

Mayer’s description of T. /aevis distinguishes this species from T. pilimana on

the basis that 7. pilimana has body spines and T. /aevis does not. It is probable

that the specimens described by Mayer were subadult or immature males, in

which the body spines are either absent or very small; in the adult they are

noticeable, particularly on pereonite II (see Figure 5).

Freshly preserved specimens showed two color variations, one having a

reddish-brown body with non-pigmented areas, the pereopods being paler and

striped with dark pigment bands; the other being translucent with pink color-

ation developed dorsally on the body and antennae.

Distribution

Type locality: Santa Catalina, California.

Other localities: Pacific Grove, Dillon Beach, Moss Beach, and Monterey

Bay, California.

New records: Pacific coasts of Oregon, Vancouver Island, and Queen Charlotte

Islands; Washington and B.C. coasts of Juan de Fuca Strait; mainland coast

of Queen Charlotte Sound, and Queen Charlotte Strait, B.C.

Tritella pilimana Mayer 1890 (Figure 6, Map 2)

Synonymy

Tritella pilimana Mayer 1890; 1903— Dougherty and Steinberg 1953;

1954 — McCain 1968

Aeginella hirsuta La Follette 1915

Material Examined

Prince William Sound: one male, one female, stations A80, A133, 1961;

NMC 10808, 10809.

Queen Charlotte Islands: seven males, one female, stations H14, W9, W11,

1957; NMC 10804-10806.

Hecate Strait: four males, one female, stations H8, H31, 1964; NMC 10810,

10811.

Queen Charlotte Sound: ten males, three females, station H37, 1964; NMC

10812.

Vancouver Island: sixteen males, eleven females, stations G2, P6c, 1955; V3,

1959; NMC 10802, 10803, 10807; USNM 148064.

Juan de Fuca Strait: two males, station W44, 1966; NMC 10815.

97949—3

Washington Coast: seven males, twelve females, stations W22, W40, 1966;

NMC 10813, 10814.

Oregon Coast: eighteen males, eighteen females, stations W57, W63, W66,

1966; NMC 10816-10818.

Description

Male: Body smooth dorsally, but with laterally pointing spines antero-

laterally on pereonites II, HI, and IV, and medially over gills. Length 11 mm.

Cephalon equals pereonite II, pereonite III equal to IV, but longer than II, V

shorter than IV and longer than VI plus VII.

Antenna 1 nearly as long as cephalon plus pereonites IJ and III, flagellum

with twelve articles. Antenna 2 shorter than peduncle of antenna 1, flagellum

six-sevenths of preceding article, slender with long swimming setae.

Left mandible with five-toothed incisor, five-toothed lacinia mobilis, and three

setae; right mandible with denticulate but not five-toothed lacinia mobilis, and

two setae. Maxilla I outer lobe with seven serrate spines. Maxilliped inner lobe

flat distally, with plumose setae and one tooth; outer lobe concave apically,

with one apical plumose seta, median margin serrate, and few marginal and

surface setae.

Propodus of gnathopod 2 twice as long as it is broad, palm slightly concave,

densely setose with proximal grasping spine and poison spine. Dactylus with

distal and slight proximal thickening, gently tapering distally, and densely

setose along inner margin. Basis attached anteriorly on pereonite II.

Pereopods 3 and 4 as in T. Jaevis; pereopods 5, 6, and 7 six-segmented, propodus

with proximal grasping spines; palmar knobs supporting grasping spines may

be present on pereopod 7.

Abdomen as in T. Jaevis.

Female: Length 7.9 mm. Differs from male in showing no lengthening of

pereonite I with maturity; the gnathopod 2 propodus is heavier, with median

minute poison spine and very few hairs on palm. Brood plates and abdomen as

in T. laevis.

Remarks

Freshly preserved specimens show two different color types; one is trans-

lucent with aggregations of dark pigment spots giving the appearance of

blotches and stripes over all the body except the pereopods; the other is trans-

lucent pink with isolated pigment blotches along the dorsal surface only.

It is probable that this species is the same animal as Aeginella hirsuta La

Follette 1915, to whose description it appears to bear a greater resemblance

than does T. Jaevis.

Distribution

Type locality: Mendocino, California.

Other localities: Humboldt Bay, Laguna Beach, Dillon Beach, and Moss

Beach, California; San Juan area, Washington.

New records: Pacific coasts of Oregon, Vancouver Island, and Queen Char-

lotte Islands; Washington coast of Juan de Fuca Strait; southern Prince

William Sound, Alaska; Dixon Entrance, and mainland coast south to John-

stone Passage, B.C.

FIGURE 6 — Tritella pilimana. Male, lateral view and appendages. Female gnathopod 2.

Dorsal view of male pereonite II, showing orientation of lateral spines, and

lateral view of anterior pereonites of female, not to scale.

97949—334

Discussion

The two Tritella species described here are very similar and are hard to

distinguish. The appendages, e.g., antennae and pereopods, of JT. pilimana

appear to be slightly slimmer than those of T. /aevis; the most noticeable points

of distinction are the flagellum of antenna 2 and the lateral body spines, which

point anteriorly in T. /aevis and laterally in T. pilimana; also, in fresh specimens,

the color differences are noticeable.

No specimens of Mayer’s Tritella sp. have been found. The specimen in the

USNM was examined and appears to differ from T. pilimana only in the spina-

tion of pereonite II and in the fact that the claw of gnathopod 2 is not setose.

One specimen of a Tritella sp. was found, which bears a strong resemblance to

T. pilimana except that it has a long, slender head spine; as the specimen is in

poor condition and is noticeably parasitised, it was decided not to call it a new

species until adequate specimens are collected.

Mayer (1903) indicates that in Tritella the females have no abdominal appen-

dages. There are, however, obvious setose lobes on the female abdomen (see

Figure 26 (2.)), and the genus diagnosis has been altered accordingly.

Genus METACAPRELLA Mayer 1903

Antenna 2 with swimming setae, flagellum biarticulate, mandibular palp

absent; molar present; outer lobe of maxilliped larger than inner lobe; gills on

pereonites III and IV; pereopods 3 and 4 absent; pereopod 5 six-segmented;

abdomen of male with one pair of appendages and one pair of lobes, of female

with one pair of appendages.

This genus was provisionally established by Mayer (1903) and subsequently

adopted by Dougherty and Steinberg (1953). Although no differences between

the species in this genus and genus Caprella have been observed, except for the

character of the female abdomen, it has been decided to allow this genus to stand

because it conforms to Mayer’s original standards of generic distinction.

At the time genus Metacaprella was adopted, it was decided to include C.

ferrea on the basis of its resemblance to M. anomala and M. kennerlyi; however,

there is no sign of rudimentary limbs on the abdomen of the female of this

species, and it has accordingly been restored to Caprella.

Metacaprella anomala (Mayer 1903) (Figures 7, 26, Map 2)

Synonymy

Caprella anomaia Mayer 1903 — Arimoto 1934

Metacaprella anomala — Dougherty and Steinberg 1953; 1954

Material Examined

Southern Alaska: six males, nine females, stations JWS 85, 1965, A173,

1961; NMC 11003, 11004; USNM acc. 85557.

Puget Sound: two males, two females, station W8, 1966; NMC 11005.

Pacific Grove: two individuals from the Zoologische Staatssammlung,

Munich.

Description

Male: Head with one pair anteriorly pointing spines, pereonites I to IV

smooth dorsally except for an unpaired spine posteriorly on IV; V has three

pairs of spines, VI and VII two pairs each; lateral spines may be present over

the bases of the second gnathopods, antero-laterally on pereonites III and IV,

and over the gills. No mature male was found; largest specimen was 9.9 mm

(Mayer’s largest male specimen was 13 mm).

Antenna 1 longer than cephalon plus pereonites II, III, and IV, flagellum

longer than peduncle and having at least twenty-one articles. Antenna 2 equals

peduncle of antenna 1; flagellum has long swimming setae.

Mouthparts typical of genus; lacinia mobilis of right mandible toothed, but

not five-toothed.

Gnathopod | propodus and dactylus with serrate grasping margin. Gnathopod

2 propodus less than half as broad as it is long, palm with proximal grasping

spine and accessory spine, distal poison spine and more distal triangular projec-

tion. Dactylus evenly tapered. Basis attached just posterior to the middle of

pereonite II, and with antero-lateral ridge.

Gills oval.

Pereopods 5, 6, and 7 with proximal grasping spines on propodus.

Abdomen typical of genus.

MX2

FicurE 7— Metacaprella anomala. Subadult male, lateral view and appendages. Female

gnathopod 2 and anterior brood plate.

Female: Length 13.2 mm. Differs from male in showing no elongation of

pereonite I; antenna 1 flagellum with twenty-three articles; some spines present

on anterior body segments. Abdomen typical of genus.

Distribution

Type locality: Pacific Grove, California.

Other localities: Japan; Moss Beach and Monterey Bay, California.

New records: Alexander Archipelago and off Sumdum Glacier, Alaska;

Puget Sound, Washington.

Discussion

Knowledge of the distribution of M. anomala is limited, but the species

appears to be highly eurytopic. It has been found intertidally and down to

50+ fm and at temperatures ranging from 14.8°C to 6.3°C. From knowledge

of its range along the North American coast and in Japan, it is suggested that

this may be an amphi-pacific species.

Metacaprella kennerlyi (Stimpson 1864) (Figure 8, Map 3)

Synonymy

Caprella kennerlyi Stimpson 1864 — Mayer 1882; 1903 — Holmes 1904 —

Johnson and Snook 1927— Wailes 1931 — Light 1941 — Ricketts and

Calvin 1952

Metacaprella kennerlyi — Dougherty and Steinberg 1953; 1954 — Gardella

1962 — Saunders 1966

Material Examined

Oregon, Washington, Puget Sound, San Juan Islands, Juan de Fuca Strait,

Strait of Georgia, Vancouver Island, Queen Charlotte Sound, Queen Charlotte

Islands, Southern Alaska, and Prince William Sound: thirty-two lots comprising

over two hundred individuals, NMC 11007-11023, 11026, 11028-11035;

USNM 50564, 173540, 187867, 102730, USNM Alaska King Crab Expedition

9-40, 12-40.

British Columbia (no other data): three lots comprising over fifty individuals,

NMC 11024, 11025, 11027.

Monterey, California, from the Zoologische Staatssammlung, Munich: one

lot comprising twenty individuals.

Description

Male: Head with one pair anteriorly pointing spines; body spiny, size and

disposition of spines varying, mature females and immature males usually

exhibiting longer spines than mature males; in some individuals the body spines

may be so much reduced as to be hardly noticeable. Typical arrangement

shown in Figure 8.

Length 22.5 mm; large specimens can be up to 30 mm in length.

Antenna 1 longer than cephalon plus pereonite II, flagellum equals peduncle

article 2 and has nineteen articles; peduncle articles 1, 2, and 3 stout and

densely setose. Antenna 2 shorter than two basal articles of antenna 1, flagellum

with swimming setae.

Mouthparts typical of genus, lacinia mobilis of right mandible denticulate,

not five-toothed.

Gnathopod 1 propodus and dactylus grasping margin slightly serrate. Gnatho-

pod 2 propodus large, more than twice as long as it is broad, palm with proximal

grasping spine, small distal poison spine, and large triangular distal projection.

