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UNITED STATES NATIONAL MUSEUM BULLETIN 278

The Caprellidae
(Crustacea: Amphipoda)
of the
Western North Atlantic

JOHN C. McCAIN

Research Associate, Department of Oceanography, Oregon State University

SoMITHSONIAN INSTITUTION PRESS
WASHINGTON, D.C.
1968

Publications of the United States National Museum

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This work forms number 278 of the Bulletin series.

Frank A. TAYLor
Director, United States National Museum

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Contents

Page
CI NGWIGMPINGIIN 0052 a ete ae eka ee eee soa vI
ECNCNTSCTRIRSTDUO ERED ae Di pe Bs ls aed ye ne ey ee ee ene 1
ENA ONOMIC RO ChlON Se ee RSE A a as ee ek etd Sas hes 2
SAK ONOMIC COSTA CUClS meee a aie ee eee ee ee ae 2
Phisirations and MeasiremiCntes 2.12 cone nas See eeee es ase = 6
Key to the Caprellidae of the western North Atlantic__.__..-__------ 6
AeuImeuG Spr0sd DOCK (ES. 2,304) oo seas aes eo eee 8
Aeginina longicornis (Kroyer) (figs. 4-7, 54)__-------- See ae 13
Caprella andreae Mayer, new rank (figs. 8, 9, 55).------------------- 19
Coprelin Dern asin Rel mens 2 er ot eee ecamcnweas 22
Caprella danilevskii Czerniavski (figs. 10, 11, 55)--------------------- 22
Conrellacquiliora Say (fitse 12519, Oo) < ono be Sanco eca elds k- 25
Capreila inearis (Linnaeus) (figs. 14,22, 51)... ...2.-.2-4---2--.- 30
Canreila langvmanus Dulin psON.. 25.4 Sacaeee ase eo oo3. foe aes 33
Caonrella penaniis..Leacn: (es) 10, 16) Ol) 2sos-24 Leechs osu ede ees 33
(Caprelin sAnguined GOUM onsite nots aa he et ee eS 40
Caprelia tcaura: Templeton: fige. 1%, 13,00). 5 5 a ects ease 40
Caprella septentrionalis Krgyer (figs. 19-22, 51)_--------------------- 44
Caprella unica Mayer (figs. 23, 24, 55).....------------------------ 49
Dewtellacatijornica Mayer (fig. 52)i22_ 2. 22a ose eee ase lease sls 54
Deutella mayert Stebbing (figs. 25, 26, 52)__--.-_---------------- oe 54
Fallotritella biscaynensis, new genus, new species (figs. 27, 28, 53)_-_--- 58
Hemiaegina minuta Mayer (figs. 29, 30, 50).------------------------ 61
Hemiproto wigleyi, new genus, new species (figs. 31, 32c-e, 50)__-----_- 65
Luconacia incerta Mayer (figs. 33-35, 52).__------------------------ 68
Mayerella limicola Huntsman (figs. 36, 50)..__---------------------- 73
Mayerella redunca, new species (figs. 37, 38, 50)---------- Joos Laseeeis 75
Metaprotella hummelincki, new species (figs. 39, 40, 50)------------- ae 78
Paracaprella pusilla Mayer (figs. 32a—-b, 41, 42, 53)_---.-------------- 82
Paracaprella tenuis Mayer (figs. 43, 44, 53)_....--------------------- 86
Parvamous tunics 1. cumang WAV Cl cc ooo bSo coe tS sash sees 89
Phtisica antillensis (Mayer) (figs. 45, 54)____.------------ i cass asters 89
Phtisica marina Slabber (figs. 46, 47, 54, 56).._---------------------- 90
Proaeginina norvegica (Stephensen) (figs. 48, 54)--------------------- 97
Pseudaeginella antiguae Barnard__.-.-..--------------------------- 100
PE ee acl a eC ac Soe nn 100
ZOogeoeraph y=. 2-222. =... 2552.2 _- se I ee an pes fda 101
Relationship between the amphipod suborders Gammaridea and Caprel-
eaves ey a eee re re Naa 107
eG iat ie CU ae ee ee ee ee oe 112

Acknowledgments

I am indebted to many persons for study material of caprellids;
to them I offer my sincere gratitude. In particular I should like to
thank Dr. Torben Wolff (Universitetets Zoélogiske Museum, Copen-
hagen) for lending me many of Mayer’s types, without which the
value of this paper would be greatly diminished. I am grateful to
Dr. Roland Wigley (Bureau of Commercial Fisheries Laboratory,
Woods Hole) for the use of his caprellid collections by the RV Gosnold
from the entire Atlantic coast of the United States. These provided
invaluable information on the distribution of many species and
contributed several new records and a new genus. To Dr. P. Wagenaar
Hummelinck (Zoélogisch Laboratorium, Utrecht) special thanks are
due for making available his collection of the virtually unstudied
West Indian caprellids. Dr. E. L. Bousfield (National Museum of
Canada, Ottawa) has aided a great deal by lending me his collection
from the Atlantic coast of the United States and by making helpful
suggestions during this research.

Drs. T. E. Bowman, H. H. Hobbs, Jr., and F. A. Chace, Jr. of
the Division of Crustacea at the Smithsonian Institution were es-
pecially helpful by reviewing the manuscript and acting as members
of my consultative committee at The George Washington University.

This study was made possible by a grant from the Smithsonian
Institution Research Foundation and by a grant-in-aid-of-research
from the Society of Sigma Xi.

vI

The Caprellidae (Crustacea: Amphipoda) of
the Western North Atlantic’

Introduction

The most important publications on the systematics of the caprel-
lids are the three monographic treatments by Paul Mayer (1882,
1890, 1903). These indispensable works summarized all that was
known of caprellid taxonomy at that time, including many western
Atlantic species. Unfortunately, Mayer’s work complicated caprellid
taxonomy by recognizing in some species, such as the composite
species Caprella acutifrons, large numbers of varieties which appear
by modern concepts to be full species. In other instances he took a
rather narrow view of species.

Although caprellid amphipods are abundant and familiar members
of the marine benthos, western Atlantic species have received only
limited and superficial study by taxonomists. Seventeen valid species
have been reported from this area. Holmes (1905) briefly characterized
the five species known from New England and Kunkel (1918) did the
same for four Connecticut species, including one species not mentioned
by Holmes. Recently, Steinberg and Dougherty (1957) reported nine
species from the Gulf of Mexico, one of which was new. In my paper
(1965) five species occurring in Virginia waters are reviewed. Both
Pearse (1908) and Stebbing (1895) described new species from the
West Indies and Huntsman (1915) described a new species from the
Bay of Fundy. Including the papers mentioned above, some 50 pub-
lications deal with the western Atlantic caprellids, the majority being
faunal lists of local areas.

This paper deals primarily with the systematics of the Caprellidae
occurring from the tropical to boreal areas of the western North
Atlantic, roughly from the Equator to Nova Scotia. The paucity of
available material from the east coast of South America prevents
consideration of species south of the Equator. North of Nova Scotia
the caprellid fauna changes abruptly and it is, therefore, desirable to
defer treatment of the caprellids from this area until the Arctic caprel-
lid fauna can be considered in its entirety. In all, 28 species of caprellids
are treated in this paper with 2 new genera, 4 new species, and 1 new
rank described.

1 Modified from a dissertation submitted to The George Washington University
in partial satisfaction of the requirements for the degree of Doctor of Philosophy.

1

2 U.S. NATIONAL MUSEUM BULLETIN 278

This work is based primarily on the collections of the Division of
Crustacea, Smithsonian Institution. Extensive unidentified collections
were obtained from the National Museum of Canada, Woods Hole
Marine Biological Laboratory, University of Cape Town, Zodlogische
Laboratorium in Utrecht, Duke University Marine Laboratory, and
the Smithsonian Oceanographic Sorting Center. In addition, I was
able to spend 4 months collecting along the Gulf of Mexico and south-
eastern Atlantic coasts of the United States. Through the cooperation
of the U.S. Coast Guard, I was allowed to accompany the USCGC
Madrona (buoy tender) on a cruise servicing buoys along the Virginia
and North Carolina coasts. These buoys yielded large numbers of
several species which aided in the study of intraspecific variation.

‘Taxonomic Section
Taxonomic Characters

Mayer (1882, 1890, 1903) usually used 11 characters to delineate
caprellid genera. These were the number of articles in the flagellum of
antenna 2, the presence or absence of swimming setae on antenna 2,
the number of articles in the mandibular palp and the setal formula for
the terminal article,the number of gill pairs, the number of appendage
pairs of both the male and female abdomens, the number of articles
in pereopods 3-5, the number of gill pairs, and the length ratio of the
inner and outer lobes of the maxilliped. Occasionally he resorted to
other characters such as the fusion of pereonites 6 and 7 in Meta-
protella and Orthoprotella. This paper adds the position of the insertion
of pereopod 5 and the presence or absence of a molar on the mandible.

Body spination varies considerably within the same species and
its value as a specific character is questionable. In Aeginina longicornis
this variation is quite pronounced and has caused a considerable
proliferation of names for what appear to be only infrasubspecific
variants. Harrison (1940) found that body spination did not appear
on Pseudoprotella phasma before the 10th instar, which lends support
to my opinion that body spination is a questionable specific character.
It should be noted that those species which are spinose are frequently
covered with large amounts of detritus. Body spination may, there-
fore, offer some protective advantage and could possibly be correlated
with predatory pressure.

The peduncle of antenna 1 is a useful character for the delineation
of some species. The presence of setules sometimes distinguishes males
of Caprella linearis (fig. 14b) from other related species. Inflation of the
peduncular articles is exhibited in several species and is quite useful
for the separation of Caprella andreae from other members of the
Caprella acutifrons group.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 3

The number of articles in the flagellum of antenna 1 varies consider-
ably, depending upon the size and sex of the individual. The greatest
number of articles may be useful for characterizing some species with
an unusually long flagellum. The relative lengths of the proximal and
distal flagellar articles have been used to characterize some species but
probably do not have generic significance. Mayer divided Caprella
scaura into groups on the basis of the number of fused proximal articles
of the flagellum; however, in most species this number is proportional
to the size of the individual and is of little value.

Most caprellid genera have 2 articles in the flagellum of antenna
2; however, some genera such as Phtisica and Hemiproto have more
than 2. Phtisica lacks a molar on the mandible. The lack of a molar
and the multiarticulate flagellum on antenna 2 may be correlated
with the fact that Phtisica is frequently found in plankton samples and
therefore subject to dietary habits different from those of benthic
forms.

Mayer (1903, p. 47) used the presence of swimming setae on antenna
2 to characterize TZritella. Later Dougherty and Steinberg (1953)
described Tritella tenuissima which lacked swimming setae, and they
expressed the opinion that the presence or absence of swimming setae
was not a good generic character. Since 7. tenwissima shares more
characters with Triliropus (p. 57) than with the members of 77itella,
Mayer’s belief that the presence of swimming setae is a valid generic
character seems justified.

The mouthparts offer, in my opinion, some of the best, although
virtually neglected, taxonomic characters. The mouthparts reflect
feeding habits and thereby, at least to some extent, the niche of an
organism. Mayer (1903, p. 13) admitted that he neglected the mouth-
parts except for the proportions of the maxillipedal lobes and the
mandibular palp. Regrettably, he (1890, p. 107; 1903, p. 73) believed
that the mouthparts of the Caprella species were all quite similar and of
little value in classification. Most of the mouthparts of Caprella
species are similar; however, the lacinia mobilis of the right mandible
offers a useful character for subdividing this large genus. The left
lacinia mobilis is usually 5-toothed apically, while the right lacinia
mobilis may be either 5-toothed, serrate, or smooth. Phtisica and
allied genera have developed ? several accessory plates in addition to
the lacinia mobilis, and these genera usually lack a molar (fig. 47i-j).
This unusual type of mandible undoubtedly reflects genetic relation-
ships and will probably form a basis for separation of the Caprellidae
into subfamilies or other higher taxa.

2 Assuming that the caprellids arose from a podocerid type gammaridean having
a typical mandible with incisor, lacinia mobilis, setal row, and molar.

4 U.S. NATIONAL MUSEUM BULLETIN 278

As stated above Mayer used the mandibular palp as a generic
character. Those caprellids which lack a mandibular palp such as
Caprella have considerably more setation on the mouthparts, antennae,
and gnathopods 1. This increase in setation could compensate for the
loss of the cleaning ability of the mandibular palp. Very little is known
of the dietary habits of those forms which bear a mandibular palp and
only a little more is known of those that do not. The setal formula for
the palp refers to the number of long, intermediate, and short setae on
the terminal article. For example, the setal formula 1-x-y-1 indicates
the presence of 1 long seta at either end of a row of a variable number
of short setae (x) and also a variable number of intermediate setae (y).

In addition to the mandible the maxilliped also offers some ne-
glected characters which may be of generic significance. The distal
margin of the inner lobe varies in shape from rounded to flattened
and may bear a variety of tooth types, spines, and setae. As examples
of the extremes of inner lobe diversity, the paired inner lobes of
Phtisica and allied genera are as large as the outer lobes, almost com-
pletely fused, and are armed with several unusually large teeth (fig.
47h) while the inner lobes of Paracaprella are much smaller than the
outer lobes, not fused, and bear only a few setae (fig. 43e).

The outer lobe of the maxilliped shows considerable variation in
spination, serration, and setation. Such characters as the subterminal
notch in Luconacia (fig. 35d) or the large serrations on the medial
margin in Mayerella (fig. 37e) may have generic importance. However,
in too few of the caprellid species have maxillipeds been adequately
illustrated and described to permit evaluation.

The palp of the maxilliped offers several characters which may
possibly be of generic significance. Schurin (1935) used the reduction
of the dactylus as one of the characters to separate his new genus,
Eugastraulaz, from the genus Caprella; however, the value of this
character is questionable. In Paracaprella and Deutella the distal end
of the terminal article bears several large setae (fig. 43e). These large
setae are not present in most other caprellid genera. Another character
which may have generic importance is the presence of a distal pro-
jection on the penultimate article as in Paracaprella (fig. 43e).

The number of spines on the outer lobe of maxilla 1 varies among
some genera. In Phtisica there are 6 spines (fig. 47f) whereas in Caprella
there are 7 spines (fig. Se). Again it is regrettable that this character
has not been examined in enough genera to comment on its value.

Gnathopod 1 has several characters which may be of generic or at
least specific value. In the western Atlantic species, the number of
grasping spines on the propodus varies from 1-5 (compare figs. 38d,
8h, 47c) and seems to be fairly constant within each genus. Another
character of gnathopod 1 which may prove to be important is the

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 5

presence or absence of serrations on the grasping margins of the
dactylus and propodus. In Paracaprella both margins are serrate
(fig. 41h) whereas in contrast, those of Hemiaegina are completely
smooth (fig. 30b).

The shape and ornamentation of gnathopod 2 has long been used
as a specific character in the Amphipoda, and it is needless to comment
on it here except that one must take into consideration the degree of
variability this appendage shows at different growth stages. Mayer
used the term “poison tooth” to refer to the large tooth on the palm
of the propodus. There is evidence that glandular material is present
near this tooth, and it appears to be venomous in nature (Wetzel,
1932, p. 387). I have used the term grasping spine when the major
“tooth” of the propodus is a spine and have restricted the use of
poison tooth to an eminence which is not delimited at its base or
which has previously been designated a poison tooth. Usually grasping
spines occur in pairs and when closed the tip of the dactylus fits
between them. These spines are found on the gnathopods and the
pereopods.

The number of gill pairs was used by Mayer as a generic character.
Undoubtedly this is an important character but perhaps too much
value is placed on it since the gills show various stages of reduction.
Some genera with 3 pairs of gills show a closer relation to genera
with 2 pairs than to other genera with 3 pairs, as for example Dodecas
and Dodecasella.

The pereopods on pereonites 3-5 are reduced in many caprellid
genera. Although the number of articles of these rudimentary append-
ages is presently important for generic identification, it is often
difficult to count the articles, particularly when the terminal article
is small or shows some degree of fusion with the penultimate article.
Since these appendages are rudimentary and show all degrees of
reduction, their value as a generic character is questionable. In
Mayerella redunca (p. 75) a female has 2 articles in pereopod 5 instead
of the usual 3 and there can be no question that this specimen belongs
to this species. The use of these pereopods as a generic character
seems to mask the phylogenetic relationships of the genera, and it is
my belief that the mouthparts provide a better concept of relation-
ships. It should be noted that I follow the system of naming the
pereopods according to the pereonites upon which they occur; i.e.,
pereopod 3 occurs on pereonite 3. This practice has not been consist-
ently followed in the past; various authors (Barnard, Briggs, Guiler,
and Huntsman) preferred to number these appendages beginning
with pereonite 3.

The abdomen presents one of the most difficult characters to use
for identification. It is extremely small and hard to illustrate accu-

6 U.S. NATIONAL MUSEUM BULLETIN 278

rately. Unfortunately, it has been one of the most important charac-
ters and the correct generic determination usually depends upon
elucidating its structure. Mayer stressed the importance of the
abdomen by separating closely related genera such as Deutella and
Luconacia primarily by differences in the abdomen. Mayer’s emphasis
on this character is justified; however, due to its vestigial nature it
suffers from the same criticism as pereopods 3-5. In dealing with the
many stages of reduction of the appendages on the abdomen, Mayer
was inconsistent in what he considered to be a ‘‘Klappe” or vanished
appendage. This is especially true in those genera which do not
bear true appendages but which have several setae or even a single
seta borne on a type of flap or lobe. For an example of this, compare
Mayer’s (1903) figures of the abdomen of the Triliropus male (pl. 9
fig. 70), which he says bears one-half pair of appendages, with that
of the Pseudoproto male (pl. 9 fig. 52) which he claims to be without
appendages. Both abdomens have lobes with several setae; therefore,
due to this inconsistency I have refrained from using Mayer’s termi-
nology of one-half appendage pairs but have instead given the number
of recognizable appendages and have described the lobes.

Illustrations and Measurements

Illustrations of the whole mounts were made by the use of a micro-
projector and those of dissected appendages with a camera lucida.
Pencil sketches were first made which were later copied on Ethulon
tracing film. All scales on the figures equal 1 mm for the whole mount.

Measurements of the total length refer to the length of a line
drawn from the anterior portion of the cephalon between the insertions
of antennae 1 and 2, through the midlateral portion of each pereonite,
to the posterior tip of the abdomen.

Key to the Caprellidae of the Western North Atlantic

(See figure 1 for explanation of characters.)

1. Mandible with palp or setae representing vestige of palp ...... 2
Mandible without palp 2. a 4, <3. 4 /o 6 Je a0 Sine one See 17
2.(1). -Pereopods Sand 4:absent.. 24. so eGe ow sw ee ee ee oe ee eee 3
Pereopods 3:and:4 present .. ..2Gs b Bids) ois eee Sil ee 6
3.(2) Abdomen with only pair of lobes . . Pseudaeginella antiguae (p. 100)
Abdomen withappendages oo. 3. 3 ous Sw ss els ge sees 4

4.(3) Abdomen with pair of appendages and pair of lobes.
Aeginella spinosa (p. 8)
Abdomen with 2 pairs of appendages. . ........2-.2++-se-. 5
5.(4) Abdomen with only 2 pairs of appendages.
Proaeginina norvegica (p. 97)
Abdomen with 2 pairs of appendages and pair of lobes.
Aeginina longicornis (p. 13)

6.(2)
7.(6)
8.(6)
9.(8)

10.(9)

11.(9)

12.(11)

13.(12)

14.(12)

15.(8)

16.(15)
17.(1)
18.(17)

19.(18)

20.(19)

21.(18)

22.(21)

23. (22)

CAPRELLIDAE OF WESTERN NORTH ATLANTIC i

Pereopods: 3 and 4 I-segmented’ 5 gc. 5 «6 kee SG Awe ed
Pereopods 3 and 4 2- or 6-segmented. ............. =. 8
Pereonites 6 and 7 fused. . . . . . Metaprotella hummelincki (p. 78)
Pereonites 6 and 7 not fused. . . . . Fallotritella biscaynensis (p. 58)
Pereopods 3 and 4 2-segmented. . . . .......42++42. 9
Pereopods 3 and 4 6-segmented. . .........4.2.+.442.2.2.-. 15
Pereopod 5 2- or 3-segmented .........4...2...... 10
Pereopod 5 6-segmented. . . . . Stace: dau ak oe CEE

Abdominal appendage of male ae ei in donors. 6 bones
Mayerelia limicola (p. 73)
Abdominal appendage of male much longer than penes, recurved at tip.
Mayerella redunca (p. 75)
Pereopod 5 inserted near midlength of pereonite 5.
Luconacia incerta (p. 68)
Pereopod 5 inserted in posterior part of pereonite5........ 12
Mandibular palp reduced; when 3-segmented, terminal article minute. 13
Mandibular palp not reduced, 3-segmented, terminal article not minute.
14
Mandibular palp represented by single seta; males with large triangular
projection on anteroventral margin of pereonite 2, basis of gnathopod
2 with proximal knob on posterior margin.
Paracaprella pusilla (p. 82)
Mandibular palp represented by knob with seta or with several reduced
articles; males with small projection on anterolateral margin of pereonite
2, basis of gnathopod 2 without proximal knob.
Paracaprella tenuis (p. 86)
Cephalon with dorsalspine . . .. . . . Deutelia californica (p. 54)
Cephalon without dorsal spine. . . . . . . . Deutella mayeri (p. 54)
Abdomen of male with 2 pairs of appendages, females with 1 pair.
Hemiproto wigleyi (p. 65)
Abdomen of male with 3 pairs of appendages, females with 2 pairs . . 16
Carpus of gnathopod 2 longer than merus . . Phtisica antillensis (p. 89)

Carpus of gnathopod 2 shorter than merus. . . Phtisica marina (p. 91)
Pereopods 3 and 4 I-segmented . . . . . Hemiaegina minuta (p. 61)
PeredpoGs: o tidied BHGeite we es once ah tec agar a ice er ce. er we
Pereopods 5-7 without grasping spines ............. 19
Pereopods 5-7 with grasping spines... . ee ak, & “ad ee

Cephalon with anteriorly directed triangular Poeun:
Caprella penantis variant (p. 35)
Cephalon without anteriorly directed projection ... ana & 20
Abdomen of male with hooked papillae at tip of cppen dare: female with
small palplike appendage. . . . . . . Caprella danilevskii (p. 22)
Abdomen of male without hooked papillae on appendage, female without
palplike appendage .. . . . . . . Caprella unica (p. 49)
Cephalon with large anteriorly directed dorsal spine or projection . . 22
Cephalon without dorsal spine or with nonanteriorly directed spine . . 24
Cephalon with sharp anterodorsally directed spine, males with up to 9
fused articles in flagellum of antenna 1. . . Caprella scaura (p. 40)
Cephalon with anteriorly directed triangular projection, males with less
than 4 fused articles in flagellum of antenna . . . bate a Ao
Palm of propodus of pereopods 5-7 convex. . Caprelln gadvene (p. 19)
Palm of propodus of pereopods 5-7 concave . Caprelia penantis (p. 33)

8 U.S. NATIONAL MUSEUM BULLETIN 278

24.(21) Ventral spine present between insertions of gnathopods 2.
Caprella equilibra (p. 25)
Ventral spine not present between insertions of gnathopods 2. . . . 25
25.(24) Propodus of gnathopod 2 with small spine on inner surface near poison
tooth ..... soe eh toe, GO
Propodus of eaaihopod 2 yathout ‘amma eemen on inner surface near
poison tooth .... . . Caprella equilibra variant (p. 29)
26.(25) Ratio of total length to feet aE basis of gnathopod 2 greater than 13.0;
dorsal surface of pereonites 1—4 usually spinose, cephalon with at least 1
large spine. . . . . . Caprella septentrionalis (p. 44)
Ratio of total length io lever of basis of gnathopod 2 smaller than 13.0;
dorsal surface of pereonites 1-4 usually smooth, cephalon infrequently
with small spine. . ........ . . Caprella linearis (p. 30)

Aeginella Boeck, 1861

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 3-segmented, setal formula for terminal article
1-x-1, molar present; outer lobe of maxilliped larger than inner lobe;
gills on pereonites 3 and 4; pereopods 3 and 4 absent, pereopod 5,
6-segmented; abdomen of male and female with pair of appendages
and pair of lobes,

Type-species: Aeginella spinosa Boeck, 1861 (by monotypy).

Remarks.—The genus Aeginella is very closely related to Aeginina
and Proaeginina, differing from them only by the structure of the
abdomen. There is a gradation from 1 pair of appendages with lobes,
2 pairs of appendages, to 2 pairs of appendages with lobes in the
series Aeginella—Proaeginina—Aeginina.

Aeginella spinosa Boeck, 1861
Fieures 2, 3, 54

Aeginella spinosa Boeck, 1861, pp. 673-674; 187la, pp. 272 (192)-273 (198);
1873-76, pp. 684-686, pl. 32, fig. 4.—M. Sars, 1863, pp. 290-291.— Mayer,
1882, p. 36; 1890, pp. 36-37, pl. 1, fig. 24, pl. 5, figs. 30-33; 1903, p. 61.—
G. Sars, 1886, pp. 70, 89; 1895, pp. 653-654, pl. 235, fig. 1—Norman, 1886,
p. 26; 1905a, p. 26.—Hansen, 1887b, pp. 172-173; 1895, p. 130.—
Vanhéffen, 1897, p. 213.—Nordgaard, 1905, p. 185.—Stephensen, 1913a,
pp. 222-223; 1916, p. 295; 1929a, p. 178, fig. 332; 1929b, pp. 19, 34; 1933,
pp. 60, 77; 1935, p. 118; 1940, p. 69; 1942, pp. 429-430, 502, 503; 1944a,
pp. 48-49, map 9; 1944b, pp. 135-136, 148, 159, 162.—Derjugin, 1915, pp.
453, 456.—Schneider, 1924 (1926), p. 59.—McCain, 1966, p. 92.

DraGnosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.

DescripTrion.—Body rather robust, spinose; cephalon with anterior
projection, separated from pereonite 1 by distinct suture. Pereonite
1 with dorsal anterior spine. Dorsal surface of pereonite 2 with pair
of spines at midlength of pereonite and single posterior spine, inser-
tion of gnathopod 2 with small spine, anterolateral margin produced

CAPRELLIDAE OF WESTERN NORTH ATLANTIC

a Cephal on at
ni Si <
™. dA
\ \
—
Pre

ny H ng
: wy Yt i i)
He \ :
a -
/

Md palp
oot ay

See se

Brood pouch palp
LI 3 ——— Gill
Pn3s— SF _ d= oy) outer lobe
ee =
AAO SEPP 5
Ke RY
Pn4

appendages
o lobes
Abd

ay inner lobe

Mzx2

Md

Figure 1.—Generalized caprellid: 4 (antenna), Abd (abdomen), G (gnathopod), Md
(mandible), Mx (maxilla), Mxp (maxilliped), P (pereopod), Pn (pereonite).

10 U.S. NATIONAL MUSEUM BULLETIN 278

into triangular projection. Dorsal surface of pereonite 3 with pair
of anterior spines, sometimes reduced to small humps, pair of mid-
posterior spines, and single posterior spine; anterolateral margin
produced as in pereonite 2; lateral margin of pleura with anterior
and posterior spines and spine above gills in males, females without
posterior spine. Pereonite 4 similar to pereonite 3 in males, females
without dorsal anterior spine. Dorsal surface of pereonite 5 similar
to pereonite 3, pleura with only anterior spine and dorsally directed
spine at base of pereopod 5. Pereonite 7 with dorsally directed spine
at base of pereopod 7. Length of largest male 20 mm, female 16 mm.

Setal formula for terminal article of mandibular palp 1-10-1 to
1-12-1. Left mandible with 5-toothed incisor, 5-toothed lacinia
mobilis, setal row of 3 serrate setae, molar with plumose seta. Right
mandible with 5-toothed incisor, lacinia mobilis serrate on cutting
margin, setal row of 2 serrate setae, molar with plumose seta. Palp
of maxilla 1 usually with 5 robust apical spines and several setae;
outer lobe with 7 apical spines, usually bifid but sometimes more
branches with increase in size of individual. Inner and outer lobes
of maxilla 2 quite setose on apical margin and spines occasionally
present. Outer lobe of maxilliped with 2 apical setae, 1 long apical
spine, and up to 12 smaller marginal spines; inner lobe with 2 small
spines and up to 12 apical setae, as many as 9 of which plumose;
palp similar to that of Caprella.

Propodus of gnathopod 1 triangular with 2 proximal grasping
spines, grasping margin not distinctly serrate; grasping margin of
dactylus serrate, particularly at tip. Propodus of gnathopod 2 quite
robust, palm heavily setose with small proximal tooth, anterior margin
with distal projection; dactylus not serrate.

Gills subelliptical.

Pereopods 3 and 4 absent. Pereopods 5-7, 6-segmented, palm of
propodus with pair of proximal grasping spines.

Abdomen of male and female with 1 pair of appendages and pair
of setose lobes; in male appendage placed on raised projection and
uniarticulate; in female, appendage neither on projection nor
articulated.

VARIATION.—This species appears to be quite constant in body
spination with the exception of the first pair of spines on the dorsal
surface of pereonite 4. These may be present as fully developed
spines or as only small humps.

Distripution.—Type-locality: Haugesund, Norway.

Other records: Murman coast to Haugesund on the Norwegian
coast, Spitsbergen, Faeroe Islands, Iceland, east and west coasts
of Greenland.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 11

Ficure 2.—Acginella spinosa, male; a, lateral view; 6, right mandible; c, inner and outer
lobes of maxilliped; d, maxilla 2; ¢, abdomen; f, maxilla 1; g, gnathopod 1; h, terminal
article of mandibular palp.

New records: Off Nova Scotia, 44°01’ N., 59°02.5’ W., 43°03’ N.,
65°30’ W., and on the Banquereau Banks; off Cape Cod, 42°25’ N.,
66°05’ W.

Remarks.—This species is an Arctic one usually found in deeper
water, to 1026 m. Its distribution is comparable to that of Pro-
aeginina norvegica and Aeginina longicornis, the latter ranging far-

279-475—68——2

12 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 3.—Aeginella spinosa, female; a, lateral view; b, right mandible; c, maxilliped; d,
gnathopod 1; ¢, maxilla 1; f, coxal plate of gnathopod 2; g, abdomen; h, gnathopod 2.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 13

ther south along the western Atlantic coast and usually being found
in shallower water.

Mayer (1903, p. 61) reported Aeginella spinosa from the asteroid,
Brisinga, and this species has also been collected from red and brown
algae and hydroids.

Aeginina Norman, 1905

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandible with 3-segmented palp, setal formula for terminal article
1—x-1 or 1—x-1-x, molar present; outer lobe of maxilliped larger
than inner lobe; gills on pereonites 3 and 4; pereopods 3 and 4 absent,
pereopod 5, 6-segmented; abdomen of male and female with 2 pairs
of biarticulate appendages and 1 pair of lobes.

Type-species: Aegina longicornis Krgyer, 1842-43 (by monotypy).

Aeginina longicornis (Kréyer, 1842-43)
Ficurss 4-7, 54

Aegina longicornis Krgyer, 1842-48, pp. 509-515, pl. 7, figs. 1-12; 1846, pl. 24,
fig. 3—Boeck, 1871la, p. 270 (190); 1873-76, pp. 677-679.—Liitken, 1875,
p. 159.—Mayer, 1882, pp. 83-84, fig. 11, pl. 5, figs. 6-10; 1890, pp. 32-
35, pl. 5, figs. 27-29, pl. 6, figs. 9, 28; 1903, pp. 60-61.—Hansen,1887b,
p. 171—Norman, 1886, p. 26; 1905a, p. 26—Stuxberg, 1882, p. 764.—
Vanhoffen, 1897, p. 213—d’A. Thompson, 1901, p. 41—M. Rathbun,
1905, pp. 7, 76.—Briiggen, 1907, pp. 237-238.—Stephensen, 1913a, pp.
220-222; 1929b, p. 34.—Bousfield, 1958, p. 315.

Aegina spinosissima Stimpson 1854 (1853), pp. 44-45.— Miers, 1877a, pp. 104-
105.—Norman, 1882, pp. 671, 684; 1886, p. 26; 1905a, p. 26.—Koelbel,
1886, p. 42.—Hansen, 1887a, p. 233; 1887b, p. 172.—Ohlin, 1895a, pp.
xvii, xix, 60-62.—Vanhoffen, 1897, p. 213.—Whiteaves, 1901, p. 220.—M.
Grieg, 1907, p. 551.—Calman, 1927, p. 42 (fig. 27).

Caprella spinifera Bell, 1855, pp. 407-408, pl. 35, fig. 2.—Goés, 1866, p. 535.

Aegina (Caprella) echinata Boeck, 1861, pp. 670-672.

Aegina laevis Boecx, 1861, pp. 672-673; 1871a, p. 272 (192); 1873-76, pp. 682-
684, pl. 32, fig. 9.

Caprella spinosissima Bate, 1862, pp. 361-362, pl. 57, fig. 3.

Aegina echinata.—Boecx, 187la, pp. 271 (191)-272 (192); 1873-76, pp. 680-
682, pl. 32, fig. 6.—Liitken, 1875, p. 159.— Mayer, 1882, pp. 34-35.—Stux-
berg, 1882, pp. 764, 780; 1887, p. 73.—G. Sars, 1895, pp. 651-652, pl. 234,
fig. 2.—Stephensen, 1927a, pp. 147-148; 1928, p. 389, fig. 93 (5-7); 1929a,
p. 178, fig. 331.—Gurjanova, 1929a, pp. 40-41, 46.

Aegina spinifera.—Buchholz, 1874, pp. 270, 388.—G. Sars, 1885, pp. 228-230,
pl. 18, fig. 5; 1886, pp. 70, 89.—Ives, 1892, p. 481.—Klinckowstrém, 1892,
p. 91.

Aegina Echinata.—Meinert, 1877-78, p. 168.

