Bulletin - United States National Museum

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SMITHSONIAN
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MUSEUM
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NATURAL
HISTORY

a, Uschakoviella echinophora Gurjanova [after Shoemaker, 1964].

b, Stilipes distincta Holmes [after Shoemaker, 1964].

UNITED STATES NATIONAL MUSEUM BULLETIN 271

The Families and Genera
of Marine Gammaridean

Amphipoda

J. LAURENS BARNARD

Associate Curator of Crustacea

SMITHSONIAN INSTITUTION PRESS
CITY OF WASHINGTON
1969

Publications of the United States National Museum

The scientific publications of the United States National Museum include two
series, Proceedings of the United States National Museum and United States National
Museum Bulletin.

In these series are published original articles and monographs dealing with
the collections and work of the Museum and setting forth newly acquired facts
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of each publication are distributed to libraries and scientific organizations and
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The Proceedings, begun in 1878, are intended for the publication, in separate
form, of shorter papers. These are gathered in volumes, octavo in size, with
the publication date of each paper recorded in the table of contents of the volume.

In the Bulletin series, the first of which was issued in 1875, appear longer,
separate publications consisting of monographs (occasionally in several parts)
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either octavo or quarto in size, depending on the needs of the presentation.
Since 1902, papers relating to the botanical collections of the Museum have been
published in the Bulletin series under the heading Contributions from the United
States National Herbarium.

This work forms number 271 of the Bulletin series.

FraNK A. TAYLOR
Director, United Staies National Museum

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Contents

Introduction .

Acknowledgments P
Status of gammaridean ayctementies i
Morphological terminology .

Morphology of a gammaridean aapEinod
Geographic distribution of marine Gammaridea .
The composition of a faunule ig
Morphological evolution of the Asnatnipyades
Interfamilial gammaridean relationships .
Limnetic evolution of the Gammaridea
Identification procedures

Diagnosis of the basic marine ganmmaridean (Gene a marine Carmmeridson

Order Amphipoda :
Key to the suborders of Avaninporal ; i
Nomenclatural changes in gammaridean orenfTas :
Key to the families of the suborder Gammaridea .
Acanthonotozomatidae
Ampeliscidae .
Amphilochidae .
Ampithoidae .
Anamixidae
Aoridae .

Argissidae .
Astyridae

Atylidae .
Bateidae.
Beaudettiidae
Calliopiidae
Cheluridae
Colomastigidae .
Corophiidae .
Cressidae
Dexaminidae .
Dogielinotidae .
Eophliantidae
Eusiridae
Gammaridae (Goesaing) .
Haustoriidae . A.
Hyalellidae and Efvalidiae
Hyperiopsidae .
Isaeidae .
Ischyroceridae .
Kuriidae

al CONTENTS

Lafystiidae
Laphystiopsidae
Lepechinellidae .
Leucothoidae
Liljeborgiidae
Lysianassidae
Melphidippidae
Ochlesidae .
Oedicerotidae
Pagetinidae
Paramphithoidae .
Pardaliscidae
Phliantidae Geueed)
Phoxocephalidae
Pleustidae .
Podoceridae 4
Prophliantidae evincaye
Sebidae . ‘
Siecocepralrdne:
Stenothoidae .
Stilipedidae
Synopiidae (revised)

Superfamily Talitroidea (@auanda Be atelndae. iTalitnidae)

Thaumatelsonidae
Vitjazianidae
Incertae sedis
Supplement
Literature cited ae
Appendix I. Dissection of an i anpaipodi
Appendix II. Amphipod analytical sheet .
Appendix III. Glossary of special terms ‘
Index and familial assignment of gammaridean penera :

Page
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387
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432
435
436
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451
454
463
473
476
478
481
482
507
512
520
524

The Families and Genera of Marine Gammaridean

Amphipoda

Introduction

Keys and diagnoses, with illustrations, to the families and genera
of marine gammaridean Amphipoda are presented here in the form
of a handbook. Since 1906, when Stebbing monographed this suborder,
the number of families has been increased by 25 percent, the number
of genera by 200 percent and the number of species by nearly 200
percent (J. L. Barnard, 1959d). Stebbing’s general arrangement of
families and genera remains substantially intact, but the discovery of
many additional species has required a considerable emendation of
familial and generic limits. The near absence of phyletic monographs
and the plethora of faunal compilations widely scattered in the litera-
ture have hindered advances in classification of the group. Ready
identification of taxa not only by nonspecialists but even by experts
has become extremely difficult because of the lack of a modern focal
point.

The present work is at best a stopgap. Specialists recognize that
much exploration and discovery remain to be accomplished in the
Amphipoda (witness the numerous new haustoriid taxa discovered by
Bousfield, 1965). No specialist is nearly satisfied with current classifi-
cation at familial levels. Many genera are extremely confused, not
only for the lack of clear description of their type-species but for the
obviously erroneous inclusion of many of their congeners. Few studies
on population variability have been published and the criteria to be
applied at generic and familial evels have rarely been tested even by
dialectical means.

No radical changes in classification are proposed herein, but numerous
problems are noted. The purpose of this paper is to assemble the
families and genera within the previously arranged general scheme
and to provide rapidly usable (and thus simplified) means for their
identification. In this way the keys coincide with the published
literature and may be of greater help than would be a brief, completely
revised system, which might not stand the test of time. Hopefully
this paper will encourage recruitment of new taxonomists and relieve
some of the burdens of identification now overwhelming the few active

1

2 U.S. NATIONAL MUSEUM BULLETIN 271

specialists. Perhaps it will stimulate monographic studies of families
and genera. It may be of greatest use to those students who must
make generic determinations of their material without adequate
library facilities; it should restrict bibliographic needs to small numbers
of publications which can be borrowed more easily than can the entire
literature. An index to the species of Gammaridea (J. L. Barnard,
1958a) may be of some help. An expanded and more useful index is
now in preparation.

Brief sections on distribution, faunules, and evolution are presented
in hopes of stimulating further inquiry into these fascinating but poorly
studied topics. Our knowledge of anatomy (especially musculature),
functional morphology, behaviour, food habits, indeed general ecology
is rudimentary.

The identification system proposed herein requires familiarity with
a basic gammaridean plan similar to that exemplified by some members
of the Gammaridae. These gammarids may be the most primitive
grade of living Amphipoda. A knowledge of their structure is vital to
an understanding of the unfolding of the complex network of speciali-
zations found in other gammaridean families. The lines of evolution
from gammarid-like ancestors, so ill understood, are not only manifold
but they confuse identification procedures. At times the specialist is
confronted with the feeling that most of the ‘missing links” in
Amphipoda are still alive. Only a few families are so remote morpho-
logically from the Gammaridae that they are instantly recognizable.
With a few exceptions the specialization of gammaridean families is
a result of the loss of morphological complexity.

The basic gammaridean is illustrated in figures 1 and 2 and described
on p. 50. Families other than the Gammaridae are provided with
short diagnoses which include only definitive and exclusive differences
from the basic gammaridean. Some of these diagnoses are identical
among several families. Those taxa are further distinguished by
intercomparison of their descriptions and by reference to paragraphs
on relationship. Part of each diagnosis is a reference to a related (or
superficially similar) family.

Diagnostic characters of the families are illustrated in a group of
boxes (figs. 3-53). Instructions for identification procedures using
these tools are presented in following pages.

Acknowledgments

This work was compiled during the writer’s tenures at the Allan
Hancock Foundation, the Beaudette Foundation of California, and
the Smithsonian Institution, to which institutions and their adminis-
trative heads my gratitude is expressed. Mrs. Dorothy M. Halmos of

MARINE GAMMARIDEAN AMPHIPODA 3

the Hancock Library and Mr. Jack Marquardt of the Smithsonian
Institution have been very helpful in obtaining references. Literature
published subsequent to December 1965 has not been incorporated.
The figures were redrawn largely from the literature and some of them
have been simplified as a means of providing standardization. Four
illustrators have assisted with the work over the years, Mr. L. R.
Hales, Mrs. D. McLaughlin and, especially, Miss Jacqueline M.
Hampton of the Beaudette Foundation; and Miss Naomi D. Mano-
witz of the Smithsonian Institution. They were partially supported
by grants from the National Science Foundation.

Dr. D. E. Hurley of New Zealand Oceanographic Institute has
assisted my thinking with regard to the difficult family Lysianassidae
by letting me read his various manuscripts concerning the classifica-
tion of that family. Dr. E. L. Bousfield of the National Museum of
Canada kindly loaned the paratype of Oldevig’s Haustorioides mun-
sternjelmi for this study. Numerous colleagues have helped with
friendly advice and encouragement and among those must be men-
tioned the foregoing plus Dr. Fenner A. Chace, Jr., Dr. Thomas E.
Bowman, and Dr. Olga Hartman. Dr. Richard Cowan, Dr. Donald
Squires, Dr. I. E. Wallen, and Dr. Raymond B. Manning have created
the ideal environment at the Smithsonian Institution for this work
to be brought to its conclusion. Portions of the final draft were com-
posed while the writer worked as a Smithsonian fellow to Bishop
Museum, Honolulu; and I must thank Dr. Roland W. Force, Director,
for providing the facilities and help of that Museum.

This paper is dedicated to Mr. Clarence R. Shoemaker (1876-1958),
America’s foremost student of Amphipoda. The frontispiece represents
two of the beautiful drawings he did.

Status of Gammaridean Systematics

More than 3300 species in about 670 genera are known currently
in the Gammaridea.* The rate of description of new species has
increased recently and will presumably exceed the average descrip-
tion of 40 per year that occurred between 1906 and 1956. Outside
of the littoral north Atlantic, Arctic U.S.S.R., and the northwestern
Pacific, knowledge is highly incomplete. The magnificent work of
Sars (1895), supported by that of Chevreux and Fage (1925), brought
the west European fauna into definition at an early stage. Unfor-
tunately, the excellent taxonomic status of north Atlantic amphipods

*About 130 genera and 850 species occur in nonmarine environments and most
are not included in this handbook.

4 U.S. NATIONAL MUSEUM BULLETIN 271

has not been exploited fully in making advances in ecological, physio-
logical, and genetical studies, although most of those special studies
so far undertaken have been pursued in the northeastern Atlantic.

Intensive work has been accomplished in antarctic seas but the
fauna there needs to be placed on a Sarsian basis with full illustra-
tions and descriptions. The works of Chevreux (1906c, 1912b) are
notable for that region, but numerous sticky problems resulting from
other studies have not been clarified there.

The remaining faunas are sketchily known. An imbalanced em-
phasis on the deep-sea is occurring in this decade, but this has helped
bring into focus many problems in the higher taxa of amphipods.
Since World War II research on the marine species has been meager,
because more than 90 percent of the printed pages have been pro-
duced by fewer than 10 taxonomists (Zoological Record, years
1953-60).

Morphological Terminology

The taxonomic assessment of gammaridean morphology is almost
always stated in subjective terms: in words and phrases which,
through experience, taxonomists have come to understand and
visualize easily, but which are difficult to convey with mensurative |
precision. There is no current solution to this difficulty but taxono- —
mists may eventually be able to compile an illustrated handbook
in which all possible shapes and relative proportions can be figured,
named, and coded. Identifications may then be amenable to mechan-
ical analysis. Meanwhile one must work with terms such as “strong,
large, small, feeble, weak, minute, elongate, shortened” and various
adjectives with the modifier ‘sub,’ such as “subquadrate”’ and
“Subacute”. I have used these terms everywhere in this handbook,
but have attempted to provide illustrations as a demonstration of
their extent, for the degree varies in the different families. For in-
stance, gnathopods of the genus Apherusa are ‘feeble’ in com-
parison to those of its familial congener Calhiopius, but they are
scarcely as feeble as the gnathopods of the Acanthonotozomatidae.
The term ‘subacute’ apparently has come to mean a shape that
has the overall appearance of sharpness but which terminally is
softly rounded. The term ‘‘subconical’”’ apparently refers to a 2-
dimensional, rather than a geometric cone but the adjective is not
necessarily synonymous with “subacute,” for a subconical process
may have a subacute apex. The term ‘“quadrate”’ has often been
applied to a squared-off shape of an otherwise imperfect rectangle
or in simpler fashion as reference to one pair of sides occurring at
right angles to each other. ‘SSubquadrate”’ should refer to sides not

MARINE GAMMARIDEAN AMPHIPODA 5

precisely at right angles to each other but also seems to have reference
to a quadrate shape with softly rounded corners.

The terms ‘‘vestigial’” and “rudimentary” have their uses, for a
mathematical definition in each case might be more involved, con-
fusing, and time consuming than is the process of learning these
conditions through trial and error.

A decision as to whether a gnathopod is simple or subchelate
is occasionally difficult to make and definitions to classify all border-
line cases are almost impossible to compose. There are no terms to
describe all of the stages between fully subchelate and fully simple
and gammaridean students may eventually encounter difficulties
arising from dogmatic use of such all-or-none terms. For instance,
two subfamilies of Lysianassidae have been described which are
partially based on a subchelate or simple gnathopod 1, but there are
several borderline cases and several genera in one subfamily which
seem to have stronger affinities with genera in the other subfamily
than with members of their own group. The contrast between the
concepts of cleft telson and entire telson is far too strong to describe
all intermediate stages and the two terms tend to condition the
human observer to a reliance on the alternatives as highly significant,
whereas numerous exceptions to this significance are apparent [e.g.,
Kusiridae and Calliopiidae].

Names for the gammaridean abdomen and its parts are not stand-
ardized. All six segments of the abdomen are frequently called the
“pleon’’, but “pleonites’’ 4-6 are often called ‘“urosome” and num-
bered as “urosomites’ 1-3. Pleonites 1-3 are rarely called the
“‘metasome.”’

Shape and proportion in seemingly infinite array are thus more
important to the taxonomist than are qualitative and numerical
expressions, but one day we may be able to convert shape and pro-
portion to precise formulations.

Morphology of a Gammaridean Amphipod

Amphipods, lke tanaids and isopods, lack a carapace covering
the thorax, so that seven definitive thoracic segments (pereonites)
are visible (as in other Peracarida the first thoracic segment with its
appendage [maxilliped] has become fused to the head). Gammaridean
and hyperiidean Amphipoda may be recognized by their possession
of three pairs of pleopods (swimmerets) and two or three pairs of
uropods on the pleon (abdomen). The consistent presence of at
least six pairs of thoracic appendages, five-plus pairs of gills and
four pairs of brood lamellae in females are definitive characters of
Gammaridea and Hyperiidea.

6 U.S. NATIONAL MUSEUM BULLETIN 271

The pereon (thorax) almost invariably bears seven pairs of legs.
The first two pairs are called gnathopods (or gamopods) and usually
are prehensile, having the seventh article (dactyl) folded back on the
sixth article (propodus, hand, or palmar article). Rarely in gnathopod
1 is there sexual dimorphism but the male often has greatly enlarged
second gnathopods. Gnathopod 2 is believed to be used primarily for
erasping the female during copulatory amplexus. The male mounts the
dorsal side of the female, projects the gnathopods around her body and
hooks them into her fifth coxae. The pair of animals then swims, darts
among algae, or rests until the female molts (possibly as much as three
days after amplexus), at which time the male emits spermatophores
(sacs of spermatozoa) that pass from the ventral side of his seventh
pereon segment into the pouch formed by the female brood lamellae.
Ecdysis of the female is taking place during this time period and im-
mediately thereafter she lays eggs through two genital pores on the
sternite of thoracic segment 5. The pores normally are so small and
heavily chitinized that the eggs cannot be laid until ecdysis occurs and
the pore openings become soft and pliable. How the spermatophores
pass from male to female is unknown but they may be assisted by
gnathopods or pereopods. The ventral side of pereonite 7 in the male
bears two minute penial projections, often spinose. Occasionally they
are hidden by small gills attached to the coxae of this segment.

Within a genus of gammaridean amphipods, the taxonomic recogni-
tion of species often depends on the shape of male gnathopod 2; hence,
it is difficult to identify females specifically because taxonomists have
not studied minute differences in females and made them basic to
identification. The second gnathopods of juveniles and females often
are alike; during maturation the male second gnathopods commence an
an increase in size and a morphological differentiation, with changes
taking place during each instar, even long after the attainment of
sexual maturity. This has resulted occasionally in taxonomic confusion
because some of these instars have been described as distinct species.
A few species are known to have radically distinct phenotypes espe-
cially in the terminal male. The ubiquitous Jassa falcata has dozens of
forms, some of these even bridging the concepts of two or more genera.

All thoracic appendages have seven articles (segments), the
proximal member of which is the coxa or sideplate. A few gammarideans
(e.g., Bateidae) have reduced numbers of articles on various thoracic
appendages. Coxae are of greater taxonomic importance in gammar-
idean Amphipoda than in other Malacostraca and in many species
are structurally more an integral part of the trunk than simply an
article of the appendage. They resemble ventral pleuron-like extensions
of segments and so contribute to the appearance of lateral compression
in the body plan. They are numbered from one to seven, with numbers

MARINE GAMMARIDEAN AMPHIPODA 7

1 and 2 belonging with gnathopods 1-2 and numbers 3-7 with
pereopods 1-5.

Gills are thoracic and generally are attached to the medial surfaces
of coxae 2-7, occasionally only on 2-6, or ?3-6. In females the medial
surfaces of coxae 2—5 (?or 2-6) carry brood lamellae. They are simply
buds in young females, but as body growth proceeds they become
longer and more heavily setose and are interlocked by their setae to
form a cradle enclosing the eggs. As the female increases in age and
size, the number of eggs laid after each molt becomes larger. Some
amphipods lay such large yolky eggs that a young female can carry
only one of them; at maturity she may be able to carry three or four.
Not all of the extruded eggs hatch; mortality of about 25-50 percent
may occur even before hatching; stunted degenerating eggs often may
be seen in the broods (as well as commensals such as spherical copepods
and reniform ostracods). Some large Amphipoda lay and carry more
than 200 eggs at a time.

Commencing with leg 3, the thoracic appendages are termed pereo-
pods (walking legs), so that five pairs of thoracic legs represent
pereopods (many specialists commence numbering pereopods with
enathopod 1). There is justification in distinguishing gnathopods by
name because of their specialization in analogy to the pereopods of
other Crustacea (e.g., posterior maxillipeds of decapods). The first
two pairs of pereopods are useful in cleaning the gnathopods and other
anterior appendages and as a balance when alighting from a swim.
The last three pairs appear rather immobile and less adapted for
walking than in isopods. Amphipods are poorly balanced for walking,
hence their mobility usually depends on swimming. Their body plan,
however, permits a motility through dense masses of hydroids and
algae. Perhaps this is one of the reasons why amphipods are one of
the most abundant macroscopic crustacean groups in algae and other
anastomoses.

Some Amphipoda living on the sea-bottom have immensely elon-
gated pereopods, which are spread out in the fashion of a spider and
prevent the amphipod from sinking into the mud. To lower the center
of gravity the body of the amphipod hangs upside down in its cradle
of legs.

A few families of Gammaridea have the habit of burrowing into
benthic sediments and their pereopods are armed densely with strong
spines which aid in burrowing. Elongate flexible setae, as well as stiff
spine-like setae may occur on pereopods 3-5 of fossorial Amphipoda.
Even though a few other nonfossorial Amphipoda, such as Ampelis-
cidae, have these elongate setae, the term “fossorial pereopods’’ is
usefully applied to the condition. The glossary contains a precise
definition of fossorial pereopods.

8 U.S. NATIONAL MUSEUM BULLETIN 271

Six or seven families of amphipods construct dwelling tubes. The
cylindrical or flattened tubes are spun from strands of material
secreted and probably manipulated by pereopods 1-2. The tubes may
be limp (Ampelisca) and lay prone on the bottom or they may be
stiff and erect and attached to rocks (Photis). Mud is occasionally
used to reinforce the walls of the erect tubes.

Glands of domiciliary Amphipoda appear to be concentrated mainly,
if not exclusively, in pereopods 1-2. They are found most heavily
concentrated in article 2, often in article 4 and occasionally in other
articles. Glands usually appear to be composed of densely packed,
“volky”’ tissue, often posterior to the main muscles of article 2, often
composed of morula-like bodies, and often of different color than the
muscles or other tissues, even in specimens preserved in alcohol; they
are frequently yellow, ochre, orange, or purple. Apparently a duct
carries the secretion to a meatus located subterminally on the dactyl.
The meatus is very difficult to see and not definitive of Gammaridea
known to be domiciliary because many other Amphipoda seem to
have a dactylar meatus but lack conspicuous glands. The presence of
these glands may be a useful clue by assisting in the identification of
several genera in Isaeidae, Ischyroceridae, Corophiidae, and Am-
pithoidae that otherwise resemble nondomiciliary Amphipoda. Com-
plete reliance cannot be made on them as familial characters, for all
members of domiciliary familes do not necessarily have glandular
pereopods 1-2 and very few species have actually been examined for
their presence. Some phoxocephalids, haustoriids, and argissids have
conspicuous glands in pereopods 3-5.

