Serials

The Caprellidae

(Crustacea, Amphipoda)

of Atlantic and Arctic Canada

The Caprellidae (Crustacea, Amphipoda)

of Atlantic and Arctic Canada

National Museum of Natural Sciences

Publications in Biological

Oceanography, No. 4

Published by the

National Museums of Canada

Staff editor

Rigmore Adamson

Musée national des Sciences naturelles

Publications d’océanographie

biologique, n° 4

Publié par les

Musées nationaux du Canada

The Caprellidae (Crustacea, Amphipoda)

of Atlantic and Arctic Canada

Diana R. Laubitz

Available by mail

from the

National Museums of Canada

Marketing Services

Ottawa, Ontario

K1A OM8

Catalogue No. NM95-7/5

National Museum of Natural Sciences

National Museums of Canada

Ottawa, Canada

1972

PO0987654321

Y798765432

Litho in Canada

Contents

List of Plates, 6

List of Maps, 7

Résumé, 8

Summary, 9

Biographical Note, 10

Preface, 11

Acknowledgements, 12

Notes on the Family Caprellidae, 13

Methodology, 13

Key to the Caprellidae of Atlantic and Arctic Canada, 17

Aeginella, 19 C. septentrionalis, 48

A. spinosa, 19 C. trispinis, 53

Aeginina, 21 C. unica, 53

A. aenigmatica, 21 Cercops, 57

A. longicornis, 23 C. holbolli, 57

Caprella, 26 Hemiaegina, 59

C. andreae, 26 H. minuta, 59

C. carina, 26 Luconacia, 61

C. ciliata, 31 L. incerta, 61

C. dubia, 31 Mayerella, 61

C. equilibra, 34 M. limicola, 61

C. linearis, 35 Metacaprella, 63

C. microtuberculata, 39 M. horrida, 63

C. penantis, 41 Paracaprella, 67

C. natalensis, 47 P. tenuis, 67

C. neglecta, 47 Proaeginina, 68

C. rinki, 47 P. norvegica, 69

C. sarsi, 48

Zoogeography, 71

Table 1: Known Distribution of Canadian Arctic and Atlantic

Caprellidae, 70

Canadian Atlantic Region, 71

Canadian Arctic Region, 72

Discussion, 75

Literature Cited, 77

List of Plates

OMAN OORWNH —

10

Aeginella spinosa, 20

Aeginina aenigmatica, 22

Aeginina longicornis, 24

Caprella carina, 28

Caprella ciliata, 30

Caprella dubia, 32

Caprella linearis, 36

Caprella microtuberculata, 38

Caprella penantis, Caprella natalensis,

Caprella neglecta, 42

Caprella penantis, Caprella natalensis,

Caprella neglecta, 44

Caprella rinki, 46

Caprella septentrionalis, 50

Caprella unica, 52

Cercops holbolli, 56

Hemiaegina minuta, 58

Mayerella limicola, Metacaprella horrida, 62

Metacaprella horrida, 64

Paracaprella tenuis, 66

List of Maps

10

Known distribution of Aeginella spinosa and Aeginina

aenigmatica north of 45°N, 16

Distribution of Paracaprella tenuis and Aeginina longi-

cornis within the Canadian Atlantic region; based on

material examined, 26

Known distribution of Aeginina longicornis north of

45°N, 27

Known distribution of Capre/la carina, Caprella dubia,

and Metacaprella horrida, 34

Known distribution of Caprel/la ciliata, Caprella rinki, and

Proaeginina norvegica, 35

Distribution of Caprella linearis and Caprella penantis

within the Canadian Atlantic region; based on material

examined, 40

Known distribution of Capre/la linearis within the North

Atlantic and northern Pacific boreal and_ subarctic

regions, 41

Known distribution of Capre/lla microtuberculata and

Cercops holbolli within the North Atlantic and arctic

regions, 49

Distribution of Capre/la septentrionalis and Caprella unica

within the Canadian Atlantic region; based on material

examined, 54

Known distribution of Capre/lla septentrionalis within

the North Atlantic boreal and arctic regions, 55

Résumé

L’auteur fait état de 21 especes de caprelles provenant des

eaux arctiques et atlantiques du Canada. Les observations

couvrent la partie septentrionale de l’Amérique du Nord

depuis la Nouvelle-Ecosse jusqu’en Alaska, ainsi que l’est et

l'ouest du Groenland. On trouve dans l’ouvrage la des-

cription détaillée de 17 especes et une clé servant a leur

identification. L’une d’elles, Aeginina aenigmatica, semble

étre nouvelle. On a placé dans le genre Metacaprella,

Caprella horrida G.O. Sars. On considére Caprella sedovi

Gurjanova synonyme de C. carina Mayer. Quant a Caprella

hystrix Bate et Westwood et a C. ultima Bate, on les a

respectivement assimilées a C. linearis (Linné) et a

C. septentrionalis Kr@yer. On considére non identifiables

Caprella sarsi Honeyman et C. trispinis Honeyman. Caprella

natalensis Mayer et C. neglecta Mayer acquiérent une

identite specifique.

On observe que les mers boréales se divisent en deux

grandes régions zoogéographiques, |’une arctique-atlantique

et l'autre subarctique-pacifique. La faune arctique est en

grande partie d’origine atlantique alors que la plupart des

especes venant du Pacifique ont des répartitions tres

locales.

Summary

Twenty-one species of caprellids are reported from the

Canadian Arctic and Atlantic. The area investigated includes

northern North America from Nova Scotia to Point Barrow,

Alaska, and eastern and western Greenland. Seventeen

species are described in detail and a key to their identification

is included. One species, Aeginina aenigmatica, is described

as new. Caprella horrida G.O. Sars has been placed in genus

Metacaprella. Caprella sedovi Gurjanova has been synon-

ymized with C. carina Mayer. Caprella hystrix Bate and

Westwood and C. ultima Bate have been assigned to

C. linearis (Linnaeus) and C. septentrionalis Krgéyer respec-

tively. Two species, Capre/la sarsi Honeyman and C. trispinis

Honeyman, are reported as_ unrecognizable. Caprella

natalensis Mayer and C. neglecta Mayer have been given

specific status.

The north-polar seas are found to comprise two major

zoogeographical regions, Atlantic-arctic and _ Pacific-

subarctic. The arctic fauna is predominantly Atlantic in

Origin, and most Pacific species have very localized

distributions.

Biographical Note

Born in London, England, Diana Laubitz graduated from

Cambridge University with a B.A., specializing in Zoology. She

came to Canada in 1956, and has been associated with the

National Museum of Natural Sciences since 1964. Her

research has been concerned primarily with the taxonomy and

systematics of the Caprellidea, and in 1970 she published

Studies on the Caprellidae (Crustacea, Amphipoda) of the

American North Pacific, in the Museum's Biological Oceano-

graphy series. Recently she has started investigating some of

the Gammaridea closely related to this group.

Preface

The Caprellidae of the western North Atlantic have recently

been treated by McCain (1968). He discussed the complicated

taxonomy of this family, resolved certain systematic problems

and listed 29 species, describing 24 in detail. The area under

consideration in his paper was the western North Atlantic

from the equator to Nova Scotia.

Since Mayer’s three monographic papers (1882, 1890,

1903), caprellids have been reported only incidentally from

North America north of Nova Scotia. Faunal checklists and

ecological surveys (e.g. Bousfield 1952-62; Bousfield and

Leim 1960; Brunel 19615, 1970; Dunbar 1942, 1954) have

included caprellids found in certain areas, but no work has

dealt exclusively with these animals. In contrast, the arctic

region around Greenland and the northeastern Atlantic have

been investigated fairly thoroughly. Most of the recent Arctic

reports and new species are found in Stephensen (1912-44a).

The present paper deals with the Caprellidae of the boreal

and arctic regions of Atlantic North America. The geographical

area investigated includes North America, from Nova Scotia

northward and westward to the Canadian arctic islands and

northern Alaska, and eastern and western Greenland. Seven-

teen species are described, one of which is new to science.

Four other species are listed but not described, since they

were not present in the collections investigated. One of these

is a cold-water species (Proaeginina norvegica). Three have

. not yet been reported from the area investigated, but have

been found as far north as Connecticut (Capre/l/a andreae,

C. equilibra, and Luconacia incerta).

Acknowledgements

The major collections studied have been those of the National

Museum of Natural Sciences, Ottawa. Other unidentified

material was obtained from _ Universitetets Zoologiske

Museum, Copenhagen; Station de Biologie marine, Grande-

Riviere, Quebec; United States National Museum, Washington,

D.C.; Fisheries Research Board of Canada, Arctic Unit, Ste-

Anne-de-Bellevue, Quebec.

The large amount of borrowed material has considerably

increased the scope and value of this paper. | must offer my

sincere thanks for their contributions to: Dr. Torben Wolff

(Universitetets Zoologiske Museum, Copenhagen); Dr. A.

Marcotte and Dr. M. Ledoyer (Station de Biologie marine,

Grande-Riviere); Dr. Thomas Bowman (U.S.National Museum,

Washington, D.C.); Dr. J.W. Wacasey (Fisheries Research

Board of Canada). | am very grateful to Dr. Charlotte Holmquist

(Naturhistoriska Riksmuseet, Stockholm) and Dr. Marit

Christiansen (Zoologisk Museum, Oslo) for the loan of type

material; and to Dr. J. Forest (Muséum National d'Histoire

Naturelle, Paris) and Mr. R. Ingle (British Museum of Natural

History) for some of C. Spence Bate’s original material.

| also thank the National Museum of Natural Sciences for

supporting this study, and particularly Dr. E.L. Bousfield for his

advice and critical help, and for the use of his unpublished

collecting station data. | am especially grateful to Dr. J.C.

McCain (Hazleton Laboratories, Falls Church, Virginia) for his

advice and comments, and for the use of his manuscript

catalogue and bibliography of the Caprellidae.

12

Notes on the Family Caprellidae

McCain (1968) and Laubitz (1970) have both commented on

the various characters used in diagnosing caprellid genera.

They have emphasized that certain characters need further

investigation, and have pointed out characters on which the

taxonomic division of the Caprellidae should eventually be

based.

Vassilenko (1968) divided the family Caprellidae into four

subfamilies. These subfamilies, based extensively on super-

ficial characters (e.g. pereopods 3 and 4, number of gills),

are here considered to be unacceptable.

McCain (1970) revised Vassilenko’s subfamilies on the

basis of mandibular type, determined by the presence or

absence of the molar process and the palp. The presence or

absence of the mandibular molar is certainly more basic than

the characters used by Vassilenko. However, other mouth-

parts exhibit characters that may be of a more fundamental

nature than the presence or absence of the mandibular palp.

Thus, the number of spines on maxilla 1 outer plate is six in

some genera, seven in others; also, the outer plate of the

maxilliped sometimes bears a row of spines along its inner

edge. These characters are here considered of higher systema-

tic importance than the mandibular palp.

No further decision on the classification of the Caprellidae

can be reached until the nature of the mouthparts of as many

genera as possible can be determined. Because of the un-

satisfactory nature of the systems so far presented, the

caprellids under review here have not been placed in sub-

familes. If, however, one uses McCain's familial taxa, the

genera in this paper can be assigned to families and sub-

families as follows:

Caprogammaridae Kudrjaschov and Vassilenko 1966, emend

McCain 1970: Cercops.

Aeginellidae Vassilenko 1968, emend McCain 1970:

Aeginellinae Vassilenko 1968, emend McCain 1970: Aeginel/a,

Aeginina, Proaeginina;

Protellinae McCain 1970: Luconacia, Mayerella.

Caprellidae White 1847, emend McCain 1970: Caprell/a,

Hemiaegina, Paracaprella.

Methodology

A special terminology has evolved around the Caprellidae, and

note must be made of certain usages.

The swimming setae on antenna 2 are the many long, paired

hairs set along the lower edge of the peduncle and, frequently,

the flagellum of this antenna; these hairs are actually used in

feeding rather than in swimming.

Notes on the Family Caprellidae

The inner and outer plates of the maxillae and the maxilliped

have frequently been referred to as lobes. | have used the term

plate, which is the standard amphipod terminology (e.g. J.L.

Barnard 1969: 522).

The pereopods and gnathopods are numbered according to

the pereonite from which they arise. Because it was originally

believed that the coxal segments were absent, the appendages

have long been treated as six-segmented, the basis being

segment one and the dactylus being segment six. This

numbering is used here, despite the fact that coxae are found

in caprellids, particularly on the gnathopods and pereopods 5

to 7. A decision concerning a reversion to normal amphipod

terminology will be made when all the caprellid genera have

been reviewed.

All figures are drawn to scale, using a microprojector;

unless otherwise stated, all scales shown equal 1 mm, and

refer to the whole-mount only. The male and female whole-

mount are to the same scale unless otherwise indicated.

Measurements were made laterally along the midline of

each pereonite, from the head, between the insertions of

antennae 1 and 2, to the tip of the abdomen.

All characters used in the Key refer to both males and

females, unless specifically stated otherwise.

Those species that were described in detail by McCain

(1968) have been treated only generally; for details refer to

the above paper.

The present location of the specimens examined has been

indicated by the following abbreviations:

BM British Museum of Natural History, London

FRBC Fisheries Research Board of Canada, Arctic Unit,

Ste-Anne-de-Bellevue, Quebec

GR Station de Biologie marine, Grande-Riviere, Quebec

MN Muséum National d‘Histoire Naturelle, Paris

NMC National Museum of Natural Sciences, National

Museums of Canada, Ottawa

NR Naturhistoriska Riksmuseet, Stockholm

USNM United States National Museum, Washington, D.C.