Aegina longicornis f. nodosa Mayer, 1890, p. 33, pl. 5, fig. 29.

Aegina longicornis f. typica Mayer, 1890, p. 33.

Aegina longicornis f. spinifera Mayer, 1890, pp. 33-34.—Gurjanova, 1935, p. 78.

Aeginella spinosissima.—Mayer, 1890, p. 37; 1903, p. 61.—Ortmann, 1901,
pp. 154-155.—Stephensen, 1912, pp. 543-544; 1913b, p. 68.

14 U.S. NATIONAL MUSEUM BULLETIN 278

Aegina longicornis f. spinigera.—Hansen, 1895, p. 130.

Aeginella longicornis.—Holmes, 1904 (1905), pp. 525-526.—Paulmier, 1905, p.
169, fig. 39.—Sumner, Osburn, and Cole, 1911 (1913), pp. 132, 134, 135, 656,
chart 102.—Kunkel, 1918, pp. 175-176, fig. 53.—Allee, 1922, pp. 57, 58.—
Dexter, 1944, p. 356.—Ferguson and Jones, 1949, p. 442.

Aegina longicornis nodosa.—M. Rathbun, 1905, pp. 7, 76-77.

Aegina longicornis spinifera.—M. Rathbun, 1905, pp. 7, 77.

Aegina longicornis spinosissima.—M. Rathbun, 1905, pp. 7, 77.

Aegina langicornis.—Briggen, 1909, pp. 42-43.

Aeginina longicornis—Norman, 1905a, p. 46.—Stappers, 1911, pp. 74-76.—
Shoemaker, 1930, p. 352 (134).—Procter, 1933, p. 256.—Stephensen, 1933,
pp. 59-60, 77; 1940, pp. 69-70; 1942, pp. 430-431, 502, 503; 1944a, p. 49,
chart X; 1944b, pp. 135, 148, 159, 162.—Gurjanova, 1936, pp. 568, 580, 588,
589; 1964, p. 313.—Elton, 1937, p. 433.—Dunbar, 1954, pp. 784, 788.—
Bousfield, 1958, p. 322.—McCain, 1965, pp. 191-192, fig. 1a; 1966, p. 92.—
Cerame Vivas and Gray, 1966, p. 263.

Draenosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.

Description.—Body spination variable, smooth to quite spiny;
cephalon separated from pereonite 1 by suture. Length of largest
male 54 mm, female 34 mm, smallest ovigerous female 9 mm.

Antenna 1 usually longer than body, flagellum with up to 26
articles. Antenna 2 setose and usually shorter than articles 1 and 2 of
antenna 1.

Mouthparts quite similar to those of typical Caprella (p. 18),
lacinia mobilis of right mandible not distinctly 5-toothed but with
several teeth and serrations.

Propodus of gnathopod 1 with pair of grasping spines, grasping
margins of dactylus and propodus only slightly serrate. Propodus of
enathopod 2 with proximal poison tooth and distal notch, tooth, and
rectangular projection, anterodistal margin with triangular projection;
basis and ischium with anterodistal projections; carpus with postero-
distal projection.

Propodus of pereopods 5-7 with pair of proximal grasping spines.

Abdomen of male and female with 2 pairs of biarticulate append-
ages and pair of lobes, medial margin of appendages with numerous
minute knobs.

VariaTion.—The degree of spination of the body is variable. The
most spiny form is illustrated in figure 4j, and there are various
degrees of spination; some are almost smooth. Spination seems to
vary from spinose in northern waters to smooth in southern; however,
spiny forms have been found infrequently in the southern part of
the range of this species.

Figure 6 illustrates the relationship of pereonite length to total
body length. The solid black lines represent individuals, and the
dashed lines indicate the linear relationship between pereonite length

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 15

Ficure 4.—4eginina longicornis, male; a, lateral view; b, right mandible; c, left mandible;
d, maxilla 1; ¢, terminal article of mandibular palp; f, abdomen; g, maxilliped; h, gnathopod
2; 1, gnathopod 1; 4, diagramatic representation of dorsal (middle) to lateral (edges) body
spination, diameter of circle proportional to length of spine.

16 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 5.—Aeginina longicornis, female; a, lateral view; b, gnathopod 2; ¢, gnathopod 1;
d, abdomen..

and total length. It is evident that pereonites 1 and 2 of both males
and females increase in length at approximately the same rate. There-
fore the statement made by many authors that pereonites 1 and 2
are elongated in males seems to be invalid.

Distripution.—Type-locality: Near Frederikskiib, Greenland, at a
depth of 22-29 m.

Other records: Siberian Polar Sea to 140° E.; Kara Sea; Novaya
Zemlya; Franz Josef Land; Spitsbergen; Murmansk; Barents Sea;
Norway; Denmark; Faeroe Islands; Shetland Islands; Jan Mayen;
Iceland; eastern and western coasts of Greenland; Baffin Bay; east
coast of North America from Newfoundland to Oregon Inlet, North
Carolina.

New records: No records are available which extend the range of
this species.

Remarks.—This Arctic species is quite common in the northern
parts of eastern North America. It is generally found in deeper water
(to 2258 m) but has been collected frequently in shallow water. The
habitat does not seem to be specific because it has been collected
from green, red, and brown algae; sea grass; hydroids; bryozoans;
and from the gut of the sea bass, Centropristis.

The seasonal distribution of ovigerous females is illustrated in
figure 7. The largest number of samples containing ovigerous fe-

CAPRELLIDAE OF WESTERN NORTH ATLANTIC bz

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Cumulative pereonite length (mm)

Ficure 6.—Aeginina longicornis, cumulative pereonite length plotted against total length.
Roman numerals indicate pereonites; I includes both pereonite 1 and cephalon. Solid
horizontal lines represent individuals.

males was collected during August and September and the breeding
season appears to extend from April to December. No ovigerous
females were present in samples taken from January to March.
Whether this is due to the fact that the caprellids do not breed during
this period or simply that adequate samples were not taken, is not
known.

The varieties of this species which are found along the east coast
of North America do not appear to be geographically or bathymetri-
cally isolated from each other and so they probably do not represent
subspecies. These varieties may represent the phenotypic expression
of different degrees of spination influenced by such parameters as
substrate, breeding season, or diet; however, such data are not
available to me.

18 U.S. NATIONAL MUSEUM BULLETIN 278

50
48
» 46
ca oS
3 e
» 3 8
ev La e tat * e °
= 40 ° S
Ss”
g 38 .
> 36
Q

Jan. Feb. Mar. Apr. May June ‘July ° Aug. “Sept. ° Oct. ° Nov." Dee.

Ficure 7.—Aeginina longicornis, monthly and latitudinal distribution of ovigerous females.

Caprella Lamarck, 1801

Flagellum of antenna 2 biarticulate, swimming setae usually pres-
ent; mandibular palp absent, molar present; outer lobe of maxilliped
larger or equal to inner lobe; gills on pereonites 3 and 4; pereopods
3 and 4 absent, pereopod 5, 6-segmented; abdomen of male with pair
of appendages and pair of lobes, female with pair of lobes.

Type-species: Cancer linearis Linnaeus, 1767 (subsequent designa-
tion by Dougherty and Steinberg, 1953).

Remarxs.—Mayer (1890, p. 107; 1908, p. 73) states that it is un-
necessary to study in detail the mouthparts of members of this genus
since the specific differences stand out much more clearly in other
characters. I agree with this statement, hence, I have not included
descriptions of the mouthparts other than the lacinia mobilis of the
right mandible except for those appendages which exhibit variation.
The typical mouthparts of Caprella may be characterized as follows:
Mandible with 5-toothed incisor; left mandible with 5-toothed lacinia
mobilis, right variable; setal row of left mandible with 3 serrate
setae, right with 2 serrate setae; molar present with single small
plumose seta. Outer lobe of maxilla 1 with 7 spines, palp with variable
number of spines and setae. Lobes of maxilla 2 usually densely setose.
Outer lobe of maxilliped with row of spines on medial margin and
usually covered with numerous setae; inner lobe flattened apically
with several spines and numerous simple and plumose setae; articles
of palp usually heavily setose, grasping margin of dactylus finely
toothed or serrate.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 19

The abdomen also appears to vary little in Caprella. In the males
it bears a pair of uniramous appendages at its proximal end; laterally
it has a pair of lobes. The surface of the abdomen is usually covered
with numerous setae, and occasionally the ventral surface between
the lobes is raised to form a hump. The female abdomen is similar to
that of the male except that it lacks appendages.

The propodus of ganthopod 1 is usually triangular in outline and
invariably has a pair of proximal grasping spines.

Caprella andreae Mayer, 1890, new rank
Fieurss 8, 9, 55

Caprella acutifrons [not Latreille]—van Beneden, 1859, pp. 78-81, pl. 1, figs.
9-11; 1861, p. 145.—[?] Stock and Bolklander, 1952, p. 3.

Caprella acutifrons f. Andreae Mayer, 1890, pp. 51, 55-56, pl. 2, fig. 38, pl. 4,
fig. 56, 70-71; 1903, pp. 80-81.—Chevreux and de Guerne, 1893, p. 3.—
d’A. Thompson, 1901, p. 41.—Stephensen, 1915, p. 53.—Chevreux and
Fage, 1925, p. 452, fig. 430a.—Ruffo, 1938, p. 150 [in part].—Utinomi,
1947, pp. 71-72.

Caprella acutifrons f. andreae—Stephensen, 1929a, p. 182.

Driacnosis.—Cephalon with anteriorly directed triangular pro-
jection, peduncle of antenna 1 robust in males, palm of propodus of
pereopods 5-7 convex with medial grasping spines.

Description.—Body smooth except for anteriorly directed tri-
angular projection on cephalon, pleura developed on pereonites 3 and
4 in larger males. Length of largest male 12 mm, female 9 mm,
smallest ovigerous female 7 mm.

Peduncle of antenna 1 inflated in males, sparsely setose. Antenna
2 typical of genus.

Mouthparts typical of genus, lacinia mobilis of right mandible
5-toothed.

Gnathopod 1 typical of genus, dactylus serrate, propodus with 2
proximal grasping spines. Propodus of gnathopod 2 in males with
proximal poison tooth and distal rectangular projection, palm densely
setose; in females propodus with proximal poison tooth, distal pro-
jection and small middistal projection; dactylus strong and con-
stricted medially.

Gills oval and usually quite large and inflated in males, females
elliptical.

Propodus of pereopods 5-7 with 2 grasping spines at midlength,
palm convex.

Abdomen typical of genus.

VARIATION.—The inflation of antenna 1 and the development of
pleura increase as the size of the individual increases, large males
having an unusually enlarged antenna 1 and well-developed pleura.

20 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 8.—Caprella andreae, male lectotype; a, lateral view; b, gnathopod 2; ¢, pereopod
6; d, abdomen; e, maxilla 1; f, maxilliped; g, maxilla 2; A, gnathopod 1; 7, right mandible;
7, left mandlble.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 21

Ficure 9.—Caprella andreae, female allolectotype; a, abdomen; J, lateral view; c, gnathopod 2.

Distripution.—Type-locality: 38°10’ N., 64°20’ W. (see remarks).

Other records: North Sea; Netherlands; Belgium; Portugal; between
Portugal and the Azores; Mediterranean coast of Spain; Gulf of Lione;
St. Raphael, France; Naples, Italy; Aegean Sea; 38°20’ N., 16°04’ W.;
between Tokyo and Honolulu; Sea of Japan; Korean Strait; West
coast of Kyushu, Japan.

New records: Algeria; off Casablanca, Morocco; off Martha’s
Vineyard, Mass.; Ocean City, N.J.; Cape Hatteras and Beaufort,
N.C.; off Key West, Fla.; Havana, Cuba; 43°09’ N., 151°52’ W.

Remarxs.—Mayer (1890) described this species as a variety of his
compound species Caprella acutifrons (see p. 33). This species is
composed of 20 varieties or forms, many of which should be considered
full species by modern criteria (see Dougherty and Steinberg, 1953).
C. andreae differs from the other forms of this compound species by
the convexity of the propodus of pereopods 5-7. It appears to be
ecologically isolated from the other members of the group by its
habit of usually attaching to floating objects such as driftwood, buoys,
and plants. It has also been found among the incrustations on the
backs of the sea turtles, Thalassochelys and Chelonia, which were

27, U.S. NATIONAL MUSEUM BULLETIN 278

collected in the Mediterranean Sea, off Havana, Key West, and
Beaufort, North Carolina.

Ruffo (1938) cites this species as occurring off Brazil and Guiana;
however, in personal correspondence he has advised me that he was
referring to C. acutifrons s. lato and not specifically to C. andreae.

Mayer’s specimens from his localities 3-7 were obtained from the
Copenhagen Museum. I have selected a lectotype and an allolecto-
type from locality 4, 38°10’ N., 64°20’ W.

Caprella bermudia Kunkel, 1910
Caprella bermudia Kunkel, 1910, pp. 108-110, fig. 42.

Remarxs.—Kunkel’s description of this species is inadequate to
separate it from C. equilibra. It has not been included under C. equilibra
because I have not been able to examine the type material. Inquiries
at most of the larger museums in Europe and North America have
not revealed their location.

Caprella danilevskii Czerniavski, 1868
Figures 10, 11, 55

Caprella Danilevskii Czerniavski, 1868, pp. 92-93, pl. 6, figs. 21-34.—Mayer,
1882, p. 54; 1890, pp. 58-60, pl. 5, fig. 44, pl. 7, figs. 12-13, 54; 1903, p. 99.—
Tichy, 1911, pp. 1131, 1133, 1184.—Zernov, 1913, p. 68.—Arimoto, 1930,
pp. 50-51, fig. 5.—S. Carausu, 1956, pp. 131, 132.

Caprella Danilewskii.—Sovinskii, 1880, pp. 88, 100-101.—d’A. Thompson, 1901,
p. 41.—Chevreux and Fage, 1925, pp. 454-455, fig. 432.—Ruffo, 1941, p.
125; 1946, p. 53.

Caprella inermis [not Grube] Haswell, 1880, p. 348, pl. 23, fig. 3; 1882, p. 314;
1884 (1885), p. 1000.—Mayer, 1882, p. 71, figs. 26-29; 1890, p. 75.—Oliveira,
1940, p. 189.—Guiler, 1954, pp. 532-533, fig. 1.

Caprella danilevskii.—Stebbing, 1888, pp. 1264-1267, pl. 145; 1910b, p. 653.—
Kunkel, 1910, pp. 110-111.—Zernov, 1913, p. 233.—Barnard, 1916, pp.
280-281; 1937, pp. 134, 197.—Hale, 1929, pp. 232-233, fig. 228.—Hiro, 1937,
pp. 312-313, pl. 22, fig. 6.—Utinomi, 1943a, p. 275; 1943b, p. 284, fig. 4;
1943c, p. 289; 1947, p. 73.—Edmondson and Mansfield, 1948, pp. 216-218,
fig. 8.—Stschapova, Mokyovsky, and Pasternak, 1957, p. 87.

Caprella Danilevkit.—Monterosso, 1915, pp. 15-16.

Caprella danilewskii.—Carausu and Carausu, 1942, p. 82, fig. 8d.—Costa, 1960a,
pp. 99, 100.

DraGcnosis.—Propodus of pereopods 5-7 with numerous setae but
lacking grasping spines; in males both pairs of gills elliptical, long
axis usually parallel to body, in female gills on pereonite 3 usually as
in males; abdomen of male with hooked papillae at tip of appendage,
that of female with small palplike appendage bearing seta at medial
base.

Derscription.—Body smooth, cephalon of large males with very
small anterior projection. Length of largest male 9 mm, female 7 mm,
smallest ovigerous female 4.5 mm.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 23

Antenna 1 and 2 typical of genus.

Mouthparts typical of genus; lacinia mobilis of right mandible with
1 tooth, apical margin smooth or minutely serrate.

Propodus of gnathopod 1 with 2 proximal grasping spines, grasping
margin of dactylus serrate. Propodus of gnathopod 2 in males elongate

Ficure 10.—Caprella danilevskii, male; a, labium; b, lateral view; c, antenna 1; d, abdomen;
e, gnathopod 2; f, gnathopod 1; g, pereopod 5; h, left mandible; i, right mandible; 4,
maxilla 2; £, maxilla 1; /, maxilliped.

24 U.S. NATIONAL MUSEUM BULLETIN 278

b

Ficure 11.—Caprella danilevskit, female; a, abdomen; b, gnathopod 2; ¢, gnathopod 1; d,
lateral view.

with posion tooth at midlength, rectangular tooth distally; dactylus
less than one-half length of propodus; in females propodus with prox-
imal poison tooth and distal rectangular tooth; dactylus more than
one-half length of propodus.

Gills 3 and 4 in males and 3 in females elliptical, long axis usually
parallel to body; gill 4 in females oval or elliptical.

Propodus of pereopods 5-7 without distinct grasping spines, palmar
margin usually with numerous stout setae; grasping margin of dactylus
serrate.

Abdomen of male typical of genus except for hooked papillae at
tip of appendage, female with small palplike appendage.

VariatTion.—The shape of the gills is variable. In males the long
axis is usually parallel to the body; however, either or both may occa-
sionally be at various angles to the body. In the females either or both
of the gills may have the long axis parallel to the body or may be at
any angle. In small males the poison tooth on the propodus of gnath-
opod 2 may be more proximal than midlength; however, the dactylus
remains quite short. The palm of the propodus of pereopods 5-7
varies in setation from numerous stout setae to very few.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC ZO

Disrrisution.—Type-locality: Black Sea.

Other records: Bay of Biscay; Mediterranean coast of France;
Mediterranean and Adriatic coasts of Italy; Sicily; Ukranian and
Roumanian Black Sea; Cherchell, Algeria; Rufisque, Senegal; South
Africa; South Arabian coast; Bermuda; Rio de Janeiro, Brazil; Oahu,
Hawaii; 5. Sakhaline; Pacific coast of Hokkaido and Honshu, Japan;
Amakusa Tomioka and Okinojima, Kyushu, Japan; Sea of Japan;
Korean Straits; Southeastern Australia; Coles Bay, Tasmania.

New records: Virginia Key, Key Biscayne, and Matheson Ham-
mock, Fla.; Loggerhead Key, Tortugas; St. Croix, Virgin Islands;
‘Trinidad.

Remarxs.—This species is quite widespread and pantropical in
its distribution. It has been collected on sea grass, the phaeophytes
Cystoseira and Sargassum, and the bryozoan Bugula.

C. danilevskw is easily distinguished from the other species of
Caprella in the western North Atlantic by its unusual gill shape, its
distinctive abdomen, and the short dactylus of the male gnathopod 2.

Caprella equilibra Say, 1818
FicurREs 12, 13, 55

Caprella equilibra Say, 1818, pp. 391-392.—de Kay, 1844, p. 41.—White, 1847,
p. 92.—Gibbes, 1848, p. xvi; 1849, p. 23.—Stebbing, 1888, pp. 1254-1256;
1910a, p. 466; 1910b. p. 653.—Kunkel, 1910, pp. 106-108, fig. 4.—Barnard,
1916, p. 281; 1930, p. 440; 1932, p. 300.—Schellenberg, 1928, p. 678.—
Procter, 1933, p. 256.—Edmonson and Mansfield, 1948, pp. 214-216, fig.
7.—Ricketts and Calvin, 1952, p. 68.—Dougherty and Steinberg, 1953, pp.
44, 47; 1954, pp. 170, 171.—Day and Morgan, 1956, p. 303.—Steinberg and
Dougherty, 1957, pp. 273-274, figs. 1-2.—Johnson, 1965, appendix 2, p. 2,
appendix 3, p. 4; 1966, appendix 2, p. 2.—McCain, 1965, pp. 193-194, fig. 1b,
f; 1966, p. 92.—Johnson and Juskevice, 1965, p. 39.

Caprella Januarii Kréyer, 1842-43, pp. 499-504, pl. 6, figs. 14-20.—Dana, 1853,
pp. 819-820; 1855, pl. 55, fig. 2.—Herklots, 1861, p. 43.

Caprella Esmarkit Boeck, 1861, pp. 674-675; 1871la, p. 275 (195); 1873-76, pp.
693-694, pl. 32, fig. 5.

Caprella laticornis Boeck, 1861, pp. 675-676; 1871la, p. 274 (194); 1873-76, pp.
689-691, pl. 32, fig. 10.

Caprella aequilibra.—Bate, 1862, pp. 362-363, pl. 57, fig. 5; 1887, pl. 175.—Bate
and Westwood, 1868, pp. 71-73.—Parfitt, 1873, p. 251.—Gamroth, 1878,
pp. 101-126, pls. 8-10.—Haller, 1879a, p. 232; 1879b, p. 404.— Mayer, 1882,
pp. 45-48, pl. 1, fig. 7, pl. 2, figs. 1-11, pl. 4, figs. 20-25, pl. 5, figs. 16-18;
1890, pp. 48-50, pl. 2, figs. 42-43, pl. 4, figs. 35-37, pl. 6, figs. 18a, 37; 1903,
pp. 89-92, pl. 3, figs. 29-34, pl. 7, figs. 66-69; 1912, pp. 4, 5.—Marion,
1883, p. 49.—Miers, 1884, p. 320.—Carus, 1885, p. 388.—Haswell, 1884
(1885), pp. 999-1000.—de Guerne, 1886, p. xliii—Norman, 1886, p. 26;
1905a, p. 26; 1905b, p. 85.—Thomson and Chilton, 1885 (1886), p. 142.—
Chevreux, 1887a, p. 335; 1898, p. 483; 1900, p. 120.—Barrois, 1888, pp. 58,
77.—G, Sars, 1895, pp. 663-664, pl. 238, fig. 3—d’A. Thompson, 1901,
p. 41.—Graeffe, 1902, p. 19.—Hutton, 1904, p. 261.—Marine Biol. Assoc.,
1904, p. 242; 1931, p. 198; 1957, p. 233—Norman and Scott, 1906, pp. ix,

26 U.S. NATIONAL MUSEUM BULLETIN 278

99.—Scott, 1906, p. 175.—Sinel, 1906 (1907), p. 222.—Tichy, 1911, p. 11384.—
Thomson, 1913, p. 245.—LaFollette, 1914, pp. 224-225, pl. 5.—Briggs,
1914 (1915), pp. 79-80.—Kunkel, 1918, pp. 180-181.—Thomson and Ander-
ton, 1921, p. 113.—Galdiano, 1924, p. 392.—Chevreux and Fage, 1925,
pp. 455-456, fig. 433.—Schellenberg, 1926, p. 470.—Johnson and Snook,
1927, pp. 280-281, fig. 235.—Stephensen, 1927a, p. 150; 1927c, p. 355;
1928, p. 386, fig. 92 (13); 1929a, pp. 180-181, figs. 43-336; 1929b, p. 34;
1942, pp. 439, 502, 503.—Fischetti, 1932, pp. 1-28, figs. 1-5.—Oldevig,
1933, p. 269, fig. 3—MacGinitie, 1935, p. 701.—Pirlot, 1939, p. 78.—Fioren-
cis, 1940, pp. 13-14, figs. 3-4, pl. 1, figs. 3, 4, 7.—Milne, 1940, p. 72.—
Oliveira, 1940, p. 1389.—Bertrand, 1941, pp. 12, 13, 14, 15, 16—McDougall,
1943, pp. 363, 370.—Hewatt, 1946, pp. 196, 199, 201, 204.—Ruffo, 1946,
p. 53.—Utinomi, 1947, p. 72.—Ellis, 1950, p. 13.—Reid, 1951, pp. 283,
289.—Guiler, 1952, p. 31; 1954, p. 582.—Tuzet and Sanchez, 1952, pp. 26-36,
fig. 1-1&2, fig. 2, fig. 3—Duke Univ. Mar. Lab., 1953, p. 22.—Belleudy,
1958, pp. 355-356.—Costa, 1960a, pp. 99, 100.—Luther and Fiedler, 1961,
p. 158, pl. 24.—Peyrot and Trilles, 1964, pp. 1-28, figs. 1-19.

Caprella ultima Bate, 1862, pp. 364-365, pl. 57, fig. 9.

Caprella monacantha Heller, 1866, pp. 54-55, pl. 4, figs. 17-19.—Stalio, 1877,
pp. 1125-1126.—Stossich, 1881, p. 230.

Caprella obtusa Heller, 1886, p. 54, pl. 4, fig. 16.—Stalio, 1877, p. 13890.—Stossich,
1881, p. 230.

Caprella megacephala A. Edwards, 1868, pp. 89-91, pl. 20, fig. 12.

Caprella aeguilibra.—Bate, 1878, p. 510.

Caprella caudata Thomson 1878 (1879), p. 246, pl. 10, fig. D-5.—Mayer, 1882,
pp. 71-72; 1890, p. 76.

Caprella obesa [not van Beneden] Haswell, 1880, pp. 348-349, pl. 24, fig. 1;
1882, p. 314.

Caprella AEquilibra.—Chevreux, 1888, p. 351.

Caprella linearis [not Linnaeus].—Barrois, 1888, pp. 56-57, 77.—Chevreux, 1899,
p. 484 [in part].—Chevreux and Fage, 1925, pp. 456-457, fig. 434 [in part].—
Pearse, 1936, p. 193.—Wells, 1961, p. 247.

Caprella mendax Mayer, 1903, p. 114, pl. 5, figs. 9-11, pl. 8, fig. 22.

Diacnosis.—Basis of gnathopod 2 less than one-half length of
pereonite 2, propodus without small proximal accessory tooth; pereo-
nite 2 usually with spine between insertions of gnathopods 2; pereo-
nites 1-2 elongated in large males.

DescriptTion.—Body smooth except for spine between insertions of
gnathopod 2, caphalon flattened anteriorly. Length of largest male
22 mm, largest female 12 mm, smallest ovigerous female 6.4 mm.

Large males with articles 2-3 of peduncle of antenna 1 slightly
shorter than antenna 2, article 3 subequal in length to article 2, article
1 less than one-half length of article 2, articles of peduncle expanded.
In females and small males peduncle of antenna 1 sometimes shorter
than antenna 2.

Mouthparts typical of genus, lacinia mobilis of right mandible
5-toothed.

Propodus of gnathopod 1 with 2 proximal grasping spines, grasping
margin of dactylus and propodus serrate. Basis of gnathopod 2 short

CAPRELLIDAE OF WESTERN NORTH ATLANTIC

27

and stout, anterodistal margin produced into triangular projection;

ischium and merus with posterodistal margin pointed in larger males;

palm of propodus with munerous setae, single proximal grasping spine,

distally with large rectangular tooth and slightly proximal tooth.
Gills ovate to elliptical, more ovate in larger males.

\
ann
7 : 'g
KO

-

Ficure 12.—Caprella equilibra, male; a, lateral view; b, labium; ¢c, maxilla 2; d, maxilla 1;

e, abdomen; f, pereopod 5; g, maxilliped; h, gnathopod 1.
279-475—68——_3

28 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 13.—Caprella equilibra, female; a, lateral view; ), gnathopod 1; ¢, maxilla 2; d,
gnathopod 2; e, maxilla 1; f, abdomen; g, inner and outer lobes of maxilliped; h, right
mandible; 7, left mandible; 7, pereopod 6.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 29

Propodus of pereopods 5-7 robust with 2 proximal grasping spines,
palm expanded slightly near grasping spines and with numerous setae.

Abdomen of male and female typical of genus.

VariaTion.—In the western North Atlantic this species is constant
in most of its characters; however, a variant occurs along the coast of
Virginia, North Carolina, and South Carolina in which the spine
between the insertions of gnathopods 2 is reduced or absent. In this
variant the propodus of the pereopods is less robust and the body
not quite so stout as in the typical C. equilibra.

Off Virginia this variant was taken on Leptoyorgia. This association
may have some relation to the reduction of the spine and stoutness
of the pereopods since C. penantis taken from Leptogorgia showed a
loss of grasping spines on the propodus of the pereopods (p. 35).

Distrisution.—Type locality: South Carolina. “. . . [found them
common in the bay of Charleston, particularly at Sullivan’s island, on
the two species of Gorgonia so common in the salt water creeks of
our southern coast” (Say, 1818).

Other records: Sweden and Norway to the Mediterranean Sea in-
cluding the British Islands; Black Sea [?]; Azores; tropical West
Africa; St. Helena Island; South Africa; Madagascar; Mid-North
Atlantic and Sargasso Sea; Bermuda; east coast of United States from
Connecticut to Georgia (Procter, 1933, cites this species from Mount
Desert Region, Maine); Port Aransas, Texas; Puerto Cabello, Vene-
zuela; Cabo Frio and Rio de Janeiro, Brazil; Mid-South Atlantic off
Brazil; Mar del Plata, Argentina; Valparaiso, Chile; Taboga Island,
Panama; between Panama and the Galapagos Islands; California;
Hawaii; Nagasaki, Mukaijima, and Saganoseki, Japan; Philippine
Islands; Cook Strait; New South Wales, Victoria, Fremantle, Aus-
tralia; New Zealand; Tasmania; Hong Kong; Singapore, Malaysia.

New records: Fernandina, entrance to St. Johns River, St. Augus-
tine, Daytona, Cape Kennedy, off Ft. Lauderdale, Biscayne Bay,
and Panama City, Fla.; Grand Isle, La.; Galveston and Port Isabel
Tex.; Trinidad; Sacco Sao Francisco and Nictherey, Brazil; Estera
de la Luna, Sonora, Mexico; Vancouver Island, British Columbia.

Remarks.—Large males of this species are easily distinguished
from the other species of Caprella by the large peduncular articles
of antenna 1 and the long pereonites 1 and 2. In both males and
females the cephalon is flattened anteriorly and in the typical form
a spine is present between the insertions of gnathopods 2. The non-
spined variant resembles other species of Caprella but can be identified
by the short stout basis of gnathopod 2 and the other characters
which are present in the typical form.

C. equilibra has been collected from various habitats including
sea grass, red and green algae, sponges, hydroids, stylasterines,

30 U.S. NATIONAL MUSEUM BULLETIN 278

alcyonarians, bryozoans, and colonial ascidians. It was also taken
from the egg mass of a blue crab and from the gut of a sea bass
Centropristis. This species ranges in depth from the surface to ?3000 m
(McCain, 1966).

In an aquarium it was preyed upon by the grass shrimp Palae-
monetes, the blenny Blennius, and the small (2 mm) snail Astyris
was observed severing the cephalon from a large male (12 mm).
When C. equilibra was offered small pieces of bivalves or bryozoans
it would accept them readily. It was also observed catching small
gcammaridean amphipods such as Ampithoe and Jassa and also several
small polychaetes. Initially the prey was seized in gnathopod 1 and
then brought to the mouthparts. Gnathopod 2 was seldom used in
the capture of prey and even when it was used, the prey was quickly
passed to gnathopod 1.

Caprella linearis (Linnaeus, 1767)
Figures 14, 22, 51

Cancer linearis Linnaeus, 1767, p. 1056; 1769, pp. 445-446; 1788, p. 2992; 1793,
p. 501.—J. Fabricius, 1793, pp. 517-518.

Onisct Scolopendroidis Pallas, 1772, p. 80, pl. 4, fig. 15a—c.

Squilla lobata Miller, 1776, p. 197.

Squilla quadrilobata Abildgaard, 1788, pp. 21-22, pl. 56, figs 4-6.

Gammarus quadrilobatus.—Abildgaard, 1789, p. 58, pl. 114, figs. 11-12.

Cancer (Gammarellus) linearis.—Herbst, 1793 ,pp. 142-144, pls. 9a, 10b.

Cancer Linnearis.—Linnaeus, 1800, p. 761.