Most domiciliary amphipods except for the Ampeliscidae have
somewhat depressed bodies, shortened pereopods, and have better
crawling ability than do nontube builders. They can be observed
emerging from their closely crowded tubes, their antennae appar-
ently being used digitally in a search for particles of food. Ampeliscid
amphipods lie upside down in their tubes and project their strongly
setose antennae as filtering organs. Only one benthic gammaridean has
been reported as a predator but some pelagic amphipods catch prey.

After hatching, young amphipods are like adults (Gammaridea do
not have larvae as do most crustaceans) and are carried about for
a few hours or days in the brood pouch. In the early stages molting
and growth are rapid; a young amphipod may first molt while still
in the brood pouch, within a day or two after hatching. As growth
proceeds rates of molting and growth decrease, so that adults may
molt every 20-30 days and in some striking instances only every
six months or so. Few amphipods have been studied for their molting
rates; of those examined it has been determined that sexual ma-
turity is reached at about the sixth molt (commencing the seventh

MARINE GAMMARIDEAN AMPHIPODA 9

instar); the animals are fertile yet the secondary sexual characters
are rudimentary; male gnathopods may be poorly developed and
brood plates are just beginning to develop in the female. Amphipods
are known to live through at least 12 instars, the females laying a
brood of eggs during the last five or six or in alternative instars; how-
ever, terminal adult females (gerontics) may lose their brood plates,
apparently fail to lay eggs and develop aberrencies of an andromorphic
nature. Occasionally, these stages have been described as distinct
species.

Amphipoda have the unfortunate habit of eating their exoskeletons
after ecdysis so that it is difficult to trace their molting sequence in
the laboratory. One must watch them continuously in order to obtain
ecdysial casts. Because the average instar appears to last about 15
days, the average maximum length of life is expected to exceed 6
months but some species in polar regions are estimated to live 5 or
6 years. Often the structure of the next instar may be seen within an
appendage. Such replication within the present organ can be confusing
to taxonomists and lead to the description of supernumerary parts.

A few species are known to be hermaphroditic, carrying both male
and female gonads, and reflecting both sexes in their secondary sexual
characteristics.

Paired pleopods on the first three segments of the pleon are biramous,
the rami multisegmented and strongly setose. Minute coupling hooks
on the medial edges of the peduncles are used to engage the pairs of
pleopods for coordinated paddling. Amphipoda usually are good
swimmers. Even burrowing amphipods swim well and phoxcephalid
males have the habit of leaving their burrows at night and swimming
to the seasurface from depths as great as 100 meters. They will swarm
around a light suspended in the water. They may be ascending in
search of females even though the latter rarely swim to a night-light.
Such swimming behavior may be a dispersal mechanism, especially
in groups having a low proportion of males. Some pelagic species
apparently undergo great vertical migration; several deep-sea amphi-
pods caught at night near the surface have been found to have ali-
mentary tracts full of benthic sediments.

Variation in pleopods is rarely of sufficient extent to be weed in
generic or familial definitions except for Phliantidae, Talitroidea and
some Corophiidae. Pleopodal morphology, nevertheless, has been
neglected and may afford some help in taxonomic distinctions.

There is justification in restricting the term pleon (=metasome) to
the first three abdominal segments bearing pleopods and utilizing the
term urosome for the last three abdominal segments bearing uropods.
“All” Malacostraca have at least one pair of uropods, that pair of
appendages on abdominal segment 6. Malacostracans generally have

10 U.S. NATIONAL MUSEUM BULLETIN 271

five pairs of pleopods on segments 1—5, but in noncaprellidean amphi-
pods the appendages of segments 4 and 5 resemble the terminal
uropod and thus are called uropods 1 and 2. Uropods in many mala-
costracans are still used for swimming although in different fashion
from pleopods, but uropods 1-2 in Amphipoda appear to be used
primarily for strengthening the caudal portion of the body to permit
jumping or flipping by rapid flexion of the urosome. In many Gam-
maridea, the third uropods still bear “‘swimming”’ setae, and may
be used for paddling or as rudders. Males especially have natatory
third uropods. But the vast majority of Gammaridea probably do
not use the third uropods for active swimming and they are often
reduced or occasionally absent in sedentary species. Caprellidea have
lost all but a vestige of the abdomen and its appendages.

The telson is a flap attached to the sixth pleonite above the anus.
It is of primary taxonomic value, depending on whether it is cleft
into two lobes, fused into a single flap, elongate, fleshy, or ornate.
Possibly, it is a vestige of paired appendages.

The head bears two pairs of antennae. The first three articles
of the first pair are known as the peduncle, the remaining smaller
articles the flagellum. In many species an accessory flagellum demon-
strates the biramous derivation of the appendage; when present,
it sprouts from the end of the third peduncular article and may be
elongate or reduced to three, two, or one articles. Although appearing
to be of minor importance, the condition of the accessory flagellum is
crucial to amphipod systematics and is useful especially at familial
and generic levels. The second antennae bear five peduncular articles,
followed by a single flagellum. The flagella of both antennal pairs
may bear, especially in males, sensory appendages, such as aesthetascs
and calceoli. Male antennae often are longer than those of females.

Frequently families and genera have been defined as lacking
accessory flagella and then have been shown to have some members
bearing extremely small 1l-articulate pieces. Allowance should be
made by the observer so as to admit to genera and families those
species with microscopic remnants of accessory flagella that have
heretofore been overlooked.

The mouthparts are composed of the following structures; they
are highly variable intergenerically and their morphology is important
for classification:

Upper lip: A single lobe or flap anterior to the mouth. In about
10 percent of known species the anterior cephalic surface above the
upper lip is produced into a point, keel, or lobe known as the epistome.
Its function is unknown. In a few families, especially Lysianassidae,
the upper lip has a keel projecting anteriorly and usually separated
from the epistomal region by a deep slit or sinus. Occasionally both

285-135 O - 69 - 2

MARINE GAMMARIDEAN AMPHIPODA 11

epistome and upper lip are produced together and occasionally they
are fully amalgamated.

Lower lip: A bilaterally symmetrical complex forming a partition
behind the mouth. It is composed of at least a pair of lateral lobes,
having their lateral extremities produced, often acutely and often
bearing apicomedially a tiny cusp enclosing the meatus of a salivary
duct. About half of the known gammarideans has a pair of medial
lobes on the lower lip.

Mandibles: A pair of appendages attached lateral to the mouth;
with the upper and lower lips they form a box around the mouth,
permitting buccal closure. The mandibles are powerful and difficult
to remove because of their large muscles. Mandibles generally have
their anterodistal ends (incisors) cut into a series of teeth for biting;
just proximal to the distal teeth may be an articulated process, also
toothed, the lacinia mobilis (accessory plate) which may occur on
only one of the mandibles. A molar with a grinding surface often
occurs on the medioventral surface of the mandible. It may be ridged
and toothed (triturative), or smooth, or be completely absent, es-
pecially in inquilinous amphipods having the mandibles elongated
for piercing and sucking. Most Gammaridea have a 3-articulate
palp attached to the dorsolateral surface of the mandible, the palp
being used to clean the bases of the antennae. Its absence is mod-
erately frequent and often of familial importance, but its reduction
to two or one articles is uncommon.

First maxillae: These are situated posterior to the lower lip.
This pair of appendages is small, each bearing a medial free lobe,
an outer lobe with heavy spines, and attached to the outer lobe a
palp composed of one or two segments, occasionally reduced in
size or absent.

Second mazillae: These are two pairs of lobes behind the first
maxillae, each composed of simple medial and lateral plates, occa-
sionally reduced to one plate or absent, rarely with the outer lobe
attached to the inner by a basal geniculation or extension. Basal
articles of maxillae and maxillipeds are present but rarely are of
taxonomic importance.

Mazxillipeds: One pair of appendages posterior to the maxillae,
each maxilliped is formed of an inner (proximal) lobe, an outer
(distal) lobe and a palp of two to four articles, rarely absent in
Gammaridea, but always absent in the pelagic and often inquilinous
Hyperiidea and in cyamid Caprellidea. The Ochlesidae, lacking
such palps, are assigned to the Gammaridea on the basis of their
resemblance to Gammaridea in other morphological features and
their supposed benthic habits. In peracaridan phylogeny the max-
ilipeds were originally the first pair of thoracic legs, but they have

12 U.S. NATIONAL MUSEUM BULLETIN 271

become incorporated into the cephalic complex; in some talitrids
such as Orchestoidea, the lines of fusion of this thoracic segment to
the head are still apparent but in most amphipods such external
demarcation is obscure.

In comparison to crustacean groups such as the Isopoda, the
body plan of the Gammaridea is conservative. Extreme deviations
from the laterally compressed body with enlarged coxal plates occur
in a few families in which the body has become dorsally depressed
(Corophiidae, Cheluridae, Podoceridae). Two or more of the urosomal
segments have become fused in the Ampeliscidae, Atylidae, Chelu-
ridae, and Kuriidae. Coxae have become deformed or enlarged in
some Lysianassidae, Astyridae, and Hyperiopsidae. The body is
shortened and puliciform in some Haustoriidae and immensely
elobular in some Lysianassidae. In the Podoceridae the first urosomal
segment has become elongated and in the EKophliantidae and Colo-
mastigidae the body becomes subcylindrical as in tanaids.

Major ornaments, of taxonomic value generally at the specific level,
include the frequent occurrence of a rostrum, the differentiation of
lateral cephalic lobes, the presence of processes on the peduncles of
the antennae (especially the Acanthonotozomatidae and Lepechinel-
lidae), and the cuspidation of the pleonal epimera. Dorsal ornamenta-
tion is most common among cold-water Gammaridea and occurs in the
form of teeth and cusps on the pereonites and pleonites. The first
urosomite is often ornamented even in genera without other display.

The absence of eyes is rarely of taxonomic concern other than at
the specific level; indeed many sublittoral oculate species are known to
have eyeless populations in bathyal depths.

The condition of the head in the Gammaridea is highly variable and
thus useful to the taxonomist, but its morphology is often left unde-
scribed. The “basic”? gammaridean generally has the head about as
long as 1.5 pereonites but it varies in different families and their genera
from much shorter than the first pereonite to as long as the first
three pereonites combined. The elongate head is especially noticeable
in the Ampeliscidae, Phoxocephalidae (including the visor-like
rostrum), Synopiidae, and Oedicerotidae. In the latter two families
the head is considered ‘‘massive”’ because it is not only elongate but
very deep. Other families, such as the Stegocephalidae may have deep
heads but they are much shorter than the first three pereonites
combined. Recognition of Synopiidae is almost fully dependent on the
final confirmation of a ‘‘massive”’ head. Most members of that group
have a massive head primarily because pereonites 1-3 are so short
that the head appears relatively large by comparison.

The presence and/or condition of the cephalic rostrum is only
occasionally conservative at the familial level (e.g., Phoxocephalidae,

MARINE GAMMARIDEAN AMPHIPODA 13

Synoplidae). Its presence is of relatively uniform value at the generic
level (example of an exception is Bathymedon) and its shape is often
of good specific value.

Gill structures have been used occasionally for specific and generic
distinctions but their conditions have been ignored in most Gamma-
ridea and require extensive study. Gills are often well known in those
genera in which accessory tube-like branchial appendages have been
discovered or where the primary gills are extraordinarily plaited or
folded. Brood lamellae of the female also have been largely ignored
although a great deal of variation occurs in their shape, setosity, and
terminal ornamentation of the setae. They and the gills may be of
assistance in tracing phylogenetic relationships among families and
superfamilies of Gammaridea. Male reproductive appendages gener-
ally occur as a small pair of projections on the seventh pereonal
sternite; occasionally they are spinose. Other sternal teeth, keels, and
flanges appear to be of rare occurrence (e.g., Aoridae, Eophliantidae).

Geographic Distribution of Marine Gammaridea

The distribution of so few species of Gammaridea is well known
enough as to offer a precise statement of their geographic distribution,
but the distribution of most genera is moderately well outlined be-
cause, of course, bits of data afforded by each species are cumulative
for genera.

Genera can be sorted out relatively easily by gross geographic zones
because faunistic monographers have concentrated primarily within
these zones. Only Sars (1895), Chevreux and Fage (1925), and Schellen-
berg (1926a) have written faunistic monographs broadly overlapping
two zones. Other large scale papers, seemingly monographic, are but
obvious collectors’ assemblages. Warm-temperate analyses have suf-
fered for the lack of discretion between that zone and either tropics
or boreal (=cold-temperate) and I have not taken the time to segre-
gate precisely the boundaries between the broad arctic-subarctic
regions and cold-temperate in describing the distribution of each
species. Arctic Amphipoda of the north polar basin are very poorly
known and probably very sparse; most of the northern Siberian,
Alaskan, Canadian, Greenlandian, and Norwegian shores are placed
in the arctic-subarctic region; Iceland, Kamchatka Peninsula, and
Okhotsk Sea are considered as boreal (cold-temperate), and that
region extends southward to the Japan Sea, middle California, Cape
Hatteras, and the Breton Capes; the warm-temperate includes the
‘southern and Baja Californias, Mediterranean Sea, much of north-
west Africa, and the northern Gulf of Mexico; south warm-temperate

14 U.S. NATIONAL MUSEUM BULLETIN 271

includes the southwestern and southeastern coasts of Australia, South
Africa, and Peru to middle Chile; antiboreal includes Tasmania and
all of New Zealand, for convenience, and in South America includes
all of Schellenberg’s (1931) Magellan and Falkland fauna. South
Georgian faunas are thrown into antarctic-subantarctic classification.
The tropics of western South America end at approximately 4° S.
Warm-temperate of eastern South America is indefinable but of no
consequence because of the absence of gammaridean studies along
most of that coast except in obvious tropical or cold-temperate regions.

Gammaridean genera have been found to fall relatively easily into
these broad classes, the genera either being confined to one class or
being of such wide distribution as to be called cosmopolitan. The
latter term, however, primarily refers to genera that radiate outwards
from tropical regions into boreal regions but not into arctic-antarctic
regions, and there has been little point in so splitting the analysis to
segregate cosmopolitan genera that extend high into polar regions.
Bathyal, abyssal, and hadal faunas are highly discrete, poorly influ-
enced by submergent polar faunas, and, thus, are recognizable as
distinct from latitudinal considerations; the few deep-sea genera that
have been found only in polar regions have been removed from their
endemic position in those regions to the deep-sea classifications.

No precise statistical methods have been used in dealing with
problem genera, those with distributions partially overlapping two
classes, because a bit of subjectivism has been applied in each case
and because the principle of ‘‘centralism’”’ has been utilized. Genera
are thus considered to be confined primarily to that region in which
‘“‘most”’ of the species occur, to wit: a genus with two boreal and one
subarctic species is considered to be boreal but a genus with eight
boreal, two warm-temperate and one tropical species is thrown into
the cosmopolitan class on the probability that more tropical species
remain to be described.

The results of this subjective analysis are presented in table 1;
on first sight the data seem to reveal mostly a relationship to study
effort, with faunas of low latitudes or southern quartospheres suffering
by comparison with well studied boreal-arctic regions. On the other
hand the data seem reasonable if one considers that antiboreal regions
are few in number, small in size, and low in habitat-diversity com-
pared with the extensive boreal regions; except for small oceanic
islands the antiboreal region is confined to Tasmania, New Zealand,
and two coasts of South America, whereas north boreal regions have
four coasts on two continents and a significant disjunct subarctic
embayment, the Okhotsk Sea. The rich antarctic shelves are a strong
contrast to the polar-arctic impoverishment but perhaps the most
striking implication in the data is the low count of tropical endemic

MARINE GAMMARIDEAN AMPHIPODA

15

TaBLE 1.—Geographic classification of gammaridean genera

Class or Zone

Cosmopolitan
Antarctic-subantarctic
Antiboreal

South warm-temperate
Tropical

North warm-temperate
Boreal
Arctic-subarctic
Bipolar !

Biboreal only
Biwarm-temperate
Bathyal only

Abyssal only

Hadal only

TOTAL
FIRST GROUP AMALGAMATION:

Arctic-boreal

Antarctic-antiboreal

North warm-temperate ?

South warm-temperate ?

Cosmopolitan (including bipolar, biboreal, bi-WT)
Deep-sea

Tropical

SECOND GROUP AMALGAMATION (genera only):

Antarctic (including bipolar)

Arctic (including bipolar)

North warm-temperate (including cosmopolitan,
bi-WT)

South warm-temperate (including cosmopolitan,
bi-WT)

Boreal (including cosmopolitan, bipolar, biboreal)

Antiboreal (including cosmopolitan, bipolar,
biboreal)

Tropical (including cosmopolitan)

Genera

104

84

183
132

105

Species in
those Genera

898
110
75
48
111
61
551
ial
488
64
7

72
4

2723

622
185
68
55
1457
239
111

1 Including arctic-boreal and antarctic-antiboreal. 2? Including biwarm-temperate (bi-WT).

16 U.S. NATIONAL MUSEUM BULLETIN 271

genera. Even though numerous species of tropical amphipods remain

to be described, my experience in sorting through quantities of

tropical materials suggests that most of these species will be described
in known genera. New genera seem to be confined primarily to
inquilinous forms probably associated with the vast numbers of

sessile tropical invertebrates. Thus Gammaridea are primarily a

cool-water group in terms of generic diversity; this is reflected in

their strong penetration of the deep-sea. Their body sizes are very
strongly associated with thermal conditions in shallow waters, the
larger bodies occurring in colder water. This does not apply to deep-

sea benthic Amphipoda, however, for there is strong indication (J. L.

Barnard, 1962d) that body size becomes smaller or remains relatively

static with increase of bottom depth along a latitudinal line.

About 40 percent of the 53 families of Gammaridea can be classified
as cosmopolitan in distribution but the other 60 percent are moderately
to strongly confined to specific regions or thermal zones (see below).
They are primarily cold-water oriented, for only 6 families with 21
genera and 53 species are confined to warm shallow waters of low
latitudes. This seems to confirm the orientation of Gammaridea to
cool waters.

The geographic distribution of noncosmopolitan gammaridean
families, or those almost wholly confined to such classification, are
as follows:

ANTARCTIC-ANTIBOREAL: Acanthonotozomatidae, Pagetinidae.

S. WARM-TEMPERATE: Ochlesidae.

TropicaL: Anamixidae, Bateidae, Kuriidae, Leucothoidae.

Boreau: Cressidae, Dogielinotidae, Lafystiidae.

BatuyaL: Astyridae.

ApyssaL: Hyperiopsidae, Lepechinellidae, Vitjazianidae.

CoLD WATER; BIPOLAR SUBMERGENTs: Atylidae, Eusiridae, Haustoriidae,
Lysianassidae, Melphidippidae, Oedicerotidae, Pleustidae, Paramphithoidae,
Pardaliscidae, Sebidae, Stegocephalidae, Stilipedidae, Synopiidae, Thaumatel-
sonidae.

WARM WATER; LOW LATITUDES: Phliantidae, Prophliantidae.

In summary, there is scarcely a coastline where an exploratory
taxonomist interested in species diversity cannot make some contribu-
tion, as Gurjanova (1962) has so ably shown in her study of boreal-
subarctic north Pacific; the taxonomist of western Europe, however,
must move into beta-taxonomy and the northwestern Atlantic
taxonomist must move in that direction in order to detect the rela-
tionships of his fauna to that of western Europe. So also, must the
northeast Pacific taxonomist orient himself to the groundwork laid
for him in Russian Pacific works. The exploratory taxonomist in-
terested in total generic diversity will find the greatest needs for
study in all warm-temperate regions, the antiboreal, the tropics,
and various islands of low latitudes.

MARINE GAMMARIDEAN AMPHIPODA 17
The Composition of a Faunule

A faunule is the aggregate of species of Gammaridea in one geo-
graphic province, region, or habitat. Five megafaunules are well
known: cold-temperate of northeast Atlantic, cold-temperate of
northwest Pacific (including the arctic-like Okhotsk region); circum-
subantarctic encompassing the Norwegian basin and the north coasts
of Siberia; the warm temperate of northeast Atlantic including the
western Mediterranean and the circum-antarctic-subantarctic faunule.
Two faunules secondarily well known are those of the South African
warm-temperate and the northeastern Pacific warm-temperate. The
faunules of individual islands or small geographic areas also have
been more intensively studied than of these larger provinces, examples
being Plymouth, England; Naples, Italy; South Georgia Island.

A good model faunule is that of southern California because it is
midlatitudinal and relatively well explored by quantitative sampling.
The littoral-sublittoral (coastal shelf 0-100 m) faunule comprises 186
reported species of which 59 species live in both littoral and shallow
sublittoral depths; thus the littoral (intertidal) faunule totals 138
species, including about 5 species of beachhoppers and the sublittoral
(coastal shelf) faunule totals 166 species.