UZM __— Universitetets Zoologiske Museum, Copenhagen

ZM Zoologisk Museum, Oslo

Detailed collection data have been given only for the less

common species. The predominantly intertidal and shallow-

water collections from eastern Canada have been summarized

by region, and by province within each region. Where used,

station numbers refer to published station data (Bousfield

19566-1962). Other major National Museums of Canada

collections, for which data is unpublished, came from the St.

Lawrence estuary, 1953 (NMC 53-52); the Gulf of St.

Lawrence, 1960 (NMC 60-180); eastern Nova Scotia, 1962

(NMC 62-116); western Nova Scotia, 1963 (NMC 63-293);

14

Methodology

the St. Lawrence estuary, north shore, 1969 (NMC 69-211).

Also deposited in NMC is a collection, gathered mainly in the

Bay of Fundy region, 1911-1916, from the Atlantic Biological

Station, St. Andrews, New Brunswick (NMC 51-15).

Information for Maps 1, 3-5, 7, 8, and 10 is based on

literature records, and on the material newly examined here.

Information for Maps 2, 6, and 9 is based mainly on the

published and unpublished data of Dr. E.L. Bousfield.

15

Map 1

-

f\

@ Aeginella spinosa

v Aeginina aenigmatica

Known distribution of Aeginel/la spinosa and Aeginina aenigmatica north of

45°N.

16

Key to the Caprellidae of Atlantic and Arctic Canada

Abdomen __ five-segmented; mandibular molar

eM ae oe cin aia pu ecidpraen nsesle days Wane Wests COMUNRT

Abdomen unsegmented; mandibular molar

WN PRIIN 622i arcide cs semaine Warde cent tye dticarad<tch sniend Cadaneieeis

Mandibular palp present, sometimes’ greatly

NSC OE ta iis LBL ld RES ae ON A

PT ATHNOULAT PEND BOSON oocenciveccdnnsrnnsnnavacindaracistedaasdve vies

Pereopods 3 and 4 present; maxilliped outer plate

without spines along inner edge .................cccceeceeeeeeees

Pereopods 3 and 4 absent; maxilliped outer plate

with spines along inner Edge ..............:cccsecceeeeeeeeeeeeees

Pereopod 5 two-segmented; maxilla 1 outer plate

Wir Seven spines: clay.) hek. U6 ile ele. ee a

Pereopod 5 six-segmented; maxilla 1 outer plate

RA SI SNES ocak cdves susinwhsste cae nvnlns copniaes totem eects ate aeen

Abdomen of male and female with one pair of ap-

pendages and one pair of lobes; spines on head and

SET ON FS 1 CWS GINC «iru isc cas cou vaesvicceweans sakeenes suesuatonweensio

Abdomen of male and female with two pairs of

appendages and one pair of lobes; spines on head

ANG PETEONite 1 PAITET «0.000. 4.c0secceseedndeareenasersceusestonroene

Flagellum of antenna 1 longer than peduncle;

pereopods 5 to 7 having segments 1, 3, and 5 sub-

equal, and longer than segment 4 ................c.:0eceeeeeeees

Flagellum of antenna 1 shorter than peduncle;

pereopods 5 to 7 having segments 1, 3, and 4 sub-

equal, and shorter than segment 5 ..........eeeeeceeesseeeees

Pereopods 3 and 4 one-segmented; swimming

BCtAS GUS OI oui. <iiincnmnareuctnniecaiadoicenty ee eed meee. See

Pereopods 3 and 4 absent; swimming setae

2 EET RG ER EERE 2 ca ati emery eee Sa eS

Cercops holbolli, 57

5

Mayerella

limicola, 61

Paracaprella

tenuis, 67

Aeginella

spinosa, 19

Aeginina

aenigmatica, 21

Aeginina

longicornis, 23

Hemiaegina

minuta, 59

1%

Key to the Caprellidae

8 Pereopods 5 to 7 without grasping spines; com-

10

11

2

13

14

18

MIGMSANVWITMIAISTETIAS SDD... icvieweceusncwsencresesesvarceudssaeseues

Pereopods 5 to 7 with grasping spines; free-living ....

Head prolonged anteriorly into a large, triangular

projection; no body spines; adult male, gnathopod 2

propodus palm with only two projections, the distal

OMe WeWIG TCO LOIIQU AD fogs <A cca scccappbeadd achounenveedbacwuntang

Head spines or tubercles discrete, if present; adult

male, gnathopod 2 propodus palm with at least

three projections, the distal one being triangular .......

Gnathopod 2 basis with anterior lateral and medial

distal spines; head and body without dorsal spines,

but with many Sinall TUDercleS «0.5.03 icccsccectescenncpercevcane

Gnathopod 2 basis with anterior lateral distal spine

only; head and body with dorsal spines, or without

PUN CISG Ore ak Tes Joke ee RNEEE ONE nd:

Antenna 1 flagellum longer than peduncle ................

Antenna 1 flagellum shorter than peduncle ...............

Anterior body ornamentation always pointed spines;

abdomen very small, barely protruding between

attachments of pereopods 7idiince. . dee a a

Anterior body ornamentation always rounded

tubercles; abdomen quite large, with median

CHRINGUS MUNID: .el es ke Se. honk QU

Pereonites 5, 6, and 7 without spines or tubercles;

in male, gnathopod 2 basis setose; in female,

pereonite 5 longer than pereonite 2 .................:cccceeee

Pereonites 5, 6, and 7 with spines or tubercles; in

male, gnathopod 2 basis not setose; in female,

pereonite 5 shorter than pereonite 2 .................::0cc0ee

Head spines or tubercles always unpaired .................

Head spines or tubercles always paired .................06605

Caprella unica, 53

9

Caprella

penantis, 41

10

Caprella

carina, 26

Ht

12

Caprella dubia, 31

Caprella micro-

tuberculata, 39

Caprella ciliata, 31

Aeginella spinosa

15

Pereonites 2, 3, and 4 with paired dorsal spines;

body strongly tuberculate; eyes very large; antenna

2 swimming setae short and Sparse ..................:000000e

Pereonites 2, 3, and 4 with unpaired dorsal spines;

body rarely tuberculate; eyes normal; antenna 2

SPITTING SETAS TOIT eile tean nee ecole tales ened

Head and anterior pereonites with few small paired

dorsal spines, frequently smooth; in male, gnathopod

2 propodus not strongly setose; in female, abdomen

with one pair of lobes, no appendages ...................6605

Head and anterior pereonites always with many

large paired dorsal and lateral spines; in male,

gnathopod 2 propodus strongly setose; in female,

abdomen with one pair of lobes and one pair of

Caprella rinki, 47

Caprella

septentrionalis, 48

Caprella

linearis, 35

Metacaprella

STN CS ie neces cen covaie nat con snes

horrida, 63

GenusAEG/NELLA Boeck 1861

Antenna 2 without swimming setae, flagellum with two articles; mandible

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

outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4;

pereopods 3 and 4 absent, pereopod 5 six-segmented; abdomen of male and

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

Type species

Aeginella spinosa Boeck 1861 (by monotypy, subsequently designated by

McCain 1968).

Aeginella spinosa Boeck 1861

Plate 1; Map 1

Synonymy

Refer to McCain 1968.

Material examined

Two females, from western Green-

land: Ingolf Station 35, 65°16'N,

55°05'W, depth 682m _ (UZM);

Ingolf Station 31, 66°35'N, 55°54'’W,

depth 166 m(UZM).

Description

Body spination of males and females

characteristic and almost constant.

Length of largest female 9.9 mm.

Antenna 1 longer than head to

pereonite 3. Antenna 2 shorter than

peduncle antenna 1.

Mouthparts similar

and Caprella.

Gnathopod 1 propodus triangular,

dactylus with serrate grasping margin.

Gnathopod 2 similar in male and

female, propodus heavy, with antero-

distal projection; palm with proximal

projection bearing a small spine, and

distally, one small and one large

projection.

Gills elliptical.

Pereopods 5 to 7 of increasing size,

to Aeginina

Plate 1

Aeginella spinosa, female

A Abdomen

B Lateral view

C Mandibular palp

D Distal segments of pereopod 6

20

Aeginina aenigmatica

anterior margin of segments 3 and 4

setose, propodus with pair of prox-

imal grasping spines, and palmar

spines.

Other localities: Norway; Spitsbergen;

Faeroe Islands; Iceland; eastern and

western Greenland; off Nova Scotia;

Banquereau Banks; off Cape Cod.

Abdomen typical of genus.

Remarks

This is an_ arctic-boreal species,

usually found in deep water, down to

1026 m.

Distribution

Type locality: Haugesund, Norway.

GenusAEGI/NINA Norman 1905

Antenna 2 without swimming setae, flagellum with two articles; mandible

with triarticulate palp, setal formula for terminal article 1+x+1, molar

present; outer plate of maxilliped larger than inner plate; gills on pereonites 3

and 4; pereopods 3 and 4 absent, pereopod 5 six-segmented; abdomen of

male and female with one pair of biarticulate and one pair of unsegmented

limbs and one pair of lobes.

Type species

Aegina longicornis Kréyer 1842-43 (by monotypy, subsequently designated

by McCain 1968).

Aeginina aenigmatica sp. n.

Plate 2; Map 1

Material examined

Seven specimens, from eastern

Greenland (UZM). Slide mount

deposited in UZM.

Description

Male holotype (immature): Body

spinose, head with one pair of dorsal

spines; pereonites 2 to 4 with three

pairs of dorsal spines, and lateral

spines over gnathopod and _ gills;

pereonite 5 with two pairs of dorsal

spines. Length 9.7 mm.

Antenna 1 as long as head to pere-

onite 5 inclusive; flagellum longer

than peduncle and with 21 articles.

Antenna 2 shorter than peduncle of

antenna 1.

Mandible with triarticulate palp,

setal formula for terminal article

1+3+1. Left mandible with five-

toothed incisor, five-toothed lacinia

mobilis, setal row of three setae; right

mandible with five-toothed incisor,

four-toothed lacinia mobilis, setal

row of two setae. Maxilla 1 palp with

several apical spines and setae; outer

plate with seven bifid and _ trifid

spines. Maxilliped inner plate with

five apical spines and many plumose

setae; outer plate with three plumose

apical spines, and a row of spines

along inner margin; palp similar to

that of Caprel/a.

Gnathopod 1 propodus triangular,

with proximal pair of grasping spines;

grasping margin of propodus and

dactylus slightly serrate. Gnathopod

2 with long basis with slight antero-

lateral projection distally; propodus

slender, rounded anterodistally, palm

with proximal projection bearing

single spine, and two small distal

projections; dactylus slender and

evenly curved.

21

Plate 2

Aeginina aenigmatica

Male

Male, gnathopod 2 propodus

Maxilliped

Maxilla 1

Maxilla 2

Gnathopod 1 propodus

m™moQW>D>

22

Fe aes AA

Lower lip

Mandibular palp

Male, abdomen

Female, abdomen

Right mandible

Left mandible

Aeginina longicornis

Gills elliptical.

Pereopods 5 to 7 increasing in

length; segments 1 and 3 com-

paratively long, segment 4 short,

propodus with proximal pair of

grasping spines.

Abdomen with one pair of biartic-

ulate appendages bearing many

minute knobs on their medial sur-

face; one pair of unsegmented ap-

pendages with an apical row of

denticulations and an apical seta;

and one pair of lobes. Penes lateral

and short.

Female: Essentially similar to male;

abdomen as in male, but lobes

setose.

Distribution

Type locality: Henry Land, eastern

Greenland, depth 40 m, 21 July

1900. One male holotype, 1 female

allotype, 2 mate and 3 female para-

types (UZM).

Remarks

The largest male found was 10 mm,

the largest female 9.4 mm. This

species was based on seven speci-

mens, all of which were so immature

that there was no sign of develop-

ment of the secondary sexual char-

acters. As there was some doubt

that they might be immature speci-

mens of A. /ongicornis, they were

compared with specimens of this

species at an approximately similar

developmental stage, and_ their

distinctiveness was apparent. Apart

from differences in antenna 1,

gnathopod 2, and the pereopods, the

abdomen shows slight differences;

the most obvious is that in A. aenig-

matica the penes are half as long as

the unsegmented appendages in the

immature specimens available, where-

as in A. /ongicornis they are of the

same length as the appendages in

both immature and adult specimens.

Aeginina longicornis (Kr¢yer 1842-43)

Plate 3; Maps 2, 3

Synonymy

Refer to McCain 1968. Also:

[?] Caprella scolopendroides (non

Lamarck) — Ross 1826

Aegina spinifera — Rodger 1894

Aegina spinosissima — G.O. Sars

1909

Aeginina longicornis — Vibe 1950 —

Steele 1961 — Brunel 1970

Material examined

Approximately 3,000 specimens.

Hudson River estuary, off New

Jersey, 12 specimens (NMC).

Atlantic Canada: Bay of Fundy, 600

specimens (NMC); Nova Scotia, 100

specimens (NMC); off Newfoundland,

18 specimens (NMC).

Gulf of St. Lawrence: Nova Scotia,

3 specimens (NMC); Prince Edward

Island, 7O specimens (NMC); New

Brunswick, 12 specimens (NMC);

Gaspé, 2,100 specimens (GR, NMC).

St. Lawrence River estuary: north

shore, 8 specimens (NMC); Saguenay

fjord, 27 specimens (GR).

Arctic Canada, west to 106°20’,

north to 76°46’, 65 specimens

(FRBC).

Greenland, western and southern,

30 specimens (UZM).

Description

Body smooth to very spinose, one

pair of dorsal head spines and lateral

spines over gnathopod 2 insertions,

usually present. Length of largest

male 50 mm, of largest female 30

mm, of smallest ovigerous female

7.7mm.