Dactylus thickened proximally, tapering distally. Basis attached posteriorly on

pereonite II, and bearing antero-lateral ridge.

Gills large, round, or oval.

Pereopods 5, 6, and 7 increasing in length posteriorly, propodus with proximal

grasping spines.

Abdomen typical of genus.

Female: Length 11.6 mm. Differs from male in showing no lengthening of

anterior pereonites; antenna 1 not greatly enlarged nor setose, flagellum with

fifteen articles; antenna 2 longer than antenna 1 peduncle; gnathopod 2 attach-

ment anterior to middle of pereonite II. Abdomen typical of genus.

Remarks

Fresh specimens, examined by the author, were translucent but had large

orange pigment spots randomly distributed over the dorsal part of the body,

and on all appendages except gills. The pattern of the spots was similar on

both sides of the body. In certain collections the juveniles were little colored,

the subadults highly colored, and mature animals less colored (See also

Gardella (1962).)

Distribution

Type locality: Puget Sound, Washington.

Other localities: California coast south to Santa Barbara; off Oregon and

Washington coasts; Friday Harbour, Washington; British Columbia; Popof

Bay, Yakutat, Dutch Harbour, and Kodiak Harbour, Alaska.

New records: Pacific coasts of Oregon, Washington, and Vancouver Island;

Queen Charlotte Islands; Washington and B.C. coasts of Strait of Georgia and

Juan de Fuca Strait; mainland coast of B.C.

Discussion

The two species of Metacaprella are readily distinguishable because of the

differences between the first antennae.

The males of these species also show a resemblance to Caprella alaskana and

C. ferrea. Normally they can be distinguished from the Caprella species because

they both have head spines rather than tubercles. Other obvious characters are

the body spination and the peduncle of antenna 1 in M. kennerlyi and the long

flagellum of antenna | in M. anomala.

YW Wy y Wh i

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FIGURE 8 — Metacaprella kennerlyi. Male, lateral view and appendages. Female gnathopod

2 and brood plates, and anterior pereonites to show spination. Diagrammatic

dorsal view of male showing typical arrangement of body spines.

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Map fig. 3 — Known distribution of (a) Metacaprella kennerlyi, (b) Caprella alaskana, and

Genus CAPRELLA Lamarck 1801

Antenna 2 usually with swimming setae, flagellum biarticulate; mandibular

palp absent; molar present; outer lobe of maxilliped larger than, or equal to,

inner lobe; gills on pereonites III and IV; pereopods 3 and 4 absent; pereopod

5 six-segmented; abdomen of male with one pair of appendages and one pair

of lobes, of female with one pair of lobes.

Caprella alaskana. Mayer 1903 (Figure 9, Map 3)

Synonymy

Caprella alaskana Mayer 1903

Material Examined

Vancouver Island: over twenty males, seventeen females, stations Fl, F2,

1955; N1, 07b, V5, 1959; H43, 1964; NMC 10913, 10914, 10921-10923, 10930;

USNM 101292.

Queen Charlotte Sound: six males, seven females, station H39, 1964; NMC

10929.

Queen Charlotte Islands: over twenty males, seventeen females, stations W4,

E21, H8a, H8b, H®, H14, 1957; NMC 10915-10920.

Alexander Archipelago: over ten males, over twenty females, stations A33,

A168, A171, A175, 1961; NMC 10926-10928, 11006.

Prince William Sound: twelve males, fourteen females, stations A129,

A151, 1961; NMC 10924, 10925.

Canoe Bay, Alaska: one juvenile, USNM Alaska King Crab Expedition

12-40. 3

Description

Male: Spination of head and body highly variable: the anterior pereonites

may be smooth with blunt dorsal spines showing only on pereonites V to VII;

or there may be dorsal spines on all pereonites and on head; antero-lateral

projections on pereonites III and IV always present. Length 13.6 mm.

Antenna 1 very long, equal to distance from head to pereonite V inclusive;

flagellum shorter than second peduncular article and having at least sixteen

articles. Antenna 2 shorter than the two basal articles of antenna 1, flagellum

with short setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Gnathopod 1 with strongly serrate grasping margin to propodus and dactylus.

Gnathopod 2 propodus length more than twice width; palm with proximal

grasping spine and accessory spine, median to distal poison spine, distal triangu-

lar projection. Basis attached posteriorly on pereonite II, slender, with small

antero-lateral ridge.

Gills oval.

Pereopods 5 to 7 increasing in length, propodus with proximal grasping spines.

EJS)

FIGURE 9 — Caprella alaskana. Subadult male, lateral view and appendages. Female gnatho-

pod 2. Lateral views of two adult females, showing variations in spination, not

to scale.

Abdomen typical of genus.

Female: Length 9.3mm. Differs from male in being spinier; shows no

lengthening of anterior pereonites; antenna | flagellum has eighteen articles;

gnathopod 2 arises anteriorly on pereonite II. Abdomen typical of genus.

Remarks :

Specimens of this species appear to show considerable variation in the degree

of spination of their bodies. They range from the very spiny ‘variety’ of Mayer

to his less spiny typical form to those, examined by the author, which are

almost spineless. The females also show a variation from moderately spiny

to extremely spiny, as shown in Figure 9. The differences are, in fact, so great

that they create a doubt that these are members of the same species. As only

one spiny female was found in the present material, and as no spiny males

have been examined, no decision can be made on the specific status of the

varieties.

All specimens examined showed multiplication of setae in the mandibular

setal row.

Distribution

Type locality: not cited, probably Alaska.

Other localities: Aleutian Islands and Kodiak Island, Alaska; Fort Rupert,

Vancouver Island, B.C.

New records: Prince William Sound and Alexander Archipelago, Alaska;

Queen Charlotte Islands, Queen Charlotte Strait, and Vancouver Island, B.C.

Caprella ferrea Mayer 1903 (Figure 10, Map 3)

Synonymy

Caprella ferrea Mayer 1903

Metacaprella ferrea — Dougherty and Steinberg 1953; 1954

Material Examined

Vancouver Island: four males, nine females, station P6c, 1955; NMC 10951;

one female, station H44, 1964; NMC 10957.

Queen Charlotte Islands: one male, station W8, 1957; four males, station W9,

1957; two males, one female, station W11, 1957; two males, station W12, 1957;

NMC 10952-10955.

Alexander Archipelago: one male, one female, station A6, 1961; NMC 10956.

British Columbia (no other data): two males, two females, NMC 10958.

Description

Male: Body spination as follows: dorsally there is one pair of small blunt

spines on the head; pereonite II has one large pair plus a variable number of

pairs; pereonite III has one pair medianly plus other pairs; pereonite IV has

median and posterior pairs; pereonite V has three pairs; and VI and VII have

two pairs each; laterally there are anterior projections and spines over the gills

on pereonites III and IV. Length of largest male found 9.4 mm; no adult male

was found.

Antenna 1 longer than cephalon plus pereonites II and III, peduncle lightly

setose, flagellum short (in adult it is shorter than third peduncular segment) and

having up to sixteen articles. Antenna 2 shorter than the two basal articles of

antenna 1 (the basal article in adult), flagellum with long swimming setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate but

not five-toothed.

Gnathopod 1 with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus length more than twice the width; palm with proximal grasping

spine and accessory spine, small distal poison spine and triangular projection.

Basis attached posteriorly on pereonite II, stout, with large antero-lateral ridge.

Gills oval.

Pereopods 5 to 7 short, propodus with proximal-medial grasping spines.

Abdomen typical of genus.

Female: Length 6.4 mm. Differs from male in having smaller head and body

spines; antenna | is not noticeably long and the flagellum has ten articles;

antenna 2 is longer than peduncle of antenna 1; gnathopod 2 arises anteriorly

on pereonite II; abdomen typical of genus.

Remarks

Caprella ferrea was provisionally referred to genus Metacaprella by Dougherty

and Steinberg (1953) because of its general resemblance to M. kennerlyi and

M. anomala, although no females had been found at that time; examination of

the female abdomen shows no evidence of rudimentary limbs, and so this species

has been restored to Caprella. It can be most easily distinguished from the two

Metacaprella species by the fact that it has paired spines postero-dorsally on

pereonite IV whereas both M. kennerlyi and M. anomala have unpaired spines

in this position; there is also the difference in the first antennae, which in

M. kennerlyi are densely setose and in M. anomala have the very long flagellum.

Distribution

Type locality: Humboldt Bay, California.

Other localities: Moss Beach and Monterey Bay, California.

New records: Pacific coasts of Vancouver Island and Queen Charlotte Islands,

B.C.; Alexander Archipelago, Alaska.

Discussion

C. ferrea and C. alaskana are extremely difficult to differentiate in the juvenile

and immature stages. As adults, C. alaskana is very much more slender than

C. ferrea, and the latter has a rougher body surface, always has headspines and

frequently has small but definite tubercles laterally on the anterior pereonites.

Pereonite V in C. alaskana is usually longer than VI plus VII and in C. ferrea is

shorter than VI plus VII. However, in the immature stages where the spination

of the body is not properly developed, and where the proportions of the body

are not established, these features are insufficient to distinguish between these

two species. Until more detailed investigation reveals more reliable characters,

it has been assumed that those immature specimens with no headspines are

C. alaskana and those with headspines are C. ferrea.

Ficure 10 — Caprella ferrea. Subadult male, lateral view and appendages. Female gnathopod

2 and brood plates. Lateral view of adult female, to show dorsal spination, not

to scale.

Caprella angusta Mayer 1903 (Figure 11, Map 4)

Synonymy (see Remarks)

Material Examined

Oregon Coast: four males, four females, stations W50, W57, W61, 1966;

NMC 10900-10902.

Juan de Fuca Strait: one male, station W34, 1966; NMC 10899.

Vancouver Island: thirteen males, seven females, stations Fl, P6c, 1955;

01, 03, 05, V3, V4b, V5, 1959; H44, 1964; NMC 10886, 10887, 10890-10896.

Queen Charlotte Sound: four males, two females, stations H50, H53, 1964;

NMC 10897, 10898.

Queen Charlotte Islands: two males, two females, stations W8, W12, 1957;

NMC 10888, 10889.

Description

Male: Head with median triangular anteriorly pointing spine; pereonites may

be smooth or may have small paired dorsal tuberculations, particularly posteri-

orly. Length 10.8 mm.

Antenna 1 as long as or longer than cephalon plus pereonite II, flagellum just

equal to peduncle and having twelve articles; antenna 2 longer than antenna 1

peduncle, flagellum with short setae.

Mouthparts typical of genus; !acinia mobilis of right mandible indistinctly

toothed.

Gnathopod 1 with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus twice as long as it is broad, palm with proximal poison spine

with small proximal accessory spine, triangular or rectangular projection dis-

tally; some setae on palm. Dactylus thickened proximally, tapered distally,

grasping margin denticulate. Basis attached anterior to middle of pereonite II,

having antero-lateral ridge.

Gills fat, oval.

Pereopods 5 to 7 increasing in length, propodus palm concave, grasping spines

proximal.

Abdomen typical of genus.

Female: Length 9.7 mm. Differs from male in having antenna 1 with nine

articles; gnathopod 2 attached anteriorly on pereonite II, with small propodus

bearing proximal grasping spine and accessory spine and evenly tapered dactylus.