TaBLE 2.—Numober of species and genera of Gammaridea in known faunules

Region Genera Species

Warm-temperate California

intertidal (compiled) 66 138
sublittoral 98 166
total, less common to both ; 112 186
Norway (Sars, 1895), boreal section, mainly sublittoral,
0-80 fms 115 221
0-50 fms 110 203
Plymouth, England (Plymouth Marine Fauna), inter- 63 92
tidal
Isle of Man (Bruce et al, 1963) intertidal 40 66
Mediterranean France (Chevreux & Fage, 1925), inter- 41 72
tidal
Atlantic France (Chevreux & Fage, 1925), intertidal 69 109
South Georgia Island 88 159
South Georgia Island, intertidal only (Schellenberg, 45 62
1931)
Falkland Islands, intertidal (Schellenberg, 1931) 48 63
Magellan continental, intertidal (Schellenberg, 1931) 57 Ut.
Indo-Pacific tropics (not including Red Sea with an 97 204
additional 22 species) (J. L. Barnard, 1965)

18 U.S. NATIONAL MUSEUM BULLETIN 271

TaBLE 3.—Dominant benthic genera of Gammaridea in sublittoral soft-bottom
faunule of southern California (+ =yes; 0=no.)

Kind of domination

Genus Number of —@=—<2-—_____—
species Specific Frequency
Diversity of Individuals
Ampelisca ! 12 =P ots
Byblis ! 1 0 +
Ericthonius 2 1 0 +
Eurystheus 2 1 0 +
Heterophoxus 2 1 0 +
Listriella 4 5 + 0
Metaphoxus 2 2 0 sta
Monoculodes 4 0 +
Paraphoxus 3 18 ste te
Photis 2 5 IP ae
(Phoxocephalus)? are 0 +
Protomedeia 2 1 0 +
Synchelidium * 5 a as
Westwoodilla 3 1 0 +

1 Forming limp tubes on or in substrate. 2 Forming stiff or limp tubes on particles or sessile infaunal
structures. 3 Burrowers. ‘4 Nestler or semicommensal.

Various other faunules are compiled in table 2. No attempt has
been made to modernize certain faunules such as Sars’ Norwegian
boreal in order to maintain congruency among the reports; thus each
faunule presumably represents a minimal statement on diversity as
a result of extensive, but not exhaustive exploration.

Generic domination within a faunule may occur through specific
diversity or through high frequencies of individuals in poorly diverse
genera. The Californian faunules of tables 2 and 3 thus comprise some
dominant genera having numerous species, only a few of which have
high frequencies of individuals and other monospecific genera, with
extraordinarily abundant individuals per square meter of habitat.
Other faunules have been compiled in table 4 on the basis only of
intrageneric diversity, as quantitative information is virtually absent
except in the California region. These few examples may give the
reader an approximation of what to expect in commencing quantita-
tive explorations of provinces.

The ecological kinds of Gammaridea within a benthic faunule are
to a large extent determined by domiciliary position rather than
food-type, as most Gammaridea are presumed to be scavengers or
inquilines and their feeding behavior is poorly known. A few true
herbivores apparently occur but only one raptorial predator has ever

MARINE GAMMARIDEAN AMPHIPODA 19

TaBLE 4.—Dominant benthic genera of Gammaridea in littoral epifaunal faunule
of southern California (+ =yes; 0=n0.)

Kind of domination
Number of @=_—-2#2______
Genus species Specific Frequency of
Diversity Individuals

*Ampithoe 11 ae at
* A oroides i 0 ate
Elasmopus 9) aig als
*Hricthonius 1 0 +
*Eurystheus 4 ap ate
Hyale 3 0 oF
*Ischyrocerus 2 0 ais
*Jassa 2 0 ae
Maera 9) ah 0
* Microdeutopus 1 0 +

Parapleustes 4 ote +0
*Photis 6 qr ate

*Forming tubes; others are nestlers.

been described. Of course there are numerous kinds of scavenger
feeding. An intertidal faunule of low latitudes in lush algae will be
comprised of extremely abundant nestlers such as Hyale, Elasmopus,
Maera; protected rock surfaces, interstices and root systems of algae
or surf-grass will contain the domicolous amphipods building tubes,
such as Ampithoe, Ericthonius, Gammaropsis, Jassa, Ischyrocerus, and
Photis. Sessile invertebrates like sponges and tunicates will harbor
nestlers and domicilary kinds as well as a few inquilines, such as
Leucothoe and Polycheria.

The sublittoral faunule on soft bottoms will be comprised of bur-
rowers such as Paraphorus (and other phoxocephalids), haustoriids,
oedicerotids; domiciliary kinds building limp tubes on the sediment
surface, primarily the Ampeliscidae; various domicilary kinds build-
ing tubes on hard particles of the substrate or on projecting tubes
of infaunal organisms, thus the Isaeidae (Photidae) ; and a few nestlers
or semicommensal organisms like Listriella.

Intertidal zones of high latitudes will have nestlers like Pontogeneia,
Paramoera.

The dominant genera of Gammaridea on sublittoral soft bottoms
by gross region are as follows (*=genera with low specific diversity
but high individual frequency):

CosMopoLitan: Ampelisca, [Gammaropsis], Idunella, Listriella, Lysianassa,

Metaphoxus, Paraphoxus, Podocerus, Synchelidium, Urothoe.

BIPOLAR, COLD-WATER: Aristias, Byblis, Epimeria, HEusirus, Liljeborgia, Mel-
phidippa, Orchomene, Podoceropsis, Tryphosella (‘‘Tryphosa, Tmetonyz’’).

20 U.S. NATIONAL MUSEUM BULLETIN 271

Antarctic: Oediceroides, Pseudorchomene*, Tryphosites*, Uristes.

ANTIBOREAL: Acontiostoma*, Amaryllis, Heterophocus.

Tropica: IJdunella, Platyischnopus.

WARM-TEMPERATE: Listrvella.

ARCTIC-BOREAL: Anonyx, Bathyporeia, Dulichia, flock of haustoriid genera,
Lepidepecreum Monoculodes, Oediceros, Onisimus, Paramphithoe, Pontocrates,
Pseudalibrotus, Westwoodilla, Unciola.

The dominant genera of Gammaridea in epifaunas of intertidal and
shallow sublittoral (0-75 m) zones by gross region (*—genera with
low specific diversity but high individual frequency; N—nestler;
I=inquiline; D=domicolous tube-dweller).

CosMopoLitaN: Allorchestes N, Ampithoe D, Cerapus D, Colomastiz I, Corophium
D, Ericthonius D, Gammaropsis D, Hyale N, Jassa D, Lembos D, Leucothoe
I, Maera N, Melita N, Photis D, Podocerus D, Polycheria I, Stenothoe I.

BIPOLAR, COLD-WATER: Aora D, Apherusa N, Halirages N, Liljeborgia N,
Paramoera N, Pontogeneia N, Proboloides (Metopoides) N.

ANTARCTIC-ANTIBOREAL: Amaryllis, Andaniotes ?1, Atyloella N, Atylopsis N,
Bovallia N*, Djerboa N*, Echiniphimedia, Gnathiphimedia, Haplocheira D,
Oradarea, Paradexamine, Pariphimediella, Pontogeniella N, Seba 1, Schraderia
N*, Stebbingia N*, Thaumatelson 1.

WARM-TEMPERATE AND TROPICAL: Amphilochus, Anamizis I*, Batea N, Ceradocus
N, Cheiriphotis D*, Chevalia D*, Cymadusa D, Elasmopus N, Guitanopsis,
Grandidierella D, Leucothoides 1*, Microdeutopus D, Microjassa D, Paragrubia
D*, Parelasmopus N, Parhyale N*.

ARCTIC-BOREAL: Amphilochus, Anisogammarus N, Aoroides D*, Atylus N,
Calliopius N, Cressa, Gammarus N, Gammarellus N, Gitanopsis, Ischyrocerus D,
Leptocheirus D, Metopa, Metopella, Microdeutopus D, Neopleustes N, Para-
pleustes N, Pleustes N, Sympleustes N.

The dominant genera of Gammaridea in the pelagic realms, in-
cluding some demersal genera, are as follows (*=dominant by
frequency of individuals only).

Neritic: Megaluropus, Paraphoxus (males), Synchelidium (pelagic phases),
Synopia.

Eprpetacic: Stenopleura*, Synopia.

BaTHY- AND ABYSSOPELAGIC: Andaniexis, Astyra, Cleonardo, Cyclocaris*,
Cyphocaris, Huandania*, Euonyz, Eusirella, Eusitrogenes*, Eusirus, Halice,
Hirondellea, Hyperiopsis, Ichnopus, Joubinella, Koroga*, Metacyphocaris*,
Orchomene, Paracallisoma*, Paralicella, Parandania*, Parargissa, Phippsiella,
Rhachotropis, Scopelocheirus.

The dominant genera of bathyal-abyssal benthos are as follows
(*=dominant by frequency of individuals only. D—=demersal?).
Conservative numbers of species in depths exceeding 200 meters
are given following each genus.

BENTHIC: Amathillopsis 6, Ampelisca 25, Anonyx 5 (Pacific boreal), Bathy-

amaryllis 4, Byblis 5, Bonnierella 5, Bathymedon 4, Bruzelia 4, Dulichia 6

(Atlantic boreal), Epimeria 6, Haploops 8, Harpinia 4 (Atlantic boreal),
Harpiniopsis 19, ‘“‘Hippomedon’”’ 12, Lepidepecreum 6, Leucothoe 4, Liljeborgia

MARINE GAMMARIDEAN AMPHIPODA 21

6, Melita 5, Metopa 7, Monoculodes 4, Oediceroides 11, Onesimoides 3, Orchomene
14, Proboloides 7, Pseudharpinia 5, Schisturella 5, Stegocephaloides 5, Stenothoe
4, Syrrhoe 5, Syrrhoites 8, Tryphosella 20, Unciola 6 (Atlantic boreal), Uristes
11, Urothoe 5.

DEMERSAL?: Aristias 8, Astyra 5, Cleonardo 8, Eurythenes* 2, Eusirus 9, Halice
8, Lepechinella 13, Pseudotiron 3, Rhachotropis 20, Valettiopsis 3.

Morphological Evolution of the Amphipoda

Amphipoda comprise four unusual suborders: (1) the Gammaridea,
primarily benthic, with perhaps 20 percent pelagic and demersal
species, but having apparently radiated a half dozen or more times
into a gradational suborder, (2) the Hyperiidea, marked by fully
pelagic (free or inquilinous) habits, the strange but universal loss of
maxillipedal palp, and a spectrum of other minor changes presumably
correlated with their habits; perhaps from podocerid gammarideans
have evolved (3) the Caprellidea, skeleton shrimps or marine praying
mantises, characterized by extremely thin tubular bodies, reduction
in abdomen, reduction in two pairs of pereopods, increased cephali-
zation and primarily adapted to a sedentary life in epifaunal anas-
tomoses; but through secondary body depression within the caprelli-
dean scheme (like Temnophlas in the gammaridean scheme) arose
the cetacean ectoparasites Cyamidae, essentially comprising a fifth
major group of Amphipoda; and finally (4) the Ingolfiellidea, appar-
ently undergoing development in association with troglobitic condi-
tions but occasionally returning to marine niches yet open to those
organisms with vestigial pleopods, often bearing cephalic ‘‘ocular’’
scales and with, perhaps, other special ecological adaptations.

Even though Amphipoda have radiated into nearly 60 families
the major diversity can probably be visualized in terms of a score
of kinds. These typological centers may be described by the following
adjectives: ingolfiellid, cyamid, caprellid, six to eight kinds of hyperiid,
gammarid, eusirid, isaeid, lysianassid, colomastigid, eophliantid, phli-
antid, talitroid, stenothoid, amphilochid-leucothoid, ampeliscid, and
acanthonotozomatid. Some outgrowths of these centers form radical
morphs but they fail to qualify as typological centers because (1)
their relationships are not discontinuous or (2) they have not radiated
strongly. The Cheluridae are an example of a radical morph with
presumed relationships to the Isaeidea and low internal diversity;
they are not considered as a typological center. The Colomastigidae,
though of low diversity, have discontinuous relationships with other
Amphipoda and are, therefore, considered as a typological center.
The eusirids (not necessarily the type-genus) have clear relationships
to another center but have radiated so strongly that they must be
considered a typological group. My concept of these centers on cur-

22 U.S. NATIONAL MUSEUM BULLETIN 271

rent knowledge is weak and open to extensive revision as we come to
understand the micromorphology, anatomy, and chemistry of the
various Amphipoda.

On numerous occasions parallel adaptation, convergent evolution,
and independent evolution of morphofunctional conditions have
occurred in Amphipoda. These concern cylindricalization of bodies
(Eophliantidae, Podoceridae, Colomastigidae), cylindricalization of
heads (Cheluridae and Eophliantidae), dorsoventral flattening of
body (Temnophlias, Podocerus and Cyamidae), development of
domitubicolous glands (Ampeliscidae and Isaeidea), loss of max-
illipedal palps (Hyperiidea, Ochlesidae, Cyamidae, and some Lysi-
anassidae), and in a host of minor ways. But, for example, the
diversity does have a measure of constriction in that Amphipoda
have never evolved as fully as have the Copepoda into numerous
parasitic modes or, to our knowledge, have the Amphipoda developed
hosts of rapacious or errant predators in the benthic realm. Predators
do occur in the nektonic Hyperiidea and Gammaridea but none
of the former and few of the latter have returned to a benthic orien-
tation. Although few Isopoda have any degree of lateral compression,
whereas many Gammaridea do have dorsoventral depression, the
Isopoda would seem to be the more highly diversified because they
have cylindrical representatives (Astacilla) and fully evolved para-
sites (Bopyridae). In contrast, the Amphipoda are far more diverse
than certain other orders of Peracarida, such as the Cumacea and
the Tanaidacea. Fourteen families of Gammaridea alone are more
or less inquilinous.

A microhabitational stress must exist between various Amphipoda
and members of the other crustacean orders and phyla, which re-
stricts a fuller display of genetic potential than now in existence.
The tendency of some Amphipoda to enter the crawling realm of
Isopoda, indicates that were Isopoda extinct, Amphipoda could
fill many of those niches, even though rudimentarily or imperfectly.

Various members of the Gammaridae have been considered as the
most primitive of living amphipods. They display most of the basic
gammaridean morphology but the strong development of the lateral
shield (Gurjanova, 1962), composed of coxae or pereopodal wings
in many gammarids, suggests that they have strongly differentiated
from a precursor lacking such a shield. If pereopodal tube-spinning
glands represent a secondary development in Amphipoda, then many
of the isaeid genera that might be considered as close to a shieldless
precursor, have probably undergone a secondary reversion by a
reduction of the lateral shield (e.g., Corophiidae). Other groups with
reduced lateral shield (Eophliantidae, Podoceridae, Colomastigidae)
apparently do not stand close to the primitive amphipod model be-

MARINE GAMMARIDEAN AMPHIPODA 23

cause of vastly modified mouthparts or of so-called pygidization, the
solidification of the urosome and its appendages by segmental coales-
cence or loss of uropodal rami and peduncles. Of course, other per-
acarids without lateral shield may also have tube-spinning glands
(tanaidaceans) and the primitive amphipod may have been in the
glandular line. There are sufficient intergrades in telsonic morphology
between isaeids and gammarids to make this suggestion very attrac-
tive. Several isaeids are almost perfect replicas of the basic gammarid-
ean except for their fleshy telsonic nobs. One has to balance at least
two alternatives, whether the coalesced telson represents a full segment
lost in phylogeny or whether the lobed telson of gammarids represents
a pair of appendages long lost. This problem is fundamental to other
crustacean orders and my impression is that the uncleft, solid telson
is the most common, thus suggesting that it is either more primitive
or at least more successful functionally. The telson in Amphipoda is
not a conservative feature by any means, and our understanding of it
as an evolutionary marker will not be clarified until we understand its
function. Several cycles of morphological development, whether by
advancement or regression are apparent. If the fleshy isaeid telson
becomes “‘falsely’’ lobed in advanced species one can almost imagine
the small step necessary to convert it into one of the poorly lobed or
entire ‘‘subfleshy’’ members of the Gammaridae. Extreme flattening,
full clefting, and elongation then occur in other gammarids and
various derived families, but coalescence of the lobes and shortening
occur again and again. In some haustoriids the telsonic lobes become
fully disjunct basally, each lobe appearing as a vestigial appendage.
The fully blown pygidization of Gammaridae, Isaeidae, and most
gammarideans, in the sense of having the last three pairs of abdominal
appendages formed into relatively inflexible, posteriorly directed
uropods, obstructs our detection of an ancestor in any other living
order of Peracarida, where only the final pair of appendages is formed
into uropods. The reduction or loss of uropods, pleopods, and most
of the abdomen in Caprellidea is clearly a secondary development.
Several good intergrades occur in this procession from Gammaridea
to Caprellidea in such taxa as the Podoceridae, Caprogammaridae
and Cercops, a caprellidean. Most Caprellidea further have thoracic
somite 2 (free segment 1 of other Amphipoda) coalesced with the
head, have a reduction in pereopods 1-2, gills and brood plates. If
the lateral shield or its functional substitute by means of tubicoly,
serve as partial protection for brood and gills in Gammaridea, then
Caprellidea, with their complete loss of lateral shield must have
some other protection, perhaps reflective of their habitats or behavior.
The brood lamellae of caprellids seem to be more strongly cornified
than those of gammarideans. The lateral shield has also been suggested

24 U.S. NATIONAL MUSEUM BULLETIN 271

to be a frictional-flotational support for those Amphipoda known to
swim in peculiar fashion on their sides. Thus the lateral shield, formed
of coxae and articular wings on pereopods, may serve as a kort-
nozzle* in reverse by channeling water ahead of the pleopods.

Good swimmers obviously occur in the Hyperiidea, for they are
all pelagonts; nevertheless, reduction of the lateral shield is a major
trend within that group. Hence, natatorial correlation to the lateral
shield is far from universal and the functional morphologist will
find many fruits to pick once he directs his attention to this problem.
The loss of pleopods in Amphipoda is a mark of the sedentary life
of Caprellidea, and the terrestrial habits of a few talitroids. Their
reduction in ingolfiellids is unexplained but may have some relation-
ship to an interstitial or troglobitic life. A few other inquilinous or
sedentary gammarideans have reduced pleopods, but they are other-
wise remarkably conservative in most domicolous, fossorial, and
inquilinous amphipods, presumably because they are often used for
creating water currents.

Function of the last pair of uropods is presumed to be in propulsion
and ruddering as in other peracaridans, but uropods 1-2 have seem-
ingly little function except as strengtheners for the urosome during
explosive flexation that is a part of jumping behavior. Were this
the sole function of uropods 1-2 the maintenance of at least a small
degree of articular flexibility of the uropodal peduncles and rami
would seem incongruous. Progressive coalescence of uropods 1-2 with
the ventral margins of their segments would afford a strong ventral
pad to be used as a jumping buffer. Perhaps the small degree of
flexibility provides a better shock absorber than would a solid uro-
some. The increase in ornamentation of uropods on certain fossorial
species indicates at least a minimal function in the digging process,
but perhaps this ornamentation primarily prevents coarse particles
from being lodged in the cracks between uropods and segmental
venters. Paddle-like expansion of rami on uropods 1-2 in a few groups
(e.¢., Synopiidae) points to a swimming function. A few sketchy
observations on swimming suggest that the uropods, despite their
restricted flexibility, can be laterally splayed and serve in balance or
braking during the end of a gliding motion.

Reduction of uropod 3 and reduction or rigidification of the uro-
some occur in various tube dwellers (e.g., Corophiidae), inquilines
(some lysianassids, cressids, thaumatelsonids, possibly kurids, pro-
phliantids, etc.), tunnel makers (Eophlantidae) and some of those
amphipods with semipermanent flexion of the abdomen (Phliantidae).
Presumably jumping, swimming, and protection from unwanted par-
ticles have been reduced in importance in these species while there

*A tube directing the propeller-wash of ships.

MARINE GAMMARIDEAN AMPHIPODA 25

is substituted an adaptation towards more favorable leverage for
burrowing into tissues (plant or animal), maintaining special positions
in tubes or channels and in streamlining. Rigidification often occurs
without decrease in urosomal or uropodal size. The tube-dwelling
Ampeliscidae and the inquilinous Dexaminidae have urosomites 2-3
coalesced while the lignivorous Cheluridae have the urosome greatly
increased in size and the uropods greatly enlarged or modified. Jumping
ability is very strong in the Cheluridae.

The ecological linkages to urosomal evolution in the Gammaridea
are manifold; no single solution to functional adaptation has been
necessary. The segmented urosome is a remarkably stable feature
of the Amphipoda; that and its three pairs of uropods are a revolu-
tionary feature unique to Amphipoda. That it has undergone only
one major reversion (Caprellidea), yet has imprinted upon it numerous
adaptive features while there is maintained a minimal structural
stability, attests to its creation as a major part of the success of
amphipods. Like the mandibles, its basic conservativeness has prob-
ably been a factor in the successful dispersal of amphipods into
many niches. The paradox of the almost complete loss of abdomen
in Caprellidea appears to be ameliorated by the recent discovery
of the Caprogammaridae (Kudrjaschov and Vassilenko, 1966) in
which a podocerid-like metasome and urosome are maintained and
by recall of the caprellidean Cercops in which the abdomen, though
distinctly vestigial, is clearly macroscopic. The Caprogammaridae
seem to demonstrate that the loss of urosome was not the trigger
for the development of caprellids; rather, the extreme development
of a tubular body with elongate segments, reduced coxae, terminal
migration of various legs, reduction in numbers of gills and brood
plates, all correlated with numerous behavioral changes were more
important. The ultimate radiation of caprellids, of course, may have
been assisted by the removal of a relatively useless abdomen and the
complete amalgamation of head and pereonite 1.