23

Plate 3

Aeginina longicornis

Antenna 1 in adult males usually

as long as body. Antenna 2 shorter

than peduncle of antenna 1.

Mouthparts typical of genus;

setation of terminal article of man-

dibular palp shows some variation.

Gnathopod 1 propodus triangular,

grasping margin of propodus and

dactylus slightly serrate. Gnathopod

2 basis with strong anterolateral

spine; propodus slender, and with

triangular projection anterodistally;

palm with proximal — projection

bearing single spine, and with two

distal projections separated by a cleft.

In large adult males, both the distal

cleft, and the notch distal to the

grasping spine become much exag-

gerated (Plate 3, fig. E).

Gills elliptical.

Pereopods 5 to 7 of increasing

length, propodus being the longest

segment and bearing proximal

grasping spines.

Abdomen typical of genus. Penes

lateral, medially directed, and long.

Distribution

Type locality: Near

Greenland.

Frederikshaab,

Other localities: Arctic Ocean from

140°E west to Baffin Bay; southern

Norway; Denmark; Faeroe Islands;

Shetland Isles; Iceland; southern

Plate 3

Aeginina longicornis

A Female

B Mandibular palp

C Male

D- Right lacinia mobilis

E Male, gnathopod 2 propodus

F Pereopod 7 propodus

G Male, abdomen

H Female, abdomen

Greenland; Newfoundland south to

North Carolina.

New records: Saguenay fjord and

north shore of St. Lawrence estuary;

various localities among the Cana-

dian arctic islands from Baffin Bay

west to 106°20'W.

Remarks

This species shows. considerable

variation in both size and spination. It

has not been possible to ascribe

reasons for these variations, although

McCain found that in general speci-

mens from northern waters were

more spinose than those from south-

ern waters.

Two female specimens were found

to be _ parasitized by nematode

worms; in one the worm was

situated within pereonites 3 and 4,

and in the other the worm was inside

one of the brood plates.

One_ intersexual specimen was

found with developing brood plates,

female genital opening, and one

penis.

This is a common North Atlantic

and Arctic species. Although it is fre-

quently found in deeper water (to

2258 m), nearly half of the Canadian

Atlantic collections were from shal-

low (less than 10 m) water.

J-N Immature:

J Male

K Mandibular palp

L Male, abdomen

M Female, abdomen

N Male, gnathopod 2 propodus

25

Caprella andreae

Map 2 Distribution of Paracaprella tenuis and Aeginina longicornis within

the Canadian Atlantic region; based on materia! examined.

Genus CAPRELLA Lamarck 1801

Antenna 2 usually with swimming setae, flagellum with two articles; man-

dibular palp absent, molar present; outer plate of maxilliped larger than, or

equal to, inner plate; gills on pereonites 3 and 4; pereopods 3 and 4 absent,

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

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

Type species

Cancer linearis Linnaeus 1767 (designated by Dougherty and Steinberg 1953).

Caprella andreae Mayer 1890

Synonymy

Refer to McCain 1968.

Remarks

This species has not been reported

north of Cape Cod. It appears to be

pelagic in habit, being attached to

floating objects, and it is therefore

possible that it will be found in the

region under consideration.

C. andreae is generally similar to

26

C. penantis, differing most obviously

in having a convex palm, with medial

grasping spines, on the propodus of

pereopods 5 to 7.

For detailed description,

McCain 1968.

consult

Caprella carina Mayer 1903

Plate 4; Map 4

Synonymy

Caprella carina — dA. Thompson

1901

Caprella carina

Map 3

Caprella carina Mayer 1903 —

Stephensen 1933

Caprella sedovi Gurjanova 1933,

1935

Material examined

Cape Parry, N.W.T., 7O°O7'N, 124°

39°;W, 23 Aug. 1962, depth 5.5m,

3 males; 7 Sept. 1962, depth 7 m,

4 males (FRBC).

Point Barrow, Alaska: Eluitkak

Passage, Elson Lagoon, 8 June 1948,

3 females; beach at Point Barrow

Base, 28 Sept. 1950, 11 males, 10

females (USNM).

Known distribution of Aeginina longicornis north of 45°N.

Kara Sea, 70°40'N, 64°17°E, depth

20 m, female lectotype (NR).

Slide mounts deposited in NMC.

Description

Head and body without dorsal spines,

but with many large and small tuber-

cles, particularly on dorsal surface;

lateral spines present over gnathopod

2 insertions, over gills, and anteriorly

on pereonite 3. Length of largest

male 34.4 mm, of largest female

19.8 mm.

Antenna 1 equal in length to

cephalon plus pereonite 2 in male,

27

Plate 4

28

Caprella carina

longer in female; flagellum of up to

19 articles, shorter than peduncle.

Antenna 2 longer than peduncle of

antenna 1, with strong swimming

setae.

Mouthparts typical of Caprel/l/a.

Left mandible with five-toothed in-

cisor, five-toothed lacinia mobilis,

setal row of three (occasionally more)

setae, molar with single plumose

seta. Right mandible with five-

toothed incisor, lacinia mobilis

toothed but not five-toothed, setal

row of two setae, molar with single

plumose seta. Maxilla 1 outer plate

with seven complex apical spines;

palp with apical spines and many

setae. Maxilla 2 plates both setose

apically. Maxilliped inner plate con-

cave apically, with four spines and

many plumose and simple setae;

outer plate with two large apical

setae, row of spines along medial

margin, and numerous’ setae on

surface; articles of palp strongly

setose, dactylus slender and bearing

numerous comb setae.

Gnathopod 1 propodus triangular,

with one pair of proximal grasping

spines; grasping margin of propodus

and dactylus_ finely denticulate.

Gnathopod 2 basis short, with lateral

and medial anterior spines; ischium

with lateral and medial anterior

spines; propodus with slight antero-

Plate 4

Caprella carina

Female (posterior carinations not shown)

Gnathopod 1 propodus

Female lectotype, gnathopod 2

Male, gnathopod 2

Male

Maxilla 1

Maxilla 2

QOummMoOdOWD>yYD

distal projection, tuberculate anterior-

ly, palm laterally setose in male,

bearing proximal projection with

single grasping spine, accessory spine

present in female, and two triangular

distal projections; dactylus strong,

evenly tapered, in male denticulate

along grasping margin.

Gills elliptical.

Pereopods 5 to 7 of increasing

length, propodus' with proximal

grasping spines.

Abdomen of male with one pair

of unsegmented appendages, and

one pair of setose lobes separated by

a median chitinous double hump;

penes medial. Abdomen of female

with one pair of setose lobes sep-

arated by chitinous hump.

Distribution

Type locality: Kara Sea (70°40’'N,

64°17’E), depth 20 m.

Other localities: Kara Sea; Disko,

western Greenland.

New records: Cape Parry, N.W.T.;

Point Barrow, Alaska (71°22’N,

156°30'W).

Remarks

Mayer based this species on two

female specimens. The specimen

deposited in the Stockholm Museum

Maxilliped

Pereopod 6

Left mandible

Male, abdomen

Female, abdomen

Lower lip

Right mandible

On Se rixtt

29

Plate 5

Caprella ciliata

Female, gnathopod 2

Male lectotype, gnathopod 2 propodus

Female

Gnathopod 1 propodus

Male, abdomen

Pereopod 7

™MONQWDYD

30

ACID

Male lectotype

Female, abdomen

Right mandible, incisor region

Maxilliped palp, terminal articles (setation

not shown)

Caprella ciliata

(No. 2336) has been designated as

lectotype.

Laubitz (1970) noted that in C. a/as-

kana Mayer, the mandibular setal row

usually contained extra setae. Extra

setae were observed in C. carina, but

it was not possible to determine

whether this condition is normal for

this species.

Caprella ciliata G.O. Sars 1883

Plate 5; Map 5

Synonymy

Caprella ciliata G.O. Sars 1883, 1894

— Norman 1886, 1905 — Mayer

1890, 1903 — Holmes 1904 —

Reibisch 1906 — Stephensen 1927,

1928, 1929a, 1940, 1942, 1944a —

Oldevig 1933 — Enequist 1949.

Material examined

Thor Station 166, 62°57'N, 19°58'W,

14 July 1903, depth 957 m, 1 male,

5 (2 ovigerous) females (UZM).

Ingolf Station 92, 64°44'N, 32°52’W

25 June 1896, depth 1838 m, 1 fe-

male (UZM).

Western Norway, 17 specimens,

including Sars’s type material and

slide mount (ZM).

Description

Body normally smooth; when dorsal

spines present, there may be paired

or single head spine, single posterior

spine on pereonite 1, paired spines

in the middle and a single spine

posteriorly on pereonite 2. Pereonite

5 comparatively long, particularly in

female. Length of largest male 13 mm,

of largest female 9.1 mm, of smallest

ovigerous female 8.8 mm.

Mouthparts apparently typical of

genus; maxilliped palp with dactylus

longer than article proximal to it;

right mandible lacinia mobilis denti-

culate. not five-toothed.

Gnathopod 1 propodus slender,

triangular, with one pair of proximal

grasping spines; not very setose;

grasping margin of propodus and

dactylus serrate. Gnathopod 2 pro-

podus_ slightly setose in female,

medial surface strongly setose in

male; palm with proximal projection

bearing grasping spine, and median

and distal projections separated by a

more or less deep cleft; dactylus of

male setose, heavy.

Gills small, elliptical.

Pereopods 5 to 7 of increasing size,

propodus with proximal pair of long

slender grasping spines.

Distribution

Type locality: Western Norway.

Other localities: Denmark; Sweden;

Iceland; Alaska.

Remarks

Our present knowledge of the distri-

bution of this species suggests that it

may be present in the region covered

by this paper; but since it has not yet

been reported from this area, it has

been treated only generally. For

further details of this species, refer to

Sars (1883, 1894) and Mayer (1903).

A lectotype has been designated

from Sars’s material deposited in the

Oslo Museum (No. F 12537a).

Caprella dubia Hansen 1888

Plate 6; Map 4

Synonymy

Caprella microtuberculata f. spinosa

— Norman 1886

Caprella microtuberculata f. spinigera

Hansen 1888 — M.J. Grieg 1907

Caprella dubia Hansen 1888 —

Vanhoffen 1897 — Scott 1899 —

d’‘A. Thompson 1901 — Mayer 1903

— Stephensen 1933, 1942, 19445 —

31

32

Plate 6

Re RD e)

XO

ia.

Caprella dubia

Gurjanova 1935, 1936, 1964

Caprella septentrionalis f. spinigera —

Mayer 1890 — Hansen 1895

Material examined

Dana Station 2361, 68°O8'N, 57°30’

W, 26 June 1925, depth 398 m,

1 male, 1 female (UZM); 65°34'N,

54°31 W, 5 July 1895, depth 128 m,

1 male, 3 (1 ovigerous) females

(UZM).

Frobisher Bay, Baffin Island, 8 Oct.

1968, depth 62 m, more than 100

specimens, all immature (FRBC).

44°30'N, 49°OO’W, 25 June 1961,

depth 120 m, 3 males, 1 female

(NMC).

Hansen’s cotypes: 66°32'N, 55°54’

W, 1884, depth 200 m, 2 males;

65°35'N, 54°50'W, 1884, depth

150 m, 1 male, 1 ovigerous female;

65°35'N, 54°50'W, 1884, depth 150 m,

2 males, 2 females (UZM).

Slide mount deposited in UZM.

Description

Body spination varies slightly, as

shown in male and female (Plate 6).

In male, pereonite 1 and anterior

portion of pereonite 2 elongate with

maturity. Length of largest male

26.1 mm, of largest female 22 mm, of

smallest ovigerous female 16.5 mm.

Antenna 1 longer than head to

pereonite 3 inclusive, flagellum with

more than 26 articles and longer than

peduncle; peduncle articles 2 and 3

setose in male. Antenna 2 with many

Plate 6

Caprella dubia

A Female

B Female, gnathopod 2 propodus

C Gnathopod 1 propodus

D Male, gnathopod 2 propodus

E Male lectotype

F Maxilla 2

G Maxilliped

strong swimming setae,

article 2 minute.

Mouthparts typical of genus; right

mandible lacinia mobilis strongly

toothed, but not five-toothed.

Gnathopod 1 propodus triangular,

with one pair of proximal grasping

spines; dactylus slightly longer than

propodus palm; grasping margin of

dactylus and_ propodus _ serrate.

Gnathopod 2 basis and ischium with

distal anterolateral spine; propodus

slender, palm setose, with proximal

projection with spine, and two distal

projections; dactylus thickened prox-

imally, strongly curved, with setae

along the inner margin in adult and

subadult males.

Gills long, oval.

Pereopods 5 to 7 with proximal

pair of grasping spines on the propo-

dus, and moderately short dactylus.

Abdomen of male and _ female

typical of genus.

flagellum

Distribution

Type locality: 65°35'N,

depth 150 m.

54°50'W,

Other localities: Eastern and western

Greenland; Franz Joseph Land; Kara

Sea.

New records: Off Newfoundland;

Baffin Island.

Remarks

A lectotype has been designated

Maxilla 1

Lower lip

Pereopod 7

Left mandible

Male, abdomen

Female, abdomen

Right mandible

S22 wor

33

Caprella equilibra

5

L :

sf

Vi

©? ° x

>

(ae

A\

M@ Caprella carina

@ C. dubia

Vv M. “iA

130° N

i

Map 4 Known distribution of Caprel/la carina, Caprella dubia, and Metacap-

rella horrida.

from Hansen’‘s cotypes deposited in

the Copenhagen Museum.

C. dubia is very similar to C. micro-

tuberculata G.O. Sars; for further

discussion, see under the latter

species, p. 39.