Caprella linearis.—Bosc, 1801-02, p. 156; 1830, p. 126, pl. 15, fig. 5.—Latreille,
1802-03, pp. 324-326, pl. 57, figs. 2-5; 1803, p. 333; 1816, p. 434.—Desmarest,
1823, p. 364; 1825, p. 278.—Johnston, 1835, pp. 671-672, fig. 71.—Drapiez,
1837, p. 353.—H. Edwards, 1840, pp. 106-107.—Goodsir, 1842, p. 190, pl. 3,
figs. 8-9.—White, 1847, pp. 91-92; 1850, pp. 59-60; 1857, pp. 214-215.—
Cocks, 1849, p. 83.—Williams, 1854, pp. 301-312, pl. 17, fig. 6.—Gosse, 1855,
p. 131, fig. 223.—Bate, 1856, p. 60; 1857, p. 151; 1862, p. 353, pl. 55, fig. 7;
1878, p. 509; 1887, p. 175.—Leydig, 1860, p. 283.—van Beneden, 1861, p.
145.—McAndrew, 1861, p. 28.—[?] Dohrn, 1866, pp. 245-250, pl. 13b.—
Bate and Westwood, 1868, pp. 52-56.—Miiller, 1869, pp. 40-41.—Metzger,
1869-70 (1871), p. 32; 1875, p. 278.—Boeck, 1871la, pp. 273 (193)-274
(194).—Iarzynsky, 1870, p. 316.—Parfitt, 1873, p. 250.—M’Intosh, 1874,
p. 272.—Maitland, 1874, p. 245.—Meinert, 1877-78, pp. 168-171; 1880,
p. 495; 1890, p. 184.—Hoex, 1879, pp. 97-161, pl. 5, figs. 1-8, 11-18, pl. 6,
fig. 2, pl. 7, figs. 1-3, 11-14; 1883-84, pp. 532, 533; 1889, p. 231.—Delage,
1881, p. 153.—Mayer, 1882, pp. 58-62, figs. 17-19, pl. 4, fig. 32; 1890, pp.
63-65; 1903, pp. 109-113, pl. 4, figs. 27-35, pl. 8, figs. 19-21.—Pelsenner,
1883, p. CX X XI; 1886, p. 218.—Schneider, 1883, p. 30; 1891, pp. 111, 122;
1924 (1926), pp. 59-60.—Blanec, 1884, pp. 88-91, pl. 5, figs. 122-129.—
Koehler, 1884 (1885), pp. 98-99, 117; 1885, pp. 27, 61.—Wagner, 1885,
p. 169.—Fowler, 1886, p. 218.—de Guerne, 1886, p. XLIII.—Norman, 1886,
p. 26; 1902, p. 483; 1905a, p. 26; 1905b, p. 85; 1907, p. 370; 1908 (1909),
p. 463.—G. Sars, 1886, pp. 69, 89; 1895, pp. 657-658, pl. 236.—[?] Thomson
and Chilton, 1886, p. 142.—Bonnier, 1887, pp. 354—356.—Chevreux, 1887,
p. 335; 1898, p. 484.—Robertson, 1886-87 (1888), pp. 71-72.—Scott, 1887

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 31

(1888), p. 250; 1897, p. 141; 1906, pp. 174-175.—Chevreux and Bouvier,
1893, p. 143.—Lameere, 1895, p. 570.—[{?] Ohlin, 1895a, pp. xvii, xix, 62-63;
1895b, p. 486.—Walker, 1895, p. 319; 1898, p. 170.—Walker and Hornell,
1896, p. 55.—Gadeau de Kerville, 1900 (1901), p. 184.—Sokolowsky, 1900,
p. 162, pl. 3, fig. 16.—Ortmann, 1901, p. 155.—d’A. Thompson, 1901, p.
41.—Whiteaves, 1901, p. 219.—Lénnberg, 1902 (1903), p. 50.—[{?] Hutton,
1904, p. 261.—Marine Biol. Assoc., 1904, p. 242; 1951, p. 198; 1957, p. 234.—
Holmes, 1904 (1905), pp. 526-527.—M. Rathbun, 1905, pp. 7, 78.—Norman
and Scott, 1906, pp. x, 98.—Reibisch, 1906, pp. 217-218, 219, 220, 221, 222,
229, 230, 233.—Sinel, 1906 (1907), p. 222.—Briiggen, 1907, p. 238.—Norman
and Brady, 1910, pp. 75-76.—Nordgaard, 1911 (1912), p. 24.—Massy, 1911
(1912), pp. 7, 22, 34, 42, 43, 45, 51, 68, 70, 73, 82, 169.—Sumner, Osburn,
and Cole, 1911 (1913), p. 657.—Tattersall, 1913, pp. 20, 22.—Derjugin, 1915,
pp. 453, 456; 1928, p. 282.—Bjoérck, 1915, p. 35; 1916, p. 9—Chumley, 1918,
pp. 52, 85, 165.— Kunkel, 1918, pp. 177-178, fig. 54.—Funke, 1922, p. 197.—
Chevreux and Fage, 1925, pp. 456-457, fig. 434 [in part].—Derjavin, 1927,
p. 14.—Stephensen, 1927a, p. 149; 1927b, p. 13; 1928, pp. 382-384, fig. 92
(1-4); 1929a, p. 179, fig. 333; 1929b, pp. 19, 34; 1935, p. 118; 1940, p. 73;
1942, pp. 436-437, 502, 503; 1944b, p. 159.—Johansen, 1930, p. 94.—Shoe-
maker, 1930, p. 353 (135).—[?] Arimoto, 1931, pp. 13-14, fig. 9.—Gurjanova,
1931, p. 201; 1964, p. 313.—Oldevig, 1933, pp. 264-266.—Procter, 1933,
p. 256.—Dons, 1935, p. 110.—Schellenberg, 1942, pp. 237-238, fig. 197.—
Dahl, 1946, p. 22.—[?] Utinomi, 1947, p. 75.—Stock and Bolklander. 1952,
pp. 3-4.—Bousfield, 1956b, p. 145; 1958, p. 315.—Brunel, 1961, p. 7.—
Toulmond and Truchot, 1964, p. 35.

Caprella Linearis—Leach, 1814, p. 404.—Risso, 1816, p. 1380.—Couch, 1864,
p. 98.

[?]Caprella Punctata Risso, 1816, pp. 130-131; 1826, p. 102.—Carus, 1885, p. 389.

Caprella laevis Goodsir, 1842, pp. 189-190, pl. 3, figs. 4-5.—White, 1847, p. 92;
1850, p. 60; 1857, p. 215.—Gosse, 1855, p. 131.—Bate, 1856, p. 60.

[?]Caprella phasma {not Montagu].—Rathke, 1843, pp. 94-96.

[?]Caprella acuminifera [not Leach].—Rathke, 1843, p. 96.

[?]Caprella scolopendroides [not Lamarck].—Rathke, 1843, p. 97.

Caprella lobata.—Bate, 1856, p. 60; 1857, p. 151; 1862, p. 354, pl. 55, fig. 8; 1878,
p. 509; 1887, p. 175.—Bate and Westwood, 1868, pp. 57-59.—Parfitt, 1873,
p. 250.

Caprella linearis f. gullmarensis Mayer, 1903, p. 112, pl. 8, fig. 20.

Caprella linearis f. distalis Mayer, 1903, p. 113, pl. 4, figs. 27-28.

Dracnosis.—Body smooth or with only few spines, peduncle of
antenna 1 usually with numerous setules, ratio of total length to
length of basis of gnathopod 2 usually less than 13.0, inner surface
of gnathopod 2 with small tooth adjacent to poison tooth.

DistrinutTion.—Type-locality: “Habitat im Oceano [uropaeo”’
(Linnaeus, 1767).

Other records: Siberian Polar Sea to 140° E.; Murman coast;
Spitsbergen; Norway to France; British Islands; Faeroe Islands;
Iceland; coast of North America from Labrador to Connecticut;
[?]Kamchatka, Japan, and New Zealand.

New records: The material available to me does not extend the
range of this species.

32 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 14.—Caprella linearis; a, male gnathopod 2; b, male lateral view; ¢, female lateral
view; d, female gnathopod 2; ¢, female pereopod 6; f, female gnathopod 1.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC ao

Remarks.—The appendages of C. linearis are quite similar to
those of C. septentrionalis and are discussed under the latter species.

C. linearis does not appear to be specific in its habitat, having been
collected from brown, green, and red algae, sea grass, sponges, hydroids,
alcyonarians, and tunicates. Ohlin (1895a) reported it from an Asterias
collected off Newfoundland. This report may refer to C. wnica since
the body form and most of the appendages of these species are quite
similar and (. wnica has been collected off Newfoundland. Mayer
(1903) also reported C. linearis from an asteroid, Solaster, which was
collected off Scotland.

C. linearis has been taken from the surface to a depth of several
hundred meters.

Caprella longimanus Stimpson, 1853

Caprella longimanus Stimpson, 1854 (1853), p. 44.—-Whiteaves, 1901, p. 220.—
Bate, 1862, pp. 360-361.
Caprella longimana.— Mayer, 1882, p. 66; 1890, p. 73.

Remarks.—A caprellid from Grand Manan was described by
Stimpson as

Body with a few spines along the back of each segment. Superior antennae rather
stout and twice as long as the inferior ones, which are very slender. Hands very
long and rather broad, with two or three teeth along the inner edge; the arms to
which they belong are placed on the thickened posterior part of the second seg-
ment. Color light-yellowish brown. Eyes red. Length about three-fourths of an
inch.

From this description it is impossible to tell to which species Stim pson

is referring. It might be C. septentrionalis because of the mention of
a few dorsal spines, but several other species bear spines.

Caprella penantis Leach, 1814
Figures 15, 16, 51

[?]Cancer Atomos Linnaeus, 1767, p. 1056; 1769, pp. 446-447; 1788, pp. 2992-
2993; 1793, p. 501; 1800, p. 761.—Pennant, 1777, p. 21, pl. 12, fig. 32.

Caprella Penantis Leach, 1814, p. 404.

Caprella acutifrons Latreille, 1816, p. 433.—[?] Desmarest, 1823, p. 363; [?] 1825,
p. 277.—[?] Drapiez, 1837, p. 353.—[?] H. Edwards, 1840, p. 108.—[?] White,
1847, p. 92; [?] 1850, p. 60; [?] 1857, p. 216.—[?] Cocks, 1849, p. 83.—[?]
Gosse, 1855, p. 131.—[?] Bate, 1856, p. 60; [?] 1862, p. 356, pl. 56, fig. 6; [?]
1878, p. 509; [?] 1887, p. 175.—[?] Bate and Westwood, 1868, pp. 60-62.—
(?] Parfitt, 1873, p. 250.—[?] Maitland, 1874, p. 245.—[?] Stalio, 1877, p.
1125.—[?] Haller, 1879a, p. 232; [?] 1879b, p. 404.—Mayer, 1882, pp. 48-
50, pl. 1, fig. 9, pl. 2, figs. 12-22, pl. 4, figs. 26-28, pl. 5, figs 15, 22, 23 [in
part]; 1890, pp. 50-57, pl. 2, figs. 34-41, pl. 4, figs. 52-71, pl. 7, figs. 16-17
{in part]; 1903, pp. 79-89, pl. 3, figs. 4-28, pl. 7, figs. 62-65 [in part].—[?]
Delage, 1881, pp. 131-132, 155, pl. 10, figs. 11-12.—-Stossich, 1881, p. 230.—
[?] Marion, 1883, p. 49.—[?] Carus, 1885, p. 388.—[?] Norman, 1886, p.
26; [?] 1905a, p. 26; [?] 1905b, p. 85; [?] 1907, p. 370; [?] 1908 (1909), p. 463.—
[?] Pelseneer, 1886, p. 218.—[?] Bonnier, 1887, p. 353.—[?] Chevreux, 1887a,

34 U.S. NATIONAL MUSEUM BULLETIN 278

pp. 318, 335; [?] 1888, p. 33; [?] 1898, p. 483; 1900, pp. 119-120.—[?] Barrois,
1888, pp. 57-58, 77.—[?] Vosseler, 1889, p. 159.—[?] Walker and Hornell,
1896, p. 54.—[?] Gadeau de Kerville, 1898, p. 348; [?] 1900 (901), p. 184.—
[?] Walker, 1898, p. 170.—[?] Beaumont, 1900, p. 795.—[?] d’A. Thompson,
1901, p. 41.—M. Rathbun, 1905, pp. 7, 77-78.—[?] Norman and Scott,
1906, pp. vii, 99.—[?] Sinel, 1906 (1907), p. 222.—Chilton, 1910 (1911),
pp. 546, 567.—[?] Monterosso, 1915, p. 15, fig. 3.—[?] Galdiano, 1924, p.
392.—Richards, 1929, p. 84; 1938, p. 213, pl. 24, fig. 7—Cowles, 1930,
p. 351.—[?] Mar. Biol. Assoc., 1931, p. 198; [?] 1957, p. 233.—Barnard, 1932,
p. 300; 1965, p. 209.—Procter, 1933, p. 256.—[?] MacGinitie, 1935, p. 701.—
Schellenberg, 1938, pp. 95, 98.—[?] Ricketts and Calvin, 1939, pp. 70-71; [?]
1952, p. 68.—[?] Bertrand, 1941, pp. 12, 13, 14.—Pearse, Humm, and Whar-
ton, 1942, p. 184.—Dexter, 1944, p. 356.—[?] MacKay, 1945, p. 205.—[?]
Hewatt, 1946, pp. 194, 196, 199, 200, 201, 202, 204.—[?] Ruffo, 1947, p. 129.—
Edmondson ad Mansfield, 1948, pp. 212-214, fig. 6—Ferguson and Jones,
1949, p. 442.—[?] Stephensen, 1949, p. 54.—Hedgpeth, 1950, pp. 77-78.—
Ellis, 1950, p. 13.—[?] Tuzet and Sanchez, 1952, pp. 26-36, figs. 1-3, 1-4, 1-5,
fig. 4.—Duke Univ. Mar. Lab., 1953, p. 22.—[?] Macnae, 1953, p. 1032.—
Bousfield, 1956b, p. 145; 1958, pp. 315, 321—Menzel, 1956, p. 41.—Pearse
and Williams, 1951, p. 143.—[?] Stschapova, Mokyovsky, and Pasternak,
1957, p. 87.—[?] Costa, 1960a, pp. 99, 100.— Wells, 1961, pp. 247, 249.—[?]
Toulmond and Truchot, 1964, p. 35.

Caprella geometrica Say, 1818, pp. 390-391.—de Kay, 914, p. 41.—White, 1847,
p. 92.—Gibbes, 1848, p. xvi; 1849, p. 23.—Bate, 1862, p. 357, pl. 56, fig. 8.—
Verrill and Smith, 1873, pp. 316-317, 480, 567, pl. 5, fig. 20.—Uhler, 1879,
pp. 26-27.—R. Rathbun, 1880 (1881), p. 121.—Norman, 1886a, p. 26;
1905, p. 26.—Holmes, 1904 (1905), p. 526.—Paulmier, 1905, p. 168, fig.
38.—Kunkel, 1918, pp. 178-180, fig. 55—Sumner, Osborn, and Cole, 1911
(1913), pp. 132, 134, 135, 657, chart 102.—Pearse, 1913, p. 378. LaFollette,
1914, pp. 222-223, pl. 1-3.—Allee, 1922, p. 58; 1923, p. 213.—Wood and
Wood, 1932, p. 18—McCain, 1965, pp. 194-196, figs. 1c,g, 2a-f.

Caprella Pennantii—I[?] Johnston, 1835, p. 671—[?] Bate, 1856, p. 60; [?] 1857,
p. 151.—[?] McAndrew, 1861, p. 28.—[?] Couch, 1864, p. 97.

[?] Caprella spinifrons Nicolet, 1849, p. 253.—Mayer, 1882, p. 70; 1890, p. 74.—
Reed, 1897, p. 11 (4).

[?] Caprella obesa van Beneden, 1861, pp. 99, 146.

Caprella Acutifrons.—[?] Herklots, 1861, p. 43.

[?] Caprella novae-zealandiae Kirk, 1878, pp. 465-466; 1878 (1879), p. 393.—Thom-
son, 1879, p. 330.

[?] Caprella Novae-Zealandiae.—Mayer, 1882, pp. 71-72; 1890, p. 76.

[?] Caprella penantiit.—Bate, 1887, p. 175.

Caprella acutifrons f. tabida Mayer, 1890, pp. 54-55, pl. 2, fig. 36, pl. 4, figs. 52, 61.

Caprella acutifrons f. neglecta Mayer, 1890, p. 55, pl. 2, fig. 37, pl. 4, figs. 57-58,
67; 1903, p. 80.—Utinomi, 1943a, pp. 273-274, figs. 2a, 3a; 19438b, pp.
282-283, fig. 2; 1943c, p. 284, fig. 1; 1947, p. 72.

[?] Caprella acutifrons f. gibbosa Mayer, 1890, p. 55, pl. 2, fig. 39, pl. 4, figs. 55, 69.

Caprella acutifrons f. carolinensis Mayer, 1890, p. 56, pl. 2, fig. 40, pl. 4, figs. 59, 65.

[?] Caprella acutifrons f. lusitanica Mayer, 1890, p. 56, pl. 4, figs. 53, 66.

Caprella acutifrons f. virginia Mayer, 1890, p. 56, pl. 2, fig. 41, pl. 4, fig. 60.

Caprella acutifrons f. natalensis Mayer, 1903, p. 81, pl. 3, figs. 22, 23—Arimoto,
1930, pp. 48-49, fig. 3—Hiro, 1937, p. 312, pl. 22, fig. 5—Stephensen, 1949,
pp. 53-54, 56.

Caprella acutifrons f. porcellio Mayer, 1903, pp. 81-82.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC oo

Caprella acutifrons f. testudo Mayer, 1903, p. 82.—Chevreux and Fage, 1925, p.
452, fig. 430t.

Caprella acutifrons f. angusta Mayer, 1903, p. 82, pl. 3, fig. 4.

Caprella acutifrons f. tibada Mayer, 1903, p. 80.

Caprella penantis—Stebbing, 1910b, p. 653.—[?] Hale, 1929, pp. 233-234.—[?]
Schellenberg, 1931, pp. 266, 272.

Caprella penantis f. natalensis.—Stebbing, 1910a, pp. 465-466.—Barnard, 1916,
pp. 281-282.

Caprella penantis f. porcellio. —Stebbing, 1910a, p. 466.

Caprella angusta-—Dougherty and Steinberg, 1953, pp. 44, 47; 1954, p. 171.—
Johnson and Juskevice, 1965, p. 38.

Caprella carolinensis.—Steinberg and Dougherty, 1957, pp. 270-273, figs. 3-7.

Diacnosis.—Cephalon with anteriorly directed triangular pro-
jection; peduncle of antenna 1 not inflated; basis of gnathopod 2
shorter than pereonite 2; pereopods concave, grasping spines proximal.

Description.—Body smooth except cephalon with anteriorly di-
rected triangular projection. Length of largest male 14 mm, largest
female 12 mm, smallest ovigerous female 4 mm.

Peduncle of antenna 1 not inflated, flagellum with up to 15 articles.
Antenna 2 usually longer than peduncle of antenna 1.

Mouthparts typical of genus, lacinia mobilis of right mandible
toothed but indistinctly 5-toothed.

Propodus of gnathopod 1 with 2 proximal grasping spines, grasping
margin of dactylus and propodus serrate. Propodus of gnathopod
2 with proximal poison tooth, palm concave in males and slightly
convex in females with distal elevated rectangular projection; grasp-
ing margin of dactylus serrate.

Gills usually ovate, occasionally elliptical.

Propodus of pereopods 5-7 usually with pair of proximal grasping
spines, pereopods increasing in length from 5 to 7.

Abdomen of male and female typical of genus.

Variation.—In the area around Alligator Harbor, Fla., C. penantis
taken on Leptogorgia showed a reduction or loss of grasping spines
on the propodus of pereopods 5-7. Approximately 90 percent of
the specimens taken during the summer of 1966 lacked grasping
spines and remaining 10 percent had either 1 or 2 grasping spines.
Other specimens of this species taken during that summer on algae
and hydroids had the usual pair of grasping spines. It is interesting
to note that CO. equilibra taken on Leptogorgia off Virginia showed
a reduction or loss of the ventral spine between the insertions of
enathopods 2.

As J illustrated in 1965 (p. 195, fig. 2a-f) the shape of the propodus
of gnathopod 2 changes with an increase in the size of the individual.
In smaller individuals of approximately 4 or 5 mm, gnathopod 2
resembles gnathopod 1 and bears a pair of grasping spines. As the

36 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 15.—Caprella penantis, male; a, lateral view; b, gnathopod 2; ¢, gnathopod 1;
d, maxilla 1; ¢, right mandible; f, left mandible; g, abdomen; A, inner lobe of maxilliped;
1, outer lobe of maxilliped.

individual increases in size there is a progressive loss of 1, then of
both spines, and a notch develops in the palm.

Stoutness of the body and the degree of pleural development
appear to be a function of growth, larger individuals having a robust
body and well-developed pleura. '

CAPRELLIDAE OF WESTERN NORTH ATLANTIC of

DistrisuTion.—Type-locality: Devonshire Coast, England.

Other records: Atlantic coast of France, Spain, and Portugal;
British Islands; Azores; Atlantic coast of North America from Nova
Scotia and Gulf of St. Lawrence to Georgia; Alligator Harbor, St.
Georges Sound, and Apalachee Bay, Fla.; Galveston, Freeport,
Port Aransas, and Port Isabel, Tex.; Safety Islands; Tristan da
Cunha and Gough Island; South Africa; California; Hawaii; Kyushu
and Honshu, Japan; Formosa Strait; Hong Kong; New Zealand;
New South Wales, Australia.

New records: Savannah Beach and Sapelo Island, Ga.; Fernandina
Beach, Mayport, St. Augustine, Marineland, Ft. Pierce, Key West,
Sarasota Bay, Tampa Bay, Dunedin, Panama City, and Destin,
Fla.; Ocean Springs, Miss.; Cayenne, French Guiana.

Remarks.—In Mayer’s last two monographs (1890, 1903), he
names 20 varieties or forms of the Caprella acutifrons group. Of

Ficure 16.—Caprella penantis, female; a, lateral view; b, gnathopod 2; c, pereopod 7; d,
gnathopod 1; ¢, abdomen.

38 U.S. NATIONAL MUSEUM BULLETIN 278

these, C. acutifrons f. angusta (1903), borealis (1903), incisa (1903),
and verrucosa (1903) (=C.verrucosa Boeck, 1871b) have since been
given specific rank. In the present paper one other variety, C. acuti-
frons f. andreae, is accorded specific rank, leaving 14 varieties in
question.

The varieties C. acutifrons f. typica (1890), minor (1890), tabida
(1890), and tibada (1903) differ from the remaining varieties pri-
marily by the distal position of the poison tooth on the palm of
the propodus of gnathopod 2. Mayer recognized C. acutifrons f.
typica and minor from Rio de Janeiro, Brazil, the latter variety
being based on a smaller individual than the former. Krgyer (1842-
43) described CO. dilatata from Rio de Janeiro. All of the above-
mentioned varieties should be assigned to C. dilatata. Juveniles of
this species bear a small proximal tooth on the palm of gnathopod
2, which is very short and spinelike and is not present on individuals
larger than approximately 8 mm.

In 1903 Mayer changed the 1890 variety tabida to tibada and
recognized C. tabida Lucas, 1849, as a different variety. C. acutifrons
f. tabida (1903) (=C. tabida Lucas, 1849), C. acutifrons f. simulatriz
(1903), C. pilimana Dougherty and Steinberg, 1953, and C. obtusi-
frons Utinomi, 1948c, differ from the remaining varieties by the lack
of a poison tooth on the palm of gnathopod 2. Specimens of the
first two varieties and of C. obtusifrons have not been examined so
I cannot make any statement on their validity.

Caprella acutifrons f. cristibrachium (1903) lacks a triangular pro-
jection on the cephalon and the shape of gnathopod 2 is quite
different from that of the other members of the C. acutifrons group.
It is doubtful that it is a variety of this group and probably should
be given specific rank.

The remaining varieties, C. acutifrons f. carolinensis (1890), virginia
(1890), testudo (1903), gibbosa (1890), lusitanica (1890), natalensis
(1903), porcellio (1903), and neglecta (1890), are quite similar in the
shape of gnathopod 2 and general body form. In the first five varieties
the palm of the propodus of gnathopod 2 is quite setose and in the
last three varieties and C. angusta the palm is scarcely setose. I have
been unable to find other distinguishing characters for these varieties,
so I have tentatively assigned them to the species C. penantis.

Dougherty and Steinberg (1953) gave C. acutifrons f. angusta,
incisa, and verrucosa specific rank. These varieties occur together
on the California coast and have been collected simulatneously from
the same hydroid. Since they are sympatric and no morphological
intergradations were observed, all of these varieties cannot be ascribed
to the same species. I agree with Dougherty and Steinberg’s decision
to give them specific rank; however, they do not state how C.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 39

angusta differs from the typical C. penantis. Specimens of C. angusta
have been compared with specimens which Mayer identified as C.
acutifrons f. natalensis and neglecta and with specimens which I
earlier (1965) identified as C. geometrica. With the exception of the
setose versus nonsetose palm of guathopod 2, I can find no variation
which is not ascribable to size differences. I have therefore synony-
mized C. angusta with the typical C. penantis.

The specimens from Cayenne, French Guiana, belong to that
portion of C. penantis which bears almost no setae on the palm of
enathopod 2. It might well be that C. penantis could be divided
into two subspecies on the basis of the setation of gnathopod 2;
however, material from Chile, Australia, and New Zealand would
have to be examined since Nicolet’s name C. spinifrons or possibly
Kirk’s name C. novae-zealandiae would probably have priority over
one of Mayer’s varietal names. Material is not available to me from
these areas so I have refrained from naming subspecies.

Kirk’s (1878) description of C. novae-zealandiae agrees with that
of C. penantis, and he states that his species is close to C. geometrica.
Thomson and Chilton (1885, 1886) synonymized C. novae-zealandiae
with C. equilibra; however, Kirk states “Cephalon furnished with
a spinous tooth directed forwards.” It seems unlikely that Kirk
could have been referring to C. equilibra, so I have synonymized
his species with C. penantis.

In my synonymy, when a reference to the variety of C. acutifrons
or C. penantis is not indicated, this lack of designation is indicated
by a question mark in brackets before the author or date. Such
records are not included in the distribution of this species, so it is
possible that C. penantis might also be found as far north as Spits-
bergen, the Mediterranean Sea, the Falkland Islands, Chile, Cook
Strait, and the Bering Sea.

This species is probably the most common caprellid along the
east coast of the United States. It occurs in such abundance from
Long Island to Chesapeake Bay that I have had several reports
that it is a pest to swimmers. One report from Sinepuxent Bay, Md.
stated that C. penantis fastens itself to the exposed parts of swimmers’
bodies and either bites or sucks, causing an lritation that forms a
blister which lasts for a week or more. Such an irritation might be
caused by the associated hydroids and not by the caprellid itself,
but this possibility has not been verified.

C. penantis is quite nonspecific in its habitat preference and has
been taken on various red and brown algae, sea grass, sponges, hy-
droids, aleyonarians, zoantharians, and bryozoans. Chilton (1911)
reports that this species was collected from a coconut which was
washed ashore; I have found it clinging to the spines of the echinoid

40 U.S. NATIONAL MUSEUM BULLETIN 278

Arbacia and on hydroids which were attached to the carapace of the
spider crab Libinia.

Caprella sanguinea Gould, 1841

Caprella sanguinea Gould, 1841, pp. 335-336.—de Kay, 1844, p. 41.—Stimpson,
1854 (1853), p. 44.—Bate, 1862, p. 360.—Mayer, 1882, p. 67; 1890, p. 73.—
Whiteaves, 1901, p. 219.

Remarxs.—This species from Massachusetts was described by
Gould as:

. an inch in length, entirely crimson except its black eyes. The head is blunt,
the lower antennae ciliated and extending to the second segment, and the upper
ones to the third segment; first two segments nearly as long as the three next,
and about one third of the whole length; on the middle of the first is a spine;
two last segments short and heart-shaped. Hands having a long curved finger;
an imperfect thumb on the second pair of legs; a tubercule at the base of the
ovate carpus, and a small spine at the middle. This might be called C. sanguinea,
from its color, which it retains in spirits.

Like Stimpson’s C. longimanus, this species is unidentifiable and it
might also belong to C. septentrionalis.

Caprella scaura Templeton, 1836
FicureEs 17, 18, 55

Caprella scaura Templeton, 1836, pp. 191-192, pl. 20 fig. 6—H. Edwards, 1840,
p. 107.—Bate, 1862, p. 355, pl. 56, fig. 4.—Mayer, 1882, p. 65; 1890, pp.
70-73, pl. 4, figs. 40-51, pl. 6, fig. 41, pl. 7, figs. 2, 35-36 [in part]; 1903,
pp. 117-120, pl. 5, figs. 13-18, pl. 10, fig. 11 [in part].—Walker, 1916, p.
346.—Barnard, 1925, pp. 371-372.—Hale, 1927, p. 315; 1929, p. 234, fig.
229.—Arimoto, 1931, pp. 16-18, pl. 3, figs. 1-6.—Hiro, 1937, pp. 314-315,
fig. 3, pl. 22, figs. 11-12.—Day and Morgan, 1956, p. 308.

Caprella nodosa Templeton, 1836, pp. 192-194, pl. 21, fig. 7.—H. Edwards, 1840,
p. 108.—Bate, 1862, p. 357, pl. 56, fig. 7.

Caprella cornuta Dana, 1853, pp. 816-817; 1855, pl. 54, fig. 5.—Bate, 1862, p.
356, pl. 56, fig. 5.—Mayer, 1882, p. 68.—Chilton, 1921, pp. 90-91, fig. 4.—
Oliveira, 1940, p. 1389.

Caprella cornuta f. obtusirostris Dana, 18538, p. 817; 1855, pl. 54, fig. 6.

Caprella attenuata Dana, 1853, pp. 817-819; 1855, pl. 55, fig. 1.—Bate, 1862,
p. 364, pl. 57, fig. 7.—Mayer, 1882, pp. 67-68, figs. 24-25; 1890, p. 73.—
Haswell, 1885, p. 1000.

Caprella attenuata f. subtenuis Dana, 1853, pp. 818-819; 1855, pl. 55, fig. le.

Caprella scaura f. typica Mayer, 1890, p. 71, pl. 4 figs. 48-49; 1903, p. 118.—
Miyadi and Masui, 1942, p. 10.—Utinomi, 1947, p. 77.

Caprella scaura f. diceros Mayer, 1890, p. 71; 1903, p. 118.—Miyadi and Masui,
1942, p. 10.—Utinomi, 1943a, p. 279; 1943b, p. 285, fig. 5; 1947, p. 77.

Caprella scaura f. cornuta Mayer, 1890, pp. 71-72, pl. 4, figs. 50-51; 1903, pl. 118.

Caprella scaura f. undetermined Mayer, 1903, p. 120.

Caprella scaura f. hamata Utinomi, 1947, p. 77, fig. 7.

DiaGNnosis.—Cephalon with anteriorly directed spine, pereonites
1-2 elongate in males, basis of gnathopod 2 approximately length
of pereonite 2.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 41

Description.—Body with anteriorly directed cephalic § spine,
female with variously developed spines on pereonites 1-7. Length
of largest male 21 mm, female 12 mm, smallest ovigerous female
6 mm.

Antenna 1 usually longer than one-half body length, flagellum with
as many as 9 fused articles in males, up to 4 in females. Length of
antenna 2 variable.

Mouthparts typical of genus, lacinia mobilis of right mandible not
distinctly 5-toothed.

Propodus of gnathopod 1 with 2 proximal grasping spines, grasping
margin of dactylus and propodus serrate. Propodus of gnathopod 2
elongate in males, palm with 2 strong teeth and distal rectangular
projection; female propodus not so elongate as male, palm with proxi-
mal spine, small distal tooth and distal rectangular projection.

Gills elliptical.

Propodus of pereopods 5-7 with 2 proximal grasping spines.

Abdomen of male and female typical of genus except with raised
medial projection.

VartaTion.—The females with the most pronounced dorsal body
spination had 1 knob at the posterior of pereonite 1, 1 pair of knobs
above the gills on pereonites 3 and 4, 1 knob at the posterior of pereon-
ite 4, 2 pairs of knobs at midlength of pereonite 5, 1 pair of knobs at.
midlength of pereonite 6, and a pair of knobs at the posterior of
peronite 7. This spination showed varous degrees of reduction from this
pattern with the knob at the posterior of pereonite 4 usually being
present. The males occasionally bore 2 pairs of knobs at midlength of
pereonite 5 and a pair of knobs at midlength of pereonite 6.

The number of fused articles in the flagellum of antenna 1 varied
from 6-9 in males and from 2-4 in females. Mayer used this character
for separating C. scaura f. cornuta from C. scaura f. typica and diceros
since C. scaura f. cornuta does not have fused articles in the flagellum
of antenna 1.

Distrispution.—Type-locality: Riviere Noire, Mauritius.

Other records: St. Croix and St. Barthélemy, Virgin Island;
Vitoria, Rio de Janeiro, and 28° S., Brazil; South Africa; Mejillones,
Chile; Cumberland Bay, Isla Mas a Tierra; Honshu, Kyushu, and the
Inland Sea, Japan; Vladivostok; Sydney and Kangaroo Island,
Australia.

New records: Cocos Island, Costa Rica; off Mayagtiez, Puerto
Rico; ha Sao Sebastiao and Santa Catarina, Brazil; False Bay, South
Africa.

Remarxs.—Mayer (1890, 1903) described 6 varieties of C. scaura
to which Utinomi (1947) added a seventh, C. scaura f. hamata. Mayer’s
varieties C. scaura f. typica (1890), diceros (1890), cornuta (1890),

42 U.S. NATIONAL MUSEUM BULLETIN 278

h

Ficure 17.—Caprella scaura, male; a, lateral view; b, gnathopod 1; ¢, maxilla 2; d, maxilla
1; 2, gnathopod 1; f, right mandible; g, abdomen; h, left mandible; 2, inner and outer
lobes of maxilliped.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 43

Ficure 18.—Caprella scaura, female; a, lateral view; b, inner and outer lobes of maxilliped;
c, gnathopod 2; d, maxilla 1; ¢, abdomen; f, gnathopod 1; g, righ mandible; A, left
mandible.

279-475—68——4

44 U.S. NATIONAL MUSEUM BULLETIN 278

and Utinomi’s C. scaura f. hamata do not bear a ventral spine between
the insertions of gnathopods 2 as do Mayer’s varieties californica
(1890), scauroides (1903), and spinirostris (1890).

Dougherty and Steinberg (1953) separated C. scaura f. californica
as a distinct species and reestablished Stimpson’s (1857) name C.
californica. This action was justified; however, they did not state
what should be done with the other two varieties which bear the ven-
tral spine. These varieties are closely related and appear to be limited
to the North Pacific. No material of the Asian varieties is available to
me and I am not able to comment on their taxonomic position. The
synonymy, therefore, includes only those references which refer to
those varieties which do not bear the ventral spine.

Barnard (1925) considered C. laevipes Mayer, 1903, a synonym of
C. scaura. C. laevipes appears to be distinct from C. scaura since the
pereopods do not bear grasping spines and Barnard’s synonymy has
not been followed.

The Caribbean material appears to be most closely related to C.
scaura f. typica which has previoulsy been taken from St. Croix and
St. Barthélemy.

C. scaura has been taken on red and brown algae, sea grass, bry-
ozoans, and on a sea urchin.

Caprella septentrionalis Krgyer, 1838
Ficures 19-22, 51

Squilla lobata [not Miiller].—O. Fabricius, 1780, pp. 248-249.