Development of a dorsal shield in a few Gammaridea has been
discussed by Gurjanova (1962). This feature is simply an extreme
dorsoventral depression of the body and a splaying of the coxae
in the Phliantidae. The abdomen is flexed under the thorax possibly
as an additional protection to the ventrum owing to removal of the
lateral shield. Many isopods, without flexed abdomen do not appear
to require this protection, so there may be other reasons for phliantid
flexion. Gurjanova points out possible precursors to phliantids in
the calliopiid genera Chosroes and Sancho but those genera must
yet be strongly segregated from phliantids because they have fully
developed mouthparts and uropods. They probably should be allo-
cated to a new family in order to qualify the development of a

26 U.S. NATIONAL MUSEUM BULLETIN 271

dorsal shield as a major, albeit rare, adaptation of Amphipoda.
Dorsal shielding is also seen in the calliopiid Amphathopsis and it
thereby forms a strong link among other Calliopiidae, the two
quixotic genera, and is conducted into the Laphystiopsidae. To
some extent body depression also occurs in a few Podoceridae and
this seems significant in light of their presumed derivation from
tubicolous isaeids having lost the tube-spinning glands. This loss
may be correlated with such dorsal shielding because both methods
of protection would seem unnecessary together.

Temnophlias, a ‘“‘phliantid,’”’ has also been included by Gurjanova
as a member of the progression from Amphithopsis through Sancho
into the Phliantidae but another suggestion might be made: that
Temnophlias is really a cylindrical organism with secondary pleuroni-
zation of the pereonites similar to munnid isopods. It may have
affinities with the EKophliantidae, the most strongly developed of the
cylindrical gammarideans. Cylindricality and dorsal shielding are
difficult to separate as various corophiids, chelurids, and aorids have
always been considered to be depressed organisms rather than
cylindrioid. Both terms partially apply to these groups. But one
may consider that those organisms have substituted domiciliary
habits in the form of tube building or burrowing for the true lateral
shield and that dorsal depression is a consequence of cylindricaliza-
tion that cannot be carried too far without rendering the organism
positionally unstable. Some depression is required for the organism
to maintain a crawling equilibrium. Gurjanova appears to regard
Phliantidae (dorsal shields) and EKophliantidae as cohesive, but
eophliantids are strikingly cylindrical. There is now evidence that
eophliantids are phycophilous burrowers and this would correspond
with the habitats of other families living in diverse kinds of ‘‘tunnels.”’
The spherical heads and cylindrical, rotatable necks of eophliantids
are suggestive of limnoriid isopods and are presumably associated
with the tunneling habit. They strongly contrast with Phliantidae
morphologically but there is one small difficulty in completely segre-
gating the two families in that one phliantid is known to be a ligni-
vore; one must presume that phliantids are not tunneling lignivores
on morphological evidence alone.

The cylindrioid Colomastigidae again may be thought of as tun-
nelers or domiciliaries because they probably inhabit tests or tissues
of sessile invertebrates. But inquilinous behavior is scarcely confined
to cylindrioid amphipods, for anamixids and dexaminids, both with
strong lateral shields, are undoubted inquilines because of their
mouthparts (Anamixidae) or their known ascidiophilous behavior
(Polycheria, Dexaminidae). Piercing and sucking mouthparts are
common in many other gammarideans such as Acanthonotozomatidae
and various Lysianassidae.

MARINE GAMMARIDEAN AMPHIPODA 2

Stenopody is the presumed primordial condition of the arthropod
appendage. How far this logic can be carried into the higher Crustacea
as a precedent for any ordinal precursor is wholly philosophical as
far as Amphipoda are concerned. Our ‘‘basic gammaridean”’ already
has specialized anterior appendages, the gnathopods. A precursor to
Amphipoda would already have one pair of maxillipeds. The highest
Crustacea, the Decapoda, have those two pairs of gnathopodal homo-
logues also attached to the head as maxillipeds. Nonambulatory func-
tion of anterior appendages is thus a universally replicated feature of
Malacostraca and one might imagine that it was a fundamental cor-
ollary of amphipodan development. Amphipoda went one step further
than isopods or tanaids in having not just one pair, but two pairs of
functional gnathopods. With few presumably secondary exceptions,
the second pair became the dominant members. The loss of stenopody
signals the development of a nonwalking function as far as the anterior
amphipod legs are concerned. In some Amphipoda these changes occur
even in legs 3-4 (“‘pereopods 1-2”’).

The loss of stenopody was far from immutably fixed in amphipods.
In returning to a slender condition, the marks of a grasping function
have been left, with few exceptions. Return to stenopody, hence en-
feeblement, of gnathopods is a mixed bag among Amphipoda other-
wise recognized as either primitive or advanced. This suggests that
gnathopodal evolvement, especially of gnathopod 2, was a primary
crystallization of the amphipod plan, but that once upon the scene
the original enlargement alone was unnecessary for whatever function
the gnathopods were put to. Perhaps enlargement even went through
a detrimental cycle of overspecialization.

Reduction in size of gnathopods must have come fairly early in
gammaridean evolution for several genera of the Gammaridae have
the gnathopodal size reduced. Some of the functional value of size
may have been replaced by increased setosity. All but two of the eight
families standing near the Gammaridae have the gnathopods enfeebled.
Liljeborgiidae retained the enlarged gnathopods, and among other
reasons, this fact supports the odd thesis that the inquilinous line of
Amphilochidae-Leucothoidae-Stenothoidae has some relationship to
the Liljeborgiidae. Those three families, despite the presence of other
morphological degradations, have not lost the presence or potentiality
of large second gnathopods. The maintenance of enlarged gnathopods
in many members of the Eusiridae-Calliopiidae-Pleustidae is also
further confirmation of their strong relationships to the basic gam-
maridean. The position of the Oedicerotidae, unusually close to the
Gammaridae in the scheme of graph 1, reflects their enlarged gnath-
opods. But in the 10 families presumed to have evolved out of a
eusirid stock, the gnathopods have become enfeebled. Marine tali-
troideans maintain the enlarged gnathopod 2 but terrestrial members

285-135 O - 69 - 3

28 U.S. NATIONAL MUSEUM BULLETIN 271

often return to the stenopodous condition even though gnathopod 2
has obviously not returned to a walking function because of its pe-
culiar morphology.

The seemingly primitive members of the isaeid stock also maintain
the enlarged gnathopod 2 but the advanced members show either an
axial reversal, a shift of domination to gnathopod 1, or an enfeeble-
ment. Potentiality for enlarged gnathopod 2 is fully maintained in the
Ampithoidae and Ischyroceridae and the ultimate peculiarity is
reached in the Cheluridae, one species of which has gnathopod 1! ex-
panded into a fully prehensile appendage like that of Maera. Although
some of the most diverse and, thus, presumably successful shallow-
water (and primarily tropical) genera have the fully enlarged and
prehensile gnathopod 2, the trend in gammaridean evolution has been
a secondary return to stenopody. Retention of the primitively en-
larged gnathopod 2 in the Podoceridae and the Caprellidea, even the
Cyamidea, is one more mark of their relationship. Those very success-
ful tropical gammaridean genera with enlarged gnathopod 2 (in males)
occur in several distinct evolutionary lines: Elasmopus, Maera, and
Ceradocus in the Gammaridae; Hyale in the Talitroidea; Gammaropsis
in the Isaeidae; the axially reversed condition expressed in Lembos
of the Aoridae; Podocerus in the Podoceridae; the inquilinous Stenothoe
of the Stenothoidae and Leucothoe of the Leucothoidae. These genera
clearly have their highest diversity in the tropics and subtropics
whether they had their origin there or not. The ecologist’s attention
should be drawn to this curious matter.

Nontropical gammarideans with enlarged gnathopod 2 are partic-
ularly conspicuous in the demersal eusirids, the liljeborgiids, various
stenothoids and the Gammaridae.

Correlation of mandibular functions with morphology are poorly
understood. Biting, chewing, grinding, piercing, and rasping functions
are obvious, but mandibular variations are far more numerous than
just those five categories. Gammaridean amphipods have been thought
of primarily as scavengers, feeding on debris and detritus, carrion and
dead plant fragments. The basic mandible seems to be adapted to
biting off chunks with the incisors and grinding those chunks with the
molarial rasp. The lack of emphasis on herbivorous habits of Gam-
maridea in the literature is surprising in view of the properly adapted
mandibles and the strong infestation of marine plants by amphipods.
Macroscopic algae and marine grasses infested with amphipods rarely
show gross cropping or evidence of bites having been removed.
Stomach contents of a few phycophilous amphipods demonstrate that
they probably feed on microscopic epiphytes. Undoubtedly the larger
and slower growing algae have evolved mechanisms to limit the success
of marine herbivores; poorly adapted macroscopic algae must surely

MARINE GAMMARIDEAN AMPHIPODA 29

become extinct in view of the almost ubiquitous hordes of marine
amphipods that are potential macroherbivores. Microscopic epiphytes
presumably survive through rapid growth while the total amphipod
population may be restricted by the seasonality of epiphytes; probably
a balance is thus maintained in ways similar to the diatom-copepod
cycle of the pelagic realm. Amphipods may benefit the macrophytes
by cleaning their surfaces of infesting epiphytes.

The microherbivorous amphipod with biting-rasping mandible and
enlarged male gnathopod 2 may be the basic member of the am-
phipodan organization. To suggest that the level bottom scavenging
amphipod evolved first and invaded epifloras later would presume
that cellulase secretion was not an original part of the amphipod plan.
The grossly compressed bodies of the basic amphipod also attest to
a preadaptation for nestling and gliding amongst anastomoses. That
this ungainly structure later was able to invade a host of other habitats
seemingly unsuitable to a flea-like morphology suggests that in the
course of their evolution numerous ‘hidden’ adaptations accrued.
Some of these may be circumstantial: high prodigality (‘‘success by
numerical pressure’’), and low genetic plasticity that maintains a
broad adaptability to feeding conditions. A more efficient mechanism
than the basic mandible may be imagined for cropping micro-epiphytes
without radical changes in the general structure. The maintenance of
that basic mandible throughout so many families and genera of am-
phipods, which obviously have put it to numerous functions, suggests
that one key to success of amphipods is their potential omnivorous
habit. A measure of this manifold feeding potential is seen in one’s
ability to trap diatom-feeding amphipods by means of a carrion-baited
undersea trap.

In the dispersal to level bottoms, amphipodan morphofunction
almost invariably changed; the successful Ampeliscidae build tubes;
the Phoxocephalidae-Haustoriidae and Oedicerotidae dig burrows and
often become much broadened in their bodies; other genera, like
Listriella, have obtained special associations with infaunal members
of the level bottoms.

Almost all of the greater Isaeidea, the tubicolous amphipods, have
maintained the basic mandible, palp included. Even the wood “‘boring”’
Cheluridae are able to rasp wood with the basic mandible. The greater
Talitroidea have maintained the rasping mandible although the palp
has been lost. Presumably one function of the palp is the cleansing
of the anterior cephalic space between the antennae. Amphipods
without palp often have few antennal setae projecting into that
space to trap particles.

The trend to a distinct change in mandibular morphology is seen
in some Gammaridae and even more strongly in some of the families

30 U.S. NATIONAL MUSEUM BULLETIN 271

closely associated with the Gammaridae, such as the Liljeborgiidae,
Eusiridae, and Phoxocephalidae-Haustoriidae. Loss of trituration sur-
face and reduction in size of molar are universal in the Liljeborgiidae,
but the reasons are not yet apparent as the ecology of the group is
poorly known; various burrowers in the Phoxocephalidae, Haustori-
idae, and Oedicerotidae have smoothed-off molars and several of those
genera have enormously enlarged molars covered with a setular velvet.
Such molars are also seen in the Synopiidae. They must have some
relationship to the fossorial or semifossorial habit of processing mineral
grains. But in those large fossorial families the normal mandible is
retained by numerous genera.

The strongest changes in mandibular morphology occur in those
families presumed to be inquilines. In one way or another these
families have adapted to piercing and sucking or possibly to the
scraping of slime but even some inquilinous amphipods maintain the
biting and grinding functions. The mandible of Polycheria is used
chiefly for burrowing into the tests of tunicates for domiciliary purposes
rather than for feeding. Indeed, the biting adaptation is rarely lost
even in the strongest inquilines; it is maintained in many Acantho-
notozomatidae, although others of the family have those incisors
developed into stylets. The conformity of the acanthonotozomatid
and stegocephalid mouthpart bundle suggests a gross piercing func-
tion as if they normally attack some large sessile invertebrate. Pre-
sumably some of the ‘‘inquilines,” like those of the greater Steno-
thoidea, are grazing predators, biting off coelenterate polyps or
consuming sponge and tunicate tissues.

The pardaliscid and stilipedid mandibles are the most paradoxical.
They are elytriform like those of some stegocephalids, lack molars
but retain palps, yet the mouthpart bundle is rarely coniform and
no one has demonstrated an inquilinous behavior. Many of these
species are nekters or demersal members of the deep-sea fauna.

Maxillipedal changes mark one of the primary subordinal grades of
evolution within the Amphipoda. The loss of palps is a condition of
the Hyperiidea. Only two families of Gammaridea and a few genera
of two other families have a marked reduction or loss of these palps.
Such loss is associated with a nektonic, often inquilinous habit, but
numerous pelagic Gammaridea have fully developed palps. Reduction
in maxillipedal plates or palps is not perfectly correlated with the
inquilinous families or those marked by mandibular changes,but as we
should expect, all stages of the perfection of this morphology are ap-
parent and the trend is obvious. Plates and palps often evolve in-
dependently as if their functions were distinct; in some cases such as
Liljeborgiidae and throughout the greater Stenothoidea, the plates
become reduced while the palps are maintained or increased in size.

MARINE GAMMARIDEAN AMPHIPODA dl
Interfamilial Gammaridean Relationships

Through reeapitulation of the general evolutionary trends within
the Gammaridea, one may estimate the pattern of evolution and rela-
tive positions of the several families (graph 1). Formulation of this
pattern was assisted by organizing data on 3-dimensional stick charts
containing plots of characters representing grades of structure, and
simple character alternatives; on wheels and triangles so as to demon-
strate presence or absence of characters in small familial groups;
by determining the fewest logical increments of change necessary to
transform one family into another. The final pattern was influenced
strongly by ampliative inference in using our knowledge of the ex-
treme members of a family plus the implication that some changes
require more genetic input than others even though genetic ‘“‘revolu-
tions” may occur repeatedly in certain groups and that certain gross
characters are very stable.

Only position and change from one condition to another could be
incorporated in graph 1; thus each family appears falsely to have equal
importance to all others. Estimates of distance might be incorporated
as a measure of genetic input; generic and specific diversity of the
familial centers could be shown by reducing the graph to an areal
projection reflecting the harmony of certain grades of structure (at
least in widespread families tested by a wide range of environmental
selection) and the low level of diversity in familial ‘‘experiments.”’
The pattern has been organized to fit the space although distance from
Gammaridae implies specialization or advancement. At best, the
pattern is a straw man.

Graph 1 has been partitioned into more than a dozen blocks of one
or more families (or genera). Some of those blocks, like the Colo-
mastigidae or Lysianassidae, have uncertain relationships to other
Gammaridea. Solid arrows represent estimates of strong and direct
relationships; arrows commencing between two families or at long
distances from family groups imply relationship only to ancestral
pools having grades of structure similar to existing families. Dotted
lines imply weak or distant relationships; wavy lines denote families
placed close to Gammaridae because of precedential pathways
established in adjacent examples. Symbols on the chart are reminders
of certain characteristic structures and are cumulative between
arrows unless marked otherwise.

The Gammaridae form the base stock and eight other families are
enclosed within the block denoting gammarid-like structure, two
families, the Phoxocephalidae and Haustoriidae, being so close to
Gammaridae as not to warrant extra space on the diagram. They
have extreme fossorial adaptations found rudimentarily in several
Gammaridae.

32 U.S. NATIONAL MUSEUM BULLETIN 271

The major line of evolution out of the Gammaridae appears to pass
through eusirid organisms in the next block below; this trend is first
marked by reduction or loss of accessory flagellum and radiates further
by various modifications to be discussed in a later paragraph. The
point to be made is that the change from gammarids to eusirids
seems to be less revolutionary than the changes from gammarids to
various families kept within the gammarid block, and each of those
probably should be accorded the rank of a block. The mark of their
relationship to gammarids is the retention of the accessory flagellum
in multiarticulate condition. A presumption that an accessory
flagellum could be redeveloped once lost would have to be invoked
in order to bring some of those gammarid-like families down into the
eusirid block where grades of other structures show some resemblance.
I see little objection and a slight precedent to that possibility in the
fact that many Gammaridea add articles to the accessory flagellum
with body growth; is there a strong difference in metameric potential
between 1 and 2 or zero and 1?. If all the gammarid-block families
were placed in the eusirid block only two, Liljeborgiidae and Vitjazi-
anidae, could be further drawn from eusirids by reestablishing the
accessory flagellum. There is no logic in funneling all other gamma-
rideans through a eusirid block when many of them can be extended
directly from the Gammaridae. Gammaridae are no less diverse than
the three major families of the eusirid block. Each of the gammarid-
block families shows some tendency in their advanced genera towards
reduction in the accessory flagellum, thereby suggesting that the
presence of the ramus in those families is primordial.

Melphidippidae have the structure of gammarids in which the
coxae have become shortened, and the gnathopods enfeebled lke
females of the gammarid genus Cheirocratus, plus an elongation of
uropod 3. Cephalic ocular bulges and a strong trend towards reduction
of the accessory flagellum are characteristic.

Vitjazianidae have the general appearance and ecology of some
pelagic eusirids but their retention of a 3-articulate accessory flagellum
and the conjoint base of the primary flagellum indicate a mode of
evolution different from that of eusirids and thus the Vitjazianidae
are maintained in the gammarid families even though they may be
more advanced than eusirids or gammarids. Gnathopod 1 has become
simple and the most specialized vitjazianids have extremely reduced
coxae. The mouthparts maintain a basic gammarid structure.

Hyperiopsidae form another pelagic theme. The typical genus
resembles hyperiid Amphipoda but retains the maxillipedal palps;
the second genus, Parargissa, differs so remarkably in its overall
appearance that it is debatably an hyperiopsid but mouthparts,
antennae, and pereopods seem to confirm the relationships between

285-135 O - 69 (Face p. 32)

ere Oe
pereopods glandular Dy,

Isaeidae vy

ettiidae

Aoridae

Ampithoidae

ee U3
a

Tschyroceridae

hotidae

rial Dy

Metoediceros

Kuriidae

|
Eophliantidae Ee
body
cylindrical

Corophiidae

=

Cheluridae
Lysianassidae

F egcee

A-1

Phliantidae
body depressed

coxae splayed Lidymocheila

285-135 O - 69 (Face p. 32)

BS aexe SES] ——
Pardaliscidae C Melphidippidae F eS Sei Cc
U-3

v

(Leese a ere
1 a er, fh as
) M BASIC GAMMARIDEAN Bis Oedicerotidae

ees os

aot
WS i) pa!

Argissidae Be

a

Beaudettiidae pereopods glandular UN

Ca Isaeidae tf
U-3
Talitroidea 6)
M

Dogielinotidae

fossorial \

Gammaridae

Haustoriidae —
Phoxocephalidae

Synopiidae

Aoridae

Vitjazianidae

Ampithoidae

a US
x

= Cal

=

Metoediceros
Liljeborgiidae Ischyroceridae
=O) Eusiridae -Calliopiidae
Pleustidae eS
Psevdamphilochus U-3 A

< G-l
Paramphithoidae

|

Eophliantidae Ee
body

cylindrical

Cheluridae ae eis

ar i Bateidae
Va- if YALE Gn. 1 vestigial
U C

J Astyridae

Corophiidae

|
|
Bie a
|
|
|
|
|
|

U-3 Co Sebidae, Pagetinidae

Stenothoidae, Cressidae

Thaumatelsonidae TID. f, Oe Se \

=
=

Lepechinellidae Stilipedidae

GZe
UV; *

Lysianassidae

oe

Acanthonotozomatidae

VAL

ag

Ampeliscidae

Phliantidae

Colomastigidae ; p .
O a body depressed as G-2
body \ =! —> ly dep A-1
cylindrical wv 1% . s Oahisinen: P = sen clara! Didymocheile

=k

Grarx 1.—Pattern of evolution in the Gammaridea (see text for discussion). Symbols:
A-1=antenna 1; B=conical mouthpart field; C=coxae 1-3; F=accessory flagellum;
G-1 (or Gn. 1), 2=gnathopods 1, 2; H=head; M=mandible, arrows pointing to special
features; O=accessory flagellum absent; P=Maxilliped: circles=plates, appendage =
palp; p-l, 5=pereopods 1, 5; T==telson, fleshy kind, lateral view; U=urosome; U-2,
3=uropods 2, 3; X=maxilla 1.