Caprella equilibra Say 1818

Synonymy

Refer to McCain 1968

Remarks

This species is widely distributed in

34

the warm temperate waters of the

world. Although it has not been re-

corded from the area covered by this

paper, it is common along the eastern

coast of North America from Florida

to Connecticut. It has also been re-

corded from the Mount Desert region

of Maine (Procter 1933), and in

Europe is found at least as far north

as Trondheimsfjord, Norway.

C. equilibra is easily distinguished

from the other Capre//a species of the

northwestern Atlantic by the pres-

ence of a single ventral spine be-

Caprella linearis

1

@ Capreila ciliata

MSOC. «*srinki

VW Proaeginina norvegica

Ve

ip. \

Map 5 Known distribution of Caprel/la ciliata, Caprella rinki, and Proaeginina

norvegica.

tween the gnathopods 2. In the adult

male the anterior pereonites are so

elongated that the gnathopods 2 arise

at mid-length of the body.

For detailed description and figures,

refer to McCain 1968.

Caprella linearis (Linnaeus 1767)

Plate 7; Maps 6, 7

Synonymy

Refer to McCain 1968. Also:

Caprella acuminifera (non Latreille)

Bate 1862 — Mayer 1882

Caprella hystrix (non Krgéyer) Bate

and Westwood 1868

Caprella septentrionalis (non Kr@yer)

— Bousfield 19566 (in part)

Caprella acutifrons (non Latreille) —

Bousfield 1958 (in part)

Caprella linearis — Steele 1961 —

Brunel 1970

Material examined

Approximately 1300 specimens.

Hudson River estuary, off New

Jersey, 11 specimens (NMC).

Atlantic Canada: Bay of Fundy, 360

35

36

Plate 7

Caprella linearis

A Male

B Maxilliped palp, terminal articles (setation

not shown)

C Female

Caprella linearis

specimens (NMC); Nova Scotia, 60

specimens (NMC); off Nova Scotia,

44°02'N, 59°12'W, depth 180-360 m,

1 female (NMC).

Gulf of St. Lawrence: Prince

Edward Island, 25 specimens (NMC);

New Brunswick, 8 specimens (NMC);

Gaspé, 800 specimens (GR, NMC).

St. Lawrence estuary, south shore,

30 specimens (NMC). ;

Alaska: Ivik, 2 Aug. 1951, 7 juveniles

(USNM); Elson Lagoon, Point Barrow,

6-20 Oct. 1948, depth 1.5-2 m, 16

juveniles (USNM); off Point Barrow,

Aug. and Sept. 1948, depth 25-45 m,

125 specimens (USNM).

BM specimens (1952.5.7.163)

from Bate’s collection: These were

labelled “Caprella acuminifera”,

Bate’s original designation of

C. hystrix Bate and Westwood. Hoeck

(1879) suggested that C. Aystrix was

a variety of C. /inearis. Mayer reverted

to the use of C. acuminifera, and

assigned this species to C. septen-

trionalis. The specimens agree well

with Bate’s original descriptions and

figures, and are here assigned to

C. linearis.

Description

Body usually smooth; tuberculations

always paired when present. In adult

males pereonites 1 and 2 considerably

elongated. Length of largest male

22.2 mm, of largest female 13.1 mm,

of smallest ovigerous female 6.2 mm.

Male antenna 1 with’ setose

peduncle and short flagellum. Anten-

na 2 shorter than antenna 1 pedun-

cular articles 1 and 2.

Mouthparts typical of genus.

Gnathopod 2 of male with basis

longer than propodus; propodus

palm with proximal projection bearing

a spine and with one accessory spine,

and two distal projections separated

by a deep cleft; dactylus strongly

curved and setose. In female, distal

projections are smaller and closer

together; dactylus not setose.

Gills elliptical.

Propodus of pereopods 5 to 7 with

proximal grasping spines.

Variation: Typically this species has a

smooth body with usually one pair of

dorsal spines on each of pereonites

5 to 7. A small number of markedly

tuberculate specimens were found.

These cannot be considered as a

separate species, since intergrades

between the two forms exist. The

specimens figured (Plate 7) were of

the intermediate type.

Distribution

Type locality: “‘Habitat in Oceano

Europaeo”’ (Linnaeus 1767).

Other localities: Siberian Polar Sea;

Murmansk coast; Spitsbergen; Eu-

ropean coast from Norway to France;

British Isles; Faeroe Islands; Iceland;

east coast of North America from

Labrador to Connecticut; southwest

Alaska; ? Japan, Kamchatka, and

New Zealand.

New records: Hudson River estuary;

Bay of Fundy; Gulf of St. Lawrence;

Point Barrow and lvik, Alaska.

Remarks

No explanation can be offered for

the variation in body spination; such

variation, shown by a number of

caprellids, is possibly of ecological

significance. There does not appear

to be any geographical isolation of

the variants, although it was noted

that all the Alaskan specimens were

strongly tuberculate. The C. Aystrix of

Bate and Westwood, from the British

Museum, were examined, and it was

apparent that these are actually

37

Plate 8

38

Caprella microtuberculata

strongly tuberculate C. /inearis.

This common boreal species is

mainly intertidal, and only a few

collections of it were taken below

25 m.

For further discussion of C. /inearis,

see C. septentrionalis, p. 48.

Caprella microtuberculata G.O. Sars

1879

Plate 8; Map 8

Synonymy

Caprella microtuberculata G.O. Sars

1879, 1885, 1886, 1894 — Mayer

1882, 1890, 1903 — Vanhoffen

1897 — Scott 1899 — Stebbing 1900

— Bruggen 1909 — Stephensen 1918,

1929b, 1942, 1944a, 19445 —

Derjugin 1915 — Gurjanova 1931,

1964 — McCain 1966

Material examined

More than 75 specimens from east-

ern Greenland eastward.

Ingolf Station 2, depth 493 m,

3 males, 1 female; Ingolf Station 3,

depth 512 m, 14 males, 20 females,

many immatures and juveniles; In-

golf Station 44, depth 1026 m, 1 male;

Ingolf Station 93, depth 1444 m,

1 female; Ingolf Station 144, depth

520 m, 1 ovigerous female.

Eastern Greenland, 65°54'N, 1902,

1 female; Henry Land, 1900, depth

40 m, 3 males, 14 females, 25 imma-

tures; 72°35 N,. 19°35'W, 1891,

depth 200 m, 1 male, 1 female;

Plate 8

Caprella microtuberculata

Female

Gnathopod 1 propodus

Male, gnathopod 2 propodus

Female, gnathopod 2 propodus

Maxilla 2

Maxilla 1

Maxilliped

QOm7mMoOODD

Finnmark, Norway, 1898, 2 males,

1 female. All specimens in UZM.

Between Bear Island and Spits-

bergen, 6 specimens, Sars’s type

material (ZM).

Slide mounts deposited in UZM.

Description

Body tuberculation variable; large

adult males have paired head tuber-

cles and dorsal tubercles on pere-

onites 5 to 7 only; the most tuber-

culate specimen observed’ was

similar to the female in Plate 8, with

the addition of a posterior pair of

tubercles on pereonite 1. Length of

largest male 30.1 mm, of largest

female 21 mm, of smallest ovigerous

female 20.5 mm.

Antenna 1 flagellum longer than

peduncle and with more than 28

articles; peduncle articles 2 and 3

setose in male. Antenna 2 shorter

than peduncle of antenna 1, long

swimming setae, flagellum article 2

minute.

Mouthparts typical of genus, right

mandible lacinia mobilis toothed, but

not five-toothed.

Gnathopod 1 propodus with one

pair of proximal grasping spines;

grasping margin of dactylus and

propodus serrate. Gnathopod 2 pro-

podus slender, palm with proximal

projection bearing a spine, and with

one or two accessory spines, and

two distal projections; in the male,

the more distal of these is much the

Pereopod 7

Male

Male, abdomen

Female, abdomen

Right mandible

Left mandible

22S nA Cr

39

Caprella microtuberculata

C. linearis e@

C. penantis a

Map 6 Distribution of Caprella linearis and Caprella penantis within the

Canadian Atlantic region; based on material examined.

larger, but in the female they are the

same size, and appear to have a com-

mon point of origin; dactylus evenly

tapered, may be setose in large adult

males.

Gills elliptical.

Pereopods 5 to 7 of increasing

length; propodus with proximal pair

of grasping spines; dactylus long and

slender.

Abdomen of male with one pair of

unsegmented appendages, and one

pair of setose lobes separated by a

median chitinous hump, penes

medial. Abdomen of female with one

pair of lobes separated by a median

chitinous hump.

Distribution

Type locality: Northwest of Bear

Island, and southern tip of Spits-

bergen, depth 70-180 fm (128-329 m).

Other localities: Eastern Greenland;

northern Norway; Spitsbergen;

Franz Joseph Land; Kara Sea; north

40

of New Siberian Islands.

New record: Faeroe Islands (62°49'N,

7°12 Wi).

Remarks

A lectotype has been designated from

Sars’s cotypes deposited in the Oslo

Museum. The female lectotype (No.

F 12317a) was from a vial labelled

with both type localities; the para-

lectotypes are numbered F 1231 7b-e,

and F 1941.

C. microtuberculata and C. dubia

are very similar in both general body

shape and the location of the body

spines. However, the differences

between them are_ sufficient to

separate them as full species. The

abdomen of C. microtuberculata is

very much more obvious than that of

C. dubia. Also, in C. microtuberculata

the spine row on the maxilliped outer

lobe consists of about five spines, the

gnathopod 1 dactylus just reaches

the end of the propodus palm, the

Caprella penantis

Map 7 Known distribution of Capre//a linearis within the North Atlantic

and northern Pacific boreal and subarctic regions.

pereopod dactylus is long and slender.

In C. dubia the maxilliped outer lobe

spine row consists of about eight

spines, the gnathopod 1 dactylus

reaches beyond the end of the pro-

podus palm, the pereopod dactylus is

fairly short.

Hansen (1888) originally assigned

C. dubia to C. microtuberculata,

giving it the varietal name sp/nigera,

but further on in the same paper

(p. 217), he gave it specific status as

C. dubia on the basis of its greater

size and slight differences in the

gnathopods 2. It should be em-

phasized that both species may have

setae on the dactylus of gnathopod 2.

Caprella penantis Leach 1814

Plate 9, figs. A-E; Plate 10, figs. A-E;

Map 6

Synonymy

Refer to McCain 1968

Material examined

About 400 specimens.

Atlantic Canada: northeastern Nova

41

Plate 9

A-E Caprella penantis

A Male, from Florida

B Male, from Cape Cod

C Male, from Prince Edward Island

D Male, abdomen

E Female, abdomen

42

F,G Caprella natalensis

F Male, abdomen

G Female, abdomen

H-K Caprella neglecta

H Female

J Male

K Female, gnathopod 2 propodus

Caprella penantis

Scotia, 25 specimens (NMC).

Gulf of St. Lawrence: files de la

Madeleine, 5 specimens (GR, NMC);

Prince Edward Island, 300 specimens

(NMC); Nova Scotia, 10 specimens

(NMC); New Brunswick, 4 speci-

mens (NMC).

New England south to Florida,

many specimens.

The greatest depth recorded in

these collections was 11 m.

Description

Head with large, triangular, anteriorly

directed projection anterior to eye.

Pleura present on pereonites 3 and 4

in mature specimens. Pereonite 1

normally shorter than head, pereonite

5 normally shorter than pereonites

6 plus 7. Length of largest male

9.5 mm, of largest female 5.9 mm,

of smallest ovigerous female 3.5 mm.

Antenna 2 longer than peduncle of

antenna 1.

Mouthparts typical of genus.

Gnathopod 2 propodus palm setose,

with large rectangular projection

distally, and large proximal pro-

jection.

Gills circular, oval, or elliptical.

Pereopods 5 to 7 of increasing

length, propodus with concave palm

and proximal grasping spines.

Abdomen typical of genus.

Variation: It was found that there is

considerable variation within this

species, particularly in the robustness

of the body. Specimens from Florida

were larger, stouter, with stronger

pleural development and more setose

antenna 2, than specimens from

Prince Edward Island; intergradations

between these two forms were

found (Plate 9, figs. A-C). Most

Crustacea spanning this geographical

range grow to larger size in the

north than in the south. No satis-

factory explanation has been found

for the peculiar size range of

C. penantis.

Distribution

Type locality: Devonshire, England.

Other localities: Atlantic coasts of

Europe, British Isles, North and

South America; Gulf of Mexico;

Caribbean region; California; Hawaii;

eastern Asia; New Zealand; Australia;

South Africa.

Remarks

The Caprella acutifrons group, en-

compassing the twenty varieties

named by Mayer (1890, 1903), has

been analysed by McCain (1968). He

tentatively assigned eight of the

varieties, plus C. angusta (Mayer

1903) to the species C. penantis. Of

the eight varieties, five (C. acutifrons

f. carolinensis, virginia, testudo,

gibbosa, and Jusitanica) have the

setose appendages (particularly the

gnathopod 2 propodus) characteristic

of C. penantis. These five are con-

sidered to have been rightly assigned

to this species. C. angusta (which

has been considered the Pacific

coast equivalent of C. penantis) and

C. acutifrons f. natalensis, porcellio,

and neglecta are not setose. As no

other difference was observed be-

tween them and C. penantis, it was

suggested that they might, as a group,

be given subspecific status. Vassilenko

(1967) gave C. neglecta specific

status, but since her specimens ap-

pear to have been typical setose

C. penantis, this species is considered

invalid. Laubitz (1970) concluded

that C. angusta was sufficiently

different from C. penantis to be con-

sidered a separate species, and re-

instated it.