Caprella septentrionalis Krgyer, 1838, p. 318; 1842-43, pp. 590-596, pl. 8, figs.
10-19.—Boeck, 1861, p. 677; 1870, p. 276 (196); 1873-76, pp. 696-698.—
Bate, 1862, p. 355, pl. 56, fig. 3.—Goés, 1866, p. 534.—Packard, 1867, p.
297.—Liitken, 1875, p. 159.—Schigdte, 1875, p. 224, pl. 5, figs. 1-8.—
Norman, 1876, p. 209; 1886, p. 26; 1902, p. 483; 1905a, p. 26.—Miers, 1877b,
p. 139; 1880, p. 69.—Meinert, 1877-1878, pp. 171-172; 1880, p. 495; 1890,
pp. 184-185.—M. Sars, 1858 (1859), p. 150.—Hoek, 1882, p. 65.—Mayer,
1882, pp. 62-64, figs. 20-22; 1890, pp. 65-68, pl. 2, figs. 26-33, pl. 4, fig. 31,
pl. 6, fig. 38; 1903, pp. 120-123, pl. 5, figs. 19-21, pl. 8, fig. 24.—Stuxberg,
1882, p. 764; 1887, p. 73.—Schneider, 1883, p. 30; 1884, pp. 130-131; 1891,
pp. 111, 122; 1924 (1926), p. 60.—Koelbel, 1886, p. 42.—G. Sars, 1886,
pp. 69, 89; 1895, pp. 659-660, 700, pl. 237, fig. 1.—Hansen, 1887b, pp. 173-
174.—Vosseler, 1889, p. 159.—Pfeffer, 1889 (1890), pp. 87, 94.—Klinckow-
strém, 1892, p. 90.—Ohlin, 1895a, pp. 63-64, xvii, xix; 1895b, p. 486.—
Vanhéffen, 1897, pp. 202, 203, 213.—Scott, 1899, p. 81; 1901, pp. 267-
268.—Ortmann, 1901, pp. 155-156.—d’ A. Thompson, 1901, p. 42.—Lénnberg,
1902 (1903), p. 50.—Holmes, 1904 (1905), p. 527.—Nordgaard, 1905,
p. 185.—M. Rathbun, 1905, pp. 7, 78-79.—M. Grieg, 1907, p. 527.—Briggen,
1909, p. 43.—Stephensen, 1913a, pp. 223-225; 1913b, p. 68; 1916, p. 295;
1927a, pp. 148-149; 1927b, p. 13; 1928, pp. 384-3886, fig. 92 (5-10); 1929a,
pp. 179-180, fig. 334; 1929b, pp. 20, 34; 1933, pp. 60, 77; 1935, p. 188; 1940,
pp. 73-74; 1942, pp. 439-441, 502, 503; 1944b, pp. 136-1387, 148, 159, 162.—
Bjérek, 1915, p. 36.—Derjugin, 1915, pp. 453, 456; 1928, p. 282.—Oldevig,

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 45

1917, p. 40; 1933, pp. 266-269, fig. 1-2 (p. 267), figs. 1-3 (p. 268).—J. Grieg,
1925, p. 22.—Johansen, 1925, p. 204; 1930, p. 94.—Shoemaker, 1926, p. 11;
1930, pp. 353 (135)-354 (136).—Gurjanova, 1929b, p. 70; 1931, p. 201.—
Dons, 1935, p. 110.—Dunbar, 1942, p. 42; 1954, pp. 784, 788.—Schellenberg,
1942, p. 238, fig. 198.—Dahl, 1946, p. 22.—Utinomi, 1943c, pp. 296-297,
fig. 10; 1947, p. 78.—Stock and Bolklander, 1952, p. 4.—Bousfield, 1956a,
p. 32; 1956b, p. 144; 1958a, p. 321; 1962, p. 53.—Bousfield and Leim, 1958,
p. 18.—Brunell, 1961, p. 7.—Prefontaine and Brunel, 1962, p. 256.

Caprella cercopoides White, 1852, p. cevii, fig. 1.

Caprella robusta Stimpson, 1854 (1853), p. 44.—Mayer, 1882, p. 66; 1890, p. 73.

Caprella punctata [not Risso] Boeck, 1861, pp. 676-677; 187la, p. 277 (197);
1873-76, pp. 698-699, pl. 32, fig. 11.—Norman, 1886, p. 26; 1905a, p. 26.—
G. Sars, 1886, pp. 69, 89; 1895, pp. 660-661, 700-701, pl. 237, fig. 2, pl. 8,
fig. 3.— Briiggen, 1907, p. 238.—Nordgaard, 1911 (1912), p. 24.—Stephensen,
1928, p. 385, fig. 92 (8); 1933, pp. 60, 77; 1940, p. 74; 1942, pp. 442-443,
504, 505; 1944a, p. 50; 1944b, p. 159.

Caprella Septentrionalis.—Herklots, 1861, p. 43.

Caprella Stimpsoni Bate, 1862, p. 361.—Whiteaves, 1901, p. 220.

[?] Caprella hystrix [not Krgyer].—Bate and Westwood, 1868, pp. 63-64.—
M’Intosch, 1874, p. 272.—Koehler, 1884 (1885), pp. 112, 117; 1885, pp. 54,
61.—Bate, 1887, p. 175.—Bonnier, 1887, p. 354.—Robertson, 1886-87 (1888),
p. 72.—Walker, 1895b, p. 475.—Norman, 1905b, p. 85.—Norman and
Scott, 1906, p. 99.

Caprella longicornis Boeck, 1871a, pp. 274 (194)- 275 (195); 1873-76, pp. 691-693,
pl. 32, fig. 7.

Caprella Lovéni Boeck, 1871a, p. 276 (196); 1873-76, pp. 694-696, pl. 32, fig. 8.—
Meinert, 1877-78, p. 171.—G. Sars, 1895, pp. 662-663, pl. 238, fig. 2.—
Stephensen, 1928, pp. 385-386, fig. 92 (10).

[?] Caprella hystryxz.—Bate, 1878, p. 509.

Caprella septentrionalis f. typica Mayer, 1890, p. 66.

Caprella septentrionalis f. longicornis Mayer, 1890, p. 66, pl. 2, figs. 26-27, 33,
pl. 4, fig. 31.

Caprella septentrionalis f. nodigera Mayer, 1890, p. 66.

Caprella septentrionalis f. polyceros Mayer, 1890, p. 66, pl. 2, fig. 32.

Caprella septentrionalis f. parva Mayer, 1890, p. 66, pl. 2, figs. 28-31.

Caprella monocera G. Sars, 1895, pp. 661-662, pl. 238, fig. 1.—Ohlin, 1895a, pp.
villi, xiii, xvii, xix, 64-65.—Nordgaard, 1905, p. 185.—Stephensen, 1928,
p. 385, fig. 92 (9); 1933, pp. 60, 77; 1940, p. 74; 1942, pp. 442, 504, 505;
1944b, p. 159.

Caprella septentrionalis f. spinigera.—Hansen, 1895, p. 130.

Caprella stimpsoni.—Holmes, 1904 (1905), p. 527.

Caprella septentrionalis longicornis.—M. Rathbun, 1905, pp. 7, 78-79.

Caprella septentrionalis stimpsont.—M. Rathbun, 1905, p. 7, 79.

Caprella septentrionalis polyceros.—M. Rathbun, 1905, p. 7, 79.

Caprella septentrionalis lovéni.—Stephensen, 1929a, p. 180, fig. 334.

Caprella septentrionalis monocera.—Stephensen, 1929a, p. 180, fig. 334.

Caprella septentrionalis punctata.—Stephensen, 1929a, p. 180, fig. 334.

Caprella septentrionalis f. monocera.—Oldevig, 1933, p. 266, fig. 2 (p. 267).

Caprella septentrionalis f. punctata.—Oldevig, 1933, p. 266, fig. 3 (p. 268).

Caprella septentrionalis f. lovéni.—Oldevig, 1933, p. 266, fig. 2 (p. 268).

Caprella lovéni.—Stephensen, 1940, p. 74; 1942, pp. 441, 504, 505; 1944b, p. 159,

Diacnosts.—Body usually with numerous spines and _ tubercles,
peduncle of antenna 1 rarely with setules, ratio of total length to

46 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 19.—Caprella septentrionalis, male; a, lateral view; b, abdomen; c, maxilliped;
d, gnathopod 1; ¢, maxilla 1; f, right mandible; g, left mandible.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 47

Ficure 20.—Caprella septentrionalis, female; a, lateral view; b, pereopod 7; c, gnathopod 2;
d, abdomen.

length of basis of gnathopod 2 usually greater than 13.0, inner surface
of gnathopod 2 with small tooth adjacent to poison tooth.

Description.—Body spination variable. Length of largest male 20
mm, largest female 20 mm, smallest ovigerous female 9 mm.

Peduncle of antenna 1 occasionally with dense setules. Length of
antenna 2 longer or shorter than peduncle of antenna 1.

Mouthparts typical of genus, right lacinia mobilis 5-toothed.

Propodus of gnathopod 1 with 2 proximal grasping spines, grasping
margin of dactylus and propodus serrate. Palm of propodus of gna-
thopod 2 with proximal poison tooth and small tooth on inner surface,
distally with small tooth, notch, and rectangular projection, antero-
distal margin occasionally with projection; basis short and robust.

Gills usually elliptical, occasionally oval and inflated.

Propodus of pereopods 5-7 with pair of proximal grasping spines.

Abdomen of male and female typical of genus.

VARIATION.—Body spination varies from quite spinose to almost as
smooth as in (. linearis. Usually the cephalon is furnished with at
least a single spine.

48 U.S. NATIONAL MUSEUM BULLETIN 278

The ratio of the total length to the length of the basis of gnathopod
2 varies slightly as illustrated in figure 22.

DisTRIBUTION.—Type-locality: “It seems to be common in
the Greenland seas***, but is not mentioned by Sabine and
Ross**** ,,..***Sydproven, Julianchaab, Fiskenaesset, Godthaab and
Nenertalik are those places from which it has been sent.”’ (Translated
from Krgyer, 1838)

Other records: Murman coast; Novaya Zemlya; White Sea; N.
Russia; Norway to France and British Islands; Faeroe Islands; Jan
Mayen; Iceland; East and West Greenland; Baffin Bay and Davis
Straits; coast of North America from Hudson Bay to Maine.

New records: Many localities south to Georges Bank.

Remarks.—This species occupies almost the same habitat as C.
linearis. It has been taken in the stomachs of the pollock Pollachwus
(Scott, 1901) and the rock cod Gadus (Johansen, 1925). It ranges in
depth from the surface to 1026 m.

As evidenced by the long synonymy, this species exhibits a variety
of body forms. Mayer (1890, 1903) viewed these forms as variants
of the same species; however, Stephensen (1928, 1940) treated several
of these forms as distinct species or subspecies. The material available
to me of this species is quite large and intergradations between the
forms are numerous. No geographical isolation of these forms is
evident and therefore they probably represent only infrasubspecific
variants. Perhaps ecological data would reveal some other type of
isolating mechanism, but type of datum is not available to me.

Figure 21 is a variant of this species, which was collected in Casco
Bay, Maine. It corresponds to G. Sar’s (1895) concept of C. monocera
and to Mayer’s C. septentrionalis f. longicornis. In common with C.
linearis it bears numerous setules on the peduncle of antenna 1 and
has elongated pereonites 1 and 2. Since the females of C. linearis and
C. septentrionalis are frequently indistinguishable and the gnathopods
are quite similar, this variant caused concern over the distinctiveness
of these species because it might represent a hybrid. In figure 22 the
ratio of the total length to the length of the basis of gnathopod 2
is plotted. The variant and males and females of C. septentrionalis
are not significantly different in this character; however, they are
significantly different from both the males and females of C. linearis.
Relying on this character, the females of the two species can be
separated and it justifies the inclusion of the variant within C. sep-
tentrionalis. The problem of the possible species distinctiveness of
Mayer’s varieties and Stephensen’s species and subspecies is still
unresolved, and its solution will probably not be found until ecologic
and genetic studies can be carried out.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 49

Ficure 21.—Caprella septentrionalis, male variant; a, lateral view; 0, gnathopod 2; c
gnathopod 1.

?

Caprella unica Mayer, 1903
FIGURES 23, 24, 55

Caprella unica Mayer, 1903, p. 127, pl. 5, fig. 39.—M. Rathbun, 1905, pp. 7, 79.
Caprella grahami Wigley and Shave, 1966, pp. 289-296, figs. 1-5.

Draenosis.—Cephalon without anteriorly directed triangular pro-
jection, propodus of pereopods 5-7 with few setae but without grasping
spines.

Description.—Body smooth to spiny. Length of largest male 18
mm, largest female 8 mm, smallest ovigerous female 5 mm.

Peduncle of antenna 1 with or without dense setules in male,
female lacking setules.

Mouthparts typical of genus except dactylus of maxilliped palp
quite robust, right lacinia mobilis 5-toothed.

Propodus of gnathopod 1 with 2 proximal grasping spines. Gnatho-
pod 2 similar to C. linearis and C. septentrionalis.

Propodus of pereopods 5-7 without grasping spines and with only
few short setae.

Abdomen typical of genus.

50 U.S. NATIONAL MUSEUM BULLETIN 278

7.0 80 9.0 10.0 11.0 12.0 13.0 14.0 15.0 16.0 17.0 18.0 19.0

Ficure 22.—Ratio of total length divided by length of basis of gnathopod 2; a, Caprella
septentrionalis, males, n=47; b, C. septentrionalis, variant males, n=10; c, C. septentrio-
nalis, females, n= 42; d, C. linearis, females, n=16; ¢, C. linearis, males, n= 34. Vertical
line represents the range, horizontal line the mean, stippled area two standard errors
either side of the mean, and white rectangle one standard deviation either side of the
mean.

VaRIATION.—The body of this species varies in the amount of
spination, larger individuals tending to be less spinose (compare figs.
23c, 24a, 24b). The peduncle of antenna 1 shows some variability in
the density of setules; larger males tend to develop more setules.

DistrrisutTion.—Type-locality: Albatross sta. 2253, 40°34’30”" N.,
69°50'45’” W., 59 m.

Other records: Southeastern New England, Long Island Sound,
Vineyard Sound, Great Harbor in Woods Hole, passage between Vine-
yard Sound and Buzzards Bay, Cape Cod Canal; 0.3 to 25 m.

New records: St. John’s, Newfoundland; Casco Bay, Maine; off
Cape Cod, 40°-43° N., 65°-72° W.; 37°31’ N., 74°52’ W.

Remarks.—Wigley and Shave (1966) give an excellent description
of C. grahami which they thought to be distinct from C. unica due
to the presence of spines on the dorsal surface of the latter species.
Mayer based his description of C. wnica upon a single immature male
which was quite spiny; however, recent material collected from the
area near the type-locality has yielded numerous specimens which
intergrade between the smooth and spiny forms. Since the other
characters of these two species agree quite closely and intergrades
are present between the two body forms, C. grahami is suppressed
as a Junior synonym of C. unica.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC BI

J

Ficure 23.—Caprella unica, male holotype; a, maxilla 1; b, gnathopod 1; ¢, lateral view;
d, right mandible; ¢, maxilla 1; f, pereopod 6; g, maxilliped; A, right mandible.

o2 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 24.—Caprella unica; a, lateral view of male with intermediate spination; ), lateral
view of male holotype of C. grahamt; c, female gnathopod 2; d, male gnathopod 2; e,
female pereopod 7; f, lateral view of female allotype of C, grahami.

C. unica has been reported on two species of starfishes, Asterias
forbest (Desor) and Asterias vulgaris Verrill. This association with
echinoderms is unusual and has been found for Pariambus typicus
(Krgyer) on Asterias and Solaster (Chevreux and Fage, 1925),
Aeginella spinosa on Brisinga (Mayer, 1903), Caprella linearis on
Asterias (Ohlin, 1895a, probably C. unica, p. 33), Phtisica marina
on Asterias (Meinert, 1880), and I have found C. penantis clinging
to the spines of Arbacia and C. scaura on an unidentified urchin.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC aa

Deutella Mayer, 1890

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 3-segmented, setal formula for terminal article
1-x-1, 1, or 2, molar present; outer lobe of maxilliped larger than
inner lobe; gills on pereonites 3 and 4; pereopods 3 and 4, 2-segmented,
pereopod 5, 6-segmented, inserted at the posterior of pereonite 5;
abdomen of male with pair of appendages and pair of setose lobes,
female with pair of lobes, with or without setae.

Type-species: Deutella californica Mayer, 1890 (by monotypy,
subsequently designated by Dougherty and Steinberg, 1953).

Remarxs.—The genus Deutella is presently composed of 4 species;
D. californica Mayer, 1890; D. incerta (Mayer, 1903); D. mayeri
Stebbing, 1895; and D. venenosa Mayer, 1890. Steinberg and
Dougherty (1957) state they believe Luconacia incerta belongs
to Deutella because differences between the two genera seem minor.
The differences to which they refer are the lack of a definitive palmar
surface on the propodus of pereopod 5 and the presence of swimming
setae in Luconacia. There exist, however, several more important
differences which are presented in table 1. It can be seen that the
genus Luconacia differs from Deutella except for the serrate lacinia
mobilis which D. mayeri shares with LZ. incerta. The remaining dif-
ferences seem sufficient to separate Deutella and Luconacia; therefore,
the genus Luconacia has been reestablished.

Deutella mayeri and D. sp. Mayer, 1890, differ from the other
species of the genus in haying only 1 or 2 setae on the terminal ar-
ticle of the mandibular palp. D. mayeri exhibits many similarities
with the species of Paracaprella, particularly P. tenuis which some-
times bears a small 3-segmented palp with a terminal seta. D. mayeri

TABLE 1.—Taxonomic characters separating Deutella and Luconacia

Deutella Dertella Deutella Luconacia
californica mayeri venenosa incerta
Apical setae on terminal present present present absent
article of maxilliped
and projection on
penultimate article
Terminal article of no knobs no knobs no knobs knobs
mandibular palp 1-x-1 1 or 2 I-x-1 1-x-1
Tip of male abdominal no papillae no papillae no papillae — papillae
appendage fringe no fringe no fringe fringe
Female abdomen lobes lobes lobes no lobes
Insertion of pereopod 5 posterior posterior posterior midlength
Right lacinia mobilis 5-toothed serrate 5-toothed serrate

54 U.S. NATIONAL MUSEUM BULLETIN 278

and Mayer’s D. sp. may represent intergrades between the two
genera. Since the mandibular palp of D. mayeri is not reduced in
size as it is in Paracaprella, I have chosen to leave this species in
Deutella, although its position is questionable.

If D. mayeri and Mayer’s D. sp. are disregarded because of their
apparent divergence from the other species of Deutella, then Luconacia
and Deutella are separated geographically, Luconacia being restricted
to the Atlantic and Deutella to the Pacific.

Deutella californica Mayer, 1890
FIGurRE 52

Deutella californica Mayer, 1890, pp. 27-28, pl. 1, figs. 3-4, pl. 3, figs. 15-16,
pl. 5, fig. 18—Dougherty and Steinberg, 1953, p. 44; 1954, pp. 169, 171,
fig. 82f.—Steinberg and Dougherty, 1957, pp. 279-281, figs. 15, 21-23,
28.—Gardella, 1962, pp. 1-2.—Johnson and Juskevice, 1965, p. 39.—Johnson,
1966, appendix 4, p. 4.

REMARKS.—This species was originally described by Mayer from
Cape Mendocino, Calif. and later reported from Monterey Bay by
Dougherty and Steinberg. Steinberg and Dougherty (1957) reported
it as occurring at “. . . probably Port Aransas, Texas.”” The material
they examined undoubtedly belongs to D. californica; however, no
additional material of this species has been collected in the western
North Atlantic, and it is probable that this species is not a member
of the caprellid fauna of that area.

D. californica can be distinguished from D. mayeri by its 1-x-1
setal formula for the terminal article of the mandibular palp, the
5-toothed lacinia mobilis on the right mandible, and the single dorsal
cephalic spine. For a description and figures of this species, consult
Steinberg and Dougherty (1957).

Deutella mayeri Stebbing, 1895
Figures 25, 26, 52

Deutella mayeri Stebbing, 1895, pp. 400-402, pl. 15a.—Mayer, 1903, pp. 44-45.

Draanosis.—Body without spines.

Description.—Body smooth, anterolateral margin of pereonite 2
with triangular projection. Length of largest male 4 mm, female 3
mm.

Mandible with 3-segmented palp, 1 or 2 setae on terminal article,
penultimate article with single distal setae. Left mandible with 5-
toothed incisor, 5-toothed lacinia mobilis, setal row of 3 serrate setae.
Right mandible with 5-toothed incisor, apically serrate lacinia mobilis,
setal row of 2 serrate setae. Palp of maxilla 1 with 3 or 4 apical spines
and several setae, outer lobe with 4 or 5 apical spines. Inner lobe of
maxilla 2 with 4 or 5 apical setae, outer lobe with 5 or 6 apical setae.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC na

V
Ficure 25.—Deutella mayeri, male; a, lateral view; b, maxilla 1; c, gnathopod 2; d, gnatho-

pod 1; ¢, maxilliped; f, abdomen; g, pereopod 4; h, pereopod 3; i, labium; j, pereopod 5;
k, maxilla 1; /, left mandible; m, right mandible.

Outer lobe of maxilliped with 2 apical setae, 1 plumose and 1 non-
plumose seta, and several medial setae; inner lobe with 3 or 4 apical
setae of which 2 or 3 plumose; terminal article of palp with 2 or 3
distal setae, penultimate article with distal triangular projection.

56 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 26.—Deutella mayeri, female; a, lateral view; b, gnathopod 1; c, maxilliped; d,
pereopod 3; ¢, abdomen; f, maxilla 1; g, maxilla 2; h, left mandible; 7, right mandible.

Propodus of gnathopod 1 triangular with 1 proximal grasping
spine; grasping margins of dactylus and propodus serrate, dactylus
with distinct teeth on grasping margin. Palm of propodus of gnathopod
2 with proximal grasping spine, distal tooth and notch at midlength;
dactylus smooth.

Terminal article of pereopods 3 and 4 with 1 plumose and 2 non-
plumose setae. Pereopods 5-7, 6-segmented, palm of propodus with
pair of proximal grasping spines.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 57

Abdomen of male with pair of appendages and pair of setose lobes,
female with pair of nonsetose lobes.

DistrinuTion.—Type-locality: Antigua, West Indies.

New records: Bahfa Fosforescente, Puerto Rico.

Fallotritelia, new genus

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 3-segmented, setal formula for terminal article
1-x-1, molar absent; outer lobe of maxilliped larger than inner lobe;
cills on pereonites 3 and 4; pereopods 3 and 4, 1-segmented, pereopod
5, 6-segmented; abdomen of male and female with pair of lobes.

Type-species: Fallotritella biscaynensis, new species (by present
designation).

Remarxs.—This new genus is closely related to Tritella Mayer,
1890, and Triliropus Mayer, 1903, but differs from these by the lack
of a molar on the mandible, having the inner lobe of the maxilliped
rounded instead of flattened, and by the lack of setae on the dactylus
of the maxillipedal palp. Table 2 summarizes the differences between
these genera.

TaBLE 2.— Taxonomic characters separating Fallotritella, Tritella, and Triliropus

| Fallotritella Tritella laevis, Tritella tenuis- Triliropus
biscaynensis pilimana, ornata sima uncinatus
Swimming setae on absent present absent absent
antenna 2
Inner lobe of maxilliped round flat flat flat
Dactylus of maxilliped no yes yes ?
with setae
Grasping spines on pro- 5 usually midlength proximal absent
podus of pereopods 5-7 absent, 6 or proximal
and 7
proximal
Insertion of perecopod 5 posterior posterior midlength posterior to
midlength

Tritella tenuissima Dougherty and Steinberg, 1953, differs from the
other species of Tritella in that swimming setae are lacking on antenna
2 and pereopod 5 is inserted near midlength on pereonite 5. These
characters are found in Triliropus, and Tritella tenuissima should
probably be transferred to this genus; however, since I have not
examined specimens of Triliropus I will refrain from making the
transfer at this time.

The generic name is derived from the Latin term “fallo,” meaning
false, in combination with Tritella, the name of a closely allied genus.

58 U.S. NATIONAL MUSEUM BULLETIN 278

Fallotritella biscaynensis, new species

Ficures 27, 28, 53

Diacnosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.

Description.—Male holotype: Body spinose. Cephalon with
single dorsal anteriorly directed spine. Pereonite 1 with single postero-
dorsal spine. Pereonites 2-4 with trace of dorsal spine at midlength
and with posterodorsal spine. Pereonites 5-7 without spines. Length
4 mm.

Antenna 1 approximately length of pereonites 2-4, flagellum of 11
articles. Antenna 2 approximately length of peduncle of antenna 1.

Mandibular palp 3-segmented, setal formula for terminal article
1-5-1. Mandibles with 5-toothed incisor, lacinia mobilis toothed, 2
accessory plates on right mandible and 3 on left, 2 setae present.
Palp of maxilla 1 with 2 apical and 2 medial setae, outer lobe with
6 serrate spines. Outer lobe of maxilliped with 3 apical setae and 1
seta at midlength on medial margin, inner lobe rounded with 2 apical
setae; dactylus of palp slightly curved with grasping margin serrate.

Propodus of gnathopod 1 triangular with 2 proximal grasping
spines; grasping margins of dactylus and propodus not serrate, dac-
tylus with single tooth and seta at tip. Propodus of gnathopod 2 with
proximal grasping spine and distal notch, basis approximately length
of pereonite 2.

Gills elliptical and sometimes held with long axis parallel to body.

Pereopods 3 and 4, l-segmented with single seta at tip. Pereopod
5 inserted posteriorly on pereonite 5, propodus without grasping
spines. Propodus of pereopods 6 and 7 with 2 proximal grasping
spines.

Abdomen with pair of setose lobes, 2 setae on anterior end of
abdomen; penes large and medial.

Female allotype: Body spinose. Cephalon with single dorsal an-
teriorly directed spine. Pereonite 1 with single posterodorsal spine.
Pereonite 2 with trace of pair of dorsal spines at midlength and with
single posterodorsal spine. Pereonites 3 and 4 with pair of dorsal
spines at midlength and single posterodorsal spine. Pereonite 5 with
trace of dorsal spine at midlength. Pereonites 6-7 without spines.
Length 2.5 mm.

Antenna 1 approximately length of pereonites 3-5, flagellum of 5
articles. Antenna 2 approximately length of peduncle of antenna 1.

Mouthparts as in male except setal formula for terminal article of
mandibular palp 1-4-1.

Gnathopod 1 as in male. Gnathopod 2 as in male except propodal
notch absent.

Gills as in male.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 59

Ficure 27.—Fallotritella biscaynensis, male holotype; a, lateral view; b, left mandible; c,
maxilliped; d, abdomen; e, gnathopod 2; f, gnathopod 1; g, pereopod 6; h, pereopod 5;
1, pereopod 3.

279-475—68——5.

60 U.S. NATIONAL MUSEUM BULLETIN 278

3

Ficure 28.—Fallotritella biscaynensis, female allotype; a, lateral view; b, gnathopod 1;
c, gnathopod 2; d, pereopod 5; ¢, right mandible; f, left mandible; g, maxilla 1; h, abdomen.

Pereopods as in male.

Abdomen similar to male except lobes separated by slight medially
raised area.

VariaTion.—The holotype was the largest male specimen. The
smallest ovigerous female was 3 mm; however, the allotype had an
empty brood pouch at 2.5 mm.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 61

The degree of spination of the dorsal surface of the pereonites
varied considerably. The most spiny specimens had 1 posterior spine
on perenoite 1, 2 at midlength, and 1 posterior spine on pereonites
2-4, and 2 spines at midlength on pereonite 5. Some of the specimens
showed different degrees of reduction of these spines, usually losing
the spines at midlength first.

In some of the specimens, grasping spines were present on the
propodus of pereopod 5.

The mouthparts were rather consistent. The setal formula for the
terminal article of the mandibular palp varied only from 1-3-1 to
1-5-1, and the outer lobe of the maxilliped occasionally had a seta
at midlength on the medial margin in addition to 3 apical and 1 seta
on the middle of the anterior surface.

Distrinution.—Type-locality: Key Biscayne, Fla., on bridge pilings
in Bear Cut, 2 m, July 15, 1966, 1 male holotype USNM 120179.

Other records: Bermuda, Oct. 10, 1960, 1 male paratype USNM
120188. Soldier Key, Fla., Jan. 27, 1961, 1 male paratype USNM
120186. Hummelinck sta. 1413, East of Soldier Key, Fla., 2 m,
Sept. 5, 1963, 1 male paratype USNM 120184. Soldier Key, Fla.,
1-2 m, July 16, 1966, 2 male paratypes USNM 120185. Key Largo,
Fla., July 9, 1966, 1 female allotype USNM 120180. Long Key, Fla.,
May 11, 1961, 1 male paratype USNM 120183. Tortugas, 1 female
paratype USNM 120182. Freelance sta. 85-56, west shore of lagoon
near Oyster Pond Landing, Barbuda, Apr. 6, 1956, 2 female para-
types USNM 120187. Freelance sta. 47-56, off Pigeon Island, St.
Lucia, 1-3 m, Mar. 22, 1956, 1 male paratype USNM 120181.

Remarxs.—This species is named for Key Biscayne, Fla., where
the holotype was collected. The only habitat records for this species
are from red algae. It is difficult to detect this species since it is quite
small and usually is covered with detritus.

Hemiaegina Mayer, 1890

Flagellum of antenna 2 biarticulate, swimming setae absent; man-
dibular palp absent, molar present; outer lobe of maxilliped larger
than inner lobe; gills on pereonites 3 and 4; pereopods 3 and 4, 1-
segmented, pereopod 5, 6-segmented; abdomen of male and female
with pair of biarticulate appendages.

Type-species: Hemiaegina minuta Mayer, 1890 (by monotypy).

Hemiaegina minuta Mayer, 1890
Ficures 29, 30, 50

Hemiaegina minuta Mayer, 1890, p. 40, pl. 1, figs. 25-27, pl. 3, figs. 32-35, pl. 5,
figs. 52-53, pl. 6, figs. 13, 33-34, pl. 7, fig. 4; 1903, p. 65, pl. 6, fig. 75.—
Arimoto, 1930, pp. 45-47, fig. 2—? Barnard, 1937, pp. 134, 198.—Utinomi,

62 U.S. NATIONAL MUSEUM BULLETIN 278

1947, p. 70.—Edmondson and Mansfield, 1948, pp. 206-207, fig. 3.—Stein-
berg and Dougherty, 1957, pp. 281-283, figs. 8-11, 13, 29.—McCain, 1965,
. 192.
aes quadripunctata Sundara Raj, 1927, pp. 126-127, pl. 18.

DiaGNosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.

Description.—Body smooth except for pair of ventral spines
between insertions of gnathopods 2. Length of largest male 4 mm,
female 3.2 mm, smallest ovigerous female 2.7 mm.

Antenna 1 approximately length of pereonites 1-4. Antenna 2
slightly longer than peduncle of antenna 1.

Left mandible with 5-toothed incisor, 5-toothed lacinia mobilis,
setal row absent, numerous setules present between molar and lacinia
mobilis. Right mandible similar to left except lacinia mobilis apically
serrate instead of 5-toothed. Palp of maxilla 1 with 2 or 3 apical spines
and 1 seta, outer lobe with 6 spines. Outer lobe of maxilla 2 with
4 or 5 apical setae, inner lobe with 3-5 apical setae. Outer lobe of
maxilliped with 2 apical setae, 1 seta near midlength of medial margin,
and larger specimens with 1 or 2 proximal setae on medial margin;
inner lobe with 2 setae and 1 small spine; dactylus of palp with
serrate grasping margin.

Propodus of gnathopod 1 with 1 small proximal grasping spine
on medial surface and large proximal knob covered with small pro-
jections, dactylus not serrate on grasping margin. Propodus of
enathopod 2 with proximal grasping spine, distal notch with slightly
proximal spine, and distal bilobed tooth.

Gills elliptical.

Pereopods 3 and 4, 1-segmented with 2 or 3 apical setae. Palm
of propodus of pereopods 5-7 concave with 5 or 6 proximal knobs,
each with small spine.

Abdomen of male and female with pair of biarticulate appendages,
dorsal lobe bilobed.

DistRrBuTion.—Type-locality: Off Amoy, China, at a depth of
15-46 m.

Other records: Off Virginia; Port Aransas, Texas; Oahu, Hawaii,
Tateyama, Honshu, Japan; 1°42.5’ S., 130°47.5’ E.; region of Fre-
mantle, Australia; Krudadai Island, India; South Arabian coast.

New records: Several localities near Bermuda; off Cape Hatteras,
N.C.; Elliot Key, Fla.; Loggerhead Key, Tortugas; 29°44’ N., 88°
23.5’ W.; St. John, Virgin Islands; False Bay, South Africa; Bora
Bora.

Remarxs.—This species is easily identified by the unusual hex-
agonal outline of the pereonites when viewed dorsally. When viewed
in this position the gnathopods 1 are usually held in a posteriorly

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 63
Ww a

ae s.\ J a
me NS = £

SS

\ Y SX

Ficure 29.—Hemiaegina minuta, male; a, dorsal view; b, maxilliped; c, left mandible; d,
pereopod 4; ¢, gnathopod 1; f, maxilla 1; g, gnathopod 2; h, abdomen; 1, pereopod 7;
j, maxilla 2; k, right mandible.

directed position and gnathopods 2 are held away from the body
(figs. 29a, 30a).

I have designated a lectotype from Mayer’s type-series which is
deposited in the Copenhagen Museum.

H. minuta has been taken in plankton tows and from Sargassum.

64 U.S. NATIONAL MUSEUM BULLETIN 278

oO

ee

Ficure 30.—Hemiaegina minuta, female; a, dorsal view; b, gnathopod 1; ¢, gnathopod 2;
d, abdomen; ¢, left mandible; f, maxilliped; g, pereopod 7.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 65

Hemiproto, new genus

Flagellum of antenna 2, 2- to 4-segmented, swimming setae absent;
mandibular palp 3-segmented, setal formula for terminal article 1-1
or 1-x-1, molar absent; outer lobe of maxilliped equal to inner lobe;
gills on pereonites 2-4; pereopods 3 and 4, 6-segmented, pereopod
5, 5-segmented; abdomen of male with 2 pairs of small uniarticulate
appendages and pair of nonsetose lobes, female with 1 pair of
uniarticulate appendages.