(UP Ph

Cy ares
eee

MARINE GAMMARIDEAN AMPHIPODA 33

the two genera. Again, some resemblance to the eusirid grade of struc-
ture is seen in the elongate enfeebled gnathopods characteristic of some
calliopuds, but gnathopod 1 is almost completely simple, the accessory
flagellum is 3-articulate and elongate as in some vitjazianids, and,
furthermore, the palp on one member of the first maxillae is specially
modified, bent and scaly. Article 4 of pereopods 1-2 is enormously
elongate, thus giving to the pereopods a strong raptorial function
apparently surrendered by the gnathopods.

Argissidae fall as a sidebranch of the gammarid families into the
eusirid grade of structure by reduction of the accessory flagellum to
2-articles. Such reduction by itself does not imply any direct rela-
tionship to those eusirid families for some marine and nonmarine
Gammaridae also have a reduced accessory flagellum; such reduction
occurs repeatedly in other completely distinct groups (e.g., isaeids).
The peculiar coxal morphology of argissids (fig. 43) is not fully unique,
for an analogous condition occurs in a gammarid genus, Megaluropus.
Enfeeblement of gnathopods is again a feature of argissids but the
quadrilocular eyes of oculate members seem significant. Except for
those eyes, the diagnostic characters of argissids, though unique to-
gether, are drawn from diverse members of the Gammaridae.

Pardaliscidae are a very difficult puzzle. They are perhaps the most
aberrant of the gammarid families, although synopiids rival them.
Once the principle has been established that the presence of a well-
developed accessory flagellum probably marks a gammarid family,
or a noneusiridean branch, the Pardaliscidae are to be kept within the
gammarid block. But they are characterized by foliation of the man-
dibles and loss of molars, the frequently occurring conjoint condition
at the base of the primary flagellum on antenna 1, feeble gnathopods,
progressively reduced coxae, and one or more peculiarities of the maxil-
lipeds: reduction in overall size of the inner plates, often a reduction
in the outer plates, and, occasionally, an elongation of the article
carrying the outer plates. The lower lip often has the inner lobes
coalesced and forming a convex bridge between the outer lobes. A
generalized view of the maxillipeds suggests affinities with the Lil-
jeborgiidae in which the maxillipedal palps, like those of the Par-
daliscidae, are relatively dominant over the basal plates. The mandibles
of Liljeborgiidae show a strong tendency to a complete loss of the
molars but liljeborgiid gnathopods are very powerful and the lilje-
borgiid with the smallest coxae has larger coxae than any pardaliscid.
The lower lip of liljeborgiids could be precursive to that of pardaliscids.

The powerful gnathopods of the Liljeborgiidae are reminiscent of
those in the Eusiridae (Rhachotropis) but most lijeborgiids have a
strongly developed accessory flagellum and all have a reduced, non-
triturative molar. Some eusirids also have this reduced molar. The

34 U.S. NATIONAL MUSEUM BULLETIN 271

EKusiridae seem to be so broadly polyphyletic that there is cause to
investigate the interrelationships of liljeborgiids with some of the
eusirids. Several grades of structure are congruent between the
Liljeborgiidae and certain members of the Pleustidae. The labia of
the two groups are very similar and the mandibular molars of pleustids
often resemble those of lujeborgiids. Pleustid gnathopods are often
enlarged but the family differs from liljeborgiids in their uncleft telson
and vestigial accessory flagellum. The outer rami of uropods 1-3 are
shortened in Pleustidae and the outer ramus of uropod 3 is uniarticu-
late but some liljeborgiids approach these conditions. The pleustid
rostrum is a development restricted to a few members only.

Synoptidae are retained in close proximity to the basic gammaridean
in view of their elongate accessory flagellum, even though several of
their members have the articles reduced to two. Gnathopods are
feeble. All but a few synopiids have the head enlarged in relation to
the shortened pereonites 1-3. Coxa 3 dominates coxa 4 in most of the
genera, and in all but one genus, article 3 of the mandibular palp has
become very short, almost vestigial. The mouthparts are otherwise
basic except for those genera having the molars extremely enlarged
and velvety smooth. This semifossorial condition has also occurred in
some haustoriids even closer to the Gammaridae than the Synopiidae.
Uropods have undergone the eusirid shortening of the outer rami but
uropods 1-2 are far more specialized in the frequent sublamellar con-
dition of the inner rami. In all but Synopia, the telson has become
elongate. Eyes when present are dorsally coalesced or contiguous.

Oedicerotidae are another group, like the Synopiidae and Parda-
liscidae of generally obscure relationships. They have reached the
eusirid grade of structure in accessory flagellum, resemble the Syn-
opiidae in head and eyes but the telson is a short, thin, ovate or
truncato-ovate, uncleft lamina, unlike that of most Synopidae.
Uropod 3 has become fully elongate, a tendency seen in several
synopiids but the pereopods have become strongly fossorial, the first
four pairs by virtue of their long setae and the fifth pair by virtue of
its immense articular elongation. On the average, gnathopods are of
medium size, thus being more powerful than those of synopiids.
Oedicerotidae have a strong resemblance to isaeids especially in
pleonal epimera and pigmentation but lack pereopodal glands and have
a thin, nonfleshy telson.

Three families, each of them highly distinct from one another, seem
to have strong relationships to the Eusiridae-Callioptidae-Pleustidae
complex. The least distinct is the Paramphithoidae, a group character-
ized by acuminate coxae; several intergrading genera suggest that
Paramphithoidae might be incorporated within the broader eusirid
amalgam. The Laphystiopsidae carry calliopiid tendencies to a loss of

MARINE GAMMARIDEAN AMPHIPODA 35

molar grinding ridges, miniaturization of coxae, and simplicity of
gnathopods to their definitive extremes. Bateidae have the cephalic
pleustid aspect but have undergone strong divergence through reduc-
tion of gnathopod 1 to a single article or two.

Paramphithoidae have links to more advanced families, either
directly or indirectly. Generic intergradations between Paramphi-
thoidae and Astyridae are readily apparent (see those families for
elaboration). The Lepechinellidae have the acuminate coxae of par-
amphithoids, but through coalescence of two urosomal segments
resemble a parallel grade, the Atylidae. Their origin is thus plotted in
graph 1 as a mixture of characters between Atylidae and Paramphi-
thoidae, an unsatisfactory procedure indicating not phyletic affinities
but gradational structure.

The body plan of the Stilipedidae conforms remarkably to that of
the Astyridae. If the pardaliscid left mandible, adopted by the
Stilipedidae, is a character of major evolutionary significance, then
one might consider that stilipedids stand close to the Pardaliscidae.
But the astyrid mandible is in a condition precursive to the pardaliscid
right mandible. It is partially flattened and grossly toothed while
retaining a nontriturative molar. It thus resembles the pardaliscid
right mandible whereas the stilipedid mandibles resemble the broad-
ened untoothed left mandible of pardaliscids. Since pardaliscids combine
both kinds in a single individual, it does not stretch one’s imagination
to consider that the astyrid mandible evolved into that of stilipedids.
That astyrids did not also produce pardaliscids is attested to by the
strong accessory flagellum and weak coxae of pardaliscids.

Atylidae have the same general advancements of the eusirids but
the fusion of two urosomites marks a line of evolution which some
students suggest has led to the inquilinous Dexaminidae lacking
mandibular palps; this condition naturally leads to the palpless
Prophliantidae in which all urosomites have become coalesced.

A more radical inquilinous specialization in the acanthonotozomatid
line seems to have its gravitational balance near the Paramphithoidae.
Here a marked change in the mouthpart field from its basic quadrati-
form bundle to a conical or triangular field suggests increasing stages
towards the functions of piercing and sucking. The individual mouth-
parts increasingly become better adapted for those habits in the various
acanthonotozomatid genera through incisorial styliformity and reduc-
tion of molars; but most acanthonotozomatids live in south polar
waters, often reach giant body proportions, and, in many cases, the
mandibles have become broad, flattened, and very powerful as if their
bearers have evolved toward a state of subpredation, perhaps on giant
sessile organisms such as sponges. They might be called predatorial
grazers if this habit is confirmed.

36 U.S. NATIONAL MUSEUM BULLETIN 271

Direct advancement to the Lafystiidae through reduction in maxil-
lipedal palps is a consequence of acanthonotozomatid specialization
and the ultimate is reached in the Ochlesidae, technically hyperiids,
because of the complete loss of palps.

Stegocephalidae are the only other major group of Gammaridea
with the strong piercing and sucking mouthpart field of acanthono-
tozomatids and even though their coxae are not as acuminate and
their mandibular palps and molars have been lost, there is still merit
in considering a direct relationship between the two families. Stego-
cephalidae have the broad, sublaminar mandibles of several acantho-
notozomatids but may have originally evolved as a pelagic group,
some members having returned to the benthos in later stages. The
breadth of the lateral shield appears to have a relationship to midwater
suspension but acumination may still be seen in the anterior coxae.
The benthic members may also be predatorial grazers and the only
known raptorial predator in the benthic Gammaridea is a member of
the Stegocephalidae.

Another line of evolution favoring inquilinous behavior is that
commencing with the Amphilochidae and Leucothoidae. Early stu-
dents of the Gammaridea noticed similarities between Liljeborgiidae
and Leucothoidae in maxillipedal structures; other resemblances such
as retention of enlarged gnathopods are so clear that one might say
that leucothoids are liljeborgiids in which the accessory flagellum has
become vestigial and gnathopod 1 has been transformed into its fully
carpochelate condition while the outer plates of the maxillipeds
became vestigial. The rudiments of the carpochelate gnathopod may
be seen in gnathopod 2 of leucothoids, thus resembling the gnathopods
of liljeborgiids. Anamixids carry the inquilinous state to the ultimate
by the transformation of mandibles and maxillae into a piercing keel.

Amphilochidae may stand almost completely alone. Their mouth-
parts strongly resemble those of liljeborgiids and their gnathopods are
usually miniaturized editions, like those of Listriella. But their heads
have the appearance of the pleustid-paramphithoid or bateid line.
The primary mark of their advancement is the reduction of coxa 1,
not. as fully reduced as in the Bateidae. This suggests their direct
precursorial relationships to the stenothoids. The peculiar Pseudam-
philochus, through its cleft but ovato-acuminate telson, unreduced
coxa 1, large rostrum and nonelongate peduncle of uropod 3, stands
among the Amphilochidae, Pleustidae, and Liljeborgiidae. Schellen-
berg (1931) suggested that it should be assigned to a unique family,
the Pseudamphilochidae.

All five families of the stenothoid complex seem to have strong
interrelationships by virtue of the mandibular form (see figures) in
which at least one mandible has a box-like shape with deeply serrate

MARINE GAMMARIDEAN AMPHIPODA 37

incisor, a lacinia mobilis formed of a thin lamina resembling the incisor
and nearly appressed to it, plus a molar bulge clearly moved distal-
wards toward the incisor and nearly or partially encroached upon by
the spine row. One group of stenothoids has the amphilochid coxa 1,
whereas the other has the leucothoid (normal) coxa 1. Sebid
enathopods even vaguely resemble gnathopod 1 of leucothoids and
numerous other similarities are apparent. Stenothoidae, Thaumatel-
sonidae, and Cressidae, those with amphilochid coxa 1, are very
closely related among themselves, even though some extremely
pygidized species have been used as types of the Thaumatelsonidae
and Cressidae. The five stenothoid families are obviously inter-
related through the condition of uropod 3. It is uniramous, pre-
sumably through loss of the inner ramus, as the remaining ramus is
basically biarticulate. Neither the Leucothoidae nor Amphilochidae
show a tendency to this condition, except perhaps for the universal
genus Pseudamphilochus. If the Amphilochidae were the precursors
of the Stenothoidae and the Leucothoidae were precursors to the
Sebidae, then the evolution of a common uropod 3 had to occur twice.

The superfamily Talitroidea, comprising originally the Talitridae,
for nearly a century have been considered as extremely distinct
gammarideans, often worthy of even subordinal rank. They are more
diverse than they ever have been if one were to add the Dogielinotidae
and Kuriidae to their ranks and to suggest that Najna and Metoediceros
represent types of new talitroid families. Bulycheva (1957) split the
Talitridae into three families, adding the Hyalidae and the Hyalellidae.
The three main families are here treated as a superfamily, mainly for
convenience in identification. Talitroids have no universally unique
characters; several other families have a uniramous uropod 3 and no
mandibular palp but among the genera of talitroids are many unusual
morphs. Often the cephalic sclerites are clearly marked. Numerous
enathopodal, pleopodal, and antennal modifications occur in terrestrial
genera; the jumping ability is extreme in those genera. The marine
members appear more regularized, some even having vestigial inner
rami on uropod 3. Perhaps the structure of fringing setae on the female
brood lamellae will prove to be characteristic of the group. Although
they may be considered as very distant from the basic gammaridean,
their singularity is damaged by discovery of the Beaudettiidae. That
monotypic family is composed of a species with clear relationship to
Elasmopus, a member of the Gammaridae. Through loss of mandibular
palp, reduction of the inner ramus of uropod 3, and telsonic modifica-
tion, Beaudettia has come close to the talitroidean grade of structure.
Morphological distance of talitroids from gammarids, as a reflection
of time or extensive genetic change, is thus lessened by our
observation of the revolutionary changes that have occurred in
Beaudettia, possibly in relatively recent times.

38 U.S. NATIONAL MUSEUM BULLETIN 271

The final manifold group in graph 1 is the greater isaeid complex
that might be accorded superfamily rank. The primitive isaeid is
conceived of as a morphological analogue to the basic member of
Gammaridae. Until one examines the fleshy telson and pereopodal
glands of primitive isaeids, one is struck by the great similarity of
generalized Gammaridae and Isaeidae (—Photidae). Isaeids have
become very diverse in many of the same ways as have the gammaridan
stock but no highly advanced inquilines have appeared, unless one
can link up some of the eophliantids; they instead appear to be
lignivores. Even so, the piercing-sucking groups like acanthonotozo-
matids, the coelenterate loving groups like the stenothoids and
amphilochids, the spongicolous and protochordate inhabiting kinds
have not evolved within the isaeid complex, perhaps because they
have been primarily adapted to form domiciles of their own out of
the pereopodal spinning glands. Identification and relationships are
clouded by the frequent loss of those glands in the Podoceridae, some
Corophiidae, the Cheluridae, and even the nomenclatorial type,
Isaea. A more basic example of the isaeid line is Gammaropsis and its
name should ideally be the root of the stock.

The most advanced members of the isaeid group have accumulated
two morphological changes, either reversal in gnathopodal domination
or development of a partially to fully rigid urosome with loss of
uropodal structures. The gnathopodal reversal may be related to a
stronger than normal cephalic orientation required of organisms living
in tubes open to the anterior end of the animal body. Rigidization of
the urosome may assist the organism in maintaining a position within
the tube. Some of these tube dwellers have even returned to making
burrows (?internally lined) in the substrate.

Perhaps the Podoceridae have come closest to the inquilinous func-
tion; their ecology and morphology are poorly known but there has
developed the impression that all their members have lost the spinning
glands and that many of them are strongly associated with hydroid
colonies, as if they were predatorial browsers.

The stability and similarity among the mouthparts of the members
of the isaeid complex leads one to the view that podocerids belong with
the group. They are frequently mentioned as the root stock of caprel-
lids; indeed, since the time (Dec. 1965) that the contents of this paper
were formed, a new family, Caprogammaridae (Kudrjaschov and
Vassilenko, 1966) has been described with further strong evidence of
podocerid-caprellid relationships.

Distinctions between Isaeidae and Aoridae are not clear except that
enathopod 1 of aorids is either larger than gnathopod 2 or is that
member having sexual dimorphism. The Corophiidae seem to be a
polyphyletic group of aorids and isaeids with pygidization. They may

MARINE GAMMARIDEAN AMPHIPODA 39

include some ischyrocerids also. The Ampithoidae are fairly uniform
by virtue of their third uropods and the Ischyroceridae, though anal-
ogous to ampithoids because of the development of a rudimentarily
uncinate condition on the outer rami, are recognizable by the elonga-
tion of the peduncle. Ischyroceridae rarely have the reversed gnatho-
podal domination. One corophiid, Hricthonius appears to combine
features of Corophiidae, Aoridae, and Ischyroceridae, but it is not the
only “‘isaeid” genus that gives trouble to the systematist.

The lignivorous Cheluridae have numerous resemblances to the
isaeid-corophiid line, but the fully pygidized urosome reveals suture
marks indicating that urosomite 3 has become extremely enlarged, a
feature unique to this family. There is no better demonstration of the
extreme genetic potential of numerous Amphipoda in replicating long
lost structures than by considering the enormous, Maera-like
enathopod 1 of Chelura insulae Calman.

Families without clear relationships to others are the Lysianassidae,
Phliantidae, Colomastigidae, Ampeliscidae, and potential families are
represented by Didymocheila, Ceina, and Biancolina.

The Ampeliscidae are a very advanced group, apparently completely
divorced from the isaeid complex, which have developed pereopodal
glands and spinning tubes of a different form from those of isaeids.
Ampeliscid morphofunction is also discrete, as far as we know.
Ampeliscids have a mixture of characters represented by argissids and
atylids but those two families otherwise bear no relationships. Thus,
the line on graph 1 connecting Ampeliscidae to Argissidae and Atylidae
represents only a focus on structural grades.

Despite similarities to the Sancho-Chosroes members of the Calliopi-
idae, the Phliantidae bear relationships to certain dorsoventrally de-
pressed podocerids, like Podocerus and I am inclined to the view that
the latter suggestion has great merit and should be examined carefully.
Phliantids also have about as much in common with Laphystiopsidae
(mostly dorsoventral depression) as they do to Sancho and Chosroes.
Apical curls on brood lamellar setae plus many similarities in buccal
and urosomal parts suggest phliantid affinities with Talitroidea.

Relationships of the Colomastigidae are difficult to trace. The body
form is subcylindrical but the head is not of the spheroid kind found in
the Eophliantidae, the rami of uropod 3 are present and elongate, the
peduncle is elongate and the mandible apparently lacks a true incisor,
that characteristic having been replaced by an enlarged spine row.
Affinities with leucothoids are seen in maxillipeds, uropods, and telson.

The Lysianassidae are almost as fully diverse as all of the other
Gammaridea put together and comprise about 20 percent of all marine
genera and species of the suborder. They are united together by their
clearly recognizable gnathopod 2 and furthermore share an almost

40 U.S. NATIONAL MUSEUM BULLETIN 271

unique antenna 1, with very short and often telescoped articles 2-3.
Otherwise they have radiated into species that exhibit all forms of
inquilinous specialization such as piercing and sucking mouthparts,
coalesced urosomites, reduced uropod 3 and into numerous highly
successful pelagonts. The usual nestling and burrowing members also
occur in profusion. Some pelagic members are predatorial and some
may have developed narcotizing glands. Even wood boring is a pre-
sumed habit of some deep-sea lysianassids eating waterlogged coco-
nuts. But none has developed the tubicoly of isaeids.

The morphologist cannot as yet recognize any clear links between
the Lysianassidae and other gammarideans. There are vague resem-
blances to Stegocephalidae, perhaps of convergence and suggesting that
Lysianassidae may have developed, like Stegocephalidae, first as a
pelagic group that later reinvaded the benthic realm. The obligatorily
pelagic genera among the Lysianassidae are very highly specialized but
many unspecialized lysianassids of other large genera (e.g., Orchomene)
occur prominently in the nekton. Elongation of article 3 on gnathopod
2 is not fully unique to the Lysianassidae, as seen in section B of the
written key to families (p. 109). The functional morphology of this
lysianassid appendage, with its other unique characters, should be a
problem of major concern to gammaridean evolutionists. It is not
grossly dimorphic in the two sexes and seems wholly inadequate, like
gnathopod 2 in so many other families, of serving as a sexual grasping
appendage. Male gnathopod 2 does serve this function in those few
Amphipoda that have been observed in amplexus.

Limnetic Evolution of the Gammaridea

That Gammaridae collectively represent the most primitive mem-
bers of the Gammaridea and of the living Amphipoda, has some
support by knowledge of the occurrence of an enormously diverse
faunule in Lake Baikal. Nearly 250 species of Amphipoda live in that
Siberian lake (see Bazikalova, 1945; Dybowsky, 1874). One might
presume from the extent of radiation that has occurred in the Baikalian
gammarids that they entered the lake shortly after its formation in
the Triassic. All Gammaridea now living in the lake are classified
with the family Gammaridae even though there is justification in re-
garding some of the genera, like Hyalellopsis, as members of novel
families. Baikalian Gammaridae strikingly reflect many of the morpho-
logical conditions now allotted familial importance in marine amphi-
pods. Non-Baikalian epigean and troglobitic amphipods also show
some of these conditions. The diversity of Amphipoda in Baikal is
an enigma in the sense that the human observer has the tendency to
allot a great deal of time for the evolution of such a rich display of

MARINE GAMMARIDEAN AMPHIPODA 4]

TABLE 5.—Freshwater or inland saline water Gammaridea .