Collections of C. penantis from the

43

HOAR

HOOD D

DONA

Male, gnathopod 2 propodus

A-E Caprella penantis, total length H 8mm (scale =0.5 mm)

ae ae : J 11.1 mm (scale =0.5 mm)

of individual: K 12.6mm (scale =0.5 mm)

a ae ii i Ae: ee L-P Caprella neglecta, total length

6.1 mm (scale =0.5 mm) of individual:

8 mm (scale = 0.5 mm) 5 mm (scale = 0.5 mm)

9.5 mm (scale = 0.5 mm) 7.5 mm (scale = 0.5 mm)

F-K Caprella natalensis, total length ieee aT Rea uh

ree eee 12.2 mm (scale = 1mm

of individual: {

14.7 mm (scale = 1 mm)

F 4.9mm (scale =0.2 mm)

G 6.2mm (scale =0.5 mm)

mOOWD

Uw @O.2—,

44

Caprella penantis

Gulf of St. Lawrence south to Florida,

and of C. angusta from the Canadian

Pacific, have been investigated in

great detail. The results confirm that

these are indeed two separate spe-

cies. The correct determination of

these species was complicated by

the discovery of the true C. penantis

along the Pacific coast of California,

which made it extremely difficult to

determine the true identity of pre-

viously published records.

It must now-be pointed out that the

specimens referred by Mayer (1903)

to C. acutifrons f. angusta were from

California, and were essentially simi-

lar to the C. penantis from this same

area. Therefore C. angusta must fall

as a synonym of C. penantis, and the

Canadian Pacific “acutifrons” species

misidentified as C. angusta by Laubitz

(1970) will have to be renamed. As

specimens of C. acutifrons f. nata-

lensis were also examined by me

and found to be essentially similar to

the false C. angusta,* it has been

decided that the varietal name should

be used, and C. natalensis (Mayer

1903) is herewith given specific

status. Our present knowledge of the

distribution of C. nata/ensis is limited

to the south Atlantic and north

Pacific. It is therefore highly probable

that this name will eventually prove

to be a junior synonym for either

C. spinifrons Nicolet 1849, reported

from Chile, or C. novae-zealandiae

Kirk 1878, from New Zealand. Until

specimens from either of these areas

can be examined, no further decision

can be made as to the correct name

for this species.

As has already been mentioned,

* Slight differences between these two ‘‘varieties”

were found. It is felt that, in the light of the known

variability of the “acutifrons” group of species, these

small differences should not be considered of a

specific nature.

C. penantis and C. nata/ensis are very

similar in general appearance. Other-

wise, C. penantis tends to be stouter

than C. nata/ensis and, particularly in

mature specimens, to have very

obvious pleural development not

present in C. natalensis. The most

useful characters in distinguishing

these two species are the gnathopod

2 propodus palm, and the ratio of the

length of pereonite 5 to pereonites 6

plus 7. It was found that the gnathopod

2 differs in all stages of male develop-

ment (Plate 10), and that differences

are also apparent in the female

gnathopods. In C. penantis, pereonite

5 is usually shorter than pereonites 6

plus 7, whereas in C. natalensis

pereonite 5 is usually longer than

pereonites 6 plus 7. Small differences

are also present in the abdomen of

both males and females (Plate 9,

figs. D-G).

Specimens of C. penantis from

Monterey Bay, California, are less

setose and smaller than the largest

specimens found in the Atlantic

(8 mm as opposed to 14 mm), but they

show the typical stout body and strong

pleural development. They are ob-

viously more setose than C. nata-

/ensis, particularly the adult males.

Specimens of C. acutifrons f.

neglecta, as identified by Mayer, were

also examined, to try to determine

their specific status. Differences be-

tween these specimens and both

C. penantis and C. natalensis, particu-

larly in the head spine and the

gnathopod 2 (Plates 9, 10), indicate

that this variety must be considered

a valid full species. Its specific status

as C. neglecta (Mayer 1890, non

Vassilenko 1967) is herewith pro-

posed. This species has been re-

ported from Hong Kong, Japan, and

Australia.

Although not examined, C. acutif-

45

Plate 11

Caprella rinki, female

A Lateral view

B Gnathopod 2 propodus

46

Caprella natalensis

rons f. porcellio appears to be simi-

lar to C. natalensis, and is recorded

from the same area of the south

Atlantic. It is therefore tentatively

assigned to this species.

Caprella natalensis Mayer 1903

Plate 9, figs. F, G; Plate 10, figs. F-K

Synonymy |

Caprella acutifrons var. natalensis

Mayer 1903 — Stephensen 1949

Caprella acutifrons var. porcellio

Mayer 1903

Caprella penantis (non Leach 1814)

— Stebbing 1910 — [?] Penrith and

Kensley 1970a, 19706

Caprella penantis var. natalensis —

K.H. Barnard 1916

Caprella angusta (non Mayer 1903)

— [?] Dougherty and Steinberg 1953

— Laubitz 1970

Material exaniined

Specimens from Tristan da Cunha,

23 Dec. 1937, depth 40-45 m

(USNM), and from Pacific North

America (USNM, NMC). This spe-

cies has been described and figured

by Laubitz (1970). It is recorded

from South Africa (Durban, Cape

Town, Cape Peninsula), South West

Africa, Tristan da Cunha, and Pacific

North America (Queen Charlotte

Islands south to Oregon; Pacific

Grove and Santa Cruz, California).

It is probable that further investiga-

tion will show this species to be the

same as C. spinifrons Nicolet, from

Chile.

Caprella neglecta Mayer 1890

Plate 9, figs. H-K; Plate 10, figs. L-P

Synonymy

Caprella acutifrons — Mayer 1882 (in

part) — Schellenberg 1938 —

Edmondson and Mansfield 1948

Caprella acutifrons var. neglecta

Mayer 1890, 1903 — Utinomi 1943a,

19436, 1947, [?] 1968, [?] 1969

Caprella acutifrons var. natalensis —

Arimoto 1930 — Hiro 1937

Material examined

Specimens from Hong Kong, 9 March

1872, Mayer’s type specimens

(USNM).

This species is closely similar to

C. penantis, and it appears from a

study of the literature that both species

are present in the seas around Japan.

C. neglecta of Vassilenko, and C. acutif-

rons f. neglecta of Utinomi (1943c)

appear to refer to C. penantis. Uti-

nomi (1968, 1969) mentions other

Japanese “C. acutifrons” records; it

has not been possible to determine

to which species these records refer.

The probable distribution of C. neg-

lecta includes Hong Kong, Japan,

Taiwan, and Hawaii.

Remarks

It is apparent that the “C. acutifrons”

group is still in need of further clarifi-

cation. Study of specimens from all

parts of its geographical range will be

needed before it can be decided which

species are truly valid and which are

subspecies or variants.

Caprella rinki Stephensen 1917

Plate 11; Map 5

Synonymy

Caprella Rinki Stephensen 1917

Caprella rinki — Stephensen 1933,

1944a — McCain 1966 — Brunel

1970 — Laubitz and Mills 1972

Material examined

Gulf of St. Lawrence: Laurentian

channel, 10 Sept. 1959, depth 377 m,

1 female (GR); 17 August 1957,

1 female (GR).

47

Caprella sarsi

Description

Body with small tuberculations

dorsally on pereonites 1 to 5, ven-

trally on pereonites 2, 3, and 4. Un-

paired spines present dorsally at mid-

length of pereonites 2 to 7, and

laterally at the anterior of pereonites

3 and 5. Length of largest female

10.1 mm.

Antenna 1 peduncle articles 2 and

3 slightly setose. In female, antenna 2

longer than peduncle antenna 1.

Mouthparts not examined. Eyes

very large.

Gnathopod 2 of female with large

anterolateral spine on basis; propodus

with proximal projection with grasp-

ing spine and accessory spine, and

medial and distal low projections,

palm slightly setose; dactylus quite

heavy.

Gills slender, elliptical.

Pereopods 5 to 7 with proximal pair

of grasping spines on propodus.

Distribution

Type locality: Southern Greenland,

depth 460-700 m.

Other localities: 61°42'N, 9°36'W,

depth 1026 m; 65°14'N, 30°39'W,

depth 1416 m; 65°30'N, 55°26'W,

depth 550 m; Gulf of St. Lawrence,

depth 180-380 m.

Remarks

The two specimens of this species

available to me were females in

rather poor condition. They are

described only generally here, as this

species has been treated in detail by

Laubitz and Mills (1972).

Caprella sarsi Honeyman 1889

Remarks

Honeyman described two species of

Caprella, from a glacial boulder and

48

from two cable hauls. C. sars/ was

described as follows:

We have already noticed this fantastic little

Crustacean on Lawson Boulder (A)... On

Cable Il they abound. We have found them

among the Hydroida, where they had been

evidently feasting. They are of various sizes,

some very small. We regard all as one

species. All that we now note regarding

them is: Under the microscope they are

plain, without spines, and have small red

eyes. They are male and female. We would

name it Caprella Sarsii.

Lawson Boulder (A) is described as:

‘“‘a ‘glacial boulder’ of the Nova Scotia

Fishing Banks”, from a depth of 65

fathoms (119 m). C. sarsi was

attached to a sponge. Cable Il was

brought up on 26 Oct. 1888, after 19

years. Its location was 43°4.38'N,

66°14.3'W, at 48 fathoms (88 m),

in the Bay of Fundy. The description

is insufficient to identify this species,

but it is Suggested that it may be

C. linearis.

Caprella septentrionalis Kr¢yer 1838

Plate 12; Maps 9, 10

Synonymy

Refer to McCain 1968. Also:

Caprella ultima Bate 1862

Caprella_ septentrionalis — Smith

1879 — G.O. Sars 1909 — Vibe 1950

— Steele 1961 — Brunel 1970

Material examined

Total of 4,500 specimens.

Atlantic Canada: Bay of Fundy,

2,100 specimens (NMC); Nova

Scotia, 640 specimens (NMC);

Newfoundland, 1 specimen (NMC);

Labrador, 30 specimens (NMC).

Gulf of St. Lawrence: Prince

Edward Island, 33 specimens (NMC);

Gaspé, 300 specimens (GR, NMC);

Anticosti Island, 20 specimens

(NMC); Newfoundland, 1 specimen

(NMC).

Caprella septentrionalis

“A\

W Caprella microtuberculata

HM Cercops holbolli

v4

ws

Map 8 Known distribution of Capre/la microtuberculata and Cercops holbolli

within the North Atlantic and arctic regions.

St. Lawrence estuary: north shore,

100 specimens (NMC); south shore,

330 specimens (GR, NMC).

Arctic Canada: Hudson Bay, 30

specimens (NMC); Hudson _ Strait,

36 specimens (NMC); Baffin Island,

27 specimens (NMC).

Greenland: western, N of 70°, 84

specimens (UZM); western, S of 70°,

560 specimens (UZM); southern,

14 specimens (UZM); eastern, 14

specimens (UZM).

Jan Mayen Island, 13 specimens

(UZM); Iceland, 2 specimens (UZM);

Spitsbergen, 1 specimen (UZM).

Bate’s specimens of C. u/tima (MN).

The collections were intertidal, to a

depth of 15 m.

Description

Body spination extremely variable,

single spine or tubercle usually pres-

ent on head. Length of largest male

31.4 mm, of largest female 23.6 mm,

of smallest ovigerous female 6.4 mm.

Antenna 1 peduncle articles 2 and

3 normally setose in male. Antenna

2 normally at least as long as pe-

49

Plate 12

(\

Tad

v ( is

E NS Ape

Caprella septentrionalis

A-C_ Males, lateral view, showing differences

of body spination

D Pereopod 7

E Large female

50

Caprella septentrionalis

duncle of antenna 1.

Mouthparts typical of genus.

Gnathopod 1 propodus with one

pair of proximal grasping spines;

grasping margin of dactylus and pro-

podus serrate. Gnathopod 2 basis

short; propodus palm with proximal

projection bearing a spine and with

accessory spine, with distal small

projection plus a large triangular

projection in the angle of the dacty-

lus; dactylus becomes more strongly

curved with maturity.

Gills elliptical, oval, or round, oc-

casionally grossly inflated.

Pereopods 5 to 7 with proximal

pair of grasping spines on the pro-

podus.

Abdomen typical of genus.

Variation: This species shows great

variation in both body spination and

size. Adult males can be recognized

by the elongation of pereonites 1 and

2, by the situaiion of the gnathopod 2

posterior to mid-pereonite 2, and by

the setose antenna 1. They have been

found as small as 11 mm. Females

and immature specimens are usually

spinier than the adult males. Three

variations of body spination are

shown in Plate 12, and intergrades

between these variations exist.

There is also variation in the form

of the body spines, particularly on the

anterior pereonites. The spines may

be slender with rounded apex, blunt

with flat apex, or bifid. The _ bifid

spines frequently give the impression

of paired spines, particularly when on

the head.

An interesting condition was noted

in some of the female specimens.

The largest females found (Plate 12)

had small, immature brood plates,

while in the same lot were found

ovigerous females of half the size.

No explanation of these ‘super fe-

males” can be given until more is

known about the growth and life

cycle of these animals.

Distribution

Type locality: Greenland.

Other localities: Murmansk coast,

Novaya Zemlya, White Sea, and

northern Russia; Europe from Nor-

way, south to France and Britain;

Spitsbergen; Faeroe Islands; Jan

Mayen Island; Iceland; eastern and

western Greenland; Baffin Bay, Davis

Strait, and the eastern coast of North

America from Cumberland Sound

south to Georges Bank.