Abdomen typical of genus.

Remarks

Of the colored specimens examined, the females were completely pink, while

the males had pink patches.

C. angusta has been “tentatively assigned” to C. penantis by McCain (1968);

however, it appears to differ sufficiently from C. penantis, as figured and

described by him, to justify its separate specific status. In general appearance

C. angusta appears to be a slimmer and less setose animal than C. penantis; it

shows no evidence of pleura; and the first and second gnathopods, as well as

the mouthparts, have noticeably fewer setae. McCain states that the develop-

ment of pleura and the stoutness of the body appear to be related to growth,

“larger individuals having a robust body and well developed pleura.” As his

largest male and female were significantly longer than the largest specimens

examined by the author, there may appear to be grounds for his decision to

include C. angusta with C. penantis. However, there do appear to be other

characters that are different in the two species, and it has been concluded that

these are sufficiently significant to separate C. angusta from C. penantis.

C. angusta C. penantis

Antenna 1 setose not setose

Pereonite I equal to head shorter than head

Gnathopod 2 propodus longer than arm propodus equal to arm

Body spines present or absent absent

Setation (of slight strong

appendages)

If C. angusta is indeed a true species, this leaves the ‘acutifrons’ varieties

natalensis, neglecta, and porcellio in an ambiguous position. They are described

by McCain as being essentially similar to C. penantis, except for the reduction

of the setae on gnathopod 2. According to Vassilenko (1967), C. neglecta is a

strongly setose type and differs from C. angusta also in the arrangement of the

swimming setae on antenna 2. As specimens of these ‘varieties’ have not been

examined by me, and as their descriptions in the literature indicate their having

the typical C. penantis form, it has been deemed inadvisable to synonymize

them with C. angusta.

It has been decided to keep the specific name angusta, as the specimens exam-

ined appear similar to Mayer’s description and figure of his variety angusta; it

is possible that either C. spinifrons Nicolet, 1849, or C. novae-zealandiae Kirk,

1878, may have precedence over this name, but until specimens from Chile or

Australia and New Zealand can be examined, no decision about the valid name

of this species can be made.

Distribution

Type locality: not cited, California.

Other localities: Dillon Beach, Pacific Grove, Santa Catalina, and Avalon,

California.

New records: Pacific coasts of Oregon, Vancouver Island, Queen Charlotte

Islands; Washington and B.C. coasts of Juan de Fuca Strait; Queen Charlotte

Sound, and Queen Charlotte Strait, B.C.

La Follette’s ““C. geometrica’’ from Laguna Beach has not been included as

there is some doubt whether this is C. angusta.

97949—4

FiGureE 11 — Caprella angusta. Male lateral view and appendages. Female gnathopod 2.

Caprella incisa Mayer 1903 (Figure 12, Map 4)

Synonymy

Caprella acutifrons var. incisa Mayer 1903

Caprella incisa — Dougherty and Steinberg 1953; 1954 — McCain 1968

Material Examined

Oregon Coast: over forty individuals, stations W57, W66, 1966; NMC 10880,

10881.

Washington Coast: six males, two females, stations W14, W35, W39, W50,

1966; NMC 10876-10879.

Vancouver Island: three males, four females, station O7b, 1959; NMC 10871;

USNM 172360.

O°N

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Map fig. 4 — Known distribution of (a) Caprella angusta, (b) C. incisa, and (c) C. verrucosa

within the American Pacific boreal region.

Queen Charlotte Sound: one juvenile, station H1, 1964; NMC 10874.

Queen Charlotte Islands: one female, station JWS 92, 1965; NMC 10875.

Baranof Island: two males, stations A171—172, 1961; NMC 10873.

Off Dangerous River, Alaska: over forty individuals, station A156, 1961;

NMC 10872.

British Columbia (no other data): seventeen males, eleven females, NMC

10882-10885.

Description

Male: Head with median triangular, anteriorly pointing spine; rest of body

with paired and unpaired tuberculations on all pereonites; pleura present.

Length 8.7 mm.

979494}

Antenna 1 longer than cephalon plus pereonite II, flagellum just shorter than

peduncle and with eleven articles; peduncular articles 2 and 3, and distal portion

of article 1, setose; antenna 2 longer than peduncle of antenna 1, flagellum

with long swimming setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Gnathopod 1 with weakly serrate grasping margin to propodus and dactylus.

Gnathopod 2 propodus very large, breadth more than half length, palmar

surface with median ventral-pointing poison spine, and distal rounded or

rectangular projection. Dactylus thick proximally, tapered distally, grasping

margin denticulate. Basis attached at middle of pereonite II, having large

antero-lateral ridge.

Gills oval.

Pereopods 5 to 7 increasing in length, propodus palm concave, grasping spines

proximal.

Abdomen typical of genus.

Female: Length 8.8 mm. Differs from male in that antenna 1 is not setose,

flagellum having nine articles; gnathopod 2 attached anteriorly on pereonite II,

and having small propodus with proximal grasping spine and median and distal

palmar projections. Abdomen typical of genus.

Remarks

When this species showed any color, it had lateral and median dorsal bands

of orange from the head to pereonite V inclusive; the color was slight or absent

on all appendages.

Distribution

Type locality: not cited, California.

Other localities: Pacific Grove, Santa Catalina, Point Reyes, Dillon Beach,

and Moss Beach, California; San Juan area, Washington.

New records: Pacific coasts of Oregon, Washington, Vancouver Island, and

Alexander Archipelago; Washington coast of Juan de Fuca Strait, Queen

Charlotte Sound, B.C.

Caprella verrucosa Boeck 1872 (Figure 13, Map 4)

Synonymy

Caprella verrucosa Boeck 1872 — Mayer 1882 — Dougherty and Steinberg

1953; 1954 — McCain 1968.

Caprella septentrionalis var. verrucosa Mayer 1890

Caprella acutifrons var. verrucosa Mayer 1903

Caprella acutifrons — La Follette 1914 (non Latreille 1816)

FIGURE 12 — Caprella incisa. Male, lateral view and appendages. Female gnathopod 2.

Caprella tuberculata Shaw 1916 (non Bate and Westwood 1868)

Caprella acutifrons f. verrucosa — Arimoto 1930 — Utinomi 1943; 1947

Material Examined

Washington Coast: one male, station W40, 1966: NMC 10869.

Vancouver Island: twelve males, five females, stations P6c, 1955; 03, 05, V3,

1959; H44, 1964; NMC 10862, 10866-10868, 10870.

Queen Charlotte Islands: one male, five females, stations W8, W9, W12, 1957;

NMC 10863-10865.

Description

Male: Head with median triangular anteriorly pointing spine; rest of body

with large blunt paired and unpaired spines dorsally, and laterally on pereonites

II to V. Length 6.1 mm.

Antenna 1 longer than cephalon plus pereonite II, flagellum equal to peduncle

articles 1 + 2, and with nine articles; antenna 2 longer than peduncle of an-

tenna 1, flagellum with short setae.

Mouthparts typical of genus; lacinia mobilis of right mandible not strongly

denticulate.

Gnathopod 1 with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus twice as long as it is broad; palmar surface with proximal

anteriorly pointing poison spine and distal triangular or rectangular projection;

antero-dorsal surface with tuberculations. Dactylus thick proximally, tapering

distally, grasping margin denticulate. Basis attached to middle of pereonite II,

having antero-lateral ridge.

Gills fat, oval.

Pereopods 5 to 7 increasing in length, propodus palm concave, grasping spines

proximal.

Abdomen typical of genus.

Female: Length 5.1 mm. Differs from male in having seven articles to fla-

gellum of antenna 1; antenna 2 nearly equals antenna | in length; gnathopod 2

propodus small, with proximal grasping spine and accessory spine, slight distal

projection and evenly tapered dactylus. Abdomen typical of genus.

Distribution

Type locality: California, probably near San Francisco.

Other localities: Dillon Beach, Pacific Grove, Santa Catalina, Point Reyes,

and Laguna Beach, California; Misaki, Yokohama, Tateyama Bay, and

Onagawa Bay, Japan.

New records: Pacific coasts of Washington, Vancouver Island, and Queen

Charlotte Islands.

Discussion

These three species of the so-called ‘acutifrons’ group are typically similar

in their general body shape, the shape and armature of the second gnathopods,

and the proportions of the antennae. They range from the almost smooth

C. angusta through C. incisa with its tuberculations to C. verrucosa with its

large blunt spines. C. incisa can be distinguished from C. verrucosa by its

much smaller tuberculations, the finely setose antenna 1 and the very large

propodus and antero-lateral ridge on the gnathopod 2. C. angusta can be

distinguished from C. incisa by its almost smooth dorsal surface, normal sized

gnathopod 2 propodus and basal ridge, the anterior attachment of the gnatho-

pod 2, the absence of pleura, and of setae on antenna 1.

Antenna 1 setae

Gnathopod 2

attachment

basal ridge

propodus

Body spines

Pleura

C. incisa

many, short

median

large,

rectangular

width > 4

length

small

present

C. angusta

sparse

triangular

width = 4 length

minute, never

on I

absent

C. verrucosa

absent

median

triangular

width = 4 length

large

present

FiGurRE 13 — Caprella verrucosa. Male, lateral view and appendages. Female gnathopod 2

Caprella borealis Mayer 1903 (Figure 14, Map 5)

Synonymy

Caprella acutifrons var. borealis Mayer 1903

Caprella alaskensis Holmes 1904

Caprella borealis — Utinomi 1943a; 1947 — McCain 1968

Material Examined

Prince William Sound: two males, two females, stations A103, A115, A121,

1961; NMC 10859-10861.

Queen Charlotte Islands: fifteen males, four females, stations H8a, H8b,

H9, El4a, W2, 1957; NMC 10854-10858.

Description

Male: Head with median tubercle over eye; paired dorsal tubercles present

posteriorly on pereonite II, medially on pereonite III, medially and posteriorly

on pereonite IV, and on V, VI, and VII; laterally there are tubercles anteriorly

on pereonites III and IV, and over the point of attachment of the gnathopod 2

and the pereopods; other lateral and dorsal tuberculation variable. Length

7.2 mm. Pereonites I and II lengthen with maturity, pereonites III to VII are

progressively shorter, VI plus VII longer than V.

Antenna 1 approximately equal to cephalon plus pereonite II, flagellum with

seven articles, and shorter than peduncular articles 1 plus 2; antenna 2 just

shorter than the two basal articles of antenna 1, flagellum with short setae.

Mouthparts typical of genus; lacinia mobilis of right mandible finely denticulate.

Gnathopod | with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus more than twice as long as it is broad; palm convex, with

minute proximal projection, distal poison spine and more distal triangular

process. Dactylus evenly tapered, inner margin denticulate. Basis attached

posterior to the middle of pereonite II, short and with antero-lateral ridge;

ischium with large spine-like antero-lateral projection.

Gills small, oval.

Pereopods 5 to 7 progressively longer; propodus palmar surface variable:

in immature males and in all females the palm is not delineated but bears a

few small spines; in adult males there may be a progression from this type of

propodus on pereopod 5 to one showing true palmar development, with a

cluster of four or five proximal grasping spines and with palmar spines on

pereopod 7.

Abdomen typical of genus.