Family and Locality Genera Species

Gammaridae

Lake Baikal 37 230+

Palearctic—Nearctic 65+ 400+

South Africa 2 11

Australia-New Zealand 5 (+3+) 25+
Hyalellidae

Neotropical 1 28

African 1 iL

Australia-New Zealand 2 5

Miscellaneous 1 4
Bogidiellidae

Tethyan shores 1 9
Hadziidae

E. Tethyan shores 1 1
Caspiellidae

Caspian Sea 1 1
Corophiidae

Tropical—Cold-temperate, mainly river mouths 4 15
Calliopiidae

Tropics—subtropics 2 5
Haustoriidae

Cold-temperate 1 1
Eusiridae

Japan, Australia 2 3+
Metoediceros

Antiboreal 1 1

Totals 127+ 740+

morphs whereas the absence of a solid pre-Miocene fossil record leaves
us no confirmation of the group’s antiquity. Those Miocene fossils
are amber enclosures of species presumably still alive in European
freshwaters. One may become enmeshed in circular thinking that the
family Gammaridae must be primitive and the suborder Gammaridea
very old because not only is Baikal very old but that Gammaridae
came to dominate it and, therefore, Gammaridae were on the scene
first as well as very early. There are no current facts to counter that
argument but one has to maintain a reserve based on logic. Perhaps
only Gammaridae are widely preadapted to freshwater conditions.
Only the Hadziidae, Caspiellidae, Bogidiellidae, Hyalellidae, and a
few members of other families (table 5) also occupy freshwaters (or
inland waters). Gammaridae dominate the freshwaters of the world,
except for the Neotropical realm where the distantly related Hya-
lellidae replace the Gammaridae. The other minor families mentioned
above probably are closely related to the Gammaridae.

42 U.S. NATIONAL MUSEUM BULLETIN 271

The cold-water orientation of marine Gammaridea and the diversity
of brackish water and marine species (the Anisogammarus complex) in
the north Pacific Basin, with affinities to Baikalian genera, also sup-
port the thesis that Gammaridea may have had a fresh-water origin
in the vicinity of Baikal and then invaded the sea. The thesis becomes
stronger if one confines the theory only to the living members of the
family Gammaridae, for that family is generically impoverished not
only in tropical shallows but in all deep-seas of the world. Numerous
species of Maera, Elasmopus and Ceradocus are prominent in the
tropical sublittoral but generic radiation is comparatively low.
Isaeids and hyalids also dominate tropical shallows.

The importance of limnetic environments to the development of
Amphipoda is also attested to by the species-flock of Hyalellidae in
Lake Titicaca, a lake now of alpine character in the tropics but a lake
perhaps of warmer climes in earlier times. It seems more than hap-
penstance that most other talitroideans (like the Hyalellidae) are now
strongly oriented to the tropics. They include the fully (and only)
terrestrial Amphipoda that occur mainly on the islands of the Indo-
Pacific region, plus the beachhoppers that also have primarily tropical
affinities. Only the aquatic genus Hyalella has successfully invaded the
Nearctic realm dominated by Gammaridae (which also dominate
Palearctic and occur in African (Ethiopian) realms. Talitroideans are
more specialized or more advanced morphologically than members of
Gammaridae in uropods, mandibles, and antennae and no one has
been able to trace any direct relationships between the two groups,
even though one may see in the Beaudettiidae a species presumably
derived from Elasmopus (Gammaridae) that has undergone a revolu-
tionary change in the direction toward Hyale (Talitroidea).

One may consider that by the time fresh-water Gammaridae had
become dispersed to a cordilleran pathway contiguous to South
America that the latter continent had become disjunct or that the cir-
cumferential Sonoran belt afforded a major barrier. There may be a
strong competitional stress between Gammaridae and Hyalel-
lidae for the northward migration of one hyalellid into the Nearctic
realm does suggest the presence of a migratory pathway now in exist-
ence, but only one species has managed to make the crossing.

Dispersal of marine Gammaridae to the Neotropical realm also
must have come after the successful invasion by hyalellids, because it
would appear that even in tropical waters marine gammarids have been
able to invade limnetic environments.

All through the ancient Tethyan Sea marine gammarids invaded
underground waters, forming the nipharigid and other troglobitic
groups. The South African genus Paramelita may have its origin in
marine melitids and have undergone convergence towards the Baika-

MARINE GAMMARIDEAN AMPHIPODA 43

lian gammarus stock, for Schellenberg (1937a,b) has included that
genus with the ‘“gammarus” group. If South African ‘‘gammaruses’’
and the poorly known Australian members of the family have indeed
been the results of separate fresh-water invasions then the southern
hemisphere was completely blocked from an overland dispersal of
Kurasian gammarids.

These stocks of ‘‘gammarus” in the southern hemisphere may not
be so successful competitively against talitroideans as are Palearctic
stocks, for another group of talitroideans, the ‘‘chiltonias’’ have
evolved in the African and Australia-New Zealand provinces.

One need not adhere too closely to the thesis that Baikal formed an
evolutionary center for the reinvasion of the sea by its products.
Baikal and Titicaca may simply be dismissed as habitats open to
whatever contiguous marine groups were present. But this also implies,
without considering special ecological adaptations of the groups, that
if Gammaridae are indeed primitive and represent the survivors of a
base stock out of which came talitroideans, that the latter were under-
going their evolution on a tropical frontier while the tropicwards
dispersal of various marine Gammaridae was stagnating. Talitroids,
therefore, penetrated the tropical barrier in the move southward be-
fore marine or fresh-water gammarids. The importance of studying
ecological stress in the groups is apparent in the limited “‘later’’ success
of marine gammarids in partially occupying some of those southern
limnetic habitats on the tropical fringe. If there was temporal inde-
pendence among all these events in Baikal, South America, South
Africa, and Australia, and if marine Gammaridae and Talitroidea
were already fully developed before any of these limnetic invasions,
then one must entertain a strong and independent ecological success
of the Palearctic-Nearctic gammarids, for talitroideans have not been
able to populate those realms.

Terrestrial nonaquatic talitroids have never become eminently
successful. They occupy no major continent except where they have
been locally introduced into gardens. They are confined to Indo-
Pacific islands in the tropics, in moist, biotically impoverished environ-
ments. Beachhoppers on the strand of large continents have never
been able to migrate inland as far as we know.

Only two groups of Amphipoda thus have any clear relationship to
historical events that we have as yet been able to discover and on which
we may devote a great deal of refinement with promise of results other
than speculation. The meager evidence seems to indicate that Gam-
maridae had their major radiation in cold northern climes, either
marine or fresh water and that they had not reached either realm of
the southern hemisphere before those continents were blocked by some
kind of environmental barrier, leaving them open to the Talitroidea.

285-135 O - 69 - 4

44 U.S. NATIONAL MUSEUM BULLETIN 271

The question of interest is whether the Talitroidea existed simul-
taneous with these events or whether they presumably came much
later, and indeed whether they evolved directly through the gammarid
funnel; whether modern Gammaridae truly represent a base stock
to all other Gammaridea or whether they are simply a blind-alley
sidebranch coincidentally representing today the logical primitive
gvammaridean.

That Gammaridae and all other Amphipoda are heavily oriented
to cold water seems more than happenstance. We may determine
eventually that the tropics have the most numerous species of Am-
phipoda of any province but there is little current evidence that we
shall also find tropical genera to be the most numerous. Cold waters
(but not high polar) undoubtedly have the greater diversity at the
higher taxonomic levels. It is occasionally, or at least not always,
true that biotic groups have their highest diversity in their centers
of origin and I do not wish to imply that diversity and centers of
origin are correlated in the Amphipoda. Amphipoda have a bimodal
diversity anyway as cold waters obviously occur in two circum-
ferential bands. But the facts that the most primitive morphotype
has its center of diversity in cold Siberian Seas, that Gammaridae
have strongly radiated generically only in a cold Siberian lake and
not in isolated tropical lakes, that Nearctic-Palearctic fresh-water
Amphipoda clearly are more diverse generically than in tropical
fresh waters, that marine gammarid genera are most numerous in
cool waters, that a whole suborder of amphipods, the Hyperiidea is
confined primarily to cool pelagic waters, that the largest family of
marine amphipods, the Lysianassidae is confined largely to cool
waters, all suggest that Amphipoda are preadapted to cool waters
and have undergone the major “post-amphipodan”’ radiation in such
climes.

Identification Procedures

The identification of a gammaridean amphipod even at familial
levels so often requires a complete dissection and analysis of all
appendages and mouthparts that the procedure is considered to be
mandatory. This handbook cannot be utilized successfully by a non-
specialist without dissecting appendages (Appendix I) and observing
minute characters (checklist of Appendix II). Once the student has
gained some experience, however, portions of these procedures may
be skipped, for some families and many genera can often be recog-
nized without extensive analysis.

The procedures for identification described herein are manifold.
They include the memorization of a basic gammaridean plan, illus-

MARINE GAMMARIDEAN AMPHIPODA 45

trated in figures 1 and 2 and diagnosed on p. 50. Almost all other
families are defined by combinations of characters expressing either
minor specializations or minor simplifications of the basic gammari-
dean plan.

THE DIAGRAMMATIC KEY TO FAMILIES.—This key (figs. 1-53) is
composed of 54 boxes, 51 of which represent a family or superfamily
containing one or more illustrated characters in solid lines that dis-
tinguish it from the basic gammaridean (figs. 1-2b). For example, if
mouthparts are not illustrated they either resemble those of the
basic gammaridean or are so highly variable that they have no diag-
nostic differences from the basic gammaridean and are thus not
mutually exclusive. Each of these concordant families is noted in the
captions of the illustrated key and character differences in broken
lines are figured and explained where necessary. The diagrammatic
key is, therefore, not an absolute endpoint, for the taxonomist must
also check the illustrated boxes of families resembling the first pro-
visional identification.

The diagrammatic key is arranged in a way to deal first with those
families which have highly characteristic and easily observed morphol-
ogy. Thus, Lysianassidae (fig. 3), with their characteristic gnathopod
2 are presented first, followed by the similar Stegocephalidae (fig. 4),
which have characteristic coxae, head, mouthparts and mandibles,
followed by Ochlesidae (fig. 5) without maxillipedal palp, Lafystiidae
(fig. 6) with 2-articulate maxillipedal palp, and Acanthonotozomatidae
(fig. 7) with characteristic coxae. Following those groups is a pair of
families with cylindrical bodies, succeeded by seven families with
uniramous uropod 3. Succeeding that point (Amphilochidae, fig. 19)
all families have a biramous uropod 3, rarely with exceptional genera
or species. Other characters are abnormal to the basic gammaridean,
however, such as the small coxa 1 of Amphilochidae and some Leuco-
thoidae and the characteristic gnathopod 1 of Leucothoidae and
Anamixidae. There follows a group of families with various combina-
tions of coalesced urosomites, and degraded mandibles or acuminate
coxae. Commencing with the Oedicerotidae (fig. 27) the identifications
become increasingly difficult as the differentiations from the basic
gammaridean become less apparent: the Oedicerotidae have elongated
pereopod 5 and short, uncleft telson; Paradaliscidae (fig. 28) have
characteristic mandibles; Bateidae (fig. 30) have degraded gnathopod
1; Synopiidae (fig. 31) have massive, rostrate heads; the next four
families (figs. 32-35) have various combinations of feeble gnathopods,
degraded mandibles, special shapes of lower lips and maxillae and
various telsonic lengths and clefts; the next three families differ from
the basic gammaridean by poorly cleft telsons and degraded accessory
flagella; the next group has cleft telesons but degraded accessory

46 U.S. NATIONAL MUSEUM BULLETIN 271

flagella; then a group with normal accessory flagella but with various
combinations of degraded molars, characteristic rostra, coxae, and
pereopods, until finally the Gammaridae are reached.

After the Gammaridae is the group of isaeid-like families, most of
which form tubes, have spinning glands in the pereopods, bear short,
uncleft fleshy telsons, posteriorly unexcavate coxa 4, and normal
mouthparts but which often have increasingly complex or reduced
uropods difficult to separate from the network of families anterior to
the Gammaridae. The identifier must keep them in mind. Some
genera of the isaeid-like families are very difficult to recognize and
all of these family keys and diagnoses should be examined when
identifying presumed Corophiidae, Aoridae, Isaeidae, Ischyroceridae,
Ampithoidae, and Podoceridae. Perhaps it is wise to become familiar
with known variations in characters of these families for it is often
difficult to recognize spinning glands in pereopods (and some genera
of isaeid-like families lack them). Shallow-water isaeids are usually
strongly pigmented in browns and purples but this is not a reliable
character, because other families occasionally have strongly pigmented
species.

Example 1: Perhaps the observer determines from the completed
checklist (Appendix II), that the characters of the box Liljeborgiidae
(fig. 41) fit the specimen being identified. All characters match those
of the basic gammaridean except for the mandibular molar, which is
nontriturative; the accessory flagellum is multiarticulate, the mandible
has a 3-articulate palp, the maxillae are normal, maxillipeds have well
developed lobes and 4-articulate palps, gnathopod 1 is of normal or
enlarged size and is subchelate, gnathopod 2 is enlarged and sub-
chelate, all pleonites are free, pereopods are generalized, uropod 3 is
large and biramous, and the telson cleft. The caption lists related
families or those families with which Liljeborgiidae might be confused
and their boxes should be examined for special combinations of char-
acters not shared with the specimen at hand. For instance, the
Liljeborgiidae caption lists Gammaridae and Eusiridae as congruent
families, but the box of Eusiridae (fig. 39) indicates (1) by the absence
of a drawing of the mandible that it is either like the basic gam-
maridean and therefore has a triturative molar, or is of variable char-
acter; and (2) that the accessory flagellum is 0-2 articulate. If the
specimen being identified has an accessory flagellum of 3-plus articles
then it is not a eusirid; if it has a 2-articulate accessory flagellum and
triturative molar then it is not a liljeboriid; if it has a 2-articulate
accessory flagellum and a nontriturative molar it may be either a
eusirid or liljeborgiid and the reader would turn to the descriptions of
those families (p. 213 and p. 291) and compare them with the speci-
men. He would read the sections on Relationships to discover that a

MARINE GAMMARIDEAN AMPHIPODA 47

few genera of Eusiridae also have a nontriturative molar and thus
resemble Liljeborgiidae; but those genera lack an accessory
flagellum entirely; furthermore their gnathopods are unlike those of
Liljeborgiidae.

Example 2: Eusiridae (fig. 39) differ from the basic gammaridean
only by the reduction of the accessory flagellum to 0-2 articles shown
in solid lines and by the elongation of the telson. Also figured in the
box is the lower lip in broken lines to show a distinction from the
Pleustidae and the Astyridae; uropods 1 and 2 in broken lines show
a distinction from the Liljeborgiidae and the illustrated telsons differ
from those of the Calliopiidae.

Thus, the vestigial eusirid accessory flagellum is shared with many
other families but the special combination of basic gammaridean
morphology plus the telson, uropods, and lower lip is distinctive.

Also, an inference is made on the caption that the student refer to
the figures of the Vitjazianidae (fig. 34), and the Oedicerotidae (fig. 27)
for characters distinguishing those families from the Eusiridae, i.e.,
the simple first gnathopod and conjoint primary flagella of antenna 1
in the Vitjazianidae and the disproportionately long fifth pereopod in
the Oedicerotidae.

Not mentioned are other deviations, such as the occasional reduc-
tion of mandibular molars and 1-articulate first maxillary palps.
Several other families share these characters and may be partially
defined by them, but those families may be distinguished from the
Eusiridae by more important characteristics.

Example 3: Calliopiidae (fig. 38) differ from the basic gammaridean
only in the reduction of the accessory flagellum to 0-1 article and the
coalescence of the telsonic lobes. Also illustrated are the lower lip
which differs from that of the Pleustidae, coxa 4 to show a distinction
from the Isaeidae (= Photidae) and telsons to show faint distinctions
from the Eusiridae. Some calliopiids have isaeid-like coxa 4 but their
telsons are not fleshy and their pereopods lack glands.

Example 4: The diagnosis of Amphilochidae (p. 132) includes
only: ‘“‘Accessory flagellum absent; coxa 1 very small, partially hidden
by following coxae.”’ Next is stated ‘‘See [the family] Pleustidae. .. .”’
The description of that family implies that coxa 1 is ‘‘normal’’ in size
and placement. In sequence, the description of the mentioned Steno-
thoidae should be examined to see that uropod 3 is uniramous and
therefore distinct from uropod 3 of the Amphilochidae.

The Diagrammatic Key to Families is, at best, a method of narrow-
ing the search for a familial identification to a few possibilities, each
textual diagnosis and description of which must be examined for good-
ness of fit. A flexibility has been maintained in the diagrammatic key
because of space limitations in presenting a compact visual impression

48 U.S. NATIONAL MUSEUM BULLETIN 271

of a group as well as the possibility that the observer will handle
undescribed genera and families. The need to cross-check diagnoses and
test keys to genera in several families is a part of the identification
process.

The textual diagnoses, descriptions and relationships of the 53
families are presented in alphabetical sequence. The Hyalidae, Hyal-
llidae, and Talitridae are combined into the superfamily Talitroidea
for ease of identification. Each family has a written key to its genera
and each family is accompanied by several sets of figures illustrating:
(1) the characters of the generic keys, and (2) the variation in lateral
aspects and appendages of the several genera.

Perhaps the key to success in using the diagrammatic keys is the
need to memorize the plan of the basic gammaridean and to keep an
account of generalized characters, especially the generalization of
heads, feeble and strong gnathopods, anterior coxae, and elongation of
telsons and pereopods. Thus, Synopiidae are extremely difficult to
recognize because the subtle shapes of their heads are primary to their
identification. Acanthonotozomatidae, Paramphithoidae, and Stego-
cephalidae are recognized mainly by their anterior coxae.

WritTrEN KEys.—If the Diagrammatic Key to Families does not
produce results one may utilize the written key to families (pp. 106-116).
The written key is not as satisfactory as the Diagrammatic Key be-
cause the observer must repeatedly make ‘‘yes or no” decisions on
individual characters and no accomodation for undescribed genera and
families can be made. Simplicity of the keys requires occasional
repetition of taxa in order to account for morphological diversity.

Extraordinarily long keys are occasionally divided into sections in
order to improve their usefulness. The establishment of subgroups in
this manner is believed to be of assistance in retaining conceptual
images while the observer is proceeding through the maze of an
extremely complicated key.

DiaGnosrs.—Familial diagnoses are compatible with the Diagram-
matic Key to Families and are not mutually exclusive, for the diagnoses
only differentiate the families from the basic gammaridean. Inter-
related families are listed as a part of the diagnoses and are discussed
under ‘‘Relationships.’”’ These discussions are not necessarily repeated
in all possible places and the reader may have to turn from the
Synopiidae to the Liljeborgiidae, for example, in order to find a dis-
cussion of the interrelationships between the two families.

Genera are diagnosed in as brief and consistent form as possible, in
direct relationship to the keys. Where two genera are mutually dis-
tinguished by a character of no taxonomic importance to other genera
of a family, the alternatives of the character are italicized and the
related genus is stated. Such character alternatives are omitted from

MARINE GAMMARIDEAN AMPHIPODA 49

the diagnoses of other genera. Character alternatives that are known as
“combining characters,’’ because of their usage in keys, are occasion-
ally listed in diagnoses and italicized; they reflect the necessity to state
a general character that is diagnostic only because it occurs in unique
combination with all other characters stated for the genus under
question; the character is not necessarily restricted to the taxon in
question.

Each diagnosis has as its framework the conditions known for the
type-species or type-genus. In some families with few genera, the
diagnoses are confined to the keys. No doubt, the range of variation
permitted in the diagnoses is too narrow; time has not been allotted to
analyze the literature of each species in each genus in order to widen
the generic limitations because the literature is occasionally so defec-
tive that one would waste time to ponder, analyze, and argue possible
truths or to speculate on omissions. Many described species are un-
doubtedly misclassified.

Brief synonymies of recently established families are given but other-
wise such information may be found in Stebbing (1906) or J. L.
Barnard (1958a).

Type-species of each genus and their modes of selection are listed
below the generic synonymies. A modern reference, if available, is
given for the type-species. The number of species is included, along
with generalized distributional notes on the genus. ‘‘Littoral’’ includes
sublittoral, to 300 m; ‘‘bathyal” includes depths from 300 to 2000 m
(here) ; ‘‘abyssal’”’ exceeds 2000 m, and “‘hadal’’ 6000 m. Some species
in depths exceeding 300 m are pelagic but have not been so designated
because of uncertain data on collecting methods. Reference to terms
such as ‘arctic, antarctic, boreal,’’ is very imprecise because no exact
definitions are followed. If a genus is centered generally in the antarctic
as well as the subantarctic the terminology is simplified to “antarctic.”
The “arctic” includes the Norwegian Basin and its fringes as well as
the polar basin. ‘“‘Biboreal’’ denotes occurrence in northern and south-
ern hemispheres; ‘‘amphiboreal”’ denotes occurrence in both oceans of
the northern hemisphere.