Remarks

As has already been stated, C. septen-

trionalis exhibits a variety of body

spination. Many of the varieties have

at one time been treated as distinct

species, particularly in the immature

stages. G.O. Sars (1894) treated

the main variations (C. punctata,

C. lovéni, and C. monocera) as species,

while Mayer (1903) appeared to

consider them as variants. Stephen-

sen (1928-19446) treated them

sometimes as species and sometimes

as subspecies. McCain (1968) con-

cluded that, because of the existence

of intergradations between the var-

ious forms, they should be con-

sidered infra-subspecific variants of

C. septentrionalis. During the pres-

ent investigation of this species, a

large number of specimens were

examined. It became apparent that

McCain’s so-called ‘‘monocera-like

variant’ is actually a typical adult

male, and that all the so-called

varieties can indeed by assigned to

C. septentrionalis, with the exception

of C. monocera. No specimens of

this variety were found. Examination

of type material showed that, while

51

Plate 13

Caprella unica

A Male

B Maxilliped palp, terminal articles (setation

not shown)

C Pereopod 5 propodus

D Female

52

Caprella trispinis

this variety is extremely similar to

C. septentrionalis, the antenna 1 is

obviously different, in that it has a

flagellum with at least 30 articles,

which is subequal to the peduncle.

Until specimens of C. monocera can

be examined in greater detail and

numbers, it should probably be con-

sidered a valid species.

C. septentrionalis and C. linearis,

particularly the females and im-

mature stages, are very similar in

form. McCain has shown that they

can readily be distinguished by the

ratio of total length to length of basis

of gnathopod 2 (greater than 13.0

for C. septentrionalis, less than 13.0

for C. linearis). Another distinction

is that in C. /inearis the head and

anterior body spines are always

paired, in C. septentrionalis they are

always unpaired.

The literature on these two species

is very confused, and McCain (1968)

has sorted out their synonymies

most helpfully. However, as_ has

already been mentioned, C. /Aystrix

Bate and Westwood 1868 (non

Krgyer) is to be assigned to C. /inearis.

The original specimens of C. ultima

Bate 1862 were also examined and

as Mayer stated (1903), these are

synonymous with C. septentrionalis,

not with C. eqgui/ibra as suggested by

McCain.

Caprella trispinis Honeyman 1889

Remarks

This is the second species from cable

hauls in Nova Scotia, as described by

Honeyman:

On Cable |, we found among the Hydroida

three complete specimens of another

species. These have a long spine on the back

of the head curved backward, and two on the

back, opposite the Che/ae, curved forward;

also granulation. These are blind. They are,

one male and two females. We would give

this the provisional name - Caprella

trispinis.

Cable | was brought up on 11 July

1888, after 19 years. Its location was

44°38'N, 54°6'W, at a depth of 570

fathoms (1042.5 m), “‘on the side

of the Grand Bank near the Gulf

Stream”.

It has not been possible to identify

this species. There are a number of

deep-water caprellids, such as Pro-

tellina ingolfi and Caprella rinki, that

have both spines and ‘‘granulations”,

but they also have eyes. The identity

of Caprella trispinis must therefore

remain in doubt.

Caprella unica Mayer 1903

Plate 13; Map 9

Synonymy

Caprella linearis (non Linnaeus) —

[?] Ohlin 1895 — Bousfield 1956a

(in part), 19566

Caprella unica Mayer 1903 — Rath-

bun 1905 — McCain 1968

Caprella grahami Wigley and Shave

1966 — Patton 1968

Material examined

Hudson River estuary, off

Jersey, 2 specimens (NMC).

Atlantic Canada: Newfoundland,

1954, A 14, Ferryland, 2 specimens;

B 8, Lord’s Cove, 4 specimens (NMC).

St. Lawrence estuary: north shore,

4 August 1969, Ile Grande Basque,

1 specimen (NMC); south’ shore,

23 August 1953, La Lorraine Cove,

30 specimens, Pte. Ste-Anne-des-

Monts, 20 specimens (NMC).

All collections are intertidal, tem-

perature range 8.8°-14.5°C.

New

Description

Body smooth to quite spiny. Length

of largest male 9.2 mm, of largest

53

Caprella unica

/ 60° C.unica |

C. septentrionalis ®

45°n _ | 55°W

Map 9 _ Distribution of Capre//a septentrionalis and Caprella unica within the

Canadian Atlantic region; based on material examined.

female 7.2 mm, of smallest ovigerous

female 4.9 mm.

Antenna 1 of male with short

flagellum and setose peduncle. An-

tenna 2 shorter than, or subequal to,

peduncle of antenna 1.

Mouthparts typical of genus;

dactylus of maxillipedal palp heavy.

Gnathopod 1 propodus with one

pair of proximal grasping spines;

grasping margin of dactylus and pro-

podus serrate. Gnathopod 2 propodus

similar to C. /inearis, but basis shorter

and heavier.

Gills elliptical.

Pereopods 5 to 7 with propodus

palm not delineated, grasping spines

absent, few palmar spines.

Abdomen typical of genus.

Distribution

Type

45"’W, depth 59 m.

54

locality: 40°34'°30"N, 69°50’

Other localities: Long Island Sound;

southern Cape Cod and off Cape Cod;

Casco Bay, Maine; St. John’s, New-

foundland.

New records: Hudson River estuary;

north and south shores, St. Lawrence

estuary; Burin Peninsula, New-

foundland.

Remarks

Patton (1968) has shown. that

C. unica appears to be an obligate

commensal with starfish (Asterias

spp.), and that in New England waters

it shows a definite, strong prefer-

ence for Asterias forbesi (Desor) over

A. vulgaris Verrill. A. forbesi has not

been recorded north of Maine while

A. vulgaris is common in the south-

ern Gulf of St. Lawrence, but is re-

placed in the northern Gulf and in the

estuary by Leptasterias polaris

Caprella unica

Map 10 Known distribution of Capre//a septentrionalis within the North

Atlantic boreal and arctic regions.

(Muller and Troschel). The collection

data on the specimens examined is

not specific, but it seems probable that

in this region it is A. vulgaris and

probably also L. polaris that act as

host to C. unica.

C. unica is very similar to C. /inearis,

differing in being more robust in both

body and appendages. The most

distinctive features are the absence

of grasping spines on the pereopods,

and the heavy dactylus to the maxil-

liped palp in C. unica. In C. linearis

the dactylus is quite slender, and

grasping spines are present on the

pereopods.

55

Plate 14

Cercops holbdlli

Genus CERCOPS Kroyer 1842-43

Antenna 2 without swimming setae, flagellum with two or three articles;

mandible with triarticulate palp, setal formula for terminal article 3, molar

absent; outer and inner plates of maxilliped very small, subequal; gills on

pereonites 2, 3, and 4; pereopods 3 and 4 one-segmented, pereopod 5 six-

segmented; abdomen five-segmented, with two pairs of biarticulate uropods

and in male, two pairs of very rudimentary pleopods.

Type species

Cercops holbolli Kréyer 1842-43 (by monotypy).

Cercops holbolli Kr¢éyer 1842-43

Plate 14; Map 8

Synonymy

Cercops Holbolli Kréyer 1842-43,

1846 — Bate 1862 — Boeck 1871,

1872-76 — Mayer 1882, 1890, 1903

— Stuxberg 1882 — Hansen 1888 —

Vanhoffen 1897 — Utinomi 1947

Cercops holboelli — Lutken 1875 —

Stephensen 1918

Cercops holbolli — d'A. Thompson

1901

Cercops holbolli — Stephensen 1933,

1944b

Material examined

Alaska: Ivik, 2 Aug. 1951, 1 male

(USNM); off Point Barrow’ Base,

9 Sept. 1948, 15 Sept. 1948, depth

33-38 m, 4 males, 1 female (USNM),

Plate 14

Cercops holbolli, immature

Male, gnathopod 2 propodus

Gnathopod 1

Male

Female

Maxilliped

Maxilla 1

Maxilla 2

QOmmooooapyp

depth 37-38 m, 1 female (USNM).

Description

Head with anteriorly directed median

spine and paired spines anterior to

eyes. Position and size of body spines

variable, dorsal unpaired spines or

humps may be present on pereonites

1 to 6. Length of largest male 5.7 mm,

of largest female 7.5 mm.

Antenna 1 setose, just longer than

head to pereonite 2; flagellum shorter

than peduncle, having about 12

articles. Antenna 2 setose and shorter

than peduncle of antenna 1, flagellum

with two articles.

Mandible with triarticulate palp

having three setae on the terminal

article; left mandible with six-toothed

incisor and lacinia mobilis, setal row

of seven setae; right mandible with

Pereopod 6

Lower lip

Pereopod 3

Male, abdomen

Left mandible

Right mandible

me rR Ge

57

Plate 15

Hemiaegina minuta, male

Pereonites 1 and 2, lateral view

B Gnathopod 1 propodus

C Pereopod 7 propodus

D Abdomen

>

58

Hemiaegina minuta

six-toothed incisor, finely denticulate

lacinia mobilis, setal row of six setae.

Maxilla 1 palp with few spines and

setae; outer plate with six complex

spines. Maxilla 2 very small, with few

apical setae on each plate. Maxilliped

inner plate small, with three apical

setae; outer plate small, with two

apical setae; palp slender.

Gnathopod 1 propodus with prox-

imal, single grasping spine, and

strongly serrate grasping margin;

dactylus grasping margin serrate.

Gnathopod 2 basis and ischium with

anterolateral spine, propodus palm

with proximal projection bearing a

spine, and small distal projection. In

adult male, palm is setose and bears

two proximal and two distal pro-

jections.

Gills small, elliptical.

Pereopods 3 and 4 with one apical

and one subapical seta. Pereopods 5

to 7 normal, propodus’ grasping

spines proximal.

Abdomen __five-segmented; seg-

ments 4 and 5 each bear one pair of

biarticulate appendages with a row

of fine denticulations along their

length; in the male, abdominal seg-

ments 1 and 2 each have a pair of

small humps bearing single setae.

Penes lateral.

Distribution

Type locality: Greenland.

Other localities: Near Greenland; off

Cape Raper, Baffin Island; ? Tsugaru

Strait, Japan.

New records: Off Point Barrow,

Alaska, depth 33-38 m; Ivik, Alaska.

Remarks

The seven specimens examined were

all immature. They extend our know-

ledge of the distribution of this species

to the western Arctic. The Japanese

locality record for C. holbolli is

questionable; it is probable that either

C. compactus Laubitz 1970, or per-

haps an undescribed third species of

this genus, is responsible for this

record.

Genus HEMIAEGINA Mayer 1890

Antenna 2 without swimming setae, flagellum with two articles; mandible

without palp, molar present; outer plate of maxilliped larger than inner plate;

gills on pereonites 3 and 4; pereopods 3 and 4 one-segmented, pereopod 5

six-segmented; abdomen with one pair of biarticulate appendages.

Type species

Hemiaegina minuta Mayer 1890 (by monotypy, subsequently designated by

McCain 1968).

Hemiaegina minuta Mayer 1890

Plate 15

Synonymy

Refer to McCain 1968. Also:

Hemiaegina minuta — Utinomi 1964,

1969 — Fine 1970

Material examined

One specimen from mid-Atlantic,

45°40.3'N, 36°45.5'W, 24 July 1966,

depth 180 m, midwater trawl! (NMC).

Description

Body smooth except for one pair of

59

Hemiaegina minuta

ventral spines between gnathopods

2; pereonites 2 to 6 have a hexagonal

outline when viewed dorsally. Length

of male, 4.7 mm.

Antenna 1 subequal in length to

pereonites 1 to 4. Antenna 2 just

longer than peduncle of antenna 1.

Left mandible with five-toothed

incisor, five-toothed lacinia mobilis;

right mandible with five-toothed

incisor, serrate lacinia mobilis.

Maxilla 1 with six spines on outer

plate. Maxilliped inner and _ outer

plates quite small, palp slender with

tapering dactylus having’ serrate

grasping margin.

Gnathopod 1 propodus with large

proximal knob bearing many minute

projections and one small grasping

spine, tip of dactylus complex;*

grasping margin of propodus and

dactylus not serrate. Gnathopod 2

propodus large, palm with proximal

grasping spine and medial projection

separated from distal bilobed pro-

jection by a notch.

Gills elongate.

Pereopods 3 and 4 one-segmented,

minute. Pereopods 5 to 7 with pro-

podus bearing several proximal

knobs, each with a spine; dactylus

with a few strong serrations just past

the centre of its grasping edge.

Abdomen with one pair of biarti-

culate appendages bearing rows of

minute knobs along medial edge.

Penes lateral.

Distribution

Type locality: Off Amoy, China, depth

15-46 m.

Other localities: Off ©Bermuda;

Virgina; North Carolina; Florida;

Tortuga Island; 29°44'N, 88°23'W;

* There appears to be an anterior claw and a

posterior marginal lobe (Plate 15, fig. B).

60

Texas; Virgin Islands; South Africa;

Hawaii; Bora Bora Island; Tateyama,

Nomasaki, and Sunohana, Japan;

1°42.5'S, 130°47.5’E; Fremantle,

Australia; Krudadai Island, India;

coast of South Arabia.

New record: Mid-Atlantic, 45°40.3'N,

36°45.5'W, depth 180 m.

Remarks |

The single specimen examined was

a very large male in rather poor con-

dition. Because the new locality was

much farther north than previously

recorded for this species, the speci-

men was compared with H. minuta

from Bermuda. Apart from being very

much larger and more robust, it

agrees in all essential characters with

this species. It was also noted that

the ventral spines were supported by

a large ventral protrusion in the large

male. H. minuta has not been re-

corded from the region under con-

sideration, but has been recorded in

plankton tows and from. floating

Sargassum sp. It is therefore sug-

gested that the specimen recorded

here was carried north within the

Gulf Stream.

Luconacia incerta

Genus LUCONACIA Mayer 1903

Antenna 2 without swimming setae, flagellum with two articles; mandible

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

outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4;

pereopods 3 and 4 two-segmented, pereopod 5 six-segmented; abdomen of

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

without appendages or lobes; pereopod 5 inserted near mid-length of

pereonite 5.