Type-species: Hemiproto wigleyt, new species (by present
designation).

Remarxks.—This new genus is closely related to Protomina Mayer,
1903, in that the males of both genera have 2 pairs of abdominal
appendages; however, in Protomina the appendages are biarticulate
and positioned near the posterior part of the abdomen and in Hemi-
proto the appendages are uniarticulate, one positioned near midlength
and one anteriorly.

Another character which these genera share is the elongation of
the carpus of gnathopod 2. This character is not unique to the 2
genera since a slight elongation occurs in Phtisica antillensis and is
probably not of generic significance.

Zemiproto lacks a molar on the mandible; however, Mayer did not
describe all of the mouthparts for his species, including Protomina.
Since specimens of Protomina are not available to me, I cannot be
certain that this character is useful for separating these 2 genera.

The generic name is a combination of Greek terms, “hemi” meaning
half, referring to the number of articles in the abdominal appendages
as compared with Phtisica and Protomina, and ‘‘protos’” meaning
first, which was the original name previously used for Phtisica.

Hemiproto wigleyi, new species
FicureEs 31, 32c-e, 50

DiaGnosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.

Drscription.—Male holotype: Body smooth. Length 7.5 mm.

Antenna 1 approximately length of body, flagellum of 9 articles.
Antenna 2 approximately as long as articles 1 and 2 of antenna 1,
flagellum of 4 articles.

Mouthparts as in Phtisiea marina except setal formula for terminal
article of mandibular palp 1-2-1 and outer lobe of maxilla 1 with 4
extremely toothed spines and 2 nontoothed spines.

Propodus of gnathopod | triangular with 4 proximal grasping spines,
1 large and slightly proximal to others, grasping margin finely serrate;
dactylus not serrate. Propodus of gnathopod 2 with 3 grasping spines,
1 large and slightly proximal to others, grasping margin with numerous

66 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 31.—Hemiproto wigleyi, male paratype; a, lateral view; b, left mandible; c, labium;
d, gnathopod 1; ¢, pereopod 4; f, abdomen.

small spines and 4 or 5 small elevations on distal portion; carpus
elongate, slightly shorter than basis.

Gills on pereonites 2-4, gill on pereonite 2 smallest.

Pereopods 3 and 4, 6-segmented, propodus with 2 or 3 grasping
spines. Pereopod 5, 5-segmented propodus without grasping spines.
Pereopods 6 and 7 missing.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 67

Abdomen with 2 pairs of small uniarticulate appendages and pair
of nonsetose lobes, posterior appendage pair with hooked seta at
tip, anterior appendage pair pyriform with seta at tapered tip.

Female allotype: Body smooth. Length 4.4 mm.

Antenna 1 approximately length of body, flagellum of 7 articles.
Antenna 2 extending slightly beyond articles 1 and 2 of antenna 1,
flagellum of 4 articles.

Mouthparts as in male except setal formula for terminal article
of mandibular palp 1-1-1.

Gnathopod 1 similar to male. Propodus of gnathopod 2 similar to
male except without elevations on distal part.

Gills as in male.

Pereopods as in male.

Abdomen with pair of uniarticulate appendages and pair of lobes.

VaARIATION.—The setal formula for the terminal article of the man-
dibular palp varies from 1-1 to 1-2-1. The number of articles in the
flagellum of antenna 1 varies from 6 to 11 and in antenna 2 from 2 to
4, Pereopods 3 and 4 occasionally have 3 grasping spines instead of
the usual 2.

Figure 32.—a and b, Paracaprella pusilla, large male; a, pereonites 1 and 2; b, dactylus of
gnathopod 2; c-e, Hemiproto wigleyi; c, abdomen of female allotype; d, gnathopod 2 of male
holotype; e, gnathopod 2 of female allotype.

68 U.S. NATIONAL MUSEUM BULLETIN 278

Disrripution.—Type-locality: Hancock Atlantic Exped. sta.
A32-39, 3 mi. N. of Coche Island, Venezuela, 35-60 m, Apr. 15, 1939,
1 male holotype USNM 120167, 1 female allotype USNM 120168,
7 male and 6 female paratypes USNM 120169.

Other records: Gosnold sta, 1556, off Ft. Lauderdale, Fla. (26°
10.6’ N., 80° 04.2’ W.), 32 m, May 26, 1964, 1 male paratype USNM
120170.

Remarks.—The specific name is in honor of Roland L. Wigley
of the U.S. Bureau of Commercial Fisheries Laboratory at Woods
Hole; Wigley collected one of the paratypes.

Luconacia Mayer, 1903

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 38-segmented, setal formula for terminal article
1-x-1 with knobs, molar present; outer lobe of maxilliped larger
than inner lobe; gills on pereonites 3 and 4; pereopods 3 and 4, 2-
segmented, pereopod 5, 6-segmented and inserted near midlength
on pereonite 5; abdomen of male with pair of appendages and pair
of lobes, female with no appendages or lobes.

Type-species: Luconacia imcerta Mayer, 1903 (by monotypy).

Remarxs.—For a summary of the differences separating this
genus from Deutella see the remarks under Deutella (p. 53).

Mayer (1903) states that antenna 2 of Zuconacia bears swimming
setae. Setae are present on antenna 2 but are no more developed than
in many other genera he considered as not bearing swimming setae.

Luconacia incerta Mayer, 1963
Figures 33-35, 52

Luconacia incerta Mayer, 1903, pp. 49-50, pl. 2, figs. 11-14, pl. 6, figs. 73-75,
pl. 9, fig. 21, 40, 57.—M. Rathbun, 1905, pp. 7, 76—Sumner, Osburn and
Cole, 1911 (1913), p. 656.

Protellopsis stebbingii Pearse, 1908, pp. 30-82, fig. 4; 1912, p. 379.—Kunkel,
1910, pp, 111-113, fig. 43.

Deutella incerta.—Steinberg and Dougherty, 1957, pp. 281, 285-286.

DraGnosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.
DescripTion.—Body spination variable. Length of largest male

9 mm, largest female 8 mm, smallest ovigerous female 3 mm.

Mandibular palp 3-segmented, setal formula for terminal article
from 1-3-1 to 1-10-1. Left mandible with 5-toothed incisor, 5-toothed
lacinia mobilis, setal row of 3 serrate setae. Right mandible with
5-toothed incisor, serrate lacinia mobilis, setal row of 2 serrate setae.

Palp of maxilla 1 with 4 apical spines and several medial setae, outer

lobe with 5 or 6 apical spines. Inner lobe of maxilla 2 with 4 or 5

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 69

apical setae, outer lobe with 5-7 apical setae. Outer lobe of maxil-
liped with 1 apical seta, 2 setae in notch near apical end of medial
margin, and several setae on middle of anterior surface; inner lobe
with 2 plumose and 2 nonplumose setae; grasping margin of terminal
article of palp serrate.

Propodus of gnathopod 1 triangular with 2 grasping spines; grasping
margins of dactylus and propodus serrate, dactylus usually with

Ficure 33.—Luconacia incerta, male lectotype; a, lateral view; b, left mandible; c, abdomen;
d, right mandible; e, maxilliped; f, maxilla 1; g, terminal article of mandibular palp; 4,
pereopod 7; 1, gnathopod 1.

70 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 34.—Luconacia incerta, female; a, maxilla 1; b, lateral view; c, maxilliped; d, maxilla
2; ¢, right mandible; f, abdomen; g, pereopod 3; h, gnathopod 1; 7, gnathopod 2; J, left
mandible.

distinct teeth on grasping margin. Propodus of gnathopod 2 with
proximal grasping spine and well-developed tooth at midlength, palm
setose; dactylus smooth. Smaller individuals show progressively
less spination (fig. 35), but the anterolateral projection of pereonite
2 and the pleural projections tend to remain distinct.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC a

Ficure 35.—Luconacia incerta, small male; a, maxilla 1; b, lateral view; c, gnathopod 2;
d, maxilliped; ¢, gnathopod 1; f, abdomen, ventral view; g, abdomen, lateral view; h,
maxilla 2; 1, pereopod 3; 7, left mandible.

12 U.S. NATIONAL MUSEUM BULLETIN 278

A small male from St. John’s bears the normal pair of cephalic
spines, but the body is otherwise different in spination. The pair of
dorsal spines at midlength on pereonite 2 are represented by a single
long spine. It also bears a single dorsoposterior spine and the antero-
lateral projections are absent. Pereonites 3 and 4 bear a single large
dorsal spine at midlength, a single dorsoposterior spine, and pleural
development is indistinct. Pereopod 4 is uniarticulate. The other
characters of this specimen agree with those of L. incerta.

DistrisuTion.—Type-locality: Off Mobile Bay, Ala., Albatross
sta. 2387, 29°24’ N., 88°04’ W., 59 m, 1 male lectotype, 1 male
paralectoypte USNM no. 26000.

Other records: Woods Hole, Mass.; Bermuda; Straits of Florida;
several localities between the mouth of the Mississippi River and
Cedar Keys, Fla.

New records: Many occurrences from Woods Hole to Key Largo,
Fla.; Port Aransas and Port Isabel, Tex.; off Yucatan; Virgin Islands;
Barbuda; Barbados; Isla de Margarita; Aruba.

Terminal article of pereopods 3 and 4 setose, basal article sometimes
with distal setae. Pereopods 5-7, 6-segmented, propodus without
palmar surface on pereopod 5; pereopods 6 and 7 more robust than
pereopod 5, propodus with pair of proximal grasping spines.

Abdomen of male with pair of uniarticulate appendages and pair
of setose lobes, appendage with small apical papillae surrounded by
fringe of small teeth; female abdomen without distinct lobes.

VARIATION.—This species varies considerably in the degree of body
spination. Larger individuals bear a pair of dorsal cephalic spines, a
pair of dorsal spines at midlength, and an anterolateral projection
on pereonite 2 and an anterolateral pleural projection on pereonites
3 and 4 of the male and on pereonite 3 of the female.

Remarxs.—I have designated a lectotype from Albatross sta. 2387.
The remaining syntypical series of Mayer’s are now designated as
paralectotypes and their localities are as follows: USNM 26001,
1 male and 1 female from Albatross sta. 2389, 29°28’ N., 87°56’ W.,
49 m; USNM 9709, 1 female from Albatross sta. 2390, 29°27'30’’ N.,
87°48'30’" W., 55 m; and USNM 26002, 1 juvenile female from
Vineyard Sound at Woods Hole.

This species is widely distributed in the temperate and tropical
areas of the western North Atlantic and has occasionally been taken
in plankton tows. It has been collected on mangrove roots, Sargassum,
Thalassia, sponges, hydroids, aleyonarians, and ascidians.

Mayerella Huntsman, 1915

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 3-segmented, setal formula for terminal article 1,

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 73

molar present; outer lobe of maxilliped larger than inner lobe; gills
on pereonites 3 and 4; pereopods 3 and 4, 2-segmented, pereopod
5, 2- or 3-segmented; abdomen of male with pair of uniarticulate
appendages and pair of lobes, female with pair of lobes.

Type-species: Mayerella limicola Huntsman, 1915 (by monotypy).

RemMarks.—This genus is now composed of 2 species, M. limicola
and M. redunca n. sp. The number of articles in pereopod 5 of the
new species varies from 2 to 3 and the generic definition has been
altered to include this variation.

Mayerella limicola Huntsman, 1915
Figures 36, 50

Mayerella limicola Huntsman, 1915, pp. 39-42, pls. 5-6.—Procter, 1933, p. 256.

Diaqnosis.—Abdominal appendage of male approximately length
of penes; propodus of gnathopod 2 with proximal grasping spine and
notch at midlength.

Description.—Body smooth. Length of largest male 7 mm, no
females examined.

Peduncle of antenna 1 approximately length of pereonite 3. Antenna
2 somewhat longer than peduncle of antenna 1.

Mandibular palp 3-segmented, setal formula for terminal article 1.
Left mandible with 5-toothed incisor, 5-toothed lacinia mobilis, setal
row of 3 serrate setae. Right mandible with 5-toothed incisor, lacinia
mobilis toothed but not distinctly 5-toothed, setal row of 2 serrate
setae, molar present. Outer lobe of labium winglike, more robust
than in most species of Caprella. Palp of maxilla 1 with 4 apical spines,
1 medial seta; outer lobe with 7 apical spines. Outer lobe of maxilla
2 with 7 apical setae, inner lobe with 5 apical setae. Outer lobe of
maxilliped with 1 apical seta, 1 middistal seta set in notch, and
several setae on anterior surface; inner lobe with 4 apical setae;
dactylus of palp serrate on grasping margin.

Propodus of gnathopod 1 triangular with 1 proximal grasping
spine, grasping margin of propodus and dactylus serrate. Propodus
of gnathopod 2 with proximal grasping spine and notch at midlength.

Gills small, elliptical.

Pereopods 3 and 4, 2-segmented, terminal article with 1 or 2 setae.
Pereopod 5, 3-segmented, terminal article with 3 or 4 short setae.
Pereopods 6 and 7, 6-segmented, propodus with numerous setae on
palmar margin and occasionally with 1 grasping spine, dactylus
slender and tapering.

Abdomen of male with pair of uniarticulate appendages of ap-
proximately length of penes and pair of lobes, penes medial.

VaRIATION.—The males varied in the presence or absence of the
single grasping spine on pereopods 6 and 7. The number of setae on

74 U.S. NATIONAL MUSEUM BULLETIN 278

m

Ficure 36.—Mayerella limicola, male; a, gnathopod 2; b, abdomen; ¢, lateral view; d, pereopod
6; e, right mandible; f, maxilliped; g, gnathopod 1; h, pereopod 4; 1, maxilla 1; 7, pereopod
3; k, left mandible; /, maxilla 2; m, labium; , pereopod 5.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 75

the terminal articles of pereopods 3 and 4 varied from 1-4, of which 1
was sometimes plumose. The number of setae on the inner lobe of the
maxilliped varied from 3 to 4.

Distrisution.—Type-locality: St. Croix River, New Brunswick,
at a depth of 9-18 m.

Other records: Several localities in the Bay of Fundy to a depth of
91m; Mount Desert Region, Maine.

New records: 44°00’ N., 68°15’ W., 101 m; 39°54’30’’ N., 70°20’00’’
We. en.

Remarxs.—The 3 males available to me were taken from greater
depths than the material which was examined by Huntsman. I have
been unable to examine Huntsman’s type material but my specimens
agree closely with his description and figures.

Thus far this species has been collected only in the Gulf of Maine to
Cape Cod region.

Mayerella redunca, new species
Figures 37, 38, 50

Draanosis.—Abdominal appendage of male much longer than penes
propodus of gnathopod 2 with grasping spine proximal of midlength
and notch distal.

Descrietion.—Male holotype: Body smooth. Length 7 mm.

Antenna 1 approximately length of pereonites 2 and 3, flagellum
of 9 articles. Antenna 2 little more than one-half length of antenna 1.

Mandibular palp 3-segmented, terminal article and penultimate
articles each with single seta. Left mandible with 5-toothed incisor,
5-toothed lacinia mobilis, setal row of 3 serrate setae. Right mandible
with 5-toothed incisor, lacinia mobilis toothed but not distinctly
5-toothed, setal row of 2 serrate setae. Palp of maxilla 1 with 3 apical
spines and 2 setae, outer lobe with 7 serrate spines. Outer lobe of
maxilliped with 2 apical setae and several setae on anterior surface and
several proximal setae, medial margin serrate; inner lobe with 2
plumose and 2 nonplumose setae; dactylus of palp wide at base,
grasping margin serrate with subterminal seta.

Propodus of gnathopod 1 triangular with 1 proximal grasping spine;
grasping margins of propodus and dactylus serrate. Propodus of
gnathopod 2 slender and elongate, grasping spine proximal to mid-
length, notch distal, basis two-thirds length of pereonite 2.

Gills oval or elliptical, third pair larger than fourth.

Pereopods 3 and 4, 2-segmented, terminal article with single seta.
Pereopod 5, 3-segmented, terminal article with 4 setae. Pereopods 6
and 7 missing.

Abdomen with pair of long uniarticulate appendages recurving at
tip; lobes setose, not distinctly separated medially.

279-475—68-_6

76 U.S. NATIONAL MUSEUM BULLETIN 278

h

Ficure 37.—Mayerella redunca, male holotype; a, lateral view; b, pereopod 5; c, left man-
dible; d, pereopod 3; ¢, maxilliped; f, gnathopod 2; g, gnathopod 1; h, abdomen; i, maxilla
1; 7, right mandible.

Female allotype: Body smooth. Length 2.25 mm.

Antenna 1 approximately length of pereonites 2 and 3, flagellum of
4 articles. Antenna 2 almost length of antenna 1.

Mouthparts as in male except dactylus of maxillipedal palp without
subterminal seta.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 77

Ficure 38.—Mayerella redunca, female; a, lateral view; b, pereopod 5; c, maxilla 1; d, gnath-
opod 1; e, pereopod 6; f, pereopod 5; g, pereopod 4; h, right mandible; 7, left mandible;
j, abdomen; k, maxilla 2; /, maxilliped (d and f-/ of allotype; a-b and e of paratype).

78 U.S. NATIONAL MUSEUM BULLETIN 278

Gnathopod 1 as in male. Propodus of gnathopod 2 with proximal
grasping spine, palm smooth and slightly convex.

Gills as in male.

Pereopods 3-5 as in male except penultimate article of pereopod 5
with 1 seta and pereopod 5, 2-segmented instead of 3-segmented.
Propodus of pereopods 6 and 7 without grasping spines but with several
setae, dactylus wide at base.

Abdomen with pair of nonsetose lobes.

Vaniation.—The only additional specimen of this species which
was available is a 3.5 mm female with a well-developed brood pouch.
This specimen agrees closely with the female allotype except the
dactylus of the left side of the maxillipedal palp bears a small sub-
terminal seta while the side does not and also pereopod 5 is 3-
segmented.

Distrisution.—Type-locality: Hancock Atlantic Exped. sta.
A32-39, 3 mi. N. of Coche Island, Venezuela, 10°50’ N., 63°54’ W.,
35-60 m, Apr. 15, 1939, 1 male holotype USNM 120176, 1 female
allotype USNM 120177, 1 female paratype USNM 120178.

Remarxs.—lI have placed this species in the genus Mayerella with
some hesitation. It agrees with the generic definition of Mayerella in
all characters which are currently in use; however, the abdomen of
this species is quite different from M. limicola. The penes are not as
long and are separated from one another, the appendages are quite
long and recurved at their tips, and the lobes are not distinctly
separated.

The species name is derived from the Latin term ‘reduncus,”
meaning curved back, referring to the recurving of the tip of the
abdominal appendage.

Metaprctella Mayer, 1890

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 8-segmented, setal formula for terminal article
1-x-y-1, molar present; outer lobe of maxilliped larger than inner lobe;
gills on pereonites 3 and 4; pereopods 3 and 4, 1-segmented, pereopod
5, 6-segmented; abdomen of male with pair of appendages and pair of
lobes, female with pair of lobes; pereonites 6 and 7 fused.

Type-species: Protella haswelliana Mayer, 1882 (by monotypy).

Metaprotella hummelincki, new species
Fiaures 39, 40, 50
Metaprotella spec. Mayer, 1903, p. 48.

Draanosts.—Body smooth except for 2 small humps dorsal to inser-
tion of gnathopod 2, pereopods 3 and 4 approximately one-fourth
length of gills; male abdominal appendage with papillae at tip.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 79

Description.—Male holotype: Body smooth except for pair of
small dorsal humps above insertion of gnathopods 2. Length 7 mm.

Antenna 1 longer than body length, flagellum broken. Antenna 2
approximately length of first 2 articles of peduncle of antenna 1.

Mandibular palp 3-segmented, setal formula for terminal article
1-10-1-1. Left mandible with 5-toothed incisor, 5-toothed lacinia
mobilis, setal row of 3 serrate setae. Right mandible with 5-toothed
incisor, lacinia mobilis toothed but not distinctly 5-toothed, setal row
of 3 serrate setae, molar with large tooth on incisor side. Palp of maxilla
1 with 6 apical spines and 2 setae, outer lobe with 4 serrate spines and
3 nonserrate spines. Outer and inner lobes of maxilla 2 with 7 apical
setae. Outer lobe of maxilliped with 3 apical setae and 2 setae in notch
at midlength of medial margin, medial and lateral margins with nu-
merous setules; inner lobe with 4 plumose and 2 nonplumose apical
setae; palp with distal serrations.

Propodus of gnathopod 1 triangular with single grasping spine,
grasping margins of dactylus and propodus serrate. Propodus of gna-
thopod 2 with proximal grasping spine and notch at midlength with
shightly proximal tooth, patm moderately setose; basis approximately
length of pereonite 2.

Gills elliptical.

Pereopods 3 and 4 approximately one-fourth length of gills with
5 or 6 apical setae. Pereopods 5-7 missing.

Abdomen with pair of appendages bearing fringe of setae and
papillae at tip and with distinct pair of lobes bearing single seta.

Female allotype: Body as in male. Length 4.5 mm.

Antenna 1 longer than body, flagellum of 12 articles. Antenna 2
approximately length of peduncle of antenna 1.

Mouthparts as in male except palp of maxilla 1 with 4 apical spines
and 4 setae, inner lobe of maxilliped with 4 plumose and 1 nonplumose
setae, and outer lobe of maxilliped with 2 apical setae.

Gnathopod 1 similar to male. Propodus of gnathopod 2 with prox-
imal grasping spine, palm without notch.

Gills as in male.

Pereopods as in male.

Abdomen with pair of lobes, each with 1 seta.

VaRIATION.—In smaller individuals antenna 1 is much shorter than
the body length and antenna 2 is as long as the peduncle of antenna 1.
The setal formula for the terminal article of the mandibular palp
varies from 1—2—1—1 to 1-12-1-1. The apical setae on the outer lobe
of the maxilliped varies from 1 to 3 and on the inner lobe the number
of nonplumose apical setae may be 1 or 2.

Distripution.—Type-locality: Magueyez Canal, La Parguera,
Puerto Rico, net haul close to bottom, Feb. 22, 1959, 1 male holotype

80 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 39.—Metaprotella hummelincki, male holotype; a, lateral view; b, right mandible;
c, left mandible; d, maxilla 1; ¢, gnathopod 1; f, maxilliped; g, pereopod 3; h, maxilla 2;
1, abdomen, ventral view; j, abdomen, lateral view; k, gnathopod 2.

USNM 120171, 1 female allotype USNM no. 120172, 3 male and 2
female paratypes USNM no. 120173.

Other records: St. Croix, Virgin Islands.

New records: Smithsonian Roebling Exped. sta. 124, Siguanea Bay,
Isle of Pines, Cuba, 4-7 m, Apr. 12, 1937, 2 male and 5 female para-

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 81

:
/1

Ficure 40.—Metaprotella hummelincki, female allotype; a, lateral view; b, maxilliped; c,
pereopod 3; d, gnathopod 2; e, left mandible; f, right mandible; g, maxilla 1.

82 U.S. NATIONAL MUSEUM BULLETIN 278

types USNM 120174. Hummelinck sta. 1423a, Bahia Fosforescente,
Puerto Rico, 1 m, Sept. 17, 1963, 1 female paratype USNM 120175.
St. John, Virgin Islands, 29 m, Mar. 17, 1906, 1 female paratype
Copenhagen Mus. St. Croix, Virgin Islands, 7 m, Feb. 19, 1906, 5
male, 5 female, and 2 juvenile paratypes Copenhagen Mus.

Remarks.—The genus Metaprotella was hitherto composed of 7
species, MM. africana Mayer, 1903; AZ. danae (Kossmann, 1880); M.
excentrica Mayer, 1890; M. haswelliana (Mayer, 1882); M. makro-
dactylos Stebbing, 1910a; AZ. problematica Mayer, 1890; and M.
sandalensis Mayer, 1898. All of these species have spiny bodies or
at least a cephalic spine with the exception of MZ. problematica which
is almost smooth. Af. hummelincki differs from Af. problematica in
that the latter species does not bear the papillae at the tip of the
male abdominal appendage and the inner margin of the outer lobe
of maxilliped is toothed and not covered with setules.

Mayer (1903) gave a short description of a Metaprotella sp. from
St. Croix. He stated that perhaps this species belonged to M. prob-
lematica; it differs, however, in having oval gills instead of rod shaped.
The gills in M. hummelincki are elliptical or cylindrical. Thus, I
cannot be certain that Mf. hummelincki is the species to which Mayer’s
M. sp. should be referred. It is unlikely that another species of Meta-
protella is present in the Caribbean and since M. hummelincki has
been collected from St. Croix, Mayer’s M. sp. has provisionally been
assigned to this species.

The species name is in honor of P. Wagenaar Hummelinck of the
Zoblogisch Laboratorium in the Netherlands, who kindly supphed
me with some of the material upon which this species is based.

The only habitat record available for this species is from Thalassia.

Paracaprella Mayer, 1890

Flagellum of antenna 2 biarticulate, swimming setae absent; man-
dibular palp 0- to 3-segmented, when 3-segmented articles reduced
as compared to Deutella, setal formula for terminal article 1, molar
present; outer lobe of maxilliped larger than inner lobe; gills on
pereonites 3 and 4; pereopods 3 and 4, 2-segmented, pereopod 5,
6-seemented; abdomen of male with pair of appendages and pair of
lobes, female with pair of lobes.

Type-species: Paracaprella pusilla Mayer, 1890 (by monotypy).

Remarks.—See remarks of Deutella on p. 53.

Paracaprella pusilla Mayer, 1890
FIGURES 32a-b, 41, 42, 53

Paracaprella pusilla Mayer, 1890, p. 41, pl. 1, figs. 28-30, pl. 3, figs. 45-47, pl. 5,
figs. 48-49, pl. 6, fig. 10; 1903, p. 67, pl. 2, figs. 36-37, pl. 7, fig. 52.—Schel-

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 83

lenberg, 1928, pp. 677-678, 1939, pp. 1386-137.—Edmondson and Mansfield,
1948, pp. 208-209, fig. 4.—Barnard, 1955, p. 99—Day and Morgan, 1956,
p. 303.—Steinberg and Dougherty, 1957, pp. 283-284, figs. 16, 19, 24, 30.—-
McCain, 1965, p. 193.

Caprella nigra Reid, 1951, pp. 283-284, 289, fig. 58.

DraGnosts.—Males with large triangular projection on antero-
ventral margin of pereonite 2; basis of gnathopod 2 short and ex-
panded with proximal knob on posterior margin, propodus with
shallow rounded notch at midlength.

Description.—Male lectotype: Body smooth on dorsal surface;
anteroventral margin of pereonite 2 with large triangular projection;
pleura developed on pereonites 3 and 4, weaker on pereonite 4.
Length 3.5 mm.

Peduncle of antenna 1 setose, usually longer than antenna 2.
Antenna 2 without swimming setae but densely setose.

Mandibular palp not present, probably represented by single seta.
Left mandible with 5-toothed incisor, 5-toothed lacinia mobilis, setal
row of 3 serrate setae. Right mandible with 5-toothed incisor, apically
serrate lacinia mobilis, setal row of 2 serrate setae. Palp of maxilla 1
with 3 apical spines and 1 lateral seta, outer lobe with 6 apical spines.
Outer and inner lobes of maxilla 2 with 4 apical setae. Maxilliped
missing.

Propodus of gnathopod 1 with 1 proximal grasping spine, grasping
margin of dactylus and propodus serrate. Basis of gnathopod 2
shorter than propodus, expanded with distal knob on_ posterior
margin; propodus with proximal grasping spine and notch at mid-
length with slightly proximal tooth; dactylus with proximal and
distal knobs bearing numerous setae between knobs.

Gills missing.

Pereopods 3 and 4, 2-segmented, terminal article small with 2
setae. Pereopods 5-7 missing.

Abdomen with pair of appendages and pair of setose lobes, ap-
pendages with setae at base and on tip.

Female: Body smooth except for small projection on anteroventral
margin of pereonite 2 and pleura only slightly developed. Length of
largest female 5 mm, smallest ovigerous female 4 mm.

Antennae and mouthparts similar to male, maxilliped as in P.
tenuis.

Gnathopod 1 similar to male. Basis of gnathopod 2 short but not
expanded and without proximal knob as in male; propodus similar
to male except without notch.

Gills subovate.

Pereopods 3 and 4 as in male. Propodus of pereopods 5-7 with
pair of proximal grasping spines, pereopod 5 much smaller than
pereopods 6 and 7.

84 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 41.—Paracaprella pusilla, male lectotype; a, pereopod 4; b, maxilla 1; c, maxilla 2;

d, labium; e, lateral view; f, left mandible; g, abdomen, ventral view; A, gnathopod 1;
1, abdomen, lateral view; 7, gnathopod 2.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 85

Ficure 42.—Paracaprella pusilla, female; a, lateral view; b, right mandible; c, abdomen;
d, gnathopod 2.

Abdomen with pair of lobes bearing 1 or 2 setae.

VARIATION.—The largest male of this species was approximately
6 mm and it bore a very large projection on the anteroventral margin
of pereonite 2 (fig. 32a). This projection becomes quite large in the
larger males and may be almost absent in small males; however,
even in small males it is sharp pointed and not rounded as in P. tenuis

The depth of the notch on the propodus of gnathopod 2 varies
from very slight in small males to very deep in large males.

Distripution.—Type-locality: Rio de Janeiro, Brazil.

Other records: Suez Canal; Ft. Lauderdale, Fla.; Port Aransas,
Tex.; Kingston, Jamaica; Martinique; Brazil at 28° S.; tropical West
Africa; Malembe, Congo; Durban, South Africa; Dar es Salaam,
Tanzania; Hawail; Amoy, China.

New records: Lake Worth, Virginia Key, Key Biscayne, Sarasota
Bay, St. Petersburg, and Panama City, Fla.; Grand Isle, La.; Port
Isabel, Tex.; St. Thomas, Virgin Islands; Guadeloupe; Margarita;
Curacao.

Remarxs.—The males of P. pusilla differ markedly from those of
P. tenuis by the large sharp-pointed projection on the anteroventral
margin of pereonite 2, the proximal knob on the basis of gnathopod
2, and the presence of setae on the dactylus of gnathopod 2. Setae
are occasionally present on the dactylus of large males of P. tenwis
but this is not generally the case. A projection is present on the
anterior margin of pereonite 2 in large males of P. tenuis, but its
position is usually more dorsal, and it is more rounded than in P.
pusilla.

86 U.S. NATIONAL MUSEUM BULLETIN 278

The females of these two species are difficult to distinguish; however,
the projection on the anterior margin of pereonite 2, although small
in P. pusilla, is still smaller in P. tenuis, and the basis of gnathopod
2 is slightly longer in females of P. tenuis than in females of P. pusilla.

P. pusilla has been taken from mangrove roots, sea grass, hydroids,
and ascidians.

A lectotype has been designated from the syntypical series which
is deposited in the Copenhagen Museum.

Paracaprella tenuis Mayer, 1903
Ficures 43, 44, 53

Paracaprella tenuis Mayer, 1903, p. 68, pl. 2, figs. 34-35, pl. 7, figs. 51, 58.—M.
Rathbun, 1905, pp. 7, 77.—Sumner, Osburn, and Cole, 1911 (1913),
p. 657.—McCain, 1965, pp. 192-193, figs. 1d-e, 2g—K.

Paracaprella simplex Mayer, 1903, p. 68, pl. 2, figs. 38-39, pl. 7, figs. 53-57.—
Cowles, 1930, p. 351.—Ferguson and Jones, 1949, p. 442.

Deutella abracadabra Steinberg and Dougherty, 1957, pp. 277-279, figs. 14, 17,
18, 20, 27.

Dracnosts.—Males with small triangular projection on antero-
lateral margin of pereonite 2; basis of gnathopod 2 not expanded and
without proximal knob; propodus with large nonrounded notch at
midlength.

Description.—Body smooth on dorsal surface, anterolateral margin
of pereonite 2 with small rounded projection, pleura on pereonite 3
generally well developed, pleura on pereonite 4 present but usually
weakly developed. Length of largest male 7 mm, largest female 5
mm, smallest ovigerous female 3 mm.

Antenna 1 approximately length of pereonites 1-3; flagellum of
males with 7-10 articles, females with 5-8. Antenna 2 slightly shorter
than peduncle of antenna 1, both antennae with numerous short setae.

Mandibular palp 0- to 3-segmented, terminal article of palp usually
with 1 seta. Mandibles as in P. pusilla. Maxilla 1 as in P. pusilla
except outer lobe with 5 or 6 spines. Outer lobe of maxilliped with 1
or 2 apical setae, inner margin with numerous setae; inner lobe
apically rounded with 4-6 apical setae; penultimate article of palp
with process, terminal article with several setae at tip.

Propodus of gnathopod 1 with 1 proximal grasping spine, grasping
margin of dactylus and propodus serrate. Propodus of gnathopod 2
similar to P. pusilla but notch at midlength more triangular and
deeper in large males than in P. pusilla, dactylus usually tapered
but occasionally similar to P. pusilla.

Gills subovate.

Pereopods 3 and 4, 2-segmented, terminal article small with 2 or
3 setae, basal article with 1 or 2 setae. Propodus of pereopods 5-7

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 87

Ficure 43.—Paracaprella tenuis, male; a, pereopod 3; b, pereopod 5; ¢, gnathopod 2; d,
lateral view; ¢, maxilliped; f, maxilla 1; g, pereopod 7 (drawn to same scale as pereopod Ds
h, gnathopod 1; 7, left mandible; 7, right mandible; k, abdomen.

88 U.S. NATIONAL MUSEUM BULLETIN 278

a

Ficure 44.—Paracaprella tenuis, female; a, lateral view; 6, gnathopod 2; ¢, pereopod 4;
d, abdomen.

with pair of proximal grasping spines, pereopods 6 and 7 with several
proximal knobs on palm, each bearing small spine, pereopod 5 smaller
than 6 and 7 (fig. 43b and g, drawn at same scale).