Female: Length 5.5 mm. Differs from the male in showing no lengthening

of anterior pereonites; antenna 1 flagellum with five articles, antenna 2 longer

than antenna 1 peduncle; gnathopod 2 attached anteriorly on pereonite II,

no spine on ischium, propodus not elongate and bearing proximal grasping

spine with accessory spine and two small distal projections, dactylus not

serrate. Abdomen typical of genus.

Distribution

Type locality: Cape Lopatka, Kamchatka Peninsula, USSR.

Other localities: Copper Island, Kamchatka Peninsula, USSR; Akkeshi Bay,

Japan; Orca, Prince William Sound, Alaska.

New records: Northern Prince William Sound, Alaska; Queen Charlotte

Islands, B.C.

Discussion

Mayer included C. borealis among his varieties of “‘C. acutifrons” ; however,

this species is different from the typical ‘acutifrons’ varieties, particularly in the

type of head spine, in the point of attachment of gnathopod 2, and in the

lengthening of the anterior segments, and was given separate specific status by

Utinomi (1943a).

There is a great similarity between this species and C. alaskensis Holmes 1904.

Unfortunately Holmes did not make any mention of the size of his incomplete

specimen, but if, as appears likely, it was a large adult male, any differences

between it and C. borealis can be accounted for by its maturity: i.e., the com-

parative lengths of pereonites I and II, the more posterior position of gnathopod

2, the enlarged spine on the ischium, and the absence of the proximal projection

on the palm of gnathopod 2. It has not been possible to locate Holmes’s single

specimen for comparative study, but it seems certain that these two species are

the same, and I have, therefore, assigned alaskensis to species borealis.

Caprella californica Stimpson 1857 (Figure 15, Map 5)

Synonymy

Caprella californica Stimpson 1857; 1863 — Boeck 1871 — Mayer 1882 —

Holmes 1909 — Dougherty and Steinberg 1953; 1954 — Gardella 1962 —

Saunders 1966 — McCain 1968

Caprella scaura—Stebbing 1888— Johnson and Snook 1927 — Wailes

1931 — Light 1941 — Ricketts and Calvin 1952; (non Templeton 1836)

Caprella scaura var. californica Mayer 1890; 1903

Caprella scaura var. scauroides Mayer 1903 — Utinomi 1947

Caprella scaura var. spinirostris Mayer 1890; 1903

Material Examined

Queen Charlotte Islands: five males, four females, stations E5, E25, W3a,

W15b, 1957; 3532, 1935; NMC 10959, 10962-10965.

Hecate Strait: over twenty males, over twenty females, stations H8, H23,

1964; NMC 10973, 10974.

Queen Charlotte Sound: nine males, over twenty females, stations H53,

H57, H58, 1964; NMC 10976-10978.

Vancouver Island: over forty males, twenty-seven females, stations M2,

P2, 1955; O2b, O04, 06, O7a, O7b, O7d, O13, 1959; H44, 1964; NMC 10960,

10961, 10966-10972, 10975.

Washington coast: over twenty males, fourteen females, stations W46, W47,

1966; NMC 10979, 10980.

FiGurE 14 — Caprella borealis. Male, lateral view and appendages. Female gnathopod 2 and

propodus of pereopod 7.

Description

Male: Body spination variable, but long slender anteriorly pointing cephalic

spine, and small ventral spine between the insertions of the second gnathopods,

always present. Spination of body varies from antero-lateral spines on pereo-

nites III and IV but no dorsal spines, to antero-lateral spines on pereonites

III to VI, lateral spines over gills on pereonite III, median dorsal spine post-

eriorly on pereonite II, medianly and posteriorly on III and IV, medianly on V,

and two pairs on VI. Length 21.5 mm.

Antenna 1 longer than cephalon plus pereonites II and III, flagellum shorter

than peduncle article 2 and with nineteen articles. Antenna 2 shorter than

antenna | peduncle articles 1 plus 2, flagellum with swimming setae.

O°N

150°w A

~ = ALASKA ee

Prince William ‘

Sound

~ fs ¢

t 4

GULF OF ALASKA Fe YUKON

za - ~

Yokutat Bay A.

145°w ;

i z "a,

« ms =| x.

PACIFIC OCEAN

140°w

Queen Charlotte f :

Islands

em)

(am)

ic)

135°W ° 17

50°N iV)

%K BRITISH

S J

“ COLUMBIA

Vancouver Island

130°w

45°N

Columbia R,

150°w =f /

S15, ALASKA a

Prince William Pe

Sound

GULF OF ALASKA

ras

Yakutat Bay yy) lod

».

foe..’

Cross Sound |

PACIFIC OCEAN

140°w

Queen Chorlotte

Islands

Columbia R,

125°w “OS

150°w

Prince William

Sound

GULF OF ALASKA

x :

PACIFIC OCEAN Ni /

140° w ; C4

Queen Charlotte

Islands

45°N

Columbia R,

125°W as

Map fig. 5 — Known distribution of (a) Caprella borealis, (b) C. californica, and (c) C. dre-

panochir within the American Pacific boreal region.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five

-toothed.

Gnathopod 1 with serrate grasping margins to dactylus and propodus. Gnatho-

pod 2 propodus longer than basis, and four times longer than it is wide, narrow

proximally and widening distally; palmar surface with two strong projections,

one medial with associated accessory spine, the second halfway between the

proximal projection and a distal triangular projection; proximal and antero-

distal surfaces setose. Dactylus short. Basis attached posteriorly on pereonite IT

and bearing an antero-lateral projection distally.

Gills long, elliptical.

Pereopods 5 to 7 increasing in length; propodus palmar surface concave,

grasping spines proximal. ‘

FicureE 15 — Caprella californica. Male, lateral view and appendages. Female gnathopod 2.

Abdomen typical of genus.

Female: Length 11 mm. Differs from male in showing no lengthening of

anterior pereonites; antenna 1 flagellum has seventeen articles; antenna 2.

as long as peduncle of antenna 1; gnathopod 2 attached just anterior to middle

of pereonite II, propodus not elongated, palm with proximal grasping spine and

accessory spine and minute distal poison spine, basis short. Abdomen typical

of genus.

Remarks

According to Gardella (1962) the color of the male is “typically dark

brown, but varies to a dark greenish-brown. The claws are tipped with white

and have red polka dots. The female is usually a light greenish-brown with green

second legs and red polka dots on all the claws, but the color may vary to

nearly black.”

Distribution

Type locality: San Francisco Bay, California.

Other localities: California from San Diego to Humboldt Bay; Juan de Fuca

Strait and San Juan area, Washington; British Columbia; Caldera, Chile;

Tsugaru Strait and off southwestern Japan; Korea Strait and off Korea;

Formosa Strait and South China Sea.

New records: Pacific coasts of Washington and Vancouver Island; Queen

Charlotte Islands, Hecate Strait, Queen Charlotte Sound, and Strait of

Georgia, B.C.

Discussion

Mayer separated three varieties, having the ventral spine between the second

gnathopods, from Caprella scaura; these he distinguished from each other on

the basis of the spination of pereonite V. It has been seen that the spination of

C. californica is highly variable, and it appears probable that varieties spini-

rostris and scauroides are true members of this species. Although I have not

been able to examine Asian or South American material, I have decided to

assign these two varieties to Caprella californica.

Caprella drepanochir Mayer 1890 (Figure 16, Map 5)

Synonymy

Caprella drepanochir Mayer 1890; 1903— Shoemaker 1920 — Utinomi

1943a; 1947

Material Examined

Bristol Bay, Alaska: one male, one juvenile, USNM Alaska King Crab

Expedition D6-41.

Prince William Sound: over one hundred individuals, stations A80, A81,

A139, A140, A153, 1961; NMC 10903-10907.

Description

Male: Head and body smooth except for antero-lateral projections on pereo-

nite III. Length 12.4 mm.

Antenna 1 approximately equal to cephalon plus pereonite II; peduncular

articles 2 and 3 setose; flagellum shorter than peduncular articles 1 plus 2

and with eleven articles. Antenna 2 longer than antenna 1 peduncle, flagellum

with short setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate but

not five-toothed.

Gnathopod 1 with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus very large, width less than half length; palmar surface with

small proximal spine, large distal poison spine separated by cleft from more

distal triangular projection. Dactylus heavy and slightly curved, inner margin

FiGure 16 — Caprella drepanochir. Male, lateral view and appendages. Female gnathopod 2

and pereopod 7.

slightly denticulate. Basis attached just posterior to middle of pereonite II,

having lateral and medial anterior ridges. The whole limb is setose except for

the distal portion of the propodus, which is tuberculate, and the dactylus.

Gills round.

Pereopods 5 to 7 show slight increase in length posteriorly; propodus palm

concave with proximal grasping spines.

Abdomen typical of genus.

Female: Length 7.7 mm. Differs from male in having short pereonite I;

antenna | flagellum with ten articles; gnathopod 2 propodus palm has proximal

grasping spine and accessory spine, distal minute poison spine; pereopods

much more slender. Abdomen typical of genus.

Distribution

Type locality: Not cited, probably China.

Other localities: Vladyvostok and Beringa Island, USSR; Akkeshi Bay,

Japan; Chamisso Harbour, Eschscholtz Bay, and Port Clarence, Alaska.

New records: Southern Prince William Sound, Alaska.

Discussion

Mayer (1903) mentions a “‘northern variety” of C. drepanochir, to which the

specimens in NMC bear a greater resemblance than to the ‘typical’ form;

however, study of specimens of the “variety” indicates that there is little

difference between them and those of the typical form, except that they appear

to be sturdier. It has been concluded that both forms belong to the same

species. The specimens described by Utinomi (1943a) appear to be immature,

particularly from the description of the gnathopod 2, which is typical of the

immature male.

Caprella equilibra Say 1818 (Figure 17, Map 6)

Synonymy

Refer to McCain (1968); see also Light 1941

Material Examined

Vancouver Island: six males, one female, stations P6c, F1, 1955; Ol, O3,

O5, O11, 1959; NMC 10989-10994.

Hecate Strait: three males, two females, stations H10, H33, 1964; NMC

10995, 10996.

Queen Charlotte Sound: five males, over ten females, station H53, 1964;

NMC 10997.

Description

Male: Head and body smooth dorsally except for pereonite V which may

have paired spines; laterally there are large anteriorly pointing spines at the

bases of the second gnathopods, antero-lateral spines on pereonites III and IV;

pereonite V is distinctly sculptured anteriorly giving the appearance of large

lateral spines. Ventrally there is a large spine between the bases of the second

gnathopods. No mature male was found, largest male length 10.2 mm.

Antenna | as long as cephalon plus pereonite II, flagellum less than one-

quarter of its total length. Antenna 2 shorter than the two basal articles of

antenna 1, flagellum with short setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Ficure 17 — Caprella equilibra. Subadult male, lateral view, appendages, and dorsal view of

pereonite V. Female gnathopod 2 and brood plates.

§0°N

Q x

150° w 7 a4

S15 ALASKA Pa

Prince William ’

Sound a

x & _

= od

GULF OF ALASKA Pe YUKON

Yakutat Bay a

145°w ;

. &

ees

4 = x,

Cross Sound

PACIFIC OCEAN

40°w

x HE

{2s

g —

Queen Crhorlotte @

Islands I

135°W ° 2)

5O°N

Vancouver Island é

‘ .