Occasionally depths in meters are given where they have some inter-
est or precision. Distributional information in parentheses indicates
rarity in those situations.

50 U.S. NATIONAL MUSEUM BULLETIN 271

Diagnosis of the Basic Marine Gammaridean
(Many Marine Gammaridae)

Figures 1, 2

Accessory flagellum well developed, with four or more articles.
Primary flagellum of antenna 1 not basally conjoint.

Mouthpart field quadratiform from lateral view. Each mandible
with 3-articulate palp, article 3 longer than article 1; molar present,
with grinding surface composed of ridges and teeth (=triturative).
Lower lip with principal lobes undivided (unnotched) and not widely
separate. Each maxilla 1 bearing inner lobe, outer spinose lobe, and
strong unflexed palp of two articles. Each maxilla 2 with two well-
developed, setose lobes. Each side of maxillipeds with large inner
(proximal) lobe and outer (distal) lobe, and 4-articulate palp, article 4
claw-shaped (unguiform).

Gnathopods well developed and subchelate (‘‘powerful”), non-
lysianassid. Gnathopod 2 larger than gnathopod 1 in male, gnathopod
1 never larger than 2 in female, article 3 of gnathopod 2 short. Article 4
of pereopods 1-2 not extensively elongate. Pereopods 3-5 of con-
gruent structure and successively slightly longer.

Three pairs of large and subequal uropods present, all biramous,
rami subequal in length, lanceolate. Peduncle of uropod 3 not elongate.
Telson deeply cleft, of medium length.

Coxae forming elongate, rectangular plates with quadrate or
rounded distal edges, coxae 1—4 of uniform shape or slightly increasing
in size consecutively, coxa 4 excavate posteriorly.

All body segments free. Metasome only as long as last five pereonites
combined. Head subcuboidal, not ‘‘massive,’’ rostrum small.

MARINE GAMMARIDEAN AMPHIPODA 51

Epimeron

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Accessory Flagellum

Antennae

Ficure 1.—BASIC GAMMARIDEAN, part I, lateral aspect. Note the following char-
acteristics of the basic gammaridean: elongate antennal peduncles; head of medium
size, rostrum small or obsolete; eyes when present paired, lateral, composed of ommatidia
and lacking cuticular lenses; accessory flagellum multiarticulate; coxae 14 subequal to
each other in size, subquadrate in outline, coxa 4 posteriorly excavate; gnathopod 2 equal
to or larger than gnathopod 1; no articles of pereopods 1-2 differentially elongated;
pereopods 3-5 subequal to each other in length; all body segments free; uropods all
biramous, no uropod elongated; telson free. See diagnosis.

52 U.S. NATIONAL MUSEUM BULLETIN 271

Figure 2A.—BASIC GAMMARIDEAN. part II: a, head, lateral aspect [mouthparts ar-
ranged in quadrate bundle]; b, urosome, left lateral aspect, with uropods 1-3 and telson
[all segments free, none extremely elongate, no uropods extremely disproportionate, all
uropods biramous, telson freely articulate and not thickened dorsoventrally]; c, uropod
3 [well developed but of medium length, rami subequal to each other, no parts greatly
elongate]; d, telson [of medium length, cleft about three fourths of its length]; ¢, gnathopod
2 [subchelate, equal to or larger than gnathopod 1].

Symbols: 1, 2, 3, etc.=articles of appendage; c=coxa; |=lacinia mobilis; i=incisor;

ip=inner plate; ir—inner ramus; op outer plate; or=-outer ramus; p=palp.

MARINE GAMMARIDEAN AMPHIPODA 53

Ficure 2B.—BASIC GAMMARIDEAN. part II: f, mandible [with 3-articulate palp, molar
present and strongly triturative]; g, lower lip [lateral lobes close together, not perfectly
oval, inner lobes if distinct, not amalgamated]; h, maxilla 1 [well developed, spinose,
palp 2-articulate]; 7, maxilla 2 [well developed]; j, coxae 1-4 [all present, generally sub-
equal in size or consecutively slightly larger, quadrate in outline, coxa 4 excavate pos-
teriorly]; k, accessory flagellum [multiarticulate]; J, maxilliped [inner and outer plates
large, palp 4-articulate]. See fig. 2A for symbols.

54 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 3.—Lysiannassidae: a, antenna 1, note very short peduncle [see Sebidae]; b, gnatho-
pod 2, and an extreme variation of articles 6-7, as offset; note elongation of article 3
[highly characteristic overall shape, but see Sebidae, Stegocephalidae].

Stegocephalidae always lack molars and palps on mandibles, have some foliaceous portions

on maxillae, always have a 1—2~articulate accessory flagellum and a characteristic shape of

coxae 14. The combination of those characters never occurs in Lysianassidae.

MARINE GAMMARIDEAN AMPHIPODA 55

Ficure 4.—Stegocephalidae: a, head, left lateral aspect showing subconical grouping of
mouthparts, shortening of head, enlargement of epistomal region (epi); 6, mandible,
note absence of palp and molar and weakly developed styliform shape [see Acantho-
notozomatidae];.c, accessory flagellum [see Acanthonotozomatidae]; d,e, variation in
telson; f, coxae 1-4, left to right, note slight acumination of anterior coxae [overall shape

not occurring in Lysianassidae].

56 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 5.—Ochlesidae: a, head, left lateral aspect showing conical grouping of mouth-
parts [stippled portions are body and coxae]; b, accessory flagellum absent; c, maxillipeds,
note absence of palps [see Acanthonotozomatidae, Lafystiidae]; d, coxae 1-4, left to right,
note subacumination; e, mandible, note styliform shape; f, telson.

MARINE GAMMARIDEAN AMPHIPODA 57

Figure 6.—Lafystiidae: a. head and conical mouthpart bundle [stippled area is body];
b, accessory flagellum absent; c, mandible, molar absent, note styliform shape; d, max-
illiped, note only two palp articles [differing from nearly all other Gammaridea]; ¢, coxae
1-4, left to right [see Acanthonotozomatidae]; f, telson.

Gnathopod 1 is simple, gnathopod 2 is weakly subchelate. Similar to Acanthonotozomatidae

except for reduced maxillipedal palp, lack of pereonal and metasomal sculpture, and shapes

of coxae 1-4.

58 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 7.—Acanthonotozomatidae: a, head, note mouthparts arranged into conical bundle
[see Paramphithoidae, Stilipedidae]; b, accessory flagellum absent or vestigial; c, man-
dible, note styliform shape [see Stegocephalidae]; d,e, variation in coxae 1-4, left to right,
note acumination of one or more coxae ventrally [see Pardaliscidae, Lafystiidae, Stili-
pedidae]; f,g, variation in telson.

Ochlesidae have no maxillipedal palps whereas Acanthonotozomatidae have 3—4-articulate

palps. Lafystiidae have 2-articulate maxillipedal palps.

MARINE GAMMARIDEAN AMPHIPODA 59

Ficure 8.—Eophliantidae: body cylindrical, coxae short: a,b, mandibles, note absence
of molars and palps [see Talitroidea (except Najna), Kuriidae, Corophiidae, Podoceridae];
c, accessory flagellum absent; d, maxilliped [see Colomastigidae]; ¢,f,g, variations of uro-
pod 3 [see Prophliantidae]; A, telsons, cleft weakly or into two pieces [see Colomastigi-
dae]; 1, body, cylindrical, coxae very short [see Phliantidae, Talitroidea].

285-135 O - 69-5

60

U.S. NATIONAL MUSEUM BULLETIN 271

Je
,
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a >

Figure 9.—Colomastigidae: a, mandible, note absence of palp, molar (stippled) and deeply
serrate cutting edge [see Eophliantidae, Corophiidae, Podoceridae]; b, accessory flagellum
absent; c, uropod 3 [see Kuriidae, Eophliantidae]; d, maxilliped, note coalesced inner
lobes [see Eophliantidae]; ¢, body, subcylindrical, coxae short; f, telson [see Eophliantidae.
Prophliantidae].

MARINE GAMMARIDEAN AMPHIPODA 61

Ficure 10.—Phliantidae: a, mandible, note absence of palp and molar, projection to left
is ?tendon [see Podoceridae, Talitroidea, Kuriidae, Dexaminidae, Pagetinidae]; }, ac-
cessory flagellum absent; c, maxilla 1, note degradation of palp [see Dexaminidae]; d,
gnathopod 1 (and 2), note feeble condition (except Ceinina) [see Prophliantidae]; note
elongated article 3; e¢, body, depressed, rugose, coxae splayed [see Eophliantidae, Colomas-
tigidae]; f, uropod 3 of Phlias, probably erroneous; g,h, variation in uropod 3 [see
Acanthonotozomatidae, Prophliantidae]; 7,7, variation in telson, condition in figure 7
rarely occurring.

Kurtidae have poorly developed outer plates of the maxillipeds.

62 U.S. NATIONAL MUSEUM BULLETIN 271

Figure 11.—Dogielinotidae: a, mandible, note absence of palp [see Haustoriidae]; b, ac-
cessory flagellum absent [see Haustoriidae]; c, antenna 1, note spinosity [see Talitroidea];
d, maxilla 1, note degradation of palp; e, epistome and upper lip, left lateral aspect, note
pendant epistome [see Talitoridea]; f, any of pereopods 3-5, note heavy spinosity [see
Talitroidea]; g, uropod 3, uniramous [see Haustoriidae]; h, telson.

MARINE GAMMARIDEAN AMPHIPODA 63

Ficure 12.—Talitroidea: a, mandible, note absence of palp; b, accessory flagellum absent;
¢,d,e, variations of uropod 3, essentially uniramous, rarely with scale-like inner ramus;
f,g, extremes of telson.

Dexaminidae, Prophliantidae, and Kuriidae have coalescence of two or three urosomal

segments. Dogielinotidae have aspects of extreme fossorial adaptation such as spinose an-

tennae and pereopods, and a nasiform epistomal projection. Eophliantidae have cylindrical
bodies, short coxae and degraded mandibular molars. Phliantidae have depressed, rugose

bodies and splayed coxae. Note the convergence of Beaudettiidae, figure 42.

64 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 13.—Kurtidae: a, mandible, note absence of palp and presence of molar [see Pro-
phliantidae]; b, accessory flagellum absent; c, uropod 3, uniramous [see Prophliantidae,
Dexaminidae]; d, urosome, left lateral diagrammatic, all segments coalesced [see Tali-
troidea]; ¢, telson.

Eophliantidae have cylindrical bodies. Phliantidae have depressed, rugose bodies and
splayed coxae.

MARINE GAMMARIDEAN AMPHIPODA 65

Ficure 14.—Pagetinidae: a, mandible, note palp [see Eophliantidae. Phliantidae]; b, ac-
cessory flagellum absent [see Sebidae]; c, maxilliped, note absence of outer plate [see
Sebidae]; d, coxae 1-3, left to right [see Stenothoidae, Cressidae, Thaumatelsonidae];
eé, uropod 3, uniramous, ramus 1—2-articulate [see Pardaliscidae]; f, telson short [see Steno-
thoidae, Cressidae, Thaumatelsonidae]; g, side of urosome, segments 2-3 apparently
coalesced.

Sebidae have chelate gnathopods.

66

BULLETIN 271

U.S. NATIONAL MUSEUM

Ficure 15.—Sebidae: a, mandible, note absence of molar; b, accessory flagellum; c, uro-
pod 3, uniramous and elongate [see Lysianassidae, Leucothoidae, Anamixidae, Amphilo-

chidae]; d, maxilliped, [see Pagetinidae]; ¢, gnathopod 1, note chelateness [see Leucothoi-
dae]; f, gnathopod 2, note chelateness and elongation of article 3 [see Pagetinidae]; g,
telson; h, antenna 1, note elongation of peduncle [see Lysianassidae].

Stenothoidae, Thaumatelsonidae, Cressidae have a reduced coxa 1.

MARINE GAMMARIDEAN AMPHIPODA 67

Ficure 16.—Cressidae: a, mandible, note obsolescent molar; b, accessory flagellum absent;
c, coxae 1-4, left to right, note coxa 4 often not completely shield-like [see Thaumatel-
sonidae, Stenothoidae, Pagetinidae]; d, maxilliped, note reduced outer plate; ¢, urosome,
left, note telson (T) fused with segment 3 [see Stenothoidae, Thaumatelsonidae]; f, uropod
3, uniramous [see Amphilochidae]; g, telson.

Article 2 of pereopod 3 is always expanded in Cressidae, unlike Stenothoidae.

68 U.S. NATIONAL MUSEUM BULLETIN 271

Figure 17.—Thaumatelsonidae: a, mandible, note small, nontriturative molar; b, accessory
flagellum absent or vestigial; c, coxae 1-4, left to right [see Pagetinidae]; d, urosome,
left side (U=urosome, T=telson), note fusion of segments and thickened telson [see
Stenothoidae, Cressidae]; e¢, maxilliped, note smallness of outer plate; f, uropod 3, unira-
mous [see Amphilochidae]; g, telson.

MARINE GAMMARIDEAN AMPHIPODA 69

Ficure 18.—Stenothoidae: a, mandibles, note variations in palp, absence of triturative
molar; b, accessory flagellum vestigial or absent; c, maxilliped, note obsolescence of outer
plate [see Amphilochidae]; d, uropod 3, uniramous [see Amphilochidae]; ¢, coxae 1-4,
left to right [see Leucothoidae, Anamixidae, Pagetinidae, Phliantidae]; f, telson.

Thaumatelsonidae often have some of the urosomal segments coalesced and the telson

strongly thickened. Cressidae have the telson fused with pleonite 6, and always have an

expanded article 2 of pereopod 3, whereas Stenothoidae always have a slender article 2.

70 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 19.—Amphilochidae: a, coxae 1-4, left to right [see Leucothoidae, Pleustidae,
Calliopiidae, Stegocephalidae]; b, accessory flagellum absent [see Stegocephalidae]; c,
uropod 3, note biramous condition [Stenothoidae, Cressidae, Thaumatelsonidae]; d,
telson, entire [with one exception].

Stegocephalidae have mandibles always without molars and palps, whereas Amphilochidae

have mandibles with either one or both occurring together. Anamixidae have a ventral

cephalic keel replacing mandibles and maxillae.

MARINE GAMMARIDEAN AMPHIPODA 71

Ficure 20.—Anamixidae: a, oblique ventral view of head, showing absence of maxillae
and mandibles, presence of ventral keel (cross-lined), maxillipeds removed to show maxil-
lary protuberances [see Leucothoidae, Amphilochidae]; b, accessory flagellum absent;
c, gnathopod 1, note chelate condition [and absence of article 7 in this case] [see Amphilo-
chidae], occasionally this appendage is absent; d, maxillipeds, note amalgamated inner
lobes and absence of outer lobes; e¢, coxae 1-4, left to right [see most Leucothoidae];
f, telson.

W2 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 21.—Leucothoidae: a, mandible, note absence of molar; b, extremes of accessory
flagellum, always very small; c, maxilliped, note obsolescence of outer plate; d, gnathopod
1, note complex subchelation [see Cressidae, Thaumatelsonidae, and Stenothoidae, all
of which also have uniramous uropod 3]; e¢, telson, entire.

Sebidae have a uniramous uropod 3 and chelate gnathopod 2. Anamixidae, resembling

Leucothoidae strongly, lack maxillae and recognizable mandibles, and have a strongly

reduced coxa 1.

MARINE GAMMARIDEAN- AMPHIPODA 73

Ficure 22.—Prophliantidae: a, mandible, note absence of palp and recognizable molar [see
Atylidae]; b, accessory flagellum absent; c, uropod 3 [see Kuriidae, Talitroidea, Phliantidae];
d, urosome, left lateral aspect, note amalgamation of all segments and telson [see Tali-
troidea, Dexaminidae]; ¢, telson.

Eophliantidae have cylindrical bodies and reduced coxae.

74 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 23.—Dexaminidae: a, mandible, palp absent [molar evanescent in Anatylinae]; 3,
accessory flagellum absent; c, urosome, left lateral aspect, note amalgamation of uro-
somites 2-3 [see Prophliantidae, Talitroidea]; d, telson.

Phliantidae have the palp of maxilla 1 degraded, and depressed rugose bodies with splayed

coxae.

MARINE GAMMARIDEAN. AMPHIPODA 75

Figure 24.—Lepechinellidae: a, accessory flagellum [see Pleustidae, Ampeliscidae]; ),
coxae 1-4, left to right, note acumination [see Atylidae, Ampeliscidae]; c, urosomites 1-3,
left to right, note amalgamation of segments 2-3, presumably occurring in all lepechinel-
lids [see Pleustidae, Paramphithoidae]; d, telson.

Presumably all lepechinellids have coalesced inner lobes on the lower lip and thus differ

from Paramphithoidae. Kuriidae, Dexaminidae, and Prophliantidae lack mandibular palps,

285-135 O - 69 - 6

76 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 25.—Atylidae: a, head [see Ampeliscidae]; b, extremes of accessory flagellum;
c, urosome, left lateral view, showing amalgamation of urosomites 2-3; d, telson.
Lepechinellidae have acuminate coxae 1-4. Shallow-water Ampeliscidae have cuticular

cephalic lenses. Dexaminidae, Prophliantidae, and Kuriidae lack a mandibular palp.

MARINE GAMMARIDEAN AMPHIPODA aoe

Figure 26.—Ampeliscidae: pereopods glandular: a, head, left lateral aspect, showing
cuticular lenses and dotted lines indicating internal pigment (lenses unique in Gam-
maridea but absent in deep-sea ampeliscids) [see Atylidae]; b, accessory flagellum absent;
c, pereopod 4 [often longer than pereopod 5 and of different shape]; d, pereopod 5; ¢,f,
variations in telson; g, urosome, left lateral aspect, note amalgamation of urosomites 2-3.

Atylidae have ommatidial eyes and lack the special combination of pereopods 4 and 5

shown here.

78 U.S. NATIONAL MUSEUM BULLETIN 271

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Ficure 27.—Oedicerotidae: a,b,c, pereopods 3, 4, 5, note elongation of pereopod 5 [see
Calliopiidae, Pleustidae, Synopiidae, Isaeidae and related families, Laphystiopsidae];
d, accessory flagellum absent [see Synopiidae, Pardaliscidae]; ¢, telson; f, uropod 3, note
elongated peduncle [see Calliopiidae, Pleustidae, Laphystiopsidae].

Isaeid-like families have a thick, fleshy telson. Oculate oedicerotids mostly have dorsally
contiguous or coalesced eyes.

MARINE GAMMARIDEAN AMPHIPODA

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Ficure 28.—Pardaliscidae: a, mandibles, left with palp not illustrated, note absence of
molars and foliate cutting edges [see Laphystiopsidae]; b, accessory flagella, usually
multiarticulate, absence extremely rare [see Stilipedidae]; c, coxae 1-7, very short [see
Stilipedidae, Acanthonotozomatidae]; d, maxilliped, note obsolescence of inner plate
[see Acanthonotozomatidae, Astyridae, Laphystiopsidae]; ¢,f, extremes of telson.

79

80 U.S. NATIONAL MUSEUM BULLETIN 271

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Ficure 29.—Stilipedidae: a, mandible [see Synopiidae, Hyperiopsidae, Astyridae, Laphy-
stiopsidae]; b, extremes of accessory flagellum, vestigial or absent [see Pardaliscidae
which usually have multiarticulate condition]; c,d, maxillae 1, 2, foliaceous [see Par-
daliscidae, never having foliaceous maxilla 2, Laphystiopsidae]; ¢, gnathopod 1, simple
(gnathopod 2 also); f, telson.

Acanthonotozomatidae have their mouthparts grouped as a conical bundle and only one

acanthonotozomatid genus has foliaceous maxillae.

MARINE GAMMARIDEAN AMPHIPODA 81

Ficure 30.—Bateidae: a, head (H) and pereonites 1-3 with coxae 2-4 [coxa | vestigial or
absentl: b, accessory flagellum absent; c, gnathopod 1, composed of coxa and article 2,
other articles absent [exclusively characteristic]; d, telson.

82 U.S. NATIONAL MUSEUM BULLETIN 271

Figure 31.—Synopiidae: a,b,c, variations of head, head very large, usually galeate, with
large downturned rostrum, except a few genera with head like fig. c; head like fig. ¢ also
occurring in Astyridae [see Vitjazianidae, Eusiridae]; d, accessory flagella, absence prob-
ably erroneous, usually more than 3-articulate [see Eusiridae]; ¢, gnathopods feeble;
f, mandible, note short article 3 of palp (except in Bruzeliopsis) [see Astyridae, Par-
daliscidae, Eusiridae]; g,h, telsons [see Astyridae, Oedicerotidae].