Abdomen of male and female similar to P. pusilla.

VARIATION.—The development of the projection on the antero-
lateral margin of pereonite 2 and pleura on pereonites 3 and 4 is
related to size as in P. pusilla, larger males having larger projections
and well-developed pleura.

The mandibular palp varies from a small hump bearing a terminal
seta to a 3-segmented palp with a seta on the terminal article. In
no specimens were the palps reduced to a single seta as in P. pusilla.

Distripution.—Type-locality: Woods Hole, Massachusetts.

Other records: Gulf of St. Lawrence; Penneville, Maine; Great
Egg Harbor, N.J.; Chesapeake Bay, Va.; Sapelo Island, Ga.; Alligator
Harbor, Fla.; Ship Island, Miss.

New records: Numerous localities between the Gulf of St. Lawrence
and Sapelo Island, Ga.; Cumberland Sound, Fernandina, Tampa Bay,
and Pensacola Bay, Fla.; Galveston Bay, Corpus Christi Bay, and
Port Isabel, Tex.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 89

Remarks.—The characters which separate this species from P.
pusilla are presented under the remarks of the latter species.

P. tenuis has been collected from various red and brown algae,
sea grass, sponges, hydroids, aleyonarians, bryozoans, and from
hydroids attached to the carapace of the spider crab Libinia. This
species has the peculiar habit of curling when preserved, which makes
it relatively easy to sort from the other caprellid species.

Pariambus typicus f. cumana (Mayer, 1903)

Podalirius typicus f. cwmana Mayer, 1903, p. 65, pl. 10, figs. 6-7.

Podalirius typicus f. camana.—Monterosso, 1915, p. 7.

Pariambus typicus f. Cumana.—Chevreux and Fage, 1925, p. 442.
Remarks.—Mayer described this form of Paériambus typicus from

“Strand von Cumae” which Chevreux and Fage took to be Cuma,

Venezuela. In Mayer’s faunal list at the end of his “Siboga’’ report

(1903), he does not mention P. typicus in the material from the

West Indies and South America so it is probable that he was referring

to Cumae, Italy, instead of Cuma, Venezuela.

Phtisica Slabber, 1769

Flagellum of antenna 2, 2- to 5-segmented, swimming setae absent;
mandibular palp 8-segmented, setal formula for terminal article
1-x-1 or 1, molar absent; outer lobe of maxilliped equal to inner
lobe; gills on pereonites 2-4; pereopods 8 and 4, 6-segmented, pereopod
5, 5-segmented; abdomen of male with 8 pairs of appendages, female
with 2 pairs of appendages, pair of lobes and raised anterior projection.

Type-species: Phtisica marina Slabber, 1769 (by monotypy).

Remarxs.—The generic name was probably derived from the
Greek word ‘‘phthisikos’” meaning consumptive; however, Slabber
does not give the derivation and therefore the original orthography
should be retained.

Phtisica antillensis (Mayer, 1903)
Ficures 45, 54

Proto antillensis Mayer, 1903, p. 21, pl. 1, fig. 4, pl. 6, fig. 21.

Draenosis.—Carpus of gnathopod 2 longer than merus, propodus
broad medially with grasping spines at midlength.

Description.—Male holotype: Body smooth. Length 1.5 mm.

Antenna 1 approximately one-half body length, flagellum of 38
articles. Antenna 2, three-fourths length of antenna 1, flagellum of 2
articles.

Terminal article of mandibular palp with 2 setae. Mandibles
similar to P. marina with incisor, lacinia mobilis, and 2 additional
plates; setal row less developed than in P. marina. Outer lobe of

90 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 45.—Phitisica antillensis, male holotype; a, lateral view; b, maxilliped; c, pereopod 4;
d, gnathopod 1; ¢, abdomen; f, terminal article of mandibular palp; g, gnathopod 2.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 91

maxilliped with 2 apical setae, 2 setae distal yet slightly proximal to
apical setae, and 2 proximal setae; inner lobe almost as large as outer
with 1 large spine and several indistinct spines, anterodistal corner of
medial margin with 3 large spines.

Propodus of gnathopod 1 triangular, approximately as wide at base
as long, with 3 proximal grasping spines. Propodus of gnathopod 2
broad at midlength with 3 grasping spines at midlength.

Gills on pereonites 2-4, gill on pereonite 2 smaller than those on
3 and 4.

Pereopods 3 and 4, 6-segmented, propodus smooth except for pair
of small spines in distal notch, dactylus short and wide at base.

Abdomen with 3 pairs of appendages, posterior 2 pairs biarticulate,
anterior pair uniarticulate with single seta at tip.

Distrisution.—Type-locality: St. Thomas, Virgin Islands, at a
depth of 18-27 m.

Remarks.—The above description is based on the young male
holotype and only specimen of this species which is deposited in the
Copenhagen Museum. The mouthparts and abdomen agree with those
of P. marina, and there can be little doubt that this species belongs
to the genus Phtisica. For differences between this species and P.
marina see the discussion of the latter species.

Phtisica marina Slabber, 1769
Fiaures 46, 47, 54, 56

Squilla acaudata pedibus quatuordecim . . .—Gronovius, 1760, p. 39, pl. 5,
figs. 8-10.

Phtisica marina Slabber, 1769, pp. 79-83, pl. 10, figs. 1-3.—Stebbing, 1888, p. 32.—
Chevreux and Bouvier, 1893, p. 142.—G. Sars, 1895, pp. 646-648, pl. 223.—
Walker, 1895a, p. 319; 1898, p. 170—Walker and Hornell, 1896, p. 54.—
Scott, 1897, p. 141; 1901, p. 267; 1906, p.174—Gadeau de Kerville, 1898,
p. 348; 1900 (1901), p. 184.—Chevreux, 1898, p. 483; 1900, pp. 118-119; 1901
(1902), p. 696.—Beaumont, 1900, p. 795.—Norman, 1902, p. 483; 1907, p.
370; 1908 (1909) p. 463.— Marine Biol. Assoc., 1904, p. 241; 1931, p. 198.—
Norman and Scott, 1906, pp. ix, 98—Norman and Brady, 1910, p. 75.—
Stebbing, 190la, p. 468——Nordgaard, 1911 (1912), p. 24.—Chichkoff,
1912, p. xxxv.—Barnard, 1916, pp. 283-284.—Chumley, 1918, pp. 34, 165.—
Schneider, 1924 (1926), pp. 58-59.—Schellenberg, 1926, p. 465; 1927, pp.
721-722, fig. 104.—Stephensen, 1927a, pp. 145-146; 1927b, p. 13; 1928, pp.
378-380, fig. 91 (1-10); 1929a, pp. 176-177, fig. 328; 1929b, pp. 19, 34; 1935,
p. 118; 1942, pp. 428-429, 502, 503; 1944b, p. 159.—Borcea, 1931, p. 677.—
Schijfsma, 1931, p. 26.—Fage, 1933, pp. 113, 115, 117, 119, 125, 126, 127,
225.—Oldevig, 1933, p. 261, fig. 1—Pirlot, 1939, p. 78—Fiorencis, 1940,
pp. 11-12, fig. 1, pl. 1, figs. 1-2—Goodhart and Harrison, 1940, p. 109.—
Bertrand, 1941, p. 16—A. Carausu, 1941, p. 11—Carausu and Carausu,
1942, pp. 79-80.—Dahl, 1946, p. 22.—Ruffo, 1946, p. 53.—Enequist, 1949,
pp. 392, 402.—Rancurel, 1949, p. 168.—Spooner, 1950, p. 253.—Reid, 1951,
pp. 281-282, 289.—Ruffo and Wieser, 1952, p. 21.—S. Carausu, 1956, p.
132.—Costa, 1960a, pp. 99, 100; 1960b, p. 104; 1961, p. 271.—Gottlieb, 1960,

279—-475—68———_7

92 U.S. NATIONAL MUSEUM BULLETIN 278

pp. 161, 163, 164—Toulmond and Truchot, 1964, p. 34.—McCain, 1966,
p. 92.

Squilla ventricosa Miller, 1776, p. 197.—Abildgaard, 1788, pp. 20-21, pl. 56,
figs. 1-3.

Cancer (Gammarellus) ventricosus.—Herbst, 17938, pp. 144-145, pl. 36, fig. lla, b.

Caprella ventricosa.—Bosc, 1801-02, p. 156.—Latreille, 1802-03, p. 327; 1803,
p. 333.

Proto Pedata Leach, 1814, p. 433.

Cancer Gammarus pedatus—Montagu, 1915, pp. 6-7, pl. 2, fig. 6.

Leptomera rubra Lamarck, 1818, p. 172.

Proto .—Latreille, 1818, p. 9, pl. 336, fig. 38.

Leptomera pedata.—Lamarck, 1818, pp. 172-173.—Guérin, 1829-44, p. 25, pl.
28, fig. 3. —H. Edwards, 1840, p. 109.—Krgyer, 1842-43, pp. 607-613, pl.
7, figs. 13-23—Rathke, 1843, pp. 97-98.—Gosse, 1855, p. 131, fig. 224.—
Liljeborg, 1856, p. 182.— Maitland, 1874, p. 245.

Proto pedatus.—Fleming, 1823, pp. 296-297.—Johnston, 1835, pp. 672-674,
fig. 73.—Couch, 1864, p. 99.

Proton pedatum.—Desmarest, 1823, p. 363, pl. 46, fig. 3; 1825, pp. 276-277,
pl. 46, fig. 3.—Bouchard-Chantereaur, 1823, p. 18.

Cancer (Leptomera) rubra.—Latreille, 1836, p. 217.

Cancer (Leptomera) pedata.—Latreille, 1836, p. 217.

Leptomera ventricosa.—H. Edwards, 1840, p. 110.

Proto pedatum.—W. Thompson, 1944, p. 4385.—White, 1847, p. 92.—Cocks,
1849, p. 83.

Proto pedata.—White, 1850, p. 61; 1857, p. 218.—Bate, 1856, p. 60; 1857, p. 151;
1862, p. 382; 1862, pp. 349-350, pl. 55, fig. 1; 1878, p. 508; 1887, p. 175.—
Boeck, 1861, p. 670.—McAndrew, 1861, p. 28.—Bate and Westwood, 1868,
pp. 38-41.—Brady and Robertson, 1869, p. 361.—Parfitt, 1873, p. 250.—
Haller, 1879a, p. 230; 1879b, p. 398.—Hoek, 1879, various pp., pl. 5, fig. 10,
pl. 7, figs. 15-16, pl. 8, figs. 1-3—Delage, 1881, pp. 132-133, 153, pl. 10,
fig. 7.—Fowler, 1886, p. 217.—Schneider, 1891, pp. 111, 122.—Walker,
1895b, p. 474.— Ussing, 1952, pp. 45-47.

Proto elongatus Dana, 1853, pp. 45-47.

Proto Goodsirii Bate, 1856, p. 60; 1857, p. 151; 1862, p. 350, pl. 55, fig. 2; 1887,
p. 175.—White, 1857, p. 218—McAndrew, 1861, p. 28.—Bate and West-
wood, 1868, pp. 42-43.—Boeck, 187la, p. 268 (188); 1873-76, pp. 671-
672, pl. 32, fig. 2—Delage, 1881, pp. 182-133, 153, pl. 10, fig. 7—Bonnier,
1887, p. 349.—Robertson, 1886-87 (1888), p. 70.—Walker, 1895b, p. 475.

Naupredia tristis van Beneden, 1861, pp. 97-99, 146, pl. 17.—Maitland, 1874,
p. 246.—Pelseneer, 1886, p. 218.

Proto elongata.—Bate, 1862, p. 350, pl. 55, fig. 3.

Proto ventricosa.—Boeck, 187la, pp. 268 (188)—269 (189); 1873-76, pp. 672-
674, pl. 32, fig. 3.—Metzger, 1875, p. 278.—Meinert, 1880, pp. 494-495.—
Mayer, 1882, pp. 22-25, pl. 1, fig. 1, pl. 3, figs. 16-29, pl. 4, figs. 12-13,
pl. 5, figs. 1-5; 1890, pp. 12-13, pl. 3, figs. 4-5, pl. 5, figs. 3-6, pl. 6, fig. 1,
pl. 7, fig. 1; 1903, pp. 20-21, pl. 6, fig. 23.—Blane, 1884, pp. 85-87, pl. 5, figs.
115-121.—Collin, 1884, p. 21.—Carus, 1885, p. 387.—Henderson, 1885,
p. 311.—Guerne, 1886, p. XLIV.—Gadeau de Kerville, 1886 (1887), p.
82.—Bonnier, 1887, pp. 349-350.—Chevreux, 1887a, pp. 317, 335; 1888,
p. 34.—Barrois, 1888, pp. 55-56, 76.—Robertson, 1886-87 (1888), pp. 69-
70.—Scott, 1887 (1888), p. 250.—Hoek, 1889, pp. 232-233.—Meinert,
1890, p. 183.—Gourret, 1892, pp. 16, 25.—Lameere, 1895, p. 570.—Sovinskii,

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 93

1895, pp. 255-256; 1898, pp. 502, 511, 512, 513, 514.—Sokolwsky, 1900,
p. 161, pl. 3, fig. 15.—d’A. Thompson, 1901, p. 41.—Graeffe, 1902, p. 19.—
Loénnberg, 1902 (1903), p. 50.—Tattersall, 1913, pp. 20, 22.—Monterosso,
1915, pp. 3-6.—Oliveira, 1940, p. 140.—S. Carausu, 1956, p. 133.

Proto Goodsiri.—Stebbing, 1876, p. 78.—Haller, 1879a, p. 231; 1879b, pp. 398-
399, pl. 22, figs. 23-25.—Hansson, 1882 (1883), p. 77.—Chevreux, 1887b,
p. 579; 1888, p. 34.

Proto Ventricosa.—Meinert, 1877-78, pp. 166-168.

Leptomera pedata (Gammarus).—M. Sars, 1858 (1859), p. 150.

Proto goodsiri.—Stebbing, 1879, p. 521.—Chevreux, 1887a, pp. 318, 335.

Proto brunneovittata Haller, 1879a, p. 231; 1879b, pp. 399-400, pl. 22, figs. 19-
22.— Mayer, 1882, p. 25; 1890, p. 14.—Carus, 1885, p. 387.

Gammarus pedatus.—Abildgaard, 1789, pp. 33-34, pl. 101, figs. 1-2.

Phthisica marina.—Norman, 1886, p. 26; 1905, p. 26.—Allen and Todd, 1900,
pp. 166, 174, 203.—Bjorcx, 1915, p. 35.—Chevreux and Fage, 1925, pp.
434-436, fig. 422.—Schellenberg, 1942, pp. 234-235, fig. 192—Marine Biol.
Assoc., 1957, pp. 232-233.

Proto spec. Mayer, 1903, p. 21.

Phthisica acaudata.—Reibisch, 1906, pp. 214-216, 219, 220, 221, 222, 229, 230,
232, 233.—Zernov, 1913. pp. 68, 127, 233.

Phtisica acaudata.—Tichy, 1911, pp. 1125-1127, 1133, 1134.—Borcea, 1931,
p. 702.—Carausu, 1956, p. 132.

Phytisca marina.—Crawshay, 1912, p. 351.

Phtysica acaudata.—Borcea, 1934, p. 404.

Phiisica antillensis [not Mayer].—Pequegnat, 1966, p. 25.

DraGcnosis.—Carpus of gnathopod 2 shorter than merus, propodus
with greatest width proximal and with proximal grasping spines.

Description.—Body smooth. Length of largest male 14 mm,
largest female 13 mm, smallest ovigerous female 3.7 mm.

Antenna 1 variable in length from one-half to length of body.
Antenna 2 approximately equal in length to peduncle of antenna 1,
flagellum of 3-5 articles.

Mandibular palp 3-segmented, setal formula for terminal article
varies from 1-1-1 to 1-6-1. Left mandible with 5-toothed incisor,
5-toothed lacinia mobilis, 2 additional plates, setal row of 7 or 8 short
blunt setae and 3 longer apically serrate setae. Right mandible with
5-toothed incisor, lacinia mobilis smooth, 2 additional plates, setal
row of 6 or 7 short setae and 2 longer apically serrate setae. Outer
lobe of labium winglike, both lobes quite setose. Palp of maxilla 1
with 3 or 4 apical spines of which 2 sometimes serrate and with 3
or 4 proximal setae, outer lobe with 6 apical spines which become
bifid or serrate in larger individuals. Outer lobe of maxilliped with
1 apical seta, 1 short apical spine, and 2-5 proximal setae along
medial margin; inner lobe as long as outer lobe and nearly fused
along medial margins, mediodistal corner with 1 very large serrate
spine, 1 small spine, and 1 slightly lateral marginally serrate spine,
lateral margin of lobe sometimes toothed; article 2 of palp robust.

94 U.S. NATIONAL MUSEUM BULLETIN 278

Propodus of gnathopod 1 triangular with 4 or 5 grasping spines,
grasping margin without serrations but with numerous short spines;
dactylus not serrate. Propodus of gnathopod 2 with 1 large and 1
small proximal grasping spine, palm with numerous short spines and
few setae.

Gills elliptical, gill on pereonite 2 smallest.

Pereopods 3 and 4, 6-segmented, palm of propodus with 3 short
spines. Pereopod 5, 5-segmented, palm of propodus without spines,
dactylus approximately one-half propodus length, basis approximately
twice propodus length. Pereopods 6 and 7, 6-segmented with 2 proximal
grasping spines, 1 medial spine, and 1 distal spine.

Abdomen of male with 2 pairs of biarticulate appendages and pair
of pyriform appendages, each pyriform appendage terminating in
seta; female with 2 pairs of biarticulate appendages, pair of lobes,
and anterior raised projection.

VariaTion.—Included in description.

DistripuTion.—Type-locality: No locality given.

Other records: Northern Norway to the Mediterranean Sea;
British Islands; Azores; Canary Islands; Mediterranean and Black
Seas; tropical West Africa; South Africa; 25°26’30’’ N., 80°02’ W.;
Rio de Janeiro, Brazil.

New records: Numerous localities from off Cape Hatteras, N.C. to
the Tortugas and the west coast of Florida to Panama City; St.
John, Virgin Islands; Cubagua and Margarita Islands, Venezuela;
Cape la Vela, Colombia.

ReEmARKS.—Gronovius (1760) described a caprellid which he called
“Squilla acaudata pedibus quatuordecim . . .’”’ and included several
figures which appear to be P. marina, Reibisch (1906) believed that
Gronovius’ paper followed the rules of binomial nomenclature so he
chose to use the name P. acaudata. It appears that the term acaudata
was used in a descriptive rather than a nominal sense, so I have chosen
to use Slabber’s name which meets the requirements for validity.

The genus Phtisica is presently composed of 3 species, P. antillensis
(Mayer, 1903), P. marina Slabber, 1769, and P. tuberculata (Guiler,
1954). In his description of P. tuberculata, then Proto tuberculata,
Guiler states that his species has gills on pereonites 3 and 4 and that
the abdomen bears 2 pairs of rudimentary appendages. These char-
acters are not consistent with the generic characteristics of Phtisica,
and it would seem that P. twberculata should be placed in a different
genus, perhaps Paraproto; however, material of this Tasmanian
species is not available to me and such a change is not possible at
this time.

Mayer (1903) gives a description of a Proto spec., stating that
it differs from P. marina because the first pair of abdominal append-

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 95

d

Ficure 46.—Phtisica marina, male; a, lateral view; b, abdomen; c, gnathopod 2;
d, gnathopod 1.

ages are strikingly small. I have examined the material upon which
his description was based but I cannot agree that these appendages
are any smaller than in P. marina, so this species is included in the
synonymy of P. marina.

Phtisica marina differs from P. antillensis in that the carpus of
gnathopod 2 is longer than the merus and the propodus of gnathopod
2 is expanded at midlength in the latter species.

96 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 47.—Phtisica marina, female; a, lateral view; b, gnathopod 2; c, gnathopod 1; d,
pereopod 3; e, maxilla 2; f, maxilla 1; g, terminal articles of mandibular palp; 4, maxil-
liped; i, right mandible; 7, left mandible; k, abdomen.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 97

P. marina is widely distributed in the warmer parts of the Atlantic.
It has been collected in plankton tows (fig. 56) at the surface and
Stephensen (1927a) reports this species from a depth of 660 m. The
habitat does not seem specific because it has been reported on green
apd brown algae, sea grass, sponges, hydroids, and bryozoans. Boeck
(1873-76, p. 671) reports this species on the sea turtle Chelonia;
however, he cites van Beneden, 1861, as the authority. Van Beneden’s
record was for Caprella andreae and not Phtisica. It has also been
found among the hydroids and algae which were attached to the
carapace of the spider crab Maja. Meinert (1880) cites this species
as occurring on the starfish Asterzas. There are several records of this
species from the guts of fishes such as the blenny, Blennius, cod
Gadus dragon net, Callionymus, and the skate, Raja.

Costa (1960b) reports that P. marina is predatory on Caprella
acanthifera. Costa states the P. marina grasps the prey with gnathopod
2 and the prey seems almost incapable of violent movement. It seems,
therefore, that some venom may be introduced by the poison tooth.

Proaeginina Stephensen, 1940

Flagellum of antenna 2 biarticulate, swimming setae absent;
mandibular palp 3-segmented, setal formula for terminal article
1-x-1, molar present; outer lobe of maxilliped larger than inner
lobe; gills on pereonites 3 and 4; pereopods 3 and 4 absent, pereopod
5, 6-segmented ; abdomen of male and female with 2 pairs of appendages.

Type-species: Parvipalpus norvegicus Stephensen, 1931 (by
monotypy).

Remarxs.—Stephensen (1931, 1940, 1942) stated that the flagel-
lum of antenna 2 is uniarticulate; however, both of the Iceland speci-
mens and the Cape Cod specimen which were available to me bear a
small article at the tip of the flagellum which is clearly defined by
a suture. In Stephensen’s (1944a, p. 46) list of the important characters
of new genera described since Mayer’s (1903) last monograph, he
stated that the number of setae on the terminal article of the mandib-
ular palp of Proaeginina is greater than 2. Although true, I have altered
the generic description to include the 1-x-1 setal formula and have
also included the biarticulate flagellum on antenna 2.

Proaeginina norvegica (Stephensen, 1931)
Fiaures 48, 54
Parvipalpus norvegicus Stephensen, 1931, pp. 1-7, figs. 1-3.
Proaeginina norvegica.—Stephensen, 1940, pp. 70-72, fig. 8; 1942, pp. 433, 502,
503, figs. 68-69; 1944a, pp. 49-50; 1944b, p. 159.—McCain, 1966, p. 92,
Diaenosis.—Since this genus is monotypic, the characters of the
genus are diagnostic for the species.

98 U.S. NATIONAL MUSEUM BULLETIN 278

Description.—Body smooth and quite slender, cephalon separated
from pereonite 1 by faint suture; however, not articulated. Length
of largest male 32 mm, largest female 23 mm.

Articles in flagellum of antenna 1 variable in number, increasing
with increasing size of individual. Flagellum of antenna 2 biarticulate,
terminal article very small.

Mandible with 3-segmented palp, setal formula for terminal article
varying from 1-5-1 to 1-27-1. Left mandible with 5-toothed incisor,
5-toothed lacinia mobilis, setal row of 3 serrate setae. Right mandible
with 5-toothed incisor, lacinia mobilis apically serrate, setal row of
2 serrate setae. Palp of maxilla 1 with 4-7 short apical spines and 3
or 4 setae on anterior surface; outer lobe with 7 apical spines, tending
to bifurcate with increased size of individual. Inner and outer lobes
of maxilla 2 variably setose. Outer lobe of maxilliped with 2 apical
setae, 1 long apical spine, and 3-7 small marginal spines; inner lobe
with 2 small spines, 6-12 apical setae of which 0-10 plumose; dactylus
of palp serrate at tip.

Propodus of gnathopod 1 triangular with 2 proximal grasping spines,
grasping margins of dactylus and propodus serrate. Propodus of
gnathopod 2 with proximal poison tooth, grasping margin slightly
serrate proximally; dactylus not serrate.

Pereopods 5-7, 6-segmented, propodus of male pereopod 5 with
pair of distal grasping spines, propodus of pereopods 6-7 of male and
female without grasping spines.

Abdomen of both male and female with 2 pairs of biarticulate
appendages.

VaRIATION.—A comparison of a male and female (32 and 23 mm
respectively) from Thor sta. 57 near Iceland with the small female
(7.5 mm) from off Cape Cod showed that no variation exists which
cannot be accounted for by increased size of the animal. The setal
formula for the terminal article of the mandibular palp in the Cape
Cod female was 1-5-1 for the left mandible and 1-6-1 for the right
mandible. In the Zhor female the setal formula for both mandibles
was 1-27-1 and for the male it was 1-21-1. Six nonplumose setae
are present on the inner lobe of the maxilliped of the Cape Cod
female, 8 or 9 on the Thor male, of which 3 or 4 are plumose, and 12
on the Thor female with 10 plumose.

Disrripution.—Type-locality: Rognsund, off NE. corner of the
Island of Stjerngy (N. Norway, about 70°40’ N.), at a depth of
200-350 m.

Other records: Iceland, 63°21’ N., 16°22’ W., 500-560 m; Davis
Straits, 63°06’ N.,"56°00’ W., 2258 m, and 61°50’ N., 56°21’ W.,
2702 m.

New record: Off Cape Cod, 42°00’ N., 68°34’ W., 174 m.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 99

Ficure 48.—Proaeginina norvegica; a, male lateral view; b, pereopod 5; ¢c, female abdomen,
lateral view; d, female lateral view; ¢, pereopods 6 and 7; f, gnathopod 2; g, terminal
articles of antenna 2; h, terminal articles of antenna 2 (Thor female); 7, gnathopod 1;
j, maxilliped; k, abdomen, lateral view; /, maxilla 1; m, left mandible; m, right mandible
(a-e after Stephensen, 1942; f-g and i-n of small female).

100 U.S. NATIONAL MUSEUM BULLETIN 278

Pseudaeginella Mayer, 1890

Flagellum of antenna 2 biarticulate, swimming setae absent; man-
dible with 3-segmented palp, setal formula for terminal article 1-3-1,
molar?; outer lobe of maxilliped larger than inner lobe; gills on
pereonites 3 and 4; pereopods 3 and 4 absent, pereopod 5, 6-segmented;
abdomen of male and female without appendages.

Type-species: Aeginella tristanensis Stebbing, 1888 (by monotypy).

Pseudaeginella antiguae Barnard, 1932
Fieure 51

Pseudaeginella tristanensis [not Stebbing].—Stebbing, 1895, p. 402.—Mayer,
1903, p. 59.
Pseudaeginella antiguae Barnard, 1932, p. 301.

Remarxks.—This species was described by Mayer (1908) and Steb-
bing (1895) as P. tristanensis from specimens collected in Antigua,
West Indies. Barnard later changed the name of these specimens to
P. antiguae with the following comment:

In view of these specimens it becomes very unlikely that the sepcimens (Stebbing
says one; Mayer says two) from Antigua are conspecific. The female according
to Mayer (1903, p. 59) has the head spine and single medio-dorsal spines on
segments 1-3, but also paired rounded tubercles in the middle of segments 3
and 4. For this species I therefore propose the name antiguae.

P. tristanensis does not have paired rounded tubercles at the mid-
length of pereonites 3 and 4 and therefore Barnard is probably correct
in his separation of these two species.

Barnard’s figures of P. tristanensis bear a remarkable similarity
to Fallotritella biscaynensis; however, none of the references mentioned
appendages in Pseudaeginella on pereonites 3 and 4 and I must
presume their absence. It may be found that appendages are present
on pereonites 3 and 4 in the two species of Pseudaeginella; if so, then
Fallotritella would probably fall as a junior synonym of this genus.
Material of Pseudaeginella was not available to me, and the location
of the type material is not known to me.

Ecology

Most caprellids seem to be very nonspecific as to the substrate
upon which they live; however, most need something to which they
can cling and therefore are not found on bare sandy or muddy bot-
toms. Species such as Caprella equilibra and Caprella penantis have
been collected on various algae, sea grasses, sponges, hydroids, sty-
lasterines, alcyonarians, zoantharians, bryozoans, ascidians, and in
several unusual habitats. Some caprellids, however, do have a fairly
specific habitat preference. Caprella unica is associated with the
starfishes Asterias forbest and A. vulgaris and indeed shows an adapta-

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 101

tion for this habitat by the loss of spination and setation of pereopods
5-7, which would definitely be a hinderance when moving over the
highly pedicellate surface of the starfish. For other species associated
with echinoderms, see p. 52.

Caprella andreae is found attached to floating objects and plants
and has even been collected from the fouling on the backs of sea
turtles. This species seems to be limited to this type of environment
and to my knowledge has not been collected attached to benthic
substrates.

Phtisica marina was found in many of the U.S. Bureau of Commercial
Fisheries vessel M/V Theodore N. Gill plankton samples and appears
to be a regular member of the Carolinian coastal water plankton
from Cape Hatteras to southern Flordia (fig. 56) P. marina lacks a
molar on the mandible which may reflect an adaptation to feeding in
its planktonic environment. Several other species have occasionally
been taken in plankton tows including Hemiaegina minuta and
Luconacia incerta; however, these species have a molar on the man-
dible.

The feeding habits of several west coast species of Caprella and
Metacaprella were reviewed by Saunders (1966). She found the bulk
of their diet consisted of diatoms, but they also consumed some
crustaceans and perhaps also fed on detritus. Since the mouthparts of
most of the species of Caprella are very similar, they must have
similar feeding habits. As noted on p. 30, I have observed Caprella
equilibra feeding on several gammaridean amphipods and polychaetes.
Harrison (1939) reviewed the habits of several caprellids. He stated
that the caprellids close the antennae down over the prey and then
gnathopod 2 grasps the prey, soon paralyzing it with the poison
spines. The prey is then shredded by the mouthparts, after which the
appendages are cleaned for bits of debris.

Caprellids are not infrequent members of the food chains of many
bottom feeding fishes and have been found in the guts of the sea bass,
Centropristis, the blenny, Blennius, the pollock, Pollachius, the rock
cod, Gadus, the dragon net, Callionymus, and the skate, Raja. Aquar-
ium observations on several species showed that they were prey for
grass shrimps, anemones, and even the small gastropod, Astyris.
Caprellids will probably be found to be prey for many other bottom
feeding fishes and larger Crustacea.

Zoogeography

The western Atlantic is rather poor in number of caprellid species
compared to similar areas in the Pacific. Utinomi (1947), Mayer
(1930), and Arimoto (1934) cite over 60 species which occur in Japa-

102 U.S. NATIONAL MUSEUM BULLETIN 278

nese waters (approximately 20° of latitude) whereas only 26 species are
found in the western North Atlantic (approximately 50° of latitude).
This paucity of species in the North Atlantic was observed by Mills
(1965) for the gammaridean genus Ampelisca. He stated that inter-
tidal species may be few because of winter ice scouring and that
perhaps sharp temperature zonation and a wide range of temperatures
do not favor the differentiation of stenothermal species. Many of the
caprellid species in the temperate zone of the western North Atlantic
are almost cosmopolitan in their distribution and appear to be highly
elastic eurythermal and euryhaline species. For example, Caprella
penantis is found in most parts of the Atlantic and Pacific from the
boreal to the tropical zone and in salinities ranging from full sea water
to as low as 10 °/oo.

Table 3 summarizes the known distribution of the caprellids of the
western North Atlantic (figs. 50-55). In general they correspond to

samel

e Hemiaegina minuta
a Hemiproto wigleyr

* Mayerella limicola

Sees 2
© Mayerella redunca;

a 40

® Previous record
O New record

pe panslaes

90

Ficure 50.—Distribution records of Hemiaegina minuta, Hemiproto wigleyi, Mayerella
limicola, Mayerella redunca, and Metaprotella hummelincki in the western North Atlantic.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 103

the faunal provinces of Johnson (1934) as modified by Hedgpeth (1953)
with the exception that a truly boreal fauna is not evident. Cerame-
Vivas and Gray (1966) reviewed these zoogeographic provinces and
divided the continental shelf fauna of North Carolina into three
assemblages, extending the Caribbean province to just north of Cape
Hatteras for the outer shelf. These provinces and examples of their
caprellid composition are summarized as follows:

1. Arctic province (north of Newfoundland)—Aeginella spinosa
and Proaeginina norvegica.

2. Boreal province (Newfoundland to Cape Cod). This province
has no endemic caprellids with the possible exception of Mayerella
limicola and receives contributions from the Arctic and Transatlantic
provinces.

Caprella linearis

@ Caprella penantis

eo

x Caprella septentrionalis

© Caprella bermudia

* ee

# Pseudaeginella ant. i

© Previous record
O New record

Figure 51.—Distribution records of Caprella bermudia, Caprella linearis, Caprella penantis,
Caprella septentrionalis, and Pseudaeginella antiguae in the western North Atlantic.

104 U.S. NATIONAL MUSEUM BULLETIN 278

ea Lh
wt! z

~

ae Deutella californica

x Deutella mayeri

© Luconacia incerta

® Previous record
O New record

90

Ficure 52.—Distribution records of Deutella californica, Deutella mayeri, and Luconacia
incerta in the western North Atlantic.

3. Transatlantic province: (A) Virginian province (Cape Cod to
Cape Hatteras). No caprellid is restricted to this province, yet Cape
Hatteras does appear to be a southern barrier for the cold-water
species Aeginina longicornis and a northern barrier for the warm-
water species Phtisica marina. (B) Carolinian province (Cape Hatteras
to Cape Kennedy and Tampa Bay to the southern tip of Texas).
As in the case of the Virginian province, no caprellid species is re-
stricted to this area. If considered together the Virginian and Caro-
linian provinces can be characterized by species such as Paracaprella
tenuis and Caprella penantis. The transatlantic province seems to
be an area of mixing of the Caprellidae with only 1 endemic form.