45°N

Columbia R

ESS

O°N

e x

150°w -

Prince William

Sound

GULF OF ALASKA

Yakutat Boy

Cross Sound |

pauline

PACIFIC OCEAN

140°w

Queen Charlotte

Islands

O°N

150°w '

Prince William

Sound Pd

x <i

GULF OF ALASKA

Yokutat Boy

Cross Sound |

’

55°N oe ‘

* Vai

PACIFIC OCEAN te

0°

<)g

5 i

140° Ww /

Queen Charlotte

Islands

Columbia R,

125°w LS

Map fig. 6— Known distribution of (a) Caprella equilibra, (b) C. mendax, and (c) C.

pilidigita within the American Pacific boreal region.

Gnathopod 1 with barely serrate grasping margin to propodus and dactylus.

Gnathopod 2 propodus with proximal grasping spine, distal triangular projec-

tion with poison spine just proximal to it; size of poison spine appears to de-

crease with age. Basis attached posteriorly on pereonite II; in the adult it is

shorter than half pereonite II and is produced in an antero-lateral ridge ending

in a spine-like process.

Gills large, oval.

Pereopods 5, 6, and 7 increasing in length posteriorly; propodus with proximal

grasping spines, palm slightly concave, with few spines.

Abdomen typical of genus.

97949—5

Female: Length 7.2 mm. Differs from male in showing no lengthening of

anterior pereonites; antenna 2 equal in length to peduncle of antenna 1;

gnathopod 2 attached anteriorly on pereonite II and having a longer basis.

Abdomen typical of genus.

Remarks

There are certain small differences between the specimens examined from

the Pacific, and the descriptions of this species from the Atlantic. As this is an

extremely widespread species it is more than likely that variations will be found

within its range; until large numbers of adult males can be studied no decision

can be reached on the exact taxonomic status of these variations.

Distribution

Type locality: South Carolina.

Other localities: See McCain (1968); also San Juan area, Washington.

New records: Vancouver Island, Queen Charlotte Sound, and Hecate Strait,

B.C.

Caprella mendax Mayer 1903 (Figure 18, Map 6)

Synonymy

Caprella mendax Mayer 1903

Caprella equilibra — Dougherty and Steinberg 1953

Material Examined

Hecate Strait: seven males, one female, stations H21, H23, H37, 1964; JWS

85, 1965; NMC 10999-11002.

Vancouver Island: eleven males, eleven females, station V3, 1959; NMC

10998.

San Juan Islands: ten males, one female, USNM 187867.

Description

Male: Body smooth, except pereonite IJ, which has a single ventral spine

between the bases of the second gnathopods and a small spine at the base of

each second gnathopod. Length 16.4 mm.

Antenna 1 longer than cephalon pius pereonites II and III, flagellum shorter

than peduncle articles 1 plus 2 and with nineteen articles. Antenna 2 shorter

than the two basal articles of antenna 1, flagellum with short setae.

Mouthparts typical of genus, lacinia mobilis of right mandible denticulate,

not five-toothed.

Gnathopod 1 with slightly serrate margin to dactylus and propodus. Gnathopod

2 propodus with proximal grasping spine with small accessory spine, distal

triangular projection with poison spine just proximal to it. Basis attached |

posteriorly on pereonite II, shorter than half pereonite IJ and having antero-

lateral projection.

Gills elliptical.

Pereopods 5, 6, and 7 increasing in length posteriorly; propodus with proximal

grasping spines and few spines on the concave palm.

Abdomen typical of genus.

Female: Length 11.2 mm. Differs from male in showing less elongation of

anterior pereonites; antenna 1 flagellum longer than peduncle articles 1 plus 2

and having sixteen articles; antenna 2 shorter than peduncle of antenna 1.

Abdomen typical of genus.

Ficure 18 — Caprella mendax. Male, lateral view, and dorsal view of pereonite V. Subadult

male, appendages. Female gnathopod 2.

97949—54

Remarks:

Mayer distinguished C. mendax from C. equilibra on the basis of the structure

of the second gnathopod, and the long pereonite V in C. mendax as contrasted

with the short pereonite V in C. equilibra. Mayer’s distinction appears rather

tenuous in his monograph; however, from studying his specimens in USNM

it appears that there is a definite difference between these two species. The

specimens named here as C. mendax show slight differences from the type

in the gnathopod 2 but are otherwise similar, and it has been decided to rein-

state this species as valid and to include these specimens in it.

Distribution

Type locality: not cited, California.

Other localities: Pacific Grove, Santa Barbara, and San Diego, California.

New records: Vancouver Island and Hecate Strait, B.C.; San Juan Islands,

Washington.

Caprella pilidigita n. sp. (Figure 19, Map 6)

Material Examined

Male holotype, NMC 10985, Townsend Point, St. John Harbour, Athlone

Island, B.C., station H53, 1964.

Over thirty male paratypes, NMC 10986, Townsend Point, St. John Harbour,

Athlone Island, B.C., station H53, 1964.

Two males, NMC 10987, Goose Island, B.C., station HS0, 1964.

One male, NMC 10988, Cox Point Inlet, Trutch Island, B.C., station H26,

1964.

Two males, USNM 101292, Stuart Island, B.C.

Description

Male: Body smooth except for a small ventral spine between the bases of

the second gnathopods. Length 9.4 mm. Pereonites I and II elongated in adult.

Antenna 1 longer than cephalon plus pereonites II and III; flagellum equals

peduncle article 2 and has twelve articles; distal portion of third peduncular

article setose. Antenna 2 shorter than the two basal articles of antenna 1,

flagellum with short setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Gnathopod 1 with slightly serrate grasping margin to propodus and dactylus.

Gnathopod 2 propodus with proximal grasping spine, distal triangular projec-

tion with small poison spine proximal to it; dactylus with hairs along inner

margin. Basis attached posteriorly on pereonite IJ and having slight antero-

lateral projection; ischium and merus square, with no projection.

Gills small, elliptical.

Pereopods 5, 6, and 7 increasing in length posteriorly; propodus with proximal

grasping spines, and many spines on concave palm. Abdomen typical of genus.

Female: No female specimens were found.

Remarks

The specific name refers to the dactylus of gnathopod 2.

Distribution

Type locality: latitude 52° 12’ N, longitude 128° 29’ W, Townsend Point,

St. John Harbour, station H53/64; NMC 10985.

Other localities: Hecate Strait, Queen Charlotte Sound, and Johnstone

Passage, B.C.

Ficure 19 — Caprella pilidigita. Male holotype, lateral view, and dorsal view of pereonite V.

Male paratype, appendages.

979496

Discussion

The species Caprella equilibra, C. mendax, and C. pilidigita are very similar

in appearance, but their differences have been considered sufficient to name

them three different species rather than variants of C. equilibra.

C. equilibra C. mendax C. pilidigita

Antenna 1 length = pereonitesI+II > pereonites > pereonites

I+ 11+ I I++ 1

flagellum length, 4 total antenna > 4 total = peduncle

antenna segment 2

no. of articles 12-16 19 12

Gnathopod 2 basis, 4 of appendage 4 of appendage +4 of appendage

ventral spine, large large small

basal spines large small absent

Pereonite III and IV

lateral anterior spines no spines no spines

Pereonite V, lateral anterior spines no spines no spines

length < pereonite IV > pereonite IV > pereonite IV

The species C. piligidita appears definitely different from C. equilibra and

C. mendax, particularly in the structure of the gnathopod 2, the body spines,

and the setation of antenna 1. The difference between C. equilibra and C.

mendax has been harder to assess because of the scarcity of adult males. Such

ecological information as is available indicates that C. mendax is found mainly

in deep water while C. equilibra is found both in deep water and intertidally.

It is hoped that new material will be collected to help solve the status of these

two species; meanwhile C. mendax is being separated from C. equilibra on

the basis of the antenna 1, pereonite V, and differences between the females.

Caprella gracilior Mayer 1903 (Figure 20, Map 7)

Synonymy

Caprella linearis var. gracilior Mayer 1903

Caprella gracilior—Dougherty and Steinberg 1953; 1954

Material Examined

Prince William Sound: over fifty individuals, stations A85 (40 ft.), A102

(60 ft.), A103 (66 ft.), 1961; NMC 10908-10910.

Alexander Archipelago: two males, one female, stations A165, 1961 (120

ft.); JWS 85, 1965 (30 fm); NMC 10911, 10912.

Description

Male: Head and body usually smooth anteriorly and with two pairs of dorsal

tubercles on pereonite V. Length 16.3 mm. This animal is very slender, with

pereonites I and II much elongated.

Antenna 1 longer than cephalon plus pereonite II; peduncle with second

article longer than first plus third articles; flagellum shorter than the two ba-

sal articles of the peduncle and with at least fifteen articles. Antenna 2 shorter

than the two basal articles of antenna 1, swimming setae short and sparse.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate, not

five-toothed.

Gnathopod 1 with strongly serrate grasping margin to propodus and dactylus.

Gnathopod 2 three times longer than it is broad, extreme distal portion setose;

palm with proximo-—medial grasping spine and accessory spine, distal poison

spine separated by a cleft from a more distal triangular projection. Dactylus

curved, finely tapered, with short hairs along grasping edge. Basis attached

posteriorly on pereonite II, to which it is of approximately equal length; it

is nearly twice the length of the propodus.

FiGurE 20 — Caprella gracilior. Subadult male, lateral view. Male appendages. Female

gnathopod 2.

97949—63

Gills slim, oval.

Pereopods 5 to 7 very slender, increasing in length posteriorly; propodus

with median grasping spines and no well defined palm.

Abdomen typical of genus.

Female: Length 7.4 mm. Differs from male in showing no lengthening of

anterior pereonites; gnathopod 2 arises anteriorly on pereonite II, propodus

not particularly slender with palm similar to male but with smaller poison

spine, dactylus without hairs. Abdomen typical of genus.

Remarks

This species was separated from C. linearis by Dougherty and Steinberg in

1953, a move contemplated by Mayer and justified by the differences between

60°N

150°w =f =

Ss ALASKA eo

Prince William S

Sound ne

x y

a ae

GULF OF ALASKA ye YUKON

Yakutat Bay (

Doe

145°w as

, ~

55°N

x yh :

PACIFIC OCEAN g i f

140°w aN J

OR:

6 $30

Queen Charlotte

Islands @e

135°W 3

5O°N

% BRITISH

. y

“4 COLUMBIA

Vancouver Island

130°w

45°N

Columbia R,

le neg a ee

ro ALASKA

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Map fig. 7 — Known distribution of (a) Caprella gracilior, (b) C. irregularis, and (c) C.

laeviuscula within the American Pacific boreal region.

these animals. Mayer cites a few intertidal records for C. gracilior, but it would

appear that this is a predominantly deep water species, having been found

mainly below 30 feet, and down to 958 fathoms.

Distribution

Type locality: not cited, probably Alaska.

Other localities: Aleutian Islands, Pribiloff Islands, and Kodiak Island,

Alaska; East Cape, Siberia, USSR; Pacific Grove and Monterey Bay, and

southern California; San Juan area, Washington.

New records: Prince William Sound and Alexander Archipelago, Alaska;

Hecate Strait, B.C.