Type species

Luconacia incerta Mayer 1903 (by monotypy, subsequently designated by

McCain 1968).

Luconacia incerta Mayer 1903

Synonymy

Refer to McCain 1968.

Remarks

Although this species has not been

recorded north of Cape Cod, it has

been taken in plankton tows and from

that specimens may be found in the

region covered by this paper. L. incerta

can most readily be distinguished

from the other caprellids of this re-

gion by the fact that it has the pereo-

pod 5 insertion near the mid-length

of pereonite 5.

For further details consult McCain

1968.

Sargassum sp. It is therefore possible

Genus MAYERELLA Huntsman 1915

Antenna 2 without swimming setae, flagellum with two articles; mandible

with triarticulate palp, setal formula for terminal article 1, molar present;

outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4;

pereopods 3 and 4 two-segmented, pereopod 5 two- or three-segmented;

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

female with one pair of lobes.

Type species

Mayerella limicola Huntsman 1915 (by monotypy, subsequently designated

by McCain 1968).

Material examined

Five females from Bay of Fundy, Har-

bor de Loutre, 1913, 9-18 m, (NMC).

Mayerella limicola Huntsman 1915

Plate 16, figs. A-F

Synonymy

Mayerella limicola Huntsman 1915 —

Procter 1933 — McCain 1968 —

Laubitz and Mills 1972

Description

Body smooth.

female 4.9 mm.

Length of largest

61

Plate 16

A-F Mayerella limicola, female G,H Metacaprella horrida

A Abdomen G Male lectotype, lateral view

B Pereopod 7 H Female, semidiagrammatic dorsal view,

C_ Lateral view showing body spination

D Pereopod 3

E Pereopod 5

F Gnathopod 2

62

Metacaprella horrida

Antenna 1 longer than head to

pereonite 2. Antenna 2 longer than

peduncle of antenna 1.

Mouthparts typical of genus.

Gnathopod 17 propodus_ slender,

with single proximal grasping spine,

and serrate palm; dactylus toothed,

grasping margin serrate. Gnathopod

2 basis long and slender; propodus

with proximal grasping spine, smooth

palm with setae; dactylus slender.

Gills elliptical, anterior pair being

segmented. Pereopods 6 and 7

slender, with long, slender propodus

and dactylus, propodus’ without

obvious palm or grasping spines.

Abdomen of female with one pair

of setose lobes.

Distribution

Type locality: St. Croix estuary, New

Brunswick.

Other localities: Bay of Fundy, to

the larger. depth of 91 m; Mount Desert region,

Pereopods 3 and 4 small, two- Maine; 44°N, 68°15’'W, depth 101 m;

segmented. Pereopod 5 two- 39°54'N, 70°20'W, depth 713 m.

Genus METACAPRELLA Mayer 1903

Antenna 2 with swimming setae, flagellum with two articles; mandibular

palp absent, molar present; outer plate of maxilliped larger than inner plate;

gills on pereonites 3 and 4; pereopods 3 and 4 absent, pereopod 5 six-

segmented; abdomen of male and female with one pair of appendages and

one pair of lobes.

Type species

Caprella kennerlyi Stimpson 1864 (designated by Dougherty and Steinberg

1953).

Remarks

This genus is now composed of three species, VW. anomala (Mayer), M. ken-

nerlyi, and M. horrida (G.O. Sars). The genus was previously thought to be

endemic to the Pacific Ocean. The addition of MW. horrida to the genus extends

its range to the Arctic and North Atlantic.

The only character that separates Metacaprella from Caprella is the female

abdomen, which bears minute appendages in Metacaprella. The abdomina

of all three species were investigated, and it was found that lobes are present

in the females, in addition to the appendages. The generic definition has

therefore been altered to include this character.

Metacaprella horrida(G.O. Sars 1877) G.O. Sars 1885, 1886 — Hansen

comb. n. 1887 — Bruggen 1909 — J.A. Grieg

Plate 16, figs. G, H; Plate 17; Map 4. 1925

Caprella horrida G.O. Sars 1877 —

Mayer 1882, 1890, 1903 — Hansen

1895 — Vanhoffen 1897 — dA.

Synonymy

Caprella_ spinosissima (non Bate

1862) Norman 1874, 1886, 1905 —

Miers 1877 — Mayer 1882, 1890 —

Thompson 1901 — Stephensen 1918,

1933, 1940, 1942, 19444, 19445 —

63

Plate 17

64

Metacaprella horrida

Steele 1961 — Gurjanova 1964 —

McCain 1966

Material examined

44°30'N, 49°00'W, 25 June 1961,

depth 120 m, 3 males, 4 females

(NMC); Frobisher Bay, 8 Aug. 1970,

depth 70 m, 1 male (NMC).

Sars’s type material: Norwegian

North-Atlantic Expedition: Station

124, 66°41'N, 6°59’E, 19 June 1877,

depth 640 m, more than 60 spec-

imens; Station. 200, 71°25’N, 15°

41'E, 17 July 1877, depth 1134 m,

more than 50 specimens; Stations 18,

48, 137, 164, 200, 343, depth 547-

1359 m, more than 60 specimens

(ZM).

Description

Head and body strongly spined, size

and number of spines variable. Large

paired dorsal spines always present

on head, just behind eye, and on

pereonite 2 over gnathopods. Spines

usually also present on head just

over the eye, and at the base of an-

tenna 2. Spination for least spiny and

most spiny specimens shown in

Plates 16 and 17. The adult male of

the less spiny form shows con-

siderable reduction in the number of

spines on pereonites 1 and 2 (Mayer

1903, Pl. 4, fig. 20). In all adult males,

pereonites 1 and 2 are elongated to

nearly half of the body length. Length

Plate 17

Metacaprella horrida

Gnathopod 1 propodus

Female

Male, gnathopod 2 propodus

Female, gnathopod 2 propodus

Maxilla 2

Maxilla 1

Lower lip

QOm7moaowvp

of largest male 25 mm, of largest

female 14.3 mm.

Antenna 1 longer than head to

pereonite 3, flagellum very much

shorter than peduncle; in adult male,

peduncle articles 2 and 3 very long,

so that the antenna is subequal to

the body length. Antenna 2 shorter

than antenna 1 peduncle, and with

long swimming setae.

Mouthparts similar to Caprella.

Right mandible lacinia mobilis

strongly toothed, but not five-toothed.

Gnathopod 1 propodus heavy, very

setose, with one pair of proximal

grasping spines; grasping margin of

propodus serrate, of dactylus denti-

culate. Gnathopod 2 basis with lateral

distal spine; propodus of male setose,

with small lateral spines and a single

anterodistal projection; palm with

proximal projection bearing small

spine, and with median projection

separated by a cleft from distal, com-

plex projection in angle of dactylus;

dactylus heavy, setose. Propodus of

female similar to male, but not parti-

cularly setose and with smaller

palmar projections. In adult male, the

propodus is slender and longer than

the basis; the dactylus is thickened

proximally, but quite slender distally.

Gills elliptical.

Pereopods 5 to 7 of increasing

length, propodus with proximal pair

of grasping spines.

Maxilliped

Left mandible

Right mandible

Pereopod 7

Subadult male

Male, abdomen

Female, abdomen

Sse ae

65

Plate 18

Paracaprella tenuis

66

mooawp

Male

Right mandible

Male, gnathopod 2

Female, gnathopod 2 propodus

Male, abdomen

Cae G)e

Female, abdomen

Pereopod 4

Pereopod 5

Pereopod 7 propodus

Paracaprella tenuis

Abdomen of male with one pair of

appendages and one pair of setose

lobes; abdomen of female with one

pair of very small appendages and one

pair of lobes.

Distribution

Type locality:

depth 753.5 m.

62°44.5'N, 1°48'E,

Other localities: Siberia; Kara Sea;

Spitsbergen; Norwegian Sea; _ Ice-

land; Faeroe--\Islands; Greenland;

Baffin Island; Hudson Bay.

New record: Off Newfoundland.

Remarks

This is a high arctic species, and has

been reported to depths of 1359 m.

The record for Newfoundland is a

considerable southward extension of

the known range of this species, but

within the same zoogeographic

region.

In his description of this species,

Sars (1885) noted the female ab-

dominal appendages, but when

Mayer proposed genus Metacaprella

(1903), he made no mention that

C. horrida should be included in that

genus.

M. horrida is readily distinguished

from M. anomala and M. kennerlyi by

its very strongly spined head and body,

and by the enormously elongated

peduncle of antenna 1 and setose

gnathopod 2 of the male.

A lectotype has been designated

from Sars’s specimens deposited in

the Zoologisk Museum, Oslo (No.

F 1944a), a subadult male of the more

spiny form, as described and figured

by Sars (1885).

Genus PARACAPRELLA Mayer 1890

Antenna 2 without swimming setae, flagellum with two articles; mandibular

palp much reduced with up to three articles, setal formula for terminal article

1, molar present; outer plate of maxilliped larger than inner plate; gills on

pereonites 3 and 4; pereopods 3 and 4 two-segmented, pereopod 5 six-

segmented; abdomen of male with one pair of appendages and one pair of

lobes; abdomen of female with one pair of lobes.

Type species

Paracaprella pusilla Mayer 1890 (by monotypy, subsequently designated by

McCain 1968).

Paracaprella tenuis Mayer 1903

Plate 18; Map 2

Synonymy

Refer to McCain 1968.

Material examined

Hudson River estuary, off

Jersey, 1 male (NMC).

Gulf of St. Lawrence: Nova Scotia,

9 specimens (NMC); Prince Edward

New

Island, 130 specimens (NMC); New

Brunswick, 13 specimens (NMC).

The greatest depth recorded in

these collections was 11 m.

Description

Body dorsally smooth, head of

characteristic high-domed — shape.

Laterally there is some _ pleural

development on pereonites 2, 3, and

4. Length of largest male 6.8 mm, of

67

Paracaprella tenuis

largest female 5.1 mm, of smallest

ovigerous female 3.6 mm.

Antenna 1 as long as head to

pereonite 2, peduncle setose. Antenna

2 setose, shorter than peduncle an-

tenna 1.

Mouthparts typical of genus; man-

dibular palp present but much re-

duced, composed of from one to

three articles, and usually having

single long apical seta.

Gnathopod 1 propodus with single

proximal grasping spine and strongly

serrate margin; dactylus toothed and

with serrate grasping margin.

Gnathopod 2 with slender basis

having distal anterolateral projection;

propodus of female slightly setose,

palm serrate, grasping spine proxi-

mal. Propodus of male with small

proximal spine separated by serrate

portion of palm from very large pro-

jection with deep triangular cleft

immediately distal; remaining palm

more or less smooth, with slight,

serrate, triangular projection in the

angle of the dactylus; dactylus with

proximal thickening, setose and

tapering more or less evenly distal

to the thickening.

Gills oval.

Pereopods 3 and 4 biarticulate, the

terminal article being minute, both

articles being setose. Pereopods 5 to

7 of increasing length, propodus with

proximal grasping spines and, in

pereopods 6 and 7, having a series

of palmar knobs, each bearing a small

spine.

Abdomen of male with one pair of

small unsegmented appendages and

one pair of setose lobes; penes arise

laterally but meet distally. Abdomen

of female with one pair of setose

lobes.

Distribution

Type locality: Woods Hole, Mass.

Other localities: Southwestern Gulf

of St. Lawrence, around Prince

Edward Island; Pennellville, Maine;

coast of North America from Cape

Cod to Florida; Gulf of Mexico.

Remarks

McCain (1968) stated that one of the

differences between P. tenuis and

the very similar P. pusilla Mayer, is

that setae are ‘‘not generally’ pres-

ent on the dactylus of the gnathopod

2 of male P. tenuis. In the collections

examined by myself, however, all

mature males had a few setae on the

dactylus, but these were both shorter

and sparser than those of P. pusi/la.

Genus PROAEGININA Stephensen 1940

Antenna 2 without swimming setae, flagellum with two articles; mandible

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

outer plate of maxilliped larger than inner plate; gills on pereonites 3 and 4;

pereopods 3 and 4 absent, pereopod 5 six-segmented; abdomen of male and

female with two pairs of biarticulate appendages and one pair of lobes.

Type species

Parvipalpus norvegicus Stephensen

designated by Stephensen 1940).

68

1931 (by monotypy, subsequently

Proaeginina norvegica

Proaeginina norvegica (Stephensen

1931)

Map 5

Synonymy

Parvipalpus norvegicus Stephensen

1931

Proaeginina norvegica — Stephensen

1940, 1942, 1944a, 19446 — McCain

1966, 1968 — Laubitz and Mills 1972

Remarks

This species has been recorded from

Davis Strait and off Cape Cod, and it

is therefore to be expected that it will

eventually be recorded from. the

Canadian Atlantic. As it is primarily

a deep-water species (174-2702 m),

it is not surprising that it was not

found in the predominantly intertidal

collections examined here. P. norve-

gica can be recognized by the

distinctive abdomen of this genus,

and by its long and slender body with

elongated pereonites 4 and 5.

For detailed description, consult

Laubitz and Mills (1972).