4. Caribbean province (from Cape Kennedy and Tampa Bay into
the tropics, outer shelf to just north of Cape Hatteras, and Bermuda).
Many endemic species such as Fallotritella biscaynensis and Hemiproto
wigleyt.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 105

* Fullotritella biseaynensis ~~ <4
e Paracaprella pusilla © we AO

woe 4
= § Paracaprella tenuis ®G

®@ Previous record
O New record

Ficure 53.—Distribution records of Fallotritella biscaynensis, Paracaprella pusilla, and
Paracaprella tenuis in the western North Atlantic.

The distribution of Paracaprella tenuis is interesting since it shows
the disjunct distribution pattern which Hedgpeth (1953) cites for
Inttorina trrorata and several other organisms. Hedgpeth points out
that this disjunct distribution arose during the interglacial periods
of the Pleistocene when Florida was submerged and counter currents
probably arose across the shoals which ‘made possible the dispersal
of northern species into the Gulf of Mexico... ’’.

Hedgpeth (1953) states that the conditions are such in the northern
Gulf of Mexico to allow mixing of temperate and tropical faunas
with a low number of endemic species. This general statement is
true for the Caprellidae since no northern Gulf caprellid is endemic
and that area receives contributions from both the temperate and
tropical areas.

106 U.S. NATIONAL MUSEUM BULLETIN 278

© Phtisica antillensis

® Phtisica marina
e Aeginella spinosa
a § Acginina longicornis

* Proaeginina nor. _/

@ Previous record
O New record

50

Ficure 54.—Distribution records of Aeginella spinosa, Aeginina longicornis, Phtisica antil-
lensis, Phtisica marina, and Proaeginina norvegica in the western North Atlantic.

Mayer (1903, p. 133) expresses the opinion that Caprellidae are
quite rare in the shallow water of the West Indies. Contrary to this
opinion, 14 species of caprellids are reported from the Caribbean in
this paper. Most of the caprellids of the area are fewer in number of
individuals and smaller than those of the northern provinces. This
might explain why so few species have previously been reported
from the Caribbean.

The caprellids are remarkably widespread in their distribution.
Many species are almost cosmopolitan, a fact which seems anomalous
in view of their limited swimming ability and the absence of a plank-
tonic larval stage. Mayer (1882, p. 90; 1890, p. 101; 1903, p. 134)
attributes this wide distribution to transportation on ship bottoms
and floating objects. Several species are associated with floating
objects and plants and some, as Caprella andreae, appear to be limited

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 107

to this type of pelagic environment. Some species, particularly
Phtisica marina, are at least transient members of the plankton.
In view of these semipelagic habitats, it is not too remarkable that
these organisms are spread over very large areas.

TABLE 3.—Known distribution of Caprellidae in the western North Atlantic

3/38 n| Qh £
8i/% | Slesie |ee,
g/sz)/22|se|e |e8al -
3/ sO /)/SmH|/ahim |Seel a a
B|22|/C2|"2) a2 Mig 2 | =
21'S |85| 25 | 86 B26) = | g
Z a oO oO & [0 oD oa)
Aeginella spinosa x x
Aeginina longicornis x <i x
Caprella andreae x x x
C. bermudia x
C. danilevskit x 2 x
C. equilibra : aa Ne > ae He ca
C. linearis = x x
C. penantis x x x x x x
C. scaura x
C. septentrionalis x *
C. unica x x x
Deutella mayert x
D. californica x
Fallotritella biscaynensis x x x
Hemiaegina minuta x x ne x x x
Hemiproto wigleyt De os
Luconacia incerta x BN x x x x
Mayerella limicola rx xe
M. redunca x
Metaprotella hummelincki x
Paracaprella pusilla x x Xx
P. tenuis x x x x
Phtisica antillensis Xx
P. marina x x x x
Proaeginina norvegica x x
Pseudaeginella antiguae x

Relationship Between the Amphipod Suborders
Gammaridea and Caprellidea

The Caprellidea appear to have arisen from a podocerid type of
gammaridean. Generally considered primitive caprellids such as
Cercops share many characters in common with the Podoceridae, and
a comparison of the figures of the podocerid Neozxendice (fig. 49a)
and the caprellid Cercops (fig. 49e) shows the overall general similarity
of body forms.

279-475

68- 8

108 U.S. NATIONAL MUSEUM BULLETIN 278

Caprella equilibra

Caprella scaura

Caprella andreae

@ Previous record
O New record

Figure 55.—Distribution records of Caprella andreae, Caprella danilevskii, Caprella equilibra,
Caprella scaura, and Caprella unica in the western North Atlantic.

Recently, Kudrjaschov and Vassilenko (1966) described a new
family, Caprogammaridea, which they thought formed a link between
the Gammaridea and Caprellidea. They placed this family within the
Gammaridea due to the following characters: The cephalon is not
fused with pereonite 1; small coxal plates are present on all pereonites
except 3 and 4; the abdomen consists of 5 free somites and bears 3
pairs of biramous pleopods and 2 pairs of uniramous uropods; and
the abdomen bears a small telson. In order to place this important link
between the 2 suborders in the Gammaridea, I believe that it would
be necessary to alter significantly the definition of the Gammaridea.
Stebbing (1906) defines this suborder as Amphipoda which have the:
Head not fused with Ist segment of peraeon. Palp of maxilliped 2- to 4-jointed.

Peraeon with 7 pairs of legs; 5 or 6 segments of peraeon with branchial vesicles;
4 segments of female with marsupial plates; 1st joint of gnathopods 1, 2 and of

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 109

33)

31°

30°

“Gill” Cruise 3

@ Phtisica present |23°
© Phtisica absent

esa

26°

Ficure 56.—Distribution of Phtisica marina in plankton samples taken on cruise 3 of the

M/V Theodore N. Gill.

peraeopods 1—5 forming or united to well developed side-plates. Pleon consisting
usually of 7 free segments, carrying 3 pairs of pleopods and usually 3 pairs, at
least 1 pair, of uropods; uropod 1 always with 2 rami. Eyes varying in size and
form, 0-4 in number. Hepato-pancreatic tubes 4, rarely 2; rectal glands 2 or 1,
sometimes rudimentary. Heart with 3 pairs, rarely 1 pair, of ostia. Nerve-chain
with 4 ganglia in pleon-segments 1-4.

110 U.S. NATIONAL MUSEUM BULLETIN 278

Ficure 49.—a and b, Neoxenodice caprellinoides, male: a, lateral view; b, maxilla 1; ¢ and
d, Caprogammarus gurjanovae, male: c, lateral view; d, maxilla 1; e and f, Cercops holbolli,
male holotype: ¢, lateral view; f, maxilla 1 (a after Schellenberg, 1926; c and d after
Kudrjaschov and Vassilenko, 1966).

CAPRELLIDAE OF WESTERN NORTH ATLANTIC Ltt

Stebbing (1906) defines the Caprellidae as Amphipoda which have the:

Head fused with 1st segment of peraeon. Palp of maxilliped 1- to 4-jointed.
Peraeon often with fewer than 7 pairs of legs; 2, rarely 3, segments of peraeon
with branchial vesicles; 2 segments in female with marsupial plates; Ist joint of
gnathopods and peraeopods wanting. Pleon and its legs rudimentary. Eyes small,
1 pair. Hepato-pancreatic tubes 2, rectal glands none. Heart with 3 pairs of ostia.
Posterior ganglia of nerve-chain very small, none situated in pleon.

The following facts should be considered before placing this im-
portant link in the Gammaridea:

1. No gammaridean bears rudimentary appendages on pereonites
3 and 4; however, in Neoxenodice they are reduced though normally
segmented. Caprogammarus (fig. 49c) bears 2 [? 1]-segmented ap-
pendages on the pereonites and the reduction of these appendages is
in the Caprellidea.

2. Gammarideans have at least 4 pairs of gills with the exception
of Neorenodice which has 3 pairs. Caprogammarus bears only 2 pairs
as Is common in the Caprellidea.

3. Gammarideans have at least 4 pairs of incubatory plates, al-
though the number is not known for Neozenodice. Caprogammarus
bears only 2 pairs as is common in the Caprellidea.

4. Most caprellids do bear small coxal plates on pereonites 1 and 2
and 5-7 which, as in Cercops and Aeginella, are as well if not better
developed than in Caprogammarus (fig. 3f).

5. Some rather advanced caprellid genera such as Aeginella and
Proaeginina have only a partial fusion of the cephalon with pereonite 1
and the suture between them is quite distinct.

6. The abdomen of Cercops consists of 5 somites and bearing in the
male what could be considered 2 pairs of very rudimentary pleopods
and 2 pairs of uropods.

7. Maxilla 1 of gammarideans always bears an inner lobe (fig. 49b)
which is absent in the Caprellidea (fig. 49f). Caprogammarus was de-
scribed as bearing an inner lobe (fig. 49d). The only irreconciliable
differences between the Caprogammaridae and the Caprellidea are
the presence of a telson, biramous pleopods, and an inner lobe on
maxilla 1 in Caprogammarus. The telson of Caprogammarus is very
small and may represent the dorsal anal lobe which is sometimes
quite large in the Caprellidea. The inner lobe in the figures of maxilla 1
of Caprogammarus seems to be of quite unusual shape and is even
larger than that of Neorenodice. It might be that the figures are not
of an actual lobe but of some distortion caused during dissection.
Regrettably, I have not been able to obtain specimens of Caprogam-
marus so I must rely on Kudrjaschov and Vassilenko’s description
and figures. The biramous pleopods and the inner lobe of maxilla 1

112 U.S. NATIONAL MUSEUM BULLETIN 278

are not as important in my opinion as the other characters mentioned
above, which Caprogammarus shares with the Caprellidea.

For these reasons, I am placing Caprogammarus in the family
Caprellidae. It might be argued that since the genera Neozenodice—
Caprogammarus—Cercops form a fairly even gradation between the
Gammaridea and the Caprellidea that merging of the suborders
would be justified. To do this, however, one would have to take into
consideration splitting the family Caprellidae into several families to
place it on the same level as gammaridean families. This probably
could be done be restricting the family Caprellidae to those forms
which have quite reduced abdomens and by using the family Ca-
progammaridae for those transitional forms such as Caprogammarus
and Cercops. In my opinion the Caprellidae are too poorly known to
make such a revision at this time and consideration should be given
to details of the mouthparts of members of all the genera involved
prior to a reorganization of familial taxa.

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CAPRELLIDAE OF WESTERN NORTH ATLANTIC EL

Bosc, L. A. G.
1801-02. Histoire naturelle des crustacés, contenant leur description et
leurs mocurs; avec figures dessinées d’aprés nature. Paris, vol. 2,
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1830. Ibid, ed. 2, vol. 2, 306 pp., pls. 10-18 [Caprellidae, pp. 123-126, pl.
15].
BovUcHARD-CANTEREAUR,
1833. Catalogue des crustacés observés jusqu’dé ce jour a l’etat vivant dans
la Boulonnais. Boulogne, 24 pp. [Caprellidae, p. 18].
BousFIELD, E. L.
1956a. Malacostracan crustaceans from the shores of western Nova Scotia.
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1958. Littoral marine arthropods and mollusks collected in western Nova
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1962. Studies on littoral marine arthropods from the Bay of Fundy region.
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1958. The fauna of Minas Basin and Minas Channel. Op. cit., no. 166,
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1914. Notes on Tasmanian Caprellidae. Pap. Proc. Roy. Soc. Tasmania,
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BRuNEL, P.

1961. Liste taxonomique des invertébrés marins des Parages de la Gaspésie
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BucHuo.z, R.

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CaLMAN, W. L.

1927. Guide to the Crustacea exhibited in the Department of Zoology,
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Ciriusu, A.

1941. Note sur quelques Caprellides des eaux frangaise et monégasques.

Bull. Inst. Océanogr., no. 803, 15 pp., 4 figs.

116 U.S. NATIONAL MUSEUM BULLETIN 278

Ci riust, S.

1956. Introducere la monografia amfipodelor marii Negre (Litoralul
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CxXrivusv, S., and CXrAusu, A.

1942. Amphipodes provenent des dragages effectués dans eaux roumaines
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Carus, J. V.

1885. Coelenterata, Echinodermata, Vermes, Arthropoda. vol. 1, xi +
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CrraME-Vivas, M. J., and Gray, I. E.

1966. The distributional pattern of benthic invertebrates of the Continental
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CHEVREUX, E.

1887a. Catalogue des crustacés amphipodes marins du sud-ouest de la
Bretagne, suivi d’un apercu de la distribution geégraphique des
amphipodes sur les cétes de France. Bull. Soc. Zool. France, vol.
12, pp. 288-340, pl. 5 [Caprellidae, pp. 317-319, 335].

1887b. Crustacés amphipodes nouveaux dragtes par l Hirondelle pendant
sa campagne de 1886. Ibid., pp. 566-580 [Caprellidae, pp. 578,
579].

1888. Troisieme campagne de I’Hirondelle, 1887. Sur quelques crustacés
amphipodes du littoral des Agores. Op. cit., vol. 13, pp. 31-35
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1898. Révision des amphipodes de la céte océanique de France. Compt.
Rendu 27th Session Assoc. Francaise Avance. Sci., pt. 2, Notes
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1900. Amphipodes provenant des campagnes de I’ Hirondelle (1885-1888).
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[Caprellidae, pp. 118-121].

1901 (1902). Amphipodes recueillis par la Melita sur les cétes occidentale
et méridionale de Corse. Compt. Rendu 30th Session Assoc.
Frangaise Avance. Sci., pp. 692-699, pl. 5 [Caprellidae: p. 696].

CHEvREUX, FE. and Bouvier, E. L.

1893. Les amphipodes de Saint-Vaast-La-Hougue. Ann. Sci. Nat. Zool.,

ser. 7, zool., vol. 15, pp. 109-144, pl. 2 [Caprellidae, pp. 142-144].
CHEvREUX, E., and pE GUERNE, J.

1893. Crustacés et Cirrhipédes commensaux des tortues marines de la
Méditerranée. Compt. Rendu 21st Session Assoc. Francaise
Avance. Sci., 3 pp. [Caprellidae, pp. 2-3].

CHEVREUX, E., and Faas, L.

1925. Amphipodes, 9. Faune de France. Paris, 488 pp., 438 figs. [Caprel-

lidae, pp. 433-461, figs. 422-437].

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 117

CHICHKOFF, G.

1912. Contribution a l’étude de la faune de la Mer Noire. Animaux récoltés
sur les cOtes Bulgares. Arch. Zool. Exp., ser. 5, vol. 10, no. 2,
pp. xxix-xxxix [Caprellidae: p. xxxv].

CHILTON, C.

1910 (1911). The Crustacea of the Kermadec Islands. Trans. New Zealand
Inst., vol. 48, pp. 544-573 [Caprellidae, pp. 546, 567].

1921. A small collection of Amphipoda from Juan Fernandez. Jn Skotts-
berg, C., the natural history of Juan Fernandez and Easter Island,
vol. 3, no. 14, pp. 81-92, 4 figs. [Caprellidae, pp. 90-91, fig. 4).

CHUMLEY, J.

i918. The fuana of the Clyde Sea area, being an attempt to record the
zoological results obtained by the late Sir John Murray and his
assistants on board the S.Y. Medusa during the years 1884 to
1892. Glasgow, pp. vi + 200, 1 map [Caprellidae, pp. 34, 52,
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Cocks, W. P.

1849. Contributions to the fauna of Falmouth. 17th Ann. Rep. Roy.

Cornwall Polytech. Soc., pp. 38-102 [Caprellidae, p. 83].
Co.tuin, J.

1884. Om Limfjordens tidligere og nuvaerende Marine Fauna, med saerligt
hensyn til Bl¢dddynfaunaen. Copenhagen, 168 pp., 1 pl. [Caprellidae,
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Costa, S.

1960a. Premier apergu sur la repartition des caprelles dans la region de
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1960b. Note préliminaire sur l’ethologie alimentaire de deux caprellides
de la Rade de Villefranche-sur-Mer. Op. cit., fasc. 19, no. 20,
pp. 103-105.

1961. Capagne de la Calypso en mer d’Alboran et dans la baie Ibéro-
Marocaine (1958). 3. Amphipodes. Ann. Inst. Oceanogr., new
ser., vol. 39, pp. 269-275 [Caprellidae, p. 271).

Covucg, R. Q.

1864. On some of the rarer forms of Cornish Crustacea. Nat. Hist. Trans.
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95-99].

Cow .gs, R. P.

1930. <A biological study of the offshore water of Chesapeake Bay. Bull.
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lidae, p. 351].

CRrAWSHAY, L. R.

1912. On the fauna of the outer western area of the English Channel.
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no. 3, pp. 292-393, pl. 6, 1 map [Caprellidae, p. 351].

CzERNIAVSEY, V.

1868. Materialia ad zoographiam Ponticum comparatam. [in Russian.]
Sezd Russkykh Iestestovoispitatelei (I Vranchi) 1. St. Petersburg,
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Daa, E.

1946. Undersékningar éver Oresund. XXX. The Amphipoda of the Sound.
Part II. Aquatic Amphipoda, with notes on changes in the hydrog-
raphy and fauna of the area. Lunds Univ. Aarsskr., new ser.,
sect. 2, vol. 42, no. 16, 49 pp., 5 figs., 8 tabs. [Caprellidae, pp. 22-23].

118 U.S. NATIONAL MUSEUM BULLETIN 278

Dana, J. D.

1853. Crustacea. United States Explor. Exped., vol. 14, pt. 2, pp. 689-1618
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1855. Op. cit., Atlas. 96 pls. [Caprellidae, pl. 10].

Day, J. H., and Morean, J. F. C.

1956. The ecology of South African estuaries. Part 8. The biology of Durban
Bay. Ann. Natal Mus., vol. 13, pt. 3, pp. 259-312, 2 figs. [Caprel-
lidae, p. 303].

DELAGE, Y.

1881. Contribution a létude de Jl appareil circulatoire des crustacés
edriophthalmes marins. Arch. Zool. Exp., ser. A, vol. 9, no. 42,
172 pp., 12 pls. [Caprellidae, pp. 120-134, 153, pl. 10].

DersAvIn, A. N.

1925. Materials of the Ponto-Azoph Carcinofauna (Mysidacea, Cumacea,
Amphipoda) [in Russian, English summary]. Russischen Hydro-
boil. Zeitschr., vol. 4, nos. 1-2, pp. 1-2, 10-35 [Caprellidae, pp. 27, 29].

1927. Hyperiidea and Caprellidea Kamchatka Expedition 1908-1909
[in Russian, English summary]. Op. cit., vol. 6, no. 1, pp. 18-15
[Caprellidae, pp. 14-15].

Dersuain, K. M.

1915. Fauna des Kola-Fjords und ihre Existentbedgungen [in Russian].
Mém. Acad. Imp. Sci. Petrograd, ser. 8, vol. 34, no. 1, pp. ix+929,
14 pls., 14 maps, 55 figs. [Caprellidae, pp. 453, 456].

1928. Amphipods. Die fauna des Weissen Meeres und seine Existenzbeding-
ugen. Liningrad Gosudarstvennyi Gidrolog. Inst. Isseldovaniia
Morei, SSSR, fase. 7-8, pp. 278-285 [Caprellidae, p. 282].

DesMaAREst, A. G.

1823. Malacostracés, vol. 28, pp. 138-425, 58 pls. In Dictionnaire des
sciences naturelles, dans lequel on traite méthodiquement des
différens étres de la nature, considérés soit en eux-mémes, d’dprés
Vetat actuel de nos connoissances, soit relativement a l’utilité qu’en
peuvent retirer la médécine, l’agriculture, le commerce et Ies
arts. Paris [Caprellidae, pp. 359-364, pl. 46].

1825. Considérations générales sur la classe de crustacés, et description
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1944. The bottom community of Ipswich Bay, Massachusetts. Ecol.,

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1942. Marine macroplankton from the Canadian Eastern Arctic. I. Amphip-
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1840. Histoire naturelle des crustacés comprenant l’anatomie, la physiol-
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1950. Amphipods. Contr. Bears Bluff Lab., no. 8, 13 pp. [Caprellidae,

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1937. Marine animals collected by Mr. A. W. Moore at Etah, northwest
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Fasricius, J. C.

1793. Entomologia systematica emendata et aucta. Secundum classes,
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Fasricivus, O.

1780. Fauna groenlandica, systematice sistens animalia Groenlandiae oc-
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120 U.S. NATIONAL MUSEUM BULLETIN 278

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1940. I Caprellidi del Mare Adriatico presso Rovigno. Thalassia, vol. 4,
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1900 (1901). Op. cit. 3° Voyage. Région d’Omonville-la-Rogue (Manche)
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1878. Beitrag zur Kenntnis der Naturgeschichte der Caprellen. Zeitschr.
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together with a particular description of each county, notices of

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 121

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Gois, A.

1866. Crustacea Amphipoda maris Spetsbergiam alluentis, cum speciebus
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GoopHart, C. B. and Harrison, R.

1940. Occurrence of some off-shore amphipods in the littoral zone. Nat.,

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1842. On anew genus, and on six new species of Crustacea, with observa-
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GossE, P. H.

1855. A manual of marine zoology for the British Isles. London, pt. 1,

pp. xi + 203, 335 figs. [Caprellidae, pp. 130-131, figs. 223-224].
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1960. The benthonic Amphipoda of the Mediterranean coast of Israel. I.
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GouLp, A. A.

1841. Report on the Invertebrata of Massachusetts, comprising the Mol-
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Gourret, P.

1892. Notes zoologiques sur l’Etang des Eaux-Blanches (Cette). Ann. Mus.

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GRAEFFE, E.

1902. V. Crustacea. Uebersicht der Fauna des Golfes von Triest nebst
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Grika, J. A.

1925. Evertebrater fra bankerne ved Spitsbergen indsamlet av m.k.
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Grisra, M. J.

1907. Invertebrés du Fond. pp. 503-567, pl. 79. In d’Orléans, L. P. R. due.
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Gronovius, L. T.

1760. Observationes de animalculis aliquot marinae aquae innatantibus
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Gufrin-MENEVILLE, F, E.

1829-44. Crustacés, vol. 2, 104 pls., vol. 3, 48 pp. In Iconographie du régne
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1886. Sur quelques amphipodes marins du nord de la France. Bull. Soc.

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1952. A list of the Crustacea of Tasmania. Rec. Queen Victoria Mus., vol.
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1954. Some collections of caprellids from Tasmania. Ann. Mag. Nat. Hist.,
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1935. Contributions to the fauna of Amphipoda and Isopoda of the southern
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1936. The zoogeography of the Kara Sea (contribution to the fauna of
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Haz, H. M.
1927. The fauna of Kangaroo Island, South Australia. Trans. Roy. Soc.
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HALLER, G.
1879a. Vorlafige Notizen iiber die Sustematik der im Mittelmeer vorkom-
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1895. Pycnogonider og Malacostrake Krebsdyr. Meddel. Grgniand, vol.
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Hansson, C. A.
1882 (1883). Bidrag til kannedom om de ligre djurlifvet vid morra Bohusl-
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De atl

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 122

Harrison, R. J.

1939. Some observations on the habits of the skeleton shrimps. Ann. Rep.
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1940. On the biology of the Caprellidae. Growth and moulting of Pseudopro-
tella phasma Montagu. Journ. Marine Biol. Assoc. United Kingdom,
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HasweE tu, W. A.

1880. On some additional new genera and species of amphipodous crusta-
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1882. Catalogue of the Australian stalk- and sessile-eyed Crustacea.
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1884 (1885). Revision of the Australian Laemodipoda. Proc. Linn. Soe.
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Hepaperta, J. W.

1950. Annotated list of certain marine invertebrates found on Texas jetties.
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1953. An introduction to the zoogeography of the northwestern Gulf of
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Harrier, C:

1866. Beitrage zur naiheren Kenntniss der Amphipoden des adriatischen
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HeEenpeERsoN, J. R.

1885. Recent additions to the invertebrate fauna of the Firth of Forth.
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Hersst, J. F. W.

1793. Versuch einer Naturgeschichte der Krabben und Krebse, nebst einer
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Herxtors, J. A.

1861. Symbolae carcinologicae. I. Catalogue des crustacés qui ont servi de
base au systéme carcinologique de M. W. de Haan, rédigé d’aprés
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Hewatt, W. G.

1946. Marine ecological studies on Santa Cruz Island, California. Ecol.
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Hiro, F. [= Ut1nomy1, H.]

1937. Caprellids from Tanabe Bay. Annot. Zool. Japonenses, vol. 16, no.

4, pp. 310-317, pl. 22.
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1879. Carcinologisches, grésstentheils gearbeitet in der zoologischen
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279-475—68——9

124 U.S. NATIONAL MUSEUM BULLETIN 278

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1882. Die Crustaceen, gesammelt wachrend der Fahrten des ‘Willem
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1883-84. Schaaldieren van de Oosterschelde (Crustacés de Il’Escaut de
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1889. Crustacea Neerlandica. II. Nieuwe lijst van tot de fauna van Neder-
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2, vol. 2 pt. 3, pp. 170-234, pls. 7-10 [Caprellidae, pp. 231-233].
Homes, S. J.
1904 (1905). The Amphipoda of southern New England. Bull. Bur. Fish.,
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HuntsMAN, A. G.
1915. A new caprellid from the Bay of Fundy. Contr. Canadian Biol.,
sessional pap. no. 39b, pp. 39-42, pls. 5-6.

Horton, F. W.
1904. Index Faunae Novae Zealandiae. London, pp. viii-+372. [Caprellidae,
p. 261].

Iarzynsky, T.

1870. Praemissus catalogus Crustaceorum amphipodum, inventorum in
mari albo et in mari glaciali ad litus murmanicum (Mypmahin
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Ives, J. E.

1891 (1892). Echinoderms and crustaceans collected by the West Greenland
Expedition of 1891. Proc. Acad. Nat. Sci. Philadelphia, pp. 479-481
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JOHANSEN, F.

1925. Fishes and marine invertebrates collected during the cruise of the
Arctic in 1923. Canadian Field Nat., vol. 39, no. 9, pp. 203-204
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1930. Marine Crustacae, Malocostraca and Pantopoda (Pycnogonida),
collected in the Gulf of St. Lawrence, Newfoundland, and the Bay
of Fundy in 1919, 1922, 1923, 1925, and 1926.

JOHNSON, C. W.

1934. List of marine Mollusca of the Atlantic coast from Labrador to Texas.

Proc. Boston Soc. Nat. Hist., vol. 40, 204 pp.
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1927. Seashore animals of the Pacific Coast. New York, pp. xiv+659, 700

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JoHNSON, R. G.

1965. Research on the marine geology and biology of Tomales Bay, Cali-
fornia. Prog. Rep. Atomic Energy Comm., Contract no. AT (11-1)-
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appendix 3, p. 4].

1966. Ibid, 17 pp., appendix 1, 6 pp., appendix 2, 3 pp., appendix 3, 4 pp.,
appendix 4, 11 pp. [Caprellidae, appendix 1, p. 3, appendix 2, p. 2,
appendix 3, p. 2, appendix 4, pp. 1 and 2].

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 125

Jounson, R. G., and JusKEvicg, J. A.

1965. Check list of marine invertebrates of the Tomales Bay region, Marin
County, California. Pacific Marine Sta. Res. Rep. no. 5, 78 pp.
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JOHNSTON, G.

1835. Illustrations in British zoology. Mag. Nat. Hist., vol. 8, pp. 668-674

[Caprellidae, pp. 668-674].
DE Kay, J. E.

1844. Crustacea, pt. 6, 70 pp., 13 pls. In Zoology of New York, or the New
York fauna; comprising detailed descriptions of all the animals
hitherto observed within the state of New York, with brief notices
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appropriate illustrations. New York (State) Nat. Hist. Surv.
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Kirk, T. W.

1878. Additions to the crustacean fauna of New Zealand. Ann. Mag. Nat.
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1878 (1879). On additions to the carcinological fauna of New Zealand.
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KuINcKOwstTROM, A.

1892. Ofversigt af de zoologiska arbetena under expeditionen till Spetsbergen
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Koruuer, R.

1884 (1885). Recherches sur la faune marins des Iles Anglo-Normandes.
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1885. Contribution a l’étude de la faune littorale des Iles Anglo-Normandes
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KOELBEL, C.

1886. Crustaceen, Pycnogoiden und Arachnoideen von Jan Mayen...
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KossMann, R.

1880. Malacostraca. Zool. Ergebn. Auftrage Acad. Wiss. Berlin Reise
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Krégyer, H.

1838. Gr6énlands amphipoder beskrevne af Henrik Kroyer. Danske Vidensk.
Selskabet Nat. Math. Afhandl., pt. 7, pp. 229-326 [Caprellidae,
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1842-43. Beskrivelse af nogle Arter og Slaegter, af Caprellina: med indle-
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1846. Voyages en Scandinavie en Laponie au Spitzberg et aux Férée.
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1966. A new family Caprogammaridae (Amphipoda, Gammaridea) found
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4 figs.

KunkKEL, B. W.

1910. The Amphipoda of Bermuda. Trans. Connecticut Acad. Arts Sci.,
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1918. The Arthrostraca of Connecticut. State Geol. Nat. Hist. Surv. Bull.,
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LAFouueEtTtsE, R.

1914. Caprellidae from Laguna Beach. Journ. Entomol. Zool. (Pomona

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1801. Systéme des animaux sans vertébres, ou, tableau général des classes,
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LAMEERE, A.

1895. Manuel de la faune de Belgique. Bruxelles, vol. 1, pp. x1+639, 701

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LATREILLE, P. A.

1802-03. Histoire naturelle, génerale et particuliére des crustacés et des
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1803. Chevrolle, vol. 5, pp. 332-333. In Nouveau dictionnaire d’histoire
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1816. Chevrolle, vol. 6, pp. 433-434. Op. cit., new ed.

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1836. Crustaceen, Arachniden und die ungeflugelten Insecten, vol. 4, pp.
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Lracu, W. E.

1818-14. Crustaceology, vol. 7, pt. 1, pp. 8838-384 [1813]; pt. 2, pp. 385-437
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Lrypie, F.

1860. Ueber Geruchs- und Gehérorgane der Krebse und Insecten. Arch.

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CAPRELLIDAE OF WESTERN NORTH ATLANTIC 127

Litsesorc, W.

1856. Om Hafs-Crustaceer vid Kullaberg i SkAne. Vetensk. Akad. Forhandl.,

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LINNAEUS, CARL VON

1767. Systema naturae. Holmiac, 12th ed., vol. 1, pt. 2, pp. 533-1327
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1769. Natuurlyke historie, of Ultvoerige beschryving der dieren, planten,
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1788. Systema naturae. Lipsiae, 13th ed., vol. 1, pt. 5, pp. 2225-3020
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1793. Systema naturae, par regna tria naturae, secundum classes, ordines,
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1800. <A general system of nature, through the three grand kingdoms of
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L6nNBorG, E.

1902 (1903). Undersékningar rérande Skeldervidens och Angrinsande
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Lucas, H. P.

1849. Histoire naturelle des animaux articles. Pt. 1. Crustacés, arachnides,
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LirxKeEn, C.

1875. The Crustacea of Greenland, pp. 146-165. In Jones, T. R., Manual
of the natural history, geology and physics of Greenland and the
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LuruHer, W., and Frepumr, K.

1961. Die Unterwasserfauna der Mittelmeerkiisten. Hamburg and Berlin,

253 pp., 28 figs., 46 pls. [Caprellidae, p. 158, pl. 24].
McANpreEw, R.

1861. List of the British marine invertebrate fauna. London, 71 pp.

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McCain, J. C.

1965. The Caprellidae (Crustacea: Amphipoda) of Virginia. Chesapeake
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1966. Abyssicaprella galatheae, a new genus and species of abyssal caprellid
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MacDona.p, R.

1939. The marine Crustacea of Ardglass Harbour, C. Down. Ann. Mag.

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McDoveatt, K. D.

1943. Sessile marine invertebrates at Beaufort, North Carolina. Ecol.
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MacGinitTig, G. E.

1935. Ecological aspects of a California marine estuary. Amer. Midland

Nat., vol. 16, no. 5, pp. 629-765 [Caprellidae, p. 701].
M’Intosu, W. C.

1874. On the invertebrate marine fauna and fishes of St. Andrews. Ann.
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MacKay, D. C. G.

1945. Notes on the aggregating marine invertebrates of Hawaii. Ecol.,

vol. 26, no. 2, pp. 205-207 [Caprellidae, p. 205].
Macnak, W.

1953. On a small collection of amphipods from Tristan da Cunha. Proc.
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MairTLanp, R. T.

1874. Naamlijst van Nederlandsche Schaaldieren. Tijdschr.Nederlandsche
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Marine Brow. Las.

1904. Plymouth marine invertebrate fauna, being notes of their local dis-
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1931. Plymouth marine fauna. Plymouth, 371 pp., 1 map [Caprellidae,
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1957. Plymouth marine fauna. Plymouth. 3rd ed., pp. xliiit457, 4 maps
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Marion, A. F.

1883. Esquisse d’une topographie zoologique du Golfe de Marseille. Ann.
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Massey, A. L.

1911 (1912). Report of a survey of trawling grounds on the coasts of Counties
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Mayer, P.