Caprella irregularis Mayer 1890 (Figure 21, Map 7)

Synonymy

Caprella irregularis Mayer 1890: 1903 — Holmes 1904 — Utinomi 1947

Material Examined

Prince William Sound: over twenty males, over thirty females, stations A86,

A87, A90, A91, A92, A96, A131, 1961; NMC. 10935-10941; USNM acc.

58092.

Alexander Archipelago: nine males, over twenty females, stations A3, A8,

A168, A174, 1961; NMC 10933, 10934, 10942, 10943.

Queen Charlotte Islands: six males, two females, station El4c, 1957; NMC

10931.

Hecate Strait: over forty males, over fifteen females, stations H22, H23,

H26, H30, 1964; NMC 10944-10947.

Queen Charlotte Sound: fourteen males, fourteen females, stations H49,

H53, 1964; NMC 10948, 10949.

Vancouver Island: two males, station O4, 1959; NMC 10932; USNM

172360.

Seattle, Washington: one female, USNM acc. 58092.

Description

Male: Body smooth except for antero-lateral projections on pereonites III

and IV, and median dorsal paired spines on pereonite VI; there may also be

dorsal paired spines on pereonites V and VII. Length 15.8 mm.

Antenna 1 shorter than cephalon plus pereonite II; flagellum shorter than

peduncular articles 1 plus 2 and with fifteen articles; peduncle slightly setose.

Antenna 2 shorter than the two basal articles of antenna 1, swimming setae

sparse, and absent on flagellum.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Gnathopod | with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus nearly three times as long as broad, and longer than basis;

palmar surface with proximal grasping spine plus two accessory spines, one

on each side of palm, distal triangular projection, and large anteriorly pointing

poison spine midway between grasping spine and distal projection; antero-

distal end of propodus with two projections which jut out on either side of

base of claw. Dactylus almost straight proximally, sharply curved distally and

with denticulations along inner edge. Basis attached posteriorly on pereonite II

and having antero-lateral projection.

Gills small, oval.

Pereopods 5 to 7 increasing in size, propodus with proximal grasping spines.

Abdomen typical of genus.

Female: Length 7.7 mm. Differs from male in showing no lengthening of

anterior pereonites; antenna 1 flagellum with twelve articles; antenna 2 longer

than peduncle of antenna 1; gnathopod 2 propodus not elongated nor extended

FiGurRE 21 — Caprella irregularis. Male, lateral view and appendages. Female gnathopod 2.

anteriorly, palm having proximal grasping spine and two accessory spines, distal

minute poison spine and more distal rounded projection; dactylus heavy and

evenly curved. Abdomen typical of genus.

Remarks

The males of this species are easily distinguished by the shape of the second

gnathopod with its anterior wings. The females are very similar to the females

of C. laeviuscula, but differ from them in having two accessory spines at the

base of gnathopod 2 grasping spine, where C. /aeviuscula have only one ac-

cessory spine.

Distribution

Type locality: not cited, probably off Korea.

Other localities: Wiuliuk Harbour, Unalaska, and Prince William Sound,

Alaska.

New records: Alexander Archipelago, Alaska; mainland coast of Queen

Charlotte Sound and Pacific coast of Vancouver Island, B.C.; Puget Sound,

Washington.

Caprella laeviuscrla Mayer 1903 (Figure 22, Map 7)

Synonymy

Caprella laeviuscula Mayer 1903 — Utinomi 1943a; 1947 — Dougherty and

Steinberg 1953; 1954 — Gardella 1962 — Saunders 1966

Material Examined

Oregon; Juan de Fuca Strait; San Juan Islands; Puget Sound; Vancouver

Island; Queen Charlotte Sound; Hecate Strait; Queen Charlotte Islands;

Alexander Archipelago; Prince William Sound; and Canoe Bay, Alaska: over

one hundred lots comprising more than one thousand individuals, NMC

11051-11149; USNM_ 101292, 173540, 181641, 187867, 163372, 102730,

USNM Alaska King Crab Expedition 47-40.

Description:

Male: Head and body smooth except for antero-lateral projections on pereo-

nites IIJ and IV. Length 16 mm.

Antenna 1 equals cephalon plus pereonite II, flagellum shorter than peduncle

articles 1 plus 2 and with thirteen articles. Antenna 2 shorter than antenna 1

peduncle, flagellum with swimming setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Gnathopod 1 with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus less than twice as long as it is broad, and longer than basis;

palm with enormous medial-distal poison spine and proximal grasping spine

and accessory spine. Dactylus heavy, tapered and curved in distal one third,

inner margin denticulate. Basis attached posteriorly on pereonite II.

Gills large, oval.

Pereopods 5 to 7 short, increasing in length posteriorly; propodus with proximal

grasping spines.

Abdomen typical of genus.

Female: Length 7.7 mm. Differs from male in showing no elongation of

anterior pereonites; antenna | flagellum with ten articles; gnathopod 2 attached

anteriorly on pereonite II, propodus shorter than arm and having minute

poison spine; abdomen typical of genus.

Remarks

Among the specimens of C. /aeviuscula there are seven lots of smaller speci-

mens; these are adult males of length of 9.6 mm and are comparatively slimmer

than the typical specimens. Both male and female (5 mm) are of the smaller

size, and, except for one doubtful case, the smaller variety are not found at the

same station as the larger type. As no other difference has been observed, and

as both varieties are found within the same geographical range, the smaller

animals have been included in C. Jaeviuscula.

Of those specimens showing colour, two variations were observed: in one

the body was translucent, with varying numbers of small orange polka dots all

over body, appendages, and brood plates, but not on gills; in the other, the

males were translucent with a few small dark pinkish-brown or black dots on

the body, and many such dots on the head and the distal end of the propodus

of gnathopod 2, and the females were pink with many such dots on body,

brood plates, and distal part of gnathopod 2 propodus but not the other

appendages. A few specimens exhibited a mixture of both types of colora-

tion.

Distribution

Type locality: not cited, Pacific coast of North America.

Other localities: Humboldt Bay and (?) Monterey Bay, California; Puget

Sound, Washington; Victoria Harbour and Fort Rupert, Vancouver Island,

B.C.; Kodiak and Adakh, Alaska; Akkeshi Bay, Japan.

New records: Prince William Sound and Alexander Archipelago, Alaska;

Queen Charlotte Islands, Queen Charlotte Sound, Queen Charlotte Strait,

Strait of Georgia, Juan de Fuca Strait, and Pacific coast of Vancouver Island,

B.C.; Pacific coast of Oregon.

Discussion

C. laeviuscula is the commonest caprellid species in the geographical area

investigated. It shows great tolerance of a wide variety of temperature and

salinity. It is possible that the different forms found are merely the same species

showing adaptation to different ecological conditions. Until sufficient ecological

data can be obtained it has been decided to ignore the size differences as noted.

FIGURE 22 — Caprella laeviuscula. Male, lateral view and appendages. Female gnathopod 2.

2: small variety male, lateral view and gnathopod 2.

Caprella rudiuscula n. sp. (Figure 23, Map 8)

Material Examined

One male holotype, NMC 11036, Gudal Bay, Graham Island, Queen Char-

lotte Islands, station W9, 1957.

One female allotype, NMC 11037, Gudal Bay, Graham Island, Queen

Charlotte Islands, station W9, 1957.

Seven males, ten females, paratypes, NMC 11038, 11040-11042, Queen

Charlotte Islands, stations W8, W9, W11, W12, 1957.

Eleven males, two females, NMC 11044, 11047, Chichagof Island, stations

A22, A168, 1961.

Six males, four females, NMC 11048, Baranof Island, Stations A171-172,

1961.

Twenty-three males, seventeen females, NMC 11045, 11046, Yakutat and

Icy bays, stations A71, A73, 1961.

Twelve males, over fifty females; NMC 11039, 11043, Vancouver Island,

stations P6c, 1955; N1, 1959.

Eight males, five females, NMC 11050, Koeye Estuary, B.C., station H35,

1964.

Three males, one female, NMC 11049, Porcher Island, B.C., station H10,

1964.

Description

Male: Body with dorsal paired tubercles on head, medianly on pereonites II,

III, and IV, anteriorly and medianly on pereonite V; antero-lateral projections

on pereonites III and IV. There are also micro tuberculations over most of the

body, particularly posteriorly, and also on the second gnathopods and the

pereopods. Length 8.5 mm.

Antenna 1 shorter than cephalon plus pereonite II, flagellum shorter than

peduncle articles 1 plus 2 and with twelve articles. Antenna 2 approximately

equal to peduncle of antenna 1, flagellum with swimming setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate,

but not five-toothed.

Gnathopod 1 with serrate grasping margin to propodus and dactylus. Gnatho-

pod 2 propodus less than twice as long as it is broad, palm with giant poison

spine, grasping spine and accessory spine proximal to it; dactylus strongly

curved and tapered distally. Basis attached posteriorly on pereonite II.

Gills oval.

Pereopods stout, quite short, propodus palm concave, with proximal grasping

spines.

Abdomen typical of genus.

Female: Length 4.4 mm. Differs from male in showing no lengthening of

anterior pereonites; antenna 1 flagellum has nine articles. Gnathopod 2 pro-

podus shorter than arm, poison spine minute and distal. Abdomen typical of

genus.

Distribution

Type locality: Gudal Bay, Graham Island, Queen Charlotte Islands, B.C.

Other localities: Long Beach, Vancouver Island, and Queen Charlotte

Sound, B.C.; Alexander Archipelago, Icy Bay, and Yakutat Bay, Alaska.

Discussion |

Caprella rudiuscula bears a strong resemblance to C. /aeviuscula (the specific

name, chosen with this in mind, indicates its tuberculate appearance): both

have similar general appearance and body proportions and show striking

similarities in the form of the second gnathopods. It has been decided to

separate C. rudiuscula from C. laeviuscula mainly because of the tubercles

and microtuberculations, and also because of the slight differences in the

appendages and mouthparts.

FiGurE 23 — Caprella rudiuscula. Male holotype, lateral view. Male paratype, appendages.

Female paratype, gnathopod 2.

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Map fig. 8 — Known distribution of (a) Caprella rudiuscula, (b) C. pustulata, and (c) C.

striata within the American Pacific boreal region.

Caprella pustulata n. sp. (Figures 24, 26, Map 8)

Material Examined

One male holotype, NMC 10840, Gudal Bay, Graham Island, Queen Char-

lotte Islands, station W11, 1957.

One female allotype, NMC 10841, Gudal Bay, Graham Island, Queen

Charlotte Islands, station W11, 1957.

Thirteen male, eleven female paratypes, NMC 10843-10846, Queen Charlotte

Islands, stations H14, W9, W11, W12, 1957.

One female, NMC 10848, Prince William Sound, station A129, 1961.

One male, NMC 10849, Baranof Island, stations A171-172, 1961.

Two males, USNM Alaska King Crab Expedition 9-40, Canoe Bay, Alaska.

Nine males, four females, NMC 10850, Koeye Estuary, Fitz Hugh Sound,

B.C., station H35, 1964.

One male, two females, NMC 10842, 10847, USNM 172360, Vancouver

Island, B.C., stations P6c, 1955; O17, 1959.

Five males, six females, NMC 10851, 10852, Washington, stations W22,

W24, 1966.

One male, NMC 10853, Oregon, station W50, 1966.