Astyridae have a uniarticulate accessory flagellum, and widely spaced outer lobes of the

lower lip. Argissidae have a unique combination of shapes of coxae 1-4. Pardaliscidae have

very short coxae. Oedicerotidae have elongated fifth pereopods. Hyperiopsidae have bent
maxillary palps and elongated fourth articles of pereopods 1-2. Occasionally Astyridae and

Pardaliscidae have heads like figure b. Oculate synopiids have dorsally contiguous or co-

alesced eyes, occasionally with anteroventral accessory ommatidia.

MARINE GAMMARIDEAN AMPHIPODA

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Ficure 32.—Astyridae: a, mandible, note setose and conical but nontriturative molar
[see Synopiidae, Pardaliscidae, most Eusiridae, Vitjazianidae]; b, accessory flagellum
[see Synopiidae, Oedicerotidae, Vitjazianidae, Hyperiopsidae]; c, lower lip, note widely
separated outer lobes [highly characteristic] [see Paramphithoidae]; d, gnathopod 1

(2 is similar) [see Liljeborgiidae, most Oedicerotidae, Vitjazianidae]; e,f, extremes of
telson.

Eusiridae generally have an elongate telson, and often enlarged gnathopods. Hyperiopsidae
have bent maxillary palps, and elongated fourth articles of pereopods 1-2. Haustoriidae
generally have either a multiarticulate accessory flagellum, or a large or triturative molar.
Astyrid cephalons may resemble those of Synopiidae (see fig. 31). Two paramphithoid
genera, with nontriturative molars and weakly acuminate coxae, resemble Astyridae closely.

83

84 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 33.—Hyperiopsidae: a,b, maxilla 1, note enlarged condition of palps, one of them
bent; c, accessory flagellum [see’ Ampeliscidae and Stilipedidae]; d,e, gnathopods 1, 2;
f, pereopod 1 (and 2 similar), note elongation of article 4 and either slender or inflated;
g,h, extremes of telson.

Combination of elongate article 4 of pereopods 1-2 and bent maxillary palps differentiates

Hyperiopsidae from Astyridae, Synopiidae, Pardaliscidae, Vitjazianidae, Calliopiidae, and

Eusiridae.

MARINE GAMMARIDEAN AMPHIPODA 85

Ficure 34.—Vitjazianidae: a, peduncle (stippled) of antenna 1, very short, and base of
flagellum conjoint (a long article 1 of antenna 1 flagellum ) [see Eusiridae, Gammaridae,
Melphidippidae]; b, accessory flagellum [see Astyridae, Stilipedidae]; c, gnathopod 1 [see
Liljeborgiidae]; d, telson.

Maxilla 1 normal [see Hyperiopsidae, Stilipedidae]. Article 4 of pereopods 1-2 of normal

length [see Hyperiopsidae]. Uropod 3 of normal length [see Melphidippidae]. Mandibular

molar large and triturative [see Astyridae, Stilipedidae]. Lower lip normal [see Astyridae].

Head small and poorly rostrate [see Synopiidael].

86

U.S. NATIONAL MUSEUM BULLETIN 271

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Ficure 35.—Laphystiopsidae: a, mandible, note simple molar (general shape differing
from that of Stilipedidae) [see Isaeidae and related families]; b, accessory flagellum
[see Astyridae]; c, coxa 4, left aspect, of only type-genus; d,e, gnathopods 1, 2 (simple)
[see Oedicerotidae, Isaeidae and related families]; f, telson.

One genus perhaps erroneously included here, see text. Probably a compound and artificial

family. Lower lip resembling Pleustidae but type-genus Laphystiopsis distinguished by

having reduced coxae and simple gnathopods. Prolaphystius has elongated urosomite 1.

Family scarcely distinct from Calliopiidae but Laphystiopsis bearing large spatulate rostrum.
See lower lip of Astyridae. Oedicerotidae usually have subchelate gnathopods, large coxae,

and elongated peduncles of uropod 3.

MARINE GAMMARIDEAN AMPHIPODA 87

Ficure 36.—Paramphithoidae: (incl. Amathillopsidae): a, coxae 1-4, left to right, note
acumination [see Pleustidae, Eusiridae, Calliopiidae]; b, extremes of accessory flagellum;
c, gnathopod 2, note feeble condition; d,e, extremes of telson.

Acanthonotozomatidae lack a distinct mandibular molar and have their mouthparts ar-

ranged in a conical bundle. Lepechinellidae have pleonites 5-6 amalgamated, extremely

slender second articles of pereopods 3-5 and coalesced inner lobes of the lower lip. Haustori-
idae have multiarticulate accessory flagella and strongly spinose pereopods.

Astyridae have nontriturative mandibular molars and widely separate outer lobes of the

lower lip. Two genera of Paramphithoidae have nontriturative molars and may be examined

in the key to the genera of Astyridae.

88 U.S. NATIONAL MUSEUM BULLETIN 271

Figure 37.—Pleustidae: a, lower lip [see Calliopiidae]; b, accessory flagellum, absent or
uniarticulate [see Liljeborgiidae]; c, coxa 4, left aspect, posteriorly excavate [see Isaeidae
and related families, Laphystiopsidae]; d, telson [see Liljeborgiidae, Eusiridae, Lepechinel-
lidae].

Paramphithoidae and Lepechinellidae have one or more of coxae 1-4 cuspidate or pointed

| Mesopleustes appears to be assignable to either Paramphithoidae or Pleustidae]. Oediceroti-

dae have a disproportionately elongated pereopod 5 and short telson. Amphilochidae have

a very small coxa 1. Laphystiopsidae may be synonymous with Pleustidae.

MARINE GAMMARIDEAN AMPHIPODA

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Ficure 38.—Callioptidae: a, lower lip, with or without inner lobes [see shape of lower lip
of Pleustidae]; b, accessory flagellum either absent or uniarticulate; c, coxa 4, left aspect,
posterior edge excavate [see Isaeidae and related families]; d,e,f, variations in telson [see
Eusiridae, Gammaridae].

Oedicerotidae have disproportionately long fifth pereopods (but see Paracalliope). Laphys-

tiopsidae have a poorly developed mandibular molar. Paramphithoidae have one or more

of coxae 1-4 cuspidate or pointed. Amphilochidae have reduced first coxae. Some calliopiids

have nonexcavate coxa 4 but lack pereopodal glands of isaeid-like families ana do not have
fleshy telsons.

89

90 U.S. NATIONAL MUSEUM BULLETIN 271

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Ficure 39.—Eusiridae: a, lower lip, with or without inner lobes [see shape of lower lip of
Pleustidae, Astyridae]; b, accessory flagellum, absent, uni- or biarticulate [see Gammaridae
which have a few species with 2-articulate accessory flagella]; c,d, uropods 1, 2, note
shortened outer rami [most Gammaridae do not]; ¢,f, extremes of telson [see Calliopiidae,
Pleustidael].

Astyridae have nontriturative mandibular molars but all Eusiridae except Eusirella have

strongly triturative molars. Vitjazianidae have simple first gnathopods and basally con-

joint primary flagella of antenna 1. Synopiidae have massive heads with deflexed rostra,
usually have multiarticulate accessory flagella and feeble gnathopods whereas most Eusiridae
have powerful gnathopods. Oedicerotidae have pereopod 5 disproportionately elongate
in comparison with pereopods 3 and 4. Paramphithoidae always have the unique combina-
tion of feeble gnathopods and one or more acuminate coxae 1-4. Liljeborgiidae have the
unique combination of powerful gnathopods and nontriturative mandibular molar. Gam-
maridae have pleonites 1-3 together not exceeding length of 5 pereonites together, whereas

Eusiridae have these pleonites (metasome) exceeding six pereonites in length; all Eusiridae

except one genus have elongate telsons, whereas Gammaridae have short telsons not or

scarcely exceeding the length of pleonite 6.

MARINE GAMMARIDEAN AMPHIPODA 91

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Ficure 40.—Melphidippidae: a, extremes of accessory flagellum; b, coxae 1-7, left to right,
note shortness; c, gnathopod 1, feeble (gnathopod 2 also feeble); d,e,f, uropods 1, 2, 3,
[note extreme elongation of uropod 3, see Gammaridae]; g, telson.

Melphidippidae differ from Vitjazianidae by the normally elongated peduncle of antenna 1.

See couplet 7, Section A of written key to families (p. 107) for definition of elongate uropod 3.

285-135 O - 69 - 8

92 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 41.—Liljeborgiidae: a, mandible (molar weak and nontriturative) [see Gammaridae,
Haustoriidae and most genera of Eusiridae]; b, extremes of accessory flagellum [see
Pleustidae]; c, gnathopods 1-2 (powerful) [see Haustoriidae, Astyridae, Vitjazianidae];
d, telson [see Pleustidae].

Article 1 of mandibular palp in Liljeborgiidae often elongate.

MARINE GAMMARIDEAN AMPHIPODA 93

Figure 42.—Beaudettiidae: a, mandible (palp absent) [see Gammaridae]; b, accessory
flagellum (2-articulate); c, coxa 4, left aspect (not posteriorly excavate); d, uropod 3
(inner ramus vestigial); ¢, telson, very short, entire [see Gammaridae].

94 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 43.—Argissidae: a, accessory flagellum [see Ampeliscidae]; b, coxae 1-4, left to right
[see Gammaridae, especially Megaluropus; Haustoriidae]; c, gnathopod 1 [2 is similar],
feeble; d, telson.

Shallow-water Ampeliscidae have external cephalic cuticular lenses and amalgamated

urosomites 2-3. Shallow-water Argissidae have four internal, bigeminous, lenticular, ocular

bodies.

MARINE GAMMARIDEAN AMPHIPODA 95

Ficure 44.—Phoxocephalidae: a,b, lateral and dorsal views of head, note wide or narrow
(stippled) rostrum in fig. b; c, accessory flagellum; d, pereopod 4 (elongated) [see Gam-
maridae]; ¢, pereopod 5; f, telson.

Combination of head with large rostrum and configurations of pereopods 4 and 5 is char-

acteristic [see Haustoriidae].

96 U.S. NATIONAL MUSEUM BULLETIN 271

gnathopods feeble

Figure 45.—Haustortidae: a,b, head, dorsal views showing small and large rostra (occa-
sionally genera occur with no rostra; those with large rostra (fig. b) have pereopod 5 like
fig. e¢, thus not shorter than pereopod 4; genera with large rostra and pereopod 5 like fig. f
are assigned to Phoxocephalidae); c, variational extremes in accessory flagellum [see
Ampeliscidae, Astyridae, Paramphithoidae]; d, pereopod 4, note spinosity and occasional
absence of article 7; ¢,f, extremes of pereopod 5, note spinosity [see Paramphithoidae];
g,h, telsons; 1, gnathopods highly variable but feeble [see most Gammaridae, Liljeborgiidae].

Ampeliscidae have amalgamated urosomites 2—3. Dogielinotidae lack mandibular palps

and have a uniramous uropod 3. Astyridae have a characteristic lower lip. Argissidae have

a special configuration of coxae 1-4.

MARINE GAMMARIDEAN AMPHIPODA 97

Figure 46.—Gammaridae, marine: a,b, two kinds of coxa 4, left aspect; c,d, two extremes
of accessory flagellum, 2- or multiarticulate, rarely 1-articulate; ¢,f,g, three kinds of telson
(arrows showing possible combinations with kinds of coxae, thus poorly excavate coxa 4
never combined with uncleft telson, except in Falklandella); h, uropod 3 (common kind);
1, uropod 3 (rare kind). Figures a,d,e and h represent a combination similar to the basic

gammaridean.

98 U.S. NATIONAL MUSEUM BULLETIN 271

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Ficure 47.—Isaeidae (incl. Photidae), pereopods usually glandular: a, coxa 4, note lack
of posterior excavation (coxa 4 may be long or short) [see some Gammaridae, Pleustidae,
Calliopiidae and related families]; b, extremes of accessory flagellum; c,d,e, variations in
uropod 3; f, telson, thick, short, and fleshy [see Calliopiidae, Pleustidae, Oedicerotidae].

Five isaeid genera have uniramous uropod 3 and might be confused with numerous other

gammaridean families having uncleft telsons. But Cressidae, Stenothoidae, and Thauma-

telsonidae have reduced first coxae; Eophliantidae have cylindrical bodies; Phliantidae,

Talitroidea, and Dogielinotidae lack mandibular palps. Sebidae have subequally large,

chelate gnathopods. Pagetinidae have evanescent mandibular molars and poorly developed

outer plates of maxillipeds. See Aoridae, Ischyroceridae, Ampithoidae, Corophiidae, Podo-
ceridae, and Cheluridae to follow for diagnostic differences.

MARINE GAMMARIDEAN AMPHIPODA 99

Ficure 48.—Aoridae, pereopods glandular, like Isaeidae but gnathopod 1 larger than
gnathopod 2 and bearing male sexual modifications: a, coxa 4, note absence of posterior
excavation; b, extremes of accessory flagellum; c,d, gnathopods 1, 2, note 1 larger than 2
[see Isaeidae]; ¢,f,g,h, variation in uropod 3 [see Ischyroceridae, Corophiidae, Cheluridae];
i, telson, short, thick, fleshy.

See Isaeidae for characters differentiating genera with uniramous uropod 3 from families

with uniramous uropod 3. Podoceridae have elongate urosomite 1. Cheluridae have coalesced

urosomites 1-3. Corophiidae have depressed urosomes. Ischyroceridae and Ampithoidae
have elongate peduncles of uropod 3. Ampithoidae have characteristic lower lips.

100 U.S. NATIONAL MUSEUM BULLETIN 271

Ay NK
: We.

Figure 49.—Ischyroceridae, pereopods glandular, like Isaeidae and Aoridae but uropod 3
with elongate peduncle and outer ramus with minute distal ornaments: a, coxa 4, note
lack of posterior excavation; b, extremes of accessory flagellum; c,d, uropod 3 and enlarged
rami, note elongate peduncle and minute nonarticulate ornaments on outer ramus [see
Isaeidae, Aoridae, Corophiidae, Ampithoidae]; ¢, telson, short, thick, fleshy.

Ampithoidae have characteristic lower lips. Podoceridae have elongate urosomite 1.

MARINE GAMMARIDEAN AMPHIPODA 101

Figure 50.—Ampithoidae, pereopods glandular, like Isaeidae and Aoridae but peduncle of
uropod 3 elongated and lower lips with characteristic shape, usually with notched outer
lobes: a, coxa 4, note lack of posterior excavation; b, extremes of accessory flagellum;
c,d, lower lips, outer lobes usually notched or excavate medially [see Ischyroceridae,
Isaeidae, Aoridae, Corophiidae]; ¢, uropod 3, note elongate peduncle, short rami, presence
of one or two large distal spines on outer ramus [see Ischyroceridae, Isaeidae, Aoridae,
Corophiidae, Cheluridae]; f, telson.

Podoceridae have elongate urosomite 1. See text for Jassa (p. 279), the ischyrocerid often

confused with ampithoids.

102 U.S. NATIONAL MUSEUM BULLETIN 271 |

ae
Nee 3

b

Vine

a Ue ite:
G4ggg 438s!

Ti
2 J

OO)

Figure 51.—Corophiidae, pereopods usually glandular; like Isaeidae, Aoridae, Ischyroceri-
dae, but urosome depressed; uropod 3 almost always uniramous: a, extremes in accessory
flagellum; 5, urosome, left lateral aspect to show depression and occasional fusion of
segments; c, coxa 4, note lack of posterior excavation [shape highly variable]; d,e,f,g,h,
variations in uropod 3, usually uniramous [see Isaeidae for distinctions from distantly
related families]; 7,7, extremes in telson.

Genera with uropod 3 like figs. e and h are probably assignable to Ischyroceridae. Genera

with enlarged gnathopod 1 are probably assignable to Aoridae, and others, except Corophium,

are assignable to Isaeidae. Podoceridae have extremely elongated urosomite 1. Colo-
mastigidae and Eophliantidae have degraded mandibles.

MARINE GAMMARIDEAN AMPHIPODA 103

Ficure 52.—Podoceridae, pereopods probably never glandular: a, urosome, left lateral
aspect, showing elongated urosomite 1, typically depressed condition and frequent
amalgamation of urosomites 2 and 3 (=2) [see Isaeidae, Ischyroceridae, Aoridae, Coro-
phiidae, Cheluridae, Ampithoidae]; b, extremes of accessory flagellum; c, uropod 1; d,e,f,
variation in uropod 2; g, coxa 4, note unexcavate posterior edge; h,7, extreme variations
of uropod 3; 7, telson.

Colomastigidae and Eophliantidae have degraded mandibles.

104 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 53.—Cheluridae: a, urosome, left lateral aspect showing amalgamation of all
segments and depressed condition, uropods 1-3 (U-1, 2, 3) and telson (T) [see Podoceridae,
Corophiidae]; b, accessory flagellum; c, coxa 4, note absence of posterior excavation; d,e,f,
uropods 1, 2, 3 [see Corophiidae, Podoceridae]; g, telson.

MARINE GAMMARIDEAN AMPHIPODA 105

Order Amphipoda

Draenosis.—Peracarid Malacostraca lacking carapace, thus having
all but one or two of the thoracic segments freely visible; one thoracic
segment carrying maxillipeds fused to head, occasionally next thoracic
segment carrying gnathopods also fused to head (Caprellidea),
followed by seven (occasionally six) visibly articulated thoracic seg-
ments, each bearing paired appendages, followed by six abdominal
segments or their macroscopic remnants (except Caprellidea), first
three (pleon) usually bearing paired biramous pleopods, remaining
three (urosome) bearing paired biramous uropods; telson freely articu-
late in primitive and majority of members; head with two pairs of
antennae, first occasionally biramous; maxillipeds lacking exopodites;
heart mainly thoracic; respiration thoracic with gills attached to coxae
(or their remnants) of segments 2-7 (variable); eyes sessile or rarely
borne on unstalked cephalic scale; eggs carried in female brood pouch
on ventral thorax formed of 2 to 4 (or 5) pairs of lamellae attached
to coxae 2-6.

There are no radically degenerate and fully endoparasitic Gam-
maridae, although a number are inquilines, ectoparasites, and com-
mensals which have sucking mouthparts and prehensile mechanisms
on their appendages.

Gammaridea, especially the Gammaridae, lie closest to the logical,
primitive stem of the Amphipoda and almost all evolutionary lines
from Gammaridae-like ancestors are based on simplification of primi-
tive parts. The Caprellidea have reduced but a vestige of the abdomen
(except Cercops and Caprogammarus) and the Hyperiidea have lost
the maxillipedal palps. Reduction of coxae is common in both of those
suborders. Ingolfiellidea have lost all pleopods but some have devel-
oped a movable scale on which the eyes are borne when present.

One family of Gammaridea, the Ochlesidae, have lost the maxilli-
pedal palps but their resemblance to Gammaridea in coxae and body
shape and their presumed benthic habits have caused their assignment
to the Gammaridea.

Key to the Suborders of Amphipoda

1. Abdomen normally vestigial, usually lacking large pleopods or uropods, except
for microscopic vestiges not exceeding two pairs, gills two or 3 pairs and
brood lamellae two pairs each, head and segment bearing gnathopod 1
usually immovably coalesced.

CAPRELLIDEA (skeletonshrimps, whalelice, 250 species)

Abdomen well developed, with six segments or their gross vestiges apparent,
bearing some gross appendages, gills and brood lamellae exceeding three
pairs.each, headiand thoraxdrees e-news ete co lye copes 2

106 U.S. NATIONAL MUSEUM BULLETIN 271

2.

3.

Maxillipeds absent . . . . . . . HYPERIIDEA (pelagic, ?500 species)
Maxillipeds bearing palps (except one rare family) . ae 3
Movable compound dactyl of gnathopods 1-2 formed of antieles 6-7 tayuelnen:
head occasionally with articulate ocular scale; pleopods when present, all
reduced in size, leaf-like.
INGOLFIELLIDEA (caves and marine, 15+ species)
Dactyl of gnathopods 1—2 formed only of article 7; head never bearing ocular
scale; pleopods biramous, elongate, rarely reduced except in terrestrial
species.

GAMMARIDEA (marine and limnetic scuds; beachhoppers; 3300 species)

Nomenclatural Changes in Gammaridean Families

Amathillopsidae are combined with the Paramphithoidae.
Anatylidae are combined with the Dexaminidae.
Hyalellidae are included with the superfamily Talitroidea.
Hyalidae are included with the superfamily Talitroidea.
Photidae are combined with the Isaeidae.

Pontogeneiidae are combined with the Eusiridae.
Talitridae are included with the superfamily Talitroidea.
Tironidae are combined with the Synopiidae.

Key to the Families of the Suborder Gammaridea '!

(“‘exceptional”’ refers to rare occurrence of familial members in this

category)
SECTION A
Article 3 of gnathopod 2 elongate (fig. 3). ........ . . Section B
Article 3 of gnathopod 2 not elongate ... . ER CERE NC ENIEAN G" 30/24

Urosomites 1-2 coalesced, urosomite 3 free (fig. 105).
some species of PHLIANTIDAE; Kamaka (Corophiidae) ; and Chevalia (Isaeidae)
Urosomites 2-8 or 1-8 coalesced. . . .-.....4. .. . . Section ©
Urosomites separate .... A Vale