1882. Die Caprelliden des Golfes von Neapel und der angrenzenden Meeres-
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1890. Ibid. Nachtrag zur Monographie derselben. Op. cit., vol. 17, pp.
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1898. Metaprotella sandalensis, n. sp. Pt. 1, pp. 53-56, 6 figs. In Willey,
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1903. Die Caprellidae der Siboga-Expedition. Siboga Exped., vol. 34, 160
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CAPRELLIDAE OF WESTERN NORTH ATLANTIC 129

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1880. Ibid., Op. cit., vol. 12, no. 3, pp. 465-512 [Caprellidae, pp. 494-496].
1890. Crustacea Malacostraca. Vidensk. Udbytte Kanonbaden ‘‘Hauch’s”
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MENzEL, R. W.
1956. Annotated check-list of the marine fauna and flora of the St. George’s
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Merzcer, A.
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1875. Crustaceen aus den Ordnungen Edriophthalmata und Podophthal-
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Miers, E. J.
1877a. Report on the Crustacea collected by the naturalists of the Arctic
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1877b. List of the species of Crustacea collected by the Rev. A. E. Eaton
at Spitzbergen in the summer of 1873, with their localities and
notes. Op. cit., ser. 4, vol. 19, pp. 131-140 [Caprellidae, p. 139].
1880. On a small collection of Crustacea made by Edward Whymper,
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1884. Crustacea (The collections from Melanesia.), pp. 178-322, pls. 18-
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Mitts, E. L.
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Miyap1, D., and Masui, T.
1942. Studies on the benthic communities of the Nanao-wan [in Japanese].
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130 U.S. NATIONAL MUSEUM BULLETIN 278

Monraay, G.

1815. Descriptions of several new or rare animals, principally marine,
discovered on the south coast of Devonshire. Trans. Linn. Soe.
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Monterosso, B.

1915. Caprellidae del Golfo di Catania. Atti Acecad. Gioania Sci. Nat.

Catania, ser. 5, vol. 8, 16 pp., 3 figs.
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1776. Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae
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Nicouet, H.

1849. Crustaceos, vol. 3, pp. 115-318. In Gay, C. Histoira fisica y politica
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NoORDGAARD, O.

1905. UHydrographical and biological investigations in Norwegian Fiords,
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1911 (1912). Faunistiske og biologiske iakttagelser ved den _ biologiske
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Norman, A. M.

1876. Crustacea, Tunicata, Polyzoa, Echinodermata, Actinozoa, Foramin-
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1882. Report on the Crustacea, pp. 683-689. In Staff Commander Tizard
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1886. Museum normaniam, or a catalogue of the Invertebrata of Europe,
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1902. Notes on the natural history of East Finmark. Ann. Mag. Nat.
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1905a, Museum normanium, or a catalogue of the Invertebrata of the
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1905b. Revised nomenclature of the species described in Bate and West-
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1907. Notes on the Crustacea of the Channel Islands. Op. cit., ser. 7,
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1908 (1909). The Crustacea of the Channel Islands. Rep. Trans. Soc.
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CAPRELLIDAE OF WESTERN NORTH ATLANTIC fod

Norman, A. M., and Scort, T.

1906. The Crustacea of Devon and Cornwall. London, pp. xv+282,

24 pls. [Caprellidae, pp. vii, viii, ix, x, 98-99].
Norman, A. M., and Brapy, G. S.

1910. The Crustacea of Northumberland and Durham. Trans. Nat. Hist.
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OHLIN, A.

1895a. Bidrag till Kinnedomen om Malakostrakfaunan i Baffin Bay och
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1895b. Additional notes to my paper: “Bidrag till kinnedomen om malakos-
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OupEvIc, H.

1917. Die Amphipoden, Isopoden und Cumaceen des Eisfjords. Zoologische
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1933. Sveriges Amphipoder. Géteborgs Vetensk. Vitterhets-Simhalles
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Ouivetra, L. P. H.

1940. Contribuicao ao conhecimente des Crustaceos do Rio de Janeiro.
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ORTMANN, A. E.

1901. Crustacea and Pyenogonida collected during the Princeton Expedi-
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PackarRD, A. S., JR.

1867. Observations on the glacial phenomena of Labrador and Maine,
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PALuAS, EP. 8.

1772. Spilicilegia zoologica quibus novae imprimis et obscurae animalium
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Parritt, E.

1873. The fauna of Devon. Pt. IX. Sessile-eyed Crustacea. Rep. Trans.
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PavuULMIER, F. C.

1905. Higher Crustacea of New York City, Bull. New York State Mus.,
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Pearse, A. S.

1908. Descriptions of four new species of amphipodous Crustacea from
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12 U.S. NATIONAL MUSEUM BULLETIN 278

1912. Notes on certain amphipods from the Gulf of Mexico, with descrip-
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1936. Estuarine animals at Beaufort, North Carolina. Journ. Elisha Mitchell
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Prarssz, A. S., Hum, H. J., and Warton, G. W.

1942. Ecology of sand beaches at Beaufort, North Carolina. Ecol. Monogr.,

vol. 12, pp. 135-190, 24 figs. [Caprellidae, p. 184].
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1951. The biota of the reefs off the Carolinas. Journ. Elisha Mitchell

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1883. Etudes sur la faune littorale de la Belgique. Mollusques et autres
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1886. Note sur la présence de Caridina desmaresti dans les eaux de la
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PENNANT, T.

1777. British zoology. London, vol. 4, pp. viii + 154, 93 pls. [Caprellidae,

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Prquranat, W. E.

1966. Biofouling studies off Panama City, Florida. I. Res. Rep. Office
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Pryrort, §., and Trituus, J. P.

1964. Recherches sur la sexualité et la gland androgene de Caprella aequili-
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PFEFFER, G.

1889 (1890). Die Fauna der Insel Jeretik, Port Wladimir, an der Murman-
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Prruot, J. M.

1939. Résultats scientifiques des Croisiéres du Navire-Ecole Belge Mer-
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PREFOUNTAINE, G., and BRUNEL, P.

1962. Liste d’invertébrés marins recueillis dans l’estuaire du Saint-Laurent
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Procter, W.

1933. Biological survey of the Mount Desert Region. Philadelphia. Part 5,
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RANCUREL, P.

1949. Notes sur les amphipodes marins de la région de Marseille. Bull.
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CAPRELLIDAE OF WESTERN NORTH ATLANTIC 133

RatTuHBun, M. J.

1905. Fauna of New England. 5. List of the Crustacea. Occ. Pap. Boston

Soc. Nat. Hist., vol. 7, 117 pp. [Caprellidae, pp. 7, 76-79].
RATHBUN, R.

1880 (1881). The littoral marine fauna of Provincetown, Cape Cod, Mas-
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RaTuHKE, H.

1843. Beitrige zur Fauna Norwegens. Nov. Act. Acad. Leop.-Carol. Nat.

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REEp, E. C.

1897. Catdlogo de los Crustéceos Amfipodos i Lemodipodes de Chile. Rev.

Chilena Hist. Nat., vol. 1, pp. 9-11 [Caprellidae, p. 11].
REIBISCH, J.

1906. Faunistisch-biologische Untersuchungen iiber Amphipoden der Nord-
see. Wiss. Meeressunt., new ser., vol. 9, pp. 185-237, pls. 8-9,
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223, 229, 230, 232, 233, 235].

Rerp, D. M.

1951. Report on the Amphipoda (Gammaridea and Caprellidea) of the
coast of tropical West Africa. Atlantide Rep., no. 2 pp. 189-291,
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Ricwarps, H. G.

1929. A faunistic survey of the marine invertebrates of New Jersey, with
especial reference to the Cape May region. M.S. Thesis Univ.
Pennsylvania, 110 pp., 4 pls. [Caprellidae, p. 84].

1938. Animals of the seashore. Boston, 273 pp., 28 pls. [Caprellidae, p.
213, pl. 24, fig. 7].

Ricketts, E. F., and Catvin, J.

1939. Between Pacific Tides. Stanford, pp. xxii + 320, 112 figs., 46 pls.
[Caprellidae, pp. 70-71, 194-195, 280].

1952. Op. cit., Revised by J. W. Hedgpeth. Stanford. 3rd ed., pp. xiii + 502,
134 figs., 1 color pl., 46 pls. [Caprellidae, pp. 68-69, 248-249, 440,
fig. 28].

Risso, A.

1816. Histoire naturelle des Crustacés des environs de Nice. Paris, 176 pp.,
3 pls. [Caprellidae, pp. 129-131].

1826. Histoire naturelle des principales productions de l’ Europe meridionale
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RoBertson, D.

1886-87 (1888). A contribution towards a catalogue of the Amphipoda
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Rurro, S.

1938. Studi sui crostacei anfipodi. VIII. Gli anfipodi marini del Museo
Civico di Storia Naturale di Genova. a) Gli anfipodi del Mediter-
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1 fig. (Caprellidae, p. 150].

1941. Studi sui crostacei anfipodi. X. Contributo alla conoscenza degli
anfipodi marini Italiani. Boll. Ist. Entomol. Roy. Univ. Bologna,
vol. 11, pp. 112-126 [Caprellidae, pp. 124-125].

134 U.S. NATIONAL MUSEUM BULLETIN 278

1946. Studi sui crostacei anfipodi. XI. Gli anfipodi benthonici di Rovigno
d’Istria (Nota preventiva.). Boll. Soc. Entomol. Italiana, vol. 76,
nos. 7 and 8, pp. 46-56, 1 map [Caprellidae, p. 53).

1947. Studi sui crostacei anfipodi. XIV.Su alcune specie di anfipodi dell-
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bon, Guinea Portoghese, Africa Australe). Mem. Mus. Civ. Storia
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RuFro, S., and WiEsErR, W.

1952. Osservazioni sistematiche ed ecologiche su alcuni anfipodi delle
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Sars, G. O.

1885. Crustacea, I. Norwegian North-Atlantic Expedition, 1876-1878.
Christiania, 280 pp., 21 pls., 1 map [Caprellidae, pp. 222-230,
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1886. Crustacea, II. Op. cit., Christiana, 96 pp., 1 map [Caprellidae, pp.
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1895. Amphipoda. An account of the Crustacea of Norway with short
descriptions and figures of all the species. Christiania and Copen-
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Sars, M.

1858 (1859). Oversigt over de in den norske arctiske Region forekommende
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1863. Geologiske og zoologiske Iagttagelser, anstillede paa en Reise i en
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SAUNDERS, C. G.

1966. Dietary analysis of caprellids (Amphipoda). Crustaceana, vol. 10,

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1817-18. An account of the Crustacea of the United States. Journ. Acad.
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ScHELLENBERG, A.

1926. Die Caprelliden un Neoxenodice caprellinoidies n.g., n. sp. der Deut-
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1927. Amphipoda des nordischen Plankton. Nordisches Plankton, vol. 20,
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1928. Report on the Amphipoda. Zoological results of the Cambridge
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1931. Gammariden un Caprelliden des Magellangebietes, Stiidgeorgiens un
der Westantarktis. Further Zool. Results Swedish Antarctic
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262-266, 272].

1938. Litorale Amphipoden des Tropischen Pazifiks. Svenska Vetensk.
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CAPRELLIDAE OF WESTERN NORTH ATLANTIC 135

1942, Krebstiere oder Crustacea IV: Flohkrebse oder Amphipoda. Tier-
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233-241].

ScuiyrsMa, K.

1931. Amphipoda van de Zuiderzee (1927-1929). Nederlandsche Dier,

Vereeniging, no. 3, pp. 7-27 [Caprellidae, p. 26].
ScuidpTs, J. C.

1875. Krebsdyrenes Sugemund. Naturhist. Tidsskr., ser. 3, vol. 3, pp.

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ScHNEIDER, J. 8.

1883. Nogle zoologiske iagttagelser fra Vardg i @st-Finmarken. Troms¢
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1884, Undersggelser af dyrelivet i de arktiske fjorde. II. Crustacea og
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1891. Undersggelser af dyrlivet i de arktiske fjorde. IV. Mollusca og
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1924 (1926). Tromsgsundets Amphipoder, Isopoder og Cumaccer. Op.cit.,
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Scuurin, A.

1935. Zur Fauna der Caprelliden der Bucht Peters des Grossen (Japanisches

Meer). Zool. Anz., vol. 122, nos. 7 and 8, pp. 198-203, 4 figs.
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1887 (1888). A revised list of the Crustacea of the Firth of Forth. 6th
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p. 250].

1897. The marine fishes and invertebrates of Loch Fyne. 15th Ann. Rep.
Fish. Board Scotland, pt. 3, pp. 107-174, 3 pls. [Caprellidae, p.
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1899. Report on the marine and freshwater Crustacea from Franz-Josef
Land, collected by Mr. William 8S. Bruce, of the Jackson-
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174, pp. 60-126, pls. 3-9 [Caprellidae, p. 81].

1901. Notes on gatherings of Crustacea, collected for the most part by
the Fishery Steamer Garland and the Steam Trawler St. Andrew
of Aberdeen, and examined during the year 1900. 19th Ann. Rep.
Fish. Board Scotland, pt. 3, pp. 235-281, pls. 17-18 [Caprellidae,
pp. 267-268].

1906. A catalogue of land, fresh-water, and marine Crustacea found in
the basin of the River Forth and its estuary. Pt. 1. Malacostraca,
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SHOEMAKER, C. R.

1926. Report on the marine amphipods collected in Hudson and James
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1930. The Amphipoda of the Cheticamp Expedition of 1917. Op. cit.,
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SINEL, J.

1906 (1907). Contribution to our knowledge of the Crustacea of the Channel
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136 U.S. NATIONAL MUSEUM BULLETIN 278

SLABBER, M.

1769. [Title page reads 1778] Natuurkundige Verlustigingen, behelzende
microscopise waarneemingen van in- en Uitlandse Water- en Land-
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figs. 1-3].

Smirtu, S. T.

1873. See Verrill, A. E., 1873.

1883a. List of the Crustacea dredged on the coast of Labrador by the ex-
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1883b. Review of the marine Crustacea of Labrador. Op. cit., vol. 6, no.
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Soxotowsky, A.

1900. Die Amphipoden Helgolands. Wiss. Meeresunt., new ser., vol. 4,
no. 2, pp. 1438-166, pl. 3 [Caprellidae, pp. 161-162, pl. 3, figs.
15-16].

Sovinsku, V.

1880. On the Amphipoda of Sevastopol Bay [in Russian]. Zapiski
Kievskago Obshestva Estestvoispytatelei, vol. 6, pt. 1, pp. 87-136,
pls. 3-5 [Caprellidae, pp. 88, 89, 90, 95-101, pl. 3, fig. 1].

1895. Higher Crustacea (Malacostraca), collected by the Second Black
Sea Deep Sea Expedition of 1890 & 1891 [in Russian]. Op. cit.,
vol. 14, no. 1, pp. 225-283, pls. 4-6 [Caprellidae, pp. 255-261].

1898. The higher Crustacea (Malacostraca) of the Bosporus from the
material collected by A. A. Ostroumoff in 1892 & 1893 [in Russian].
Op. cit., vol. 15, no. 2, pp. 447-518, pls. 8-13 [Caprellidae, pp.
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Spooner, G. M.

1950. Notes on the Plymouth marine fauna. Amphipoda. Journ. Marine
Biol. Assoc. United Kingdom, vol. 29, no. 1, pp. 247-253 [Caprel-
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Srauio, L.

1877. Catalogo metidico e descrittive dei Crostacei dell’Adriatico. Atti
Reale Ist. Veneto Sci. Lett. Arti, ser. 5, vol. 3, pp. 355-385, 499-
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STaPPErs, L.

1911. Crustaces Malacostraces. Campagne Arctique de 1907, Duc d’ Orleans.
Bruxelles, vol. 7, pp. vi+152, I-XII, I-XII, pls. 1-7, 2 maps
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SresBina, T. R. R.

1876. Description of a new species of sessile-eyed crustacean, and other
notices. Ann. Mag. Nat. Hist., ser. 4, vol. 17, pp. 73-80, pls. 4-5
[Caprellidae, p. 78].

1879. Sessile-eyed Crustacea of Devonshire. Supplementary list. Rep. Trans.
Devonshire Assoc. Adv. Sci. Lit. Arts, vol. 11, pp. 516-524 [Caprel-
lidae, p. 521].

1888. Report on the Amphipoda collected by H.M.S. Challenger during
the years 1873-76. Rep. Challenger, zool., vol. 29, pt. 67, pp.
xxiv-+1737, i-xii, 210 pls. [Caprellidae, pp. 1226-1268, pls.
139-145].

1895. Two new amphipods from the West Indies. Ann. Mag. Nat. Hist.,
ser. 6, vol. 15, pp. 397-403, pls. 14-15.

1906.

CAPRELLIDAE OF WESTERN NORTH ATLANTIC 137

Amphipoda I. Gammaridea. Das Tierreich, pt. 21, 806 pp., 127 figs.

1910a. General catalogue of South African Crustacea (part V. of S. A.

Crustacea, for the marine investigations of South Africa). Ann.
South African Mus., vol. 6, pt. 4, pp. 281-593, pls. 41-48 [Caprel-
lidae, pp. 464-471, pl. 48].

1910b. Scientific results of the trawling expedition of H.M.C.S. Thetis.

Part V: Crustacea Amphipoda. Mem. Australian Mus., vol. 4,
pt. 12, pp. 567-658, pls. 47-60 [Caprellidae, pp. 629-632, 651-654,
pl. 60].

STEINBERG, J. E., and DoucuErrty, FE. C.

1957.

The skeleton shrimps (Crustacea: Caprellidae) of the Gulf of Mexico.
Tulane Stud. Zool., vol. 5, no. 11, pp. 267-288, 30 figs.

STEPHENSEN, K.

1912.

Report on the Malacostraca Pyenogonida and some Entomostraca
collected by the Danmark Expedition to North-East Greenland.
Meddel. Grgnland, vol. 45, pp. 503-630, pls. 34-43 [Caprellidae,
pp. 543-544].

1913a. Grgnlands Krebsdyr og Pyenogonider (Conspectus Crustaceorum et

1913b

1915.

1916.

1927a

1927b

1927c.

1928.

1929a

1929b

1931.

1933.

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. Account of the Crustacea and the Pyenogonida collected by Dr.
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and Giesecke Lake in West Greenland. Op. cit., vol. 51, pp. 55-77,
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Isopoda, Tanaidacea, Cumacea, Amphipoda (excl. Hyperiidea). Rep.
Danish Oceanogr. Exped. 1908-10, vol. 2, biol. D.1, 53 pp., 33
figs. [Caprellidae, pp. 4, 53].

Zoogeographical investigations of certain fjords in southern Greenland
with special reference to Crustacea, Pycnogonida and Echinoder-
mata including a list of Aleyonaria and Pisces. Meddel. Grgnland,
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. Revideret Fortegnelse over Danmarks Arter af Amphipoda (3. Del:
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. The Folden Fiord. Crustacea II. List of the Amphipoda. Troms¢g

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Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. XL.
Crustacea from the Auckland and Campbell Islands. Vidensk.
Meddel. Dansk Naturhist. Foren., vol. 83, pp. 289-390, 33 figs.
[Caprellidae, pp. 354-355].

Storkrebs II. Ringkrebs. 1. Tangloper (Amfipoder). Danmarks Fauna.
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figs. 91-93].

. Amphipoda. Die Tierwelt der Nord- und Ostsee, no. 14, 188 pp.,

43 figs. [Caprellidae, pp. 174-182, figs. 42-43].

. Marine Crustacea Amphipoda. Zoology of the Faroes, no. 23, 40 pp.
[Caprellidae, pp. 19-20, 34].

A new caprellid from N. Norway Parvipalpus norvegicus n. sp.
Norske Vidensk. Selskabs Skr., no. 5, 7 pp., 3 figs.

Amphipoda. The Godthaab Expedition 1928. Meddel. Grgnland,
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138 U.S. NATIONAL MUSEUM BULLETIN 278

1935. En del amphipoder fra Vest- og Nord-Norge. Norske Vendensk.
Selskab. Forhandl., vol. 7, no. 32, pp. 115-118 [Caprellidae, p. 118].
1940. Marine Amphipoda. Zoology of Iceland, vol. 3, pt. 26, 111 pp., 13
figs. [Caprellidae, pp. 69-75].
1942. The Amphipoda of N. Norway and Spitsbergen with adjacent waters.
Troms¢ Mus. Skr., vol. 3, pt. 4, pp. 8363-526, 26 figs. [Caprellidae,
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1944a. Crustacea Malacostraca VIII (Amphipoda IV). Danish Ingolf Exped.,
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1944b. Amphipoda. The Zoology of East Greenland. Meddel. Grgnland,
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1949. The Amphipoda of Tristan da Cunha. Res. Norwegian Sci. Exped.
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Stimpson, W.
1854 (1853). Synopsis of the marine Invertebrata of Grand Manan: or the
region about the mouth of the Bay of Fundy, New Brunswick.
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1857. On the Crustacea and Echinodermata of the Pacific shores of North
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Stock, J. H., and BoLKLanpsEr, A. E. M. H.
1952. Notes on adventive Amphipoda (Crustacea, Malacostraca) on the
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Stossicu, M.
1881. Prospetto della fauna del mare Adriatico. Boll. Soc. Adriatica Sci.
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Stscuapova, T. F., Moxyrvsky, O. B., and Pasternak, F. A.
1957. Flora and fauna of the coastal zones of Putjatin Island (Japan Sea).
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STuXBERG, A.
1882. Evertebratfaunan i Sibiriens Ishaf, vol. 1, pp. 677-812, pl. 15. In
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1887. Faunan p& och kring Novaja Semlja, vol. 5, 239 pp. In Norden-
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Sumner, F. B., Ospurn, R. C., and Cots, L. J.
1911 (1913). A biological survey of the water of Woods Hole and vicinity.
Part 1, sect. 1: Physical and zoological, pp. 11-441, 225 maps;
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SUNDARA Rag, B.
1927. Suborder Caprellidea (Laemodipoda). The littoral fauna of Krusadai
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TATTERSALL, W. M.
1913. Clare Island Survey. Part 42, Amphipoda. Proc. Roy. Irish Acad.,
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CAPRELLIDAE OF WESTERN NORTH ATLANTIC 139

TEMPLETON, R.

1836. Descriptions of some underscribed exotie Crustacea. Trans. Entomol.
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TuHompson, D’Arcy W.

1901. A catalogue of Crustacea and of Pycnogonida contained in the
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THompson, W.

1844, Additions to the fauna of Ireland. Ann. Mag. Nat. Hist., vol. 13,

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THomson, G. M.

1878 (1879). New Zealand Crustacea, with descriptions of new species.
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1879. Additions to the amphipodous Crustacea of New Zealand. Ann.
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1913. The natural history of Otago Harbour and the adjacent sea, together
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TuHomson, G. M., and Cuitton, C.

1885 (1886). Critical list of the Crustacea Malacostraca of New Zealand.

Op. cit., vol. 18, pp. 141-159 [Caprellidae, pp. 141-142].
TuHomson, G. M., and ANDERTON, T.

1921. History of the Portobello Marine Fish-Hatchery and Biological
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Ticuy, M. J.

1911. Zamietka o Caprellidae Chernago Moria [in Russian]. Bull. Acad.

Imp. Sci. St. Petersburg, ser. 6, no. 16, pp. 1125-1134.
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1964. Inventaire de la faune marine de Roscoff. Amphipodes-Cumacés.
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Tuzet, O., and SancueEz, 8S.

1952. Sur lappareil génital male et la spermatogenese de Caprella acutifrons
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Ua.er, P. R.

1879. List of animals observed at Fort Wool, Va. Chesapeake Zool. Lab.
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Ussina, H.

1952. Havedderkoppen (Nymphon grossipes) og Caprellen (Proto pedata).

Flora og Fauna, year 58, nos. 1 and 2, pp. 45-47.
Utinomi, H.

1943a. Caprellids obtained in Onagawa Bay, northern Japan. Sci. Rep.
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1943b. Report of the biological survey of Mutu Bay. 37. Caprellids from
Asamusi. Ibid., pp. 281-287.

279-475—68——10

140 U.S. NATIONAL MUSEUM BULLETIN 278

1943c. The fauna of Akkeshi Bay XIII. Caprellidea. Journ. Fac. Sci.
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1947. Caprellidae of Japan and adjacent waters [in Japanese]. Seibutu
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VANHOFFEN, E.
1897. Die Fauna und Flora Grénlands. Grénland Exped. Gesellsch.
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VERRILL, A. E.
1873. Report upon the invertebrate animals of Vineyard Sound and ad-
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VOSSsELER, J.
1889. Amphipoden und Isopoden von Spitzbergen. Arch. Naturgesch.,
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WaGne_R, N.
1885. Die wirbellosen Thiere des weissen Meeres .. . Leipzig. vol. 1, 171
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WALKER, A. O.
1895a. Revision of the Amphipoda of the L.M.B.C. District. Proc. Trans.
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1895b. The Amphipoda of Bate and Westwood’s “British sessile-eyed
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CAPRELLIDAE OF WESTERN NORTH ATLANTIC 141

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index

(Page numbers of principal entries in italies)

abracadabra, Deutella, 86
acanthifera, Caprella, 97
acaudata, Phthisica, 93
Phtisica, 93, 94
Phtysica, 93
acuminifera, Caprella, 31
acutifrons, Caprella, 1, 2, 19, 21, 23,
33-34, 37
Aegina echinata, 13
laevis, 13
langicornis, 14
longicornis, 13
longicornis nodosa, 13, 14
longicornis spinifera, 13, 14
longicornis spinigera, 14
longicornis spinosissima, 14
longicornis typica, 13
spinifera, 13
spinosissima, 13
Aeginella, 8, 111
longicornis, 14

Astyris, 30, 101
Atomos, Cancer, 33
attenuata, Caprella, 40
bermudia, Caprella, 22, 103, 107
biscaynensis, Fallotritella, 7, 57, 68-61.
100, 104, 105, 107
Blennius, 30, 97, 101
borealis, Caprella acutifrons, 38
Brisinga, 13, 52
brunneovittata, Proto, 93
Bugula, 25
californica, Caprella, 44
Caprella scaura, 44
Deutella, 7, 53, 54, 104, 107
Callionymus, 97, 101
camana, Podalirius typicus, 89
Cancer Atomos, 33
linearis, 18, 30
pedatus, 92
rubra, 92
ventricosus, 92

spinosa, 6, 8-13, 52, 103, 106, 107} Caprella, 3, 4, 14, 18-19, 101

spinosissima, 13
tristanensis, 100
Aeginina, 8, 13
longicornis, 2,
106, 107
aequilibra, Caprella, 25-26
africana, Metaprotella, 82
Ampelisca, 102
Amphithoe, 30
andreae, Caprella, 2, 7, 19-22, 97, 101,
106, 107, 108
Caprella acutifrons, 19, 38
angusta, Caprella, 35, 38, 39
Caprella acutifrons, 35, 38
antiguae, Pseudaeginella, 6, 100, 103,
107
antillensis, Phtisica, 7, 65, 89-91, 93,
94, 95, 106, 107
Proto, 89
Arbacia, 40, 52
Asterias, 33, 52, 97
forbesi, 52, 100
vulgaris, 52, 100

6, 11, 13-18, 104,

acanthifera, 97
acuminifera, 31
acutifrons, 1, 2, 19, 21, 23, 33-34,

37
acutifrons andreae, 19, 38
acutifrons angusta, 35, 38
acutifrons borealis, 38
acutifrons carolinensis, 34, 38
acutifrons cristibrachium, 38
acutifrons gibbosa, 34, 38
acutifrons incisa, 38
acutifrons lusitanica, 34, 38
acutifrons minor, 38
acutifrons natalensis, 34, 38, 39
acutifrons neglecta, 34, 38, 39
acutifrons porcellio, 34, 38
acutifrons simulatrix, 38
acutifrons tabida, 34, 38
acutifrons testudo, 35, 38
acutifrons tibada, 35, 38
acutifrons typica, 38
acutifrons verrucosa, 38
acutifrons virginia, 34, 38

143

144 U.S.

Caprella—Continued
aequilibra, 25-26
andreae, 2, 7, 19-22, 97, 101, 106,

107, 108
angusta, 35, 38, 39
attenuata, 40
attenuata subtenuis, 40
bermudia, 22, 103, 107
californica, 40
carolinensis, 35
caudata, 26
cercopoides, 45
cornuta, 40
cornuta obtusirostris, 40
danilevskii, 7, 22-25, 107, 108
danilewskii, 22
dilatata, 38
equilibra, 8, 22, 25-30, 35, 39, 100,
101, 107, 108
Esmarkii, 25
geometrica, 34, 39
grahami, 49, 50
hystrix, 45
hystryx, 45
inermis, 22
Januarii, 25
laevipes, 44
laevis, 31
laticornis, 25
linearis, 2, 8, 26, 30-33, 47, 48, 49,
52, 103, 107
linearis distalis, 31
linearis gullimarensis, 31
lobata, 31
longicornis, 45
longimana, 33
longimanus, 33, 40
lovéni, 45
megacephala, 26
mendax, 26
monacantha, 26
monocera, 45, 48
nigra, 83
nodosa, 40
novae-zealandiae, 34, 39
obesa, 26, 34
obtusa, 26
obtusifrons, 38
penantis, 7, 29, 33-40, 52, 100, 102,
103, 104, 107
penantis natalensis, 35
penantis porcellio, 35
Pennantii, 34

NATIONAL MUSEUM BULLETIN

278

Caprella—Continued
phasma, 31
pilimana, 38
punctata, 31, 45
robusta, 45
sanguinea, 40
scaura, 3, 7, 40-44, 52, 107, 108
scaura californica, 44
scaura cornuta, 40, 41
scaura diceros, 40, 41
scaura hamata, 40, 41, 44
scaura scauroides, 44
scaura spinirostris, 44
scaura typica, 40, 41, 44
scolopendroides, 31
septentrionalis, 8, 33, 40, 44-49,
103, 107
septentrionalis longicornis, 45, 48
septentrionalis lovéni, 45
septentrionalis monocera, 45
septentrionalis nodigera, 45
septentrionalis parva, 45
septentrionalis polyceros, 45
septentrionalis punctata, 45
septentrionalis spinigera, 45
septentrionalis stimpsoni, 45
septentrionalis typica, 45
spinifera, 13
spinosissima, 13
stimpsoni, 45
tabida, 38
ultima, 26
unica, 7, 33, 49-52, 100, 107
ventricosa, 92
verrucosa, 38
caprellinoides, Neoxenodice, 110
Caprogammarus, 111-112
eurjanovae, 110
carolinensis, Caprella, 35
Caprella acutifrons, 34, 38
caudata, Caprella, 26
Centropristis, 16, 30, 101
cercopoides, Caprella, 45
Cercops, 107, 111-111
Chelonia, 21, 97
cornuta, Caprella, 40
Caprella scaura, 40,41
cristibrachium, Caprella, acutifrons, 38
cumana, Pariambus typicus, 89
Podalirus typicus, 89
Cystoseira, 25
danae, Metaprotella, 82
danilevskii, Caprella, 7, 22-25, 107, 108

INDEX

danilewskii, Caprella, 22
Deutella, 4, 6, 53-54, 68, 82
abracadabra, 86
californica, 7, 53, 64, 104, 107
incerta, 53, 68
mayeri, 7, 53, 64-57, 104, 107
venenosa, 53
diceros, Caprella scaura, 40, 41
dilatata, Caprella, 38
distalis, Caprella linearis, 31
Dodecas, 5
echinata, Aegina, 13
elongatus, Proto, 92
equilibra, Caprella, 8, 22, 25-30, 35, 39
100, 101, 107, 108
Esmarkii, Caprella, 25
Eugastraulax, 4
excentrica, Metaprotella, 82
Fallotritella, 57, 100
biscaynensis, 7, 57, 58-61, 100, 104
105, 107
forbesi, Asterias, 52, 100
Gadus, 48, 97, 101
Gammarus pedatus, 93
quadrilobata, 30
geometrica, Caprella, 34, 39
gibbosa, Caprella acutifrons, 34, 38
Goodsirii, Proto, 92, 93
grahami, Caprella, 49, 50
gullimarensis, Caprella linearis, 31
gurjanovae, Caprogammarus, 110
hamata, Caprella scaura, 40, 41, 44
haswelliana, Metaprotella, 82
Protella, 78
Hemiaegina, 5, 61
minuta, 7, 61-64, 101, 102, 107
quadripunctata, 62
Hemiproto, 3, 66
wigleyi, 7, 65-68, 102, 104, 107
hummelincki, Metaprotella, 7, 78-82,
102, 107
hystrix, Caprella, 45
hystryx, Caprella, 45
incerta, Deutella, 53, 68
Luconacia, 7, 53, 68-72, 101, 104,
107
incisa, Caprella acutifrons, 38
inermis, Caprella, 22
frrorata, Littorina, 105
Jassa, 30
laevipes, Caprella, 44

145

laevis, Aegina, 13
Caprella, 31
Tritella, 57
langicornis, Aegina, 14
laticornis, Caprella, 25
Leptogorgia, 29, 35
Leptomera pedata, 92, 93
ventricosa, 92
Libinia, 40, 89
limicola, Mayerella, 7, 738-75, 78, 102,
103, 107
linearis, Cancer, 18, 30
Caprella, 2, 8, 26, 30-33, 47, 48, 49,
52, 103, 107
Littorina irrorata, 105
lobata, Caprella, 31
Squilla, 30, 44
longicornis, Aegina, 13
Aeginella, 14
Aeginina, 2, 6, 11, 13-18, 104, 106,
107
Caprella, 45
Caprella septentrionalis, 45, 48
longimana, Caprella, 33
longimanus, Caprella, 33, 40
lovéni, Caprella, 45
Caprella septentrionalis, 45
Luconacia, 4, 6, 53, 54, 68
incerta, 7, 53, 68-72, 101, 104, 107
lusitanica, Caprella acutifrons, 34, 38
Maja, 97
makrodactylos, Metaprotella, 82
marina, Phthisica, 93
Phtisica, 7, 52, 89, 91-97, 101, 104,
106, 107, 109
Phytisea, 93
Mayerella, 4, 72-73, 78
limicola, 7, 73-75, 78, 102, 103, 107
redunca, 5, 7, 73, 75-78, 102, 107
mayeri, Deutella, 7, 53, 54-67, 104, 107
megacephala, Caprella, 26
mendax, Caprella, 26
Metacaprella, 101
Metaprotella, 2, 78, 82
africana, 82
danae, 82
excentrica, 82
haswelliana, 82
hummelincki, 7, 78-82, 102, 107
makrodactylos, 82
problematica, 82
sandalensis, 82
minor, Caprella acutifrons, 38

wo an

7 a

LENIN

3 9088 0