Description

Male: Head and body, gnathopod 2, and pereopods covered with numerous

pustulations of various size, of which only the larger ones are indicated in the

figures. Large upward-directed single blunt spine on head, anterior to eyes;

aggregations of pustulations anteriorly and posteriorly on pereonites I to IV.

Laterally, anterior spines on pereonites IJ and III, and at base of second

gnathopods. The cephalon and pereonites II to V and gnathopod 2 setose.

Length 7.9 mm. Cephalon shorter than pereonite II, subsequent pereonites

successively shorter than preceding pereonite.

Antenna 1 approximately equal to cephalon plus pereonite II, flagellum with

ten articles; first peduncular segment with small dorsal knob distally, all

peduncular seyments setose; antenna 2 longer than peduncle of antenna 1,

flagellum with long swimming setae.

Lacinia mobilis of right mandible toothed, but not five-toothed. Other mouth-

parts typical of genus.

Gnathopod 1 propodus with two proximal grasping spines, margin of dactylus

and propodus serrate. Gnathopod 2 propodus more than twice as long as it

is broad, setose proximally and along palmar edge, pustulate distally; palm

only slightly concave, with small proximal grasping spine and distal poison

spine; dactylus heavy but smoothly tapering to distal end, inner margin serrate.

Basis attached posteriorly on pereonite II; short and stout, with lateral and

medial anteriorly directed tuberculated ridges. Ischium small, merus twice

the size of the ischium and circular, carpus very small and triangular.

Gills small and round.

Pereopods 5, 6, and 7 sturdy, increasing in size posteriorly; palmar surface of

propodus concave, with proximal grasping spines, and many spines along its

margin.

Abdomen typical of genus.

Female: Length 7.6 mm. Similar to the male except that antenna 1 is not

setose, and flagellum has nine articles; gnathopod 2 attached anteriorly on

pereonite II, propodus not elongate or setose, grasping spine arises from proxi-

mal projection with an accessory spine, and poison spine much reduced; no

lateral spines on pereonite II. Brood plates typical of Caprella. Abdomen

typical of genus.

Remarks

The specific name of this animal is descriptive of much of the surface of the

body and appendages.

Distribution

Type locality: Gudal Bay, Graham Island, Queen Charlotte Islands, B.C.

Other localities: Oregon; Washington; Vancouver Island, Koeye Estuary,

and Queen Charlotte Islands, B.C.; Baranof Island, Prince William Sound,

and Canoe Bay, Alaska.

Discussion

Caprella pustulata is very similar to C. pilipalma Dougherty and Steinberg

1953, and a comparison has been drawn with it and the holotype of C. pili-

palma:

C. pilipalma C. pustulata

Cephalic spine dorsally directed, dorsally directed,

pointed, slim blunt, stout

Gnathopod 2 poison spine absent poison spine distal

Body tuberculations low and small raised and large

Gills elliptical circular

These differences, along with others, are sufficient to separate these animals

into two distinct species.

C. pustulata is even more similar to C. scabra Holmes 1904; specimens of

this species have not been available to the author, but from the original de-

scription certain differences would appear to be present: no mention is made

of body nor, particularly, of gnathopod 2 and antenna 1 setation for C. scabra;

antenna 2 is implied to be much more slender than antenna 1; certain differences

are also apparent in the basis of gnathopod 2 and in the fact that C. scabra

has no proximal palmar spine on the propodus. Holmes mentions the variability

of the body spination in his species, so differences here cannot be taken into

account.

It can be argued that most of the differences between these two species may

be related to differences in the maturity of the animals described, Holmes’s

specimen being 19 mm long; as C. pustulata must be at least subadult, as

evidenced by the anterior lengthening of pereonite II, it will have to be considered

a distinct species, mainly on the basis of the setation, until evidence to the

contrary is forthcoming.

FIGURE 24 — Caprella pustulata. Male holotype, lateral view. Male paratype, appendages.

Female paratype, gnathopod 2. Male and female paratypes, anterior pereonites

showing major pustulations, not to scale.

Caprella striata Mayer 1903 (Figure 25, Map 8)

Synonymy

Caprella striata Mayer 1903

Material Examined

Queen Charlotte Strait: two males, one female, juveniles, station V3, 1959,

(30 fm); NMC 10981.

Queen Charlotte Sound: one male, one female, juveniles, station H52,

1964 (38 fm); NMC 10982.

Queen Charlotte Islands: over thirty individuals, stations JWS 85, 1965

(30 fm); WVV 1966 (82 fm); NMC 10983, 10984.

Bristol Bay, Alaska: over fifteen individuals, USNM Alaska King Crab

Expedition D4-41 (43 fm), D8-41 (27 fm).

Description

Male: Anterior pereonites normally smooth; posteriorly there may be one

or two tubercles on pereonites V, VI, and VII. Length 20.5 mm.

Antenna 1 approximately five-sevenths of the length of the body; flagellum

longer than peduncle and having at least twenty-seven articles. Antenna 2

equal in length to peduncle of antenna 1, flagellum with swimming setae.

Mouthparts typical of genus; lacinia mobilis of right mandible denticulate

but not five-toothed.

Gnathopod 1 with strongly serrate margin to propodus and dactylus. Gnatho-

pod 2 propodus less than twice as long as it is broad; palm with proximal

grasping spine and accessory spine, distal poison spine and triangular projection.

Basis attached just posterior to middle of pereonite II, having antero-lateral

projection.

Gills oval.

Pereopods 5 to 7 increasing in length, propodus with proximal grasping spines.

Abdomen typical of genus.

Female: Length 9.8 mm. Differs from male in having a shorter antenna 1

with at least seventeen articles in the flagellum; gnathopod 2 attached anteriorly

on pereonite II. Abdomen typical of genus.

Remarks

This is primarily a northern, deep water caprellid; depths at which it has

been found range from 7 to 150 fm.

Distribution

Type locality: not cited, Alaska.

Other localities: Aleutian Islands and northwestern Alaska.

New records: Dixon Entrance, Hecate Strait, Queen Charlotte Sound, and

Queen Charlotte Strait, B.C.

Discussion

No specimens of Mayer’s so-called var. glacialis were found in the collections

examined; this species shows considerable variation in the spination of the

body and g/acialis is probably just an extreme of this variation.

97949—7

Genus PSEUDOLIROPUS n. gen.

Antenna 2 without swimming setae, flagellum biarticulate; mandibular palp

three-segmented, setal formula for terminal article x + y + 1, molar absent (?);

outer lobe of maxilliped larger than inner lobe; gills on pereonites HI and

IV; pereopods 3 and 4 one-segmented; pereopod 5 two-segmented; abdomen of

female with one pair of setose lobes and one pair of appendages.

Type species: Pseudoliropus vanus n. sp. (by present designation).

Remarks

This genus is very similar to Liropus Mayer, 1890, from which it differs in

having a two- instead of three-segmented pereopod 5, no apparent molar

process, mandibular palp setal formula x + y + 1 instead of 1 + x + 1, and

possibly in the female abdomen. The most obvious and significant difference

is in the mandibular palp, and it is this character that will determine this genus

until more material becomes available.

The generic name is derived from the Greek term pseudo = false, in combina-

tion with Liropus, the genus it resembles.

Pseudoliropus vanus n. sp.

The single specimen, an immature female, was not dissected, but was

investigated as a whole mount. Length 5.5 mm.

Antenna 1 with seven articles to its flagellum. Antenna 2 just longer than

antenna | peduncle, with some long setae on all segments.

Mandibular palp terminal article with long distal seta, and one more medium-

length seta than short spine-like process. Maxilla 1 outer lobe with six apical

setae.

Gnathopod 1 propodus slim and triangular. Gnathopod 2 propodus stout,

palm with 2 proximal grasping spines and some spines and setae.

Pereopods 3 and 4 minute, with three and two apical setae respectively. Pereo-

pod 5 has two apparent segments; distal segment, with one apical and several

sub-apical setae, appears to be two fused segments. The pereopod formula

should probably be stated as: P3=4, P4=4, P5=24. Pereopod 6 armed with

spines along most of anterior edge; propodus palm with proximal knobs armed

with grasping spines, few palmar spines or setae.

Abdomen with anterior setae and posterior minute one-segmented appendages

armed with one apical seta.

Remarks

The specific name is derived from the Latin term vanus=unsubstantial,

referring to the nature of the material on which this species is based.

Distribution

Type locality: off Langara Island (latitude 54°03’N, longitude 134°00’W),

Queen Charlotte Islands, August 1965; 600 fm, one immature female, NMC

11150.

97949—-74

ECOLOGICAL AND ZOOGEOGRAPHICAL DISCUSSION

A wide variety of habitats suitable for caprellid amphipods is found along

the North American Pacific coast, and an analysis of certain ecological factors

was made to determine whether these factors have any effect on the distribution

of the caprellids within this region. The significance of this analysis is limited

by the small numbers of locality records, along with the seasonally restricted

range and non-specific nature of the collections here under consideration.

However, despite the limitations of the material, it has been possible to reach

the following tentative conclusions on the levels of temperature and salinity

tolerance of those species that were found at twelve or more localities.

It was found that whereas the major limiting factor in the distribution of

the caprellid species under review was temperature, variations in salinity

also had a significant effect.

Most of the species occurred almost exclusively in the 10°-15°C temperature

range. However, within this range it was noted that of the species discussed in

this paper, Tritella laevis, T. pilimana, Caprella angusta, C. pustulata, and

C. rudiuscula were found at the lower temperatures; Perotripus brevis and

Caprella californica were found at the higher temperatures; and Metacaprella

kennerlyi and Caprella laeviuscula were noticeably more eurythermic than the

other species.

The effect of salinity is more difficult to determine. In certain areas (e.g.,

northern Prince William Sound) the effect of low salinity is probably masked

by the effect of low temperature. In other areas, the salinity recorded may

not be the normal value for the locality. Thus the salinity recorded near an

estuary at low tide will be lower than is normal for the locality. Yearly fluctua-

tions also occur, in regions of large spring runoff (e.g., Strait of Georgia) and

glacial melt (e.g., northern Prince William Sound). However, it was found that

caprellids were more frequent along the open coast than along the protected

coast, which is attributable at least in part to the effect of salinity fluctuations.

For example, in the Queen Charlotte Islands region, caprellids were found at

49 per cent of the outer coast stations, but only at 38 per cent of the inner

coast stations. In the brackish Strait of Georgia, noticeably fewer species and

individuals were found than in adjacent more saline areas.

As the majority of the collections were made in the intertidal and shallow

water zones of shores and estuaries, it is probable that the salinity values

recorded are lower than is normal for most localities. In view of this it was

decided that the generally accepted value of 30 parts per thousand was not a

realistic figure to signify the difference between marine and brackish water,

and the lower value of 28 parts per thousand was chosen. (For more detailed

discussion of the biological boundary between polyhaline and marine waters,

see Dahl 1948.) Using this value it was found that Tritella laevis and Caprella

angusta were strongly stenohaline species, and that Perotripus brevis, Tritella

pilimana, Caprella laeviuscula, and C. rudiuscula were noticeably more eury-

haline than the other species (see Table 1).

FiGurE 25 — Caprella striata. Male lateral view and appendages. Female gnathopod 2.

FIGURE 26 — Abdomina of the various genera.