69

Table 1. Known Distributions of Canadian Arctic and Atlantic Caprellidae

to Long Island Sound

W Arctic (140°E to 125°W)

NE Spitsbergen (S)

Novaya Zemlya (N)

Franz Joseph Land (F)

D4 < < & & | KaraSea

Siberia to 140°E

Labrador to Gulf

of St. Lawrence

Faeroe Islands (F)

SW Spitsbergen

Pacific Alaska and Siberia

Gulf of St. Lawrence

<x < <x < x< x< >< | W Greenland

x< < x << XX x< SE Greenland

. x< x x D4

NW Greenland

NE Greenland (G)

Iceland (I)

N Norway and

Barents Sea

S of Cape Cod

Species

()

Caprella dubia

Metacaprella horrida

Caprella carina

C. microtuberculata

Cercops holbolli

Aeginina longicornis

Aeginella spinosa

Aeginina aenigmatica

Caprella rinki

Proaeginina norvegica

Caprella septentrionalis

C. ciliata

C. linearis

C. unica

Mayerella limicola

Caprella andreae

C. equilibra

C. penantis

Paracaprella tenuis

Hemiaegina minuta

Luconacia incerta

>< << | N American Arctic to 125°W

Xx

X

x KK

70

Zoogeography

The known distributions of the Cana-

dian Atlantic and Arctic Caprellidae,

illustrated by Maps 1 to 10, are sum-

marized in Table 1. The’ three

zoogeographical regions considered

here, arctic, subarctic, and boreal,

conform to Grainger’s (1966) modi-

fication of Dunbar’s (1951, 1953)

definitions, with the exception of the

position of the boundary between

subarctic and boreal regions. Dun-

bar’s definition of this boundary was

based on the southward penetration

of arctic water. The caprellid fauna

shows a marked change in the Gulf

of St. Lawrence region. Such a faunal

change has been noted for other

groups (Hall 1964; Powell 1968).

With division based on climate and

faunal distribution, the subarctic

province probably includes, besides

southern Greenland, most of Hudson

Bay and Labrador, northern and

eastern Newfoundland only, and the

northern Gulf of St. Lawrence, in-

cluding the St. Lawrence estuary

(Ganong 1890; Stephenson = and

Stephenson 1954; Bousfield 1956a,

19566; Brunel 1961a).

It is here considered that the

boundary between subarctic and

boreal faunal regions lies in the

northern half of the Gulf of St.

Lawrence. Although detailed ecolog-

ical knowledge of the region is

lacking, it appears probable that this

boundary runs from northern New-

foundiand (Strait of Belle Isle) near

the north shore of the Gulf, north and

west of Anticosti Island, and across

the St. Lawrence estuary to northern

Gaspé.

Canadian Atlantic Region

The Canadian Atlantic region lies

within the subarctic and_ boreal

zoogeographical provinces. These

provincial divisions are based on

coastal conditions, and pertain pri-

marily to shallow-water regions. In

deeper waters within each province,

conditions are more like those of

more northerly regions, with corres-

ponding faunal differences. Brunel

(1961a) and _ Drainville, Tiphane

and Brunel (1963) have observed

that the depths of the Saguenay fjord,

which are colder and have a different

fauna from both the St. Lawrence

estuary and the Gulf, contain an

“arctic enclave” isolated from the

arctic province. Similarly, subarctic

conditions exist within the boreal

province, particularly in the Lauren-

tian channel (Gulf of St. Lawrence),

at the mouth of the Bay of Fundy and

in its cold-water estuaries, and in the

Gulf of Maine.

Within the boreal province there

are also ‘‘pockets” of southern, or

warm-temperate, forms. The warm-

temperate Virginian province’ is

generally defined as lying between

Cape Cod and Cape Hatteras. Ganong

(1890) pointed out that typically

Virginian faunas were to be found in

variouS unconnected areas of the

boreal region. These findings have

been corroborated and expanded by

later workers. The presence of warm

marine pockets has been established

in the following areas: warm-water

bays and estuaries of the Bay of

Fundy, particularly the St. John

estuary, Chignecto Bay, Minas Basin,

St. Marys Bay; sheltered coves and

warm-water estuaries of Atlantic

Nova Scotia; Newfoundland, in

sheltered bays around the coast; the

Magdalen shallows of the south-

western Gulf of St. Lawrence

(Ganong 1890; Huntsman 1924;

Stephenson and Stephenson 1954;

Bousfield 1956a-62). These warm

pockets support relict populations of

species widespread in this region

7 /

Zoogeography

during the postglacial period of

minimum sea _ levels and warm

temperate climate (Bousfield 1962,

1967).

Seven caprellid species are here

recorded from the Canadian Atlantic.

Caprella rinki, a sublittoral species,

was found only in the Laurentian

channel, below 300 m. The distribu-

tions of the other species reflect the

zoogeographical conditions of this

region. Caprella penantis and Paracap-

rella tenuis, both warm-temperate

species, were found almost ex-

clusively within the warm Magdalen

shallows (Maps 2 and 6). Except for

a few isolated locations, these two

species are otherwise recorded

from south of Cape Cod.

Caprella linearis and C. unica, and

more particularly C. septentrionalis

and Aeginina_ longicornis, were

found mostly within the cooler

northern part of the Gulf, in the St.

Lawrence estuary, along the Atlantic

shores of Nova Scotia, and in the Bay

of Fundy (Maps 2, 6, and 9).

Canadian Arctic Region

Knowledge of the distribution of

caprellids within the Canadian Arctic,

and indeed within the Arctic as a

whole, is limited and discontinuous.

While certain areas of the Arctic have

been studied quite extensively (for

example, Greenland and northern

Norway), we have very little know-

ledge of the fauna of other regions

such as the western Canadian Arctic.

Therefore, any conclusions reached

on the distribution of caprellids within

the arctic and subarctic regions must

be of a tentative nature. More ex-

tensive collections are needed, parti-

cularly from the far eastern and far

western portions of these regions. A

number of caprellid species have

been reported from the Pacific (Mayer

72

1903), but they appear to be con-

fined to south of the Bering Strait.

As has been pointed out by Hedgpeth

(1963) and Coachman (1963), the

shallow waters of the Beaufort and

Siberian seas have reduced salinity.

Also, because of the effects of ice

during the winter, they offer a very

hostile type of environment (Mohr

and Tibbs 1963). However, caprellid

species recorded from other parts of

the north-polar seas may well be

present in the deeper, more saline

waters to the north of the Pacific

subarctic.

Within the arctic region, Caprella

dubia and Metacaprella horrida are

probably endemic species. The ma-

jority of records for these two species

within the subarctic region are at

depths below 100 m. Their dis-

tribution is therefore of the high arctic

type, as defined by Ekman (1953).

In the subarctic region, two distinct

patterns of distribution are found.

The panarctic species, Caprella

carina, C. microtuberculata, and pos-

sibly Cercops holbolli, are found in

arctic and subarctic regions, but do

not extend southward into the boreal

region. The _ arctic-boreal species,

Aeginina longicornis and Aeginella

spinosa, though primarily subarctic

in distribution, are found also in the

arctic region and as far south as Cape

Cod, and for A. /ongicornis, Cape

Hatteras.

Caprella unica and Mayerella

limicola are possibly endemic to the

boreal region. The other species

characteristic of this region show a

variety of distribution patterns.

Caprella linearis and C. ciliata have

discontinuous circumpolar distribu-

tion, being recorded from the north-

ern Atlantic and Pacific oceans, with

some occurrences in the subarctic.

C. rinki and Proaeginina norvegica

Canadian Arctic Region

are sublittoral species, found usually

below 200 m. Caprella septentriona-

lis, a boreo-arctic species, is found

mainly in boreal and_ subarctic

regions, with few records for the

Arctic.

The single record for Aeginina

aenigmatica is insufficient for con-

clusions to be reached on its distri-

bution, but it will probably prove to

be a subarctic or arctic species.

There appear to be two main zoo-

geographical regions inhabited by

north-polar caprellids: Atlantic-

arctic, reaching from approximately

125°W eastward to at least 140°E,

probably to 170°E; and Pacific sub-

arctic, which includes the Chukchi

and Beaufort seas.

The only caprellid species recorded

east of 140°E is Caprella micro-

tuberculata. \t is possibly Siberian in

origin, and has not yet been recorded

west of Greenland.

Four species have been recorded

west of 125°W. Two of these, C. ci/-

jata and C. /inearis, are circum-

polar. C. carina may prove to be a

circumpolar species, but its present

distribution record indicates a Siberian

or Pacific origin. Cercops holbolli

appears to be confined to the North

American Arctic. It may be a relict,

one of a group of invertebrates that,

according to Nesis (1962), crossed

the Arctic during the Upper Pliocene.

Certainly, the primitive nature of this

species, and its possible presence in

Japan, indicates a Pacific origin.

The remaining species have not

been recorded either east of 140°E

or west of 125°W. Although sub-

sequent collecting may show some

of them to be circumpolar in distri-

bution, it is apparent that the majority

of north-polar caprellids are Atlantic

in origin.

Study of the literature indicates that

similar distribution patterns exist in

other faunal groups. Both asteroids

(Grainger 1966) and pycnogonids

(Hedgpeth 1963) have an arctic fauna

that is predominantly Atlantic, with

a Pacific subarctic fauna limited to

the Chukchi Sea region. Filatova’s

(1957) investigation of arctic bivalve

mollusks of the U.S.S.R. showed

that, while a number of Atlantic

species are found as far east as the

northern Chukchi Sea, _ relatively

few north Pacific species were found

to have penetrated into the East

Siberian Sea. The arctic Bryozoa

show a high degree of circumpo-

larity, and Osburn (1955) and

Powell (1968) have concluded that

“there is no. significant difference

between the bryozoan faunas of the

Pacific-Arctic and Atlantic-Arctic

areas’. However, the majority of the

arctic species are Atlantic in origin,

the few Pacific-arctic species having

localized distribution. It appears that

a similar situation exists in the gam-

maridean amphipods, with a fairly

homogeneous fauna in most of the

cold-water region. There is, how-

ever, a definite Pacific element in this

fauna. There is also a smaller ele-

ment radiating from the central

Siberian Seas (Gurjanova 1951;

Steele 1961). Two caprellids that

may have originated in the latter

region are Caprella carina’ and

C. microtuberculata. Other caprellids

have penetrated it from the west.

Gurjanova’s other cold-water regions

(1951) do not appear to be sup-

ported either by amphipod or by other

faunal distributions. However, the

Chukchi-American transitional, as

modified by Steele (1961), is ap-

proximately equivalent to what has

here been called the Pacific-sub-

arctic.

Lemche that

(1941) recognized

73

Zoogeography

zoogeographical classification within

the arctic-subarctic-boreal region

cannot. satisfactorily be achieved

using only Ekman’s system. Geo-

graphical location is less important

than climatological and _ hydro-

graphical conditions. Thus, cold-

stenothermal species may be re-

corded from all three regions, though

from different depths. Lemche there-

fore based his faunal classification

on both propagative and vegetative

requirements of the species. Such a

system may be usefully applied to all

arctic faunas. Study of the compo-

sition and collecting data of the

collections of most museums would

produce much information on breed-

ing biology and_ distribution of

caprellids. Unfortunately, no analyses

have been made of the available

facts, and Lemche’s system cannot

as yet be applied to the caprellids.

In summary, we may say that

there are two major faunal regions

within the north-polar seas. The

Atlantic-arctic (including the sub-

arctic) extends from at least 125°W,

eastward to at least 140°E. The

Pacific-subarctic is confined to

140°E to 125°W at most. The ma-

jority of arctic caprellids are of

Atlantic origin, while the Pacific-

arctic species have very localized

distributions, being found for the

most part south of the Bering Strait.

74

Discussion

Many of the problems encountered

when dealing with the Caprellidae

will be solved only when detailed

ecological and _ genetical studies

have been carried out, and when we

have detailed knowledge of the life

history of these animals. Some of

these problems will be discussed

briefly.

In certain caprellid species, for

example, Aeginella spinosa, body

Spination appears to be quite con-

stant; more frequently, however, the

body spination is highly variable. In

some species, such as Aeginina

longicornis or Caprella unica, both

smooth and spiny forms are found.

In others, the size of the spines varies,

with only certain spines always

being present, as in the case of

Cercops holbolli. \t has been observed

that spination is often related to

growth, so that in many species,

immature and female specimens are

spinier than adult males. However,

this relationship does not hold for all

species, nor for all specimens of

species where it is Common. It Is

therefore probable that other factors

are involved in determining the ex-

tent of spine development. Ecological,

and perhaps geographical, conditions

may prove to influence the develop-

ment of spines in certain caprellids.

Experiments need to be carried out

to ascertain what external factors

are here involved.

A second problem is lack of know-

ledge of the growth, development,

and life cycles of caprellids. Collec-

tions of caprellids in many institutions

could yield much information on

these subjects. The data are avail-

able, but need to be analysed. For

example, until more is known we

Cannot know why Caprella penantis

is larger and more robust in the

south, i.e. Florida, than in the north,

i.e. Gulf of St. Lawrence. It may be

that the north, being colder, is less

congenial to this species, affording

a shorter season for growth. How-

ever, the explanation may not be as

simple as this. In both Aeginina

longicornis) and Caprella_ septen-

trionalis, large individuals were

found at both northern and southern

extremes of the range, and the local-

ity records for small individuals (one-

third to one-half of the usual adult

size) offered no reason for their size.

Similarly, no explanation can be

given for the large female C. sep-

tentrionalis, with undeveloped brood

plates, in a collection where the

ovigerous females were half the size.

A third problem lies in the literature

of the group. It is apparent that many

species, particularly of Capre//a, are

very similar to one another. It is not

always possible to determine to

which species of a similar pair a

record refers, and many records may

be suspected of referring to both. It

should therefore be emphasized that,

for such species as C. /inearis and

C. septentrionalis, C. microtuberculata

and C. dubia, some of the references

are questionable, and synonymy lists

should be regarded only as guides to

the literature.

75

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Tengen chy Wo

ASOT A sattteget of Crectaves wet Pyenn

porite cornated inte Muaeon of Unire tv

Codege, Durwiee Oyrdter. 56 px:

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