1. Perotripus brevis, 2. Tritella laevis, 3. Deutella californica, 4. Mayerella

banksia: A. ventral view; B. lateral view, 5. Metacaprella anomala, 6. Caprella

pustulata.

The results of this analysis of the effects of salinity were reflected in the

distribution of the species. Thus Trite/la laevis was found most frequently at

Pacific coast stations, while Perotripus brevis was found to be typical of the

inner passages and bays.

These results refer to the habitat preference of the intertidal species. Con-

sideration must also be given to those species that were found pelagically or

sublittorally. Apparently the three species (Caprella incisa, C. laeviuscula, and

Metacaprella kennerlyi) that were found attached to floating debris were not

in their normal habitat, for the collection data indicate that they were being

carried along with surface detritus. Of the species found sublittorally the

majority were primarily intertidal species (see Table 1). Four species were

collected only or mainly at deep-water stations. Caprella gracilior (Mayer 1903;

Dougherty and Steinberg 1953) and C. striata (Mayer 1903) are known to be

Table 1—ECOLOGICAL OBSERVATIONS

Temperature and

Vertical salinity tolerance Location

distribution (expressed as ratio)

S 3 3 |Temperature °C Coast type

53 . ee a ee es ey (expressed

= i elo tie} S pS <28< J|open: protected

Zh eee | bay aga sey, oe

Caprella borealis 8 Gala. daal ~ _ —

C. drepanochir 3 p) - - -

C. alaskana 18 15 2 ae | 1:4 Del

C. irregularis 20 14;2)|4 ale ee 1+-:2 PZ

C. striata 4 4 - - ~

Metacaprella anomala 3 2 1 - - -

Mayerella banksia 3) bo hes - - -

Cercops compactus 1 1 _ = -

Caprella ferrea ‘| ch - - -

C. pilidigita 4 3 1 ~ - —

C. pustulata 13 13 2 Ba, Jl 1:7 ie |

C. rudiuscula Le) 13 3: 10:..0 2:3 ya |

Perotripus brevis 13 jG AP a | i | 1 oF 0 203 iba)

Tritella laevis 19 19 1 8? 0 Pe7 4:1

T. pilimana 18 Pe ae Oy ah, G4 we ia | j Se

Caprella angusta iy 16 1 Pes) al 0:10 201

C. californica 21 16 | 5 Opn 1322 2 ea |

C. equilibra 9 9 - - -

C. gracilior 5 5 - — -

C. incisa 11 1 9 1 = - -

C. laeviuscula 89 Isa 7 1 S205 23 iz

C. mendax 4 1 Ge es - - -

C. verrucosa 8 fae | - - ~

Metacaprella kennerlyi | 28 jee ba | les 9 me aa 1—:2 ae |

Deutella californica 6 6 - - =

deep-water species; from the present records it appears that Caprella mendax

and Mayerella banksia are also predominantly sublittoral animals.

The major portion of the North American Pacific is included in the cold

temperate (Sverdrup et al. 1942) or boreal (Ekman 1953; Schenk and Keen 1936)

region. The southern part (1.e., latitudes 30°N-45°N) or ‘low boreal’ region

has not been included in detail in this paper. To the north of the boreal region

the subarctic region has also been omitted.

The general boreal region has been divided into zones on the basis of summer

surface temperatures. These zones have been further subdivided into exposed

and protected coasts. It is felt that such a breakdown is justified by the findings

already presented. It should, however, be noted that exposed and protected

coasts differ not only in salinity, but also in surf action, available habitats,

etc. (cf., Bousfield 1958). On the basis of the caprellid fauna and published

hydrographical information (e.g., Bousfield and McAllister 1962), the American

Pacific boreal region may be subdivided as follows (see Table 2, p. 83).

The subarctic zone includes three protected coast areas in which both tem-

perature and salinity are affected by local glaciers, viz: northern Prince William

Sound, the glacier coast of southeastern Alaska, and the inner coast of Alexander

Archipelago. Here summer temperatures range from 5-10°C, and surface

salinities range from 10-30 parts per thousand. This zone appears to have no

endemic species but is characterized by Caprella borealis and C. depranochir,

two subarctic species that have been able to reach this region despite the

intervening cold—temperate zone of Prince William Sound and the south coast

of Alaska.

The cold-temperate zone includes both open and protected coastal regions.

Here summer temperatures range from 10—-15°C.

The exposed cold—temperate zone includes the Pacific coasts of Alexander

Archipelago, the Queen Charlotte Islands, Vancouver Island, Washington

and Oregon, and also California south to Point Conception. Here surface

salinities are normally 30 parts per thousand or more (i.e., fully marine).

The protected cold-temperate zone includes southern Prince William Sound

and inlets of the southern Alaska coast to Seward, the mainland and inland

coasts of Hecate Strait south to Johnstone Passage, and the outer reaches

of Puget Sound. Here surface salinities are 20-30 parts per thousand and

usually 25-30 parts per thousand.

All the species described in this paper were found in the cold—temperate

zone. Moreover, the known ranges of the new species Cercops compactus,

Mayerella banksia, Caprella pilidigita, C. pustulata, and C. rudiuscula, and

also of C. ferrea and possibly Deutella californica are limited to this zone.

Perotripus brevis and possibly Caprella pilidigita appear to be protected coast

species, while Tritella laevis and Caprella pustulata, and possibly C. ferrea,

C. incisa, and C. verrucosa, appear to be exposed coast species.

The temperate zone includes three enclosed regions: the Strait of Georgia,

the inner reaches of Puget Sound, and inner San Francisco Bay. Due to the

semi land-locked nature and exposure to river discharge of these estuarine

environments, summer temperatures are higher (15—20° C.) and surface salinities

are lower (16—29°/ ) than on adjacent open coasts. Only four species have

been recorded from this zone, namely Caprella californica, C. equilibra, C.

laeviuscula, and Metacaprella kennerlyi. These species are eurytopic and are

found throughout the cold—temperate zone also. It is believed that the paucity

of caprellids in the temperate zone is more the result of lower salinities than

of higher temperatures.

Table 2—GEOGRAPHICAL DISTRIBUTION

1. Number indicates number of stations at which the species was found.

2. X_ indicates reference in literature.

3. 4/ indicates inclusion in check-list for the San Juan area.

Subarctic

glacial Cold-Temperate (Boreal) Temperate

5-10°C 10-15° C 15-20° C

10-30% 30+% 20-30% 15-30% |30+%

Protected Protected Protected | Open

coast Open coast coast coast coast

e : elsl6 2 = ee

me cea ae eee Ome = a) es | 33

g g(4/2/s/2/C] = |a| ce] 8/48] Es le] lel,| se

hires lee Le le eS les | Stee ee! eae] is oS el fe

S/2/E/4/8/ 8) 0/8] 5 | 2/88/23] oS] oe] 3s] s/ 8/8] se

BL ele] lS /SlEls| 2 Sl SS (2) SE eS) eC) B/E 3S

o |] = Seiegle]/olsilelwZ 3) ° Ons m OC) |" Oleg

Al4i8|eididizl4| elelGel el eelegqiz|8izgi ge! we

2|elelélslslels/=£/|§|so/2|22|28| 2/912) 3| 23

Bole le fais be | O ee) ore Se a pe Om Le | Slee les

aelelel2isials|s| 2 lel Seisl ss) sulelelela] os

a (= P= a = = Sa me eg aS a Ws =

SS VS HS SHS est] Gl SS Ree reper NY pee ey SN apts ee eee

S|Z2/2/0 1S SS |e OO tan) OO he la |e Pad

No. of Caprellid

Stations 3 | 3} 41]10/19] 17] 16 | 12 24| 35 14:14)]71] 3

Caprella borealis x 1 4 2) 1

C. drepanochir <i & 5

C. alaskana x By || oy le! 3 2 3 1

C. irregularis Sea OK 13 / GHG 1

C. striata x 1 3

Metacaprella anomala\| X eh x 1

Mayerella banksia 1 4

Cercops compactus 1

Caprella ferrea 1| 4] 2 x

C. pilidigita 3 1

C. pustulata 1); 4) 3 3 1 1

C. rudiuscula 2B 3} 4] 1 3

Perotripus brevis 1 At ae Fi49 x

Tritella laevis 4] 6 Seine x 2 2 x

T. pilimana Smet 1/5 x 2 3 3 x

Caprella angusta Dal eS Dates x 2 3 x

C. californica x DAO ane ol OX 7 1 xX x

C. equilibra >< 5 1 x 3 x x

C. gracilior x 2 1 a/ SC at 1 x

C. incisa 1 1 2 2) 4 x< 1 x

C. laeviuscula >< ik DS CUP Sioa yl) 2 ye 12 4 of x |15 | 19 9 3/81] 4

C. mendax “ x 3 1 x

C. verrucosa x 2|4 1 < 1 x

Metacaprella kennerlyi 1 147 sy || 22 x 1 2, 2 1S be sl ><

Deutella californica 2 iL |} PAs) oS 1 x

foe)

Study of the literature on the caprellids of the North American Pacific

shows that forty species, including the six species newly recorded here, have

been recorded from the region from Point Conception north to northwestern

Alaska. Twenty-seven of these species are apparently endemic to this region.

Of the remainder, eleven species are found also in the northwestern Pacific.

Six of these species are amphi-Pacific, of which three (Caprella borealis, C.

drepanochir, and C. paulina Mayer) are also subarctic, and two (C. irregularis

and C. laeviuscula) are cold-temperate; C. gracilior is found throughout the

northern and eastern parts of this region. Records indicate that Metacaprella

anomala may also be an amphi-Pacific species. Caprella brevirostris Mayer,

C. californica, C. equilibra, and C. verrucosa have not been recorded north

of the Queen Charlotte Islands and therefore have discontinuous pan-Pacific

distributions. The two remaining species are found sublittorally: Caprella

ciliata Sars (recorded by Holmes 1904) is found also in the North Sea (Sars

1895; Mayer 1890, 1903), and C. ungulina Mayer is found also off Tierra del

Fuego and the Galapagos Islands (Mayer 1903).

Caprella is the dominant genus in the north Pacific and is represented by

twenty-nine species in American waters. The remaining eleven species come

from seven different genera, four of which (Perotripus, Metacaprella, Tritella,

and Pseudoliropus) are endemic to the Pacific.

Conditions along the North American Pacific coast lead to a diversity of

habitats, and even non-specialized collections contain many species of caprel-

lids. Ekman’s (1953) general postulation that the North Pacific is faunistically

rich is supported by these findings, and also by those of workers in Japanese

waters. Bulycheva (1957) has suggested that the American fauna is poorer

because of differences in climate. It is suggested here that this is too great a

simplification of the differences. The Japanese seas do have a greater range

of temperatures, both seasonally and geographically, than the American

Pacific at equivalent latitudes. However, the western Pacific also has a more

varied coastline, and a broader continental shelf, which apparently gives rise

to a greater variety of intertidal and marine habitats. Thus both climatic and

geographical conditions can be expected to cause differences in the fauna

of the two regions of the North Pacific (See also Gislén 1943-44). However,

it must be emphasized that the American fauna has been far less intensively

studied than the fauna of Japan. It is expected that more specific collecting

of caprellids, particularly along the open coast from Cross Sound to Point

Conception, will lead to the discovery that this region supports a fauna of

comparable diversity to that of Japan.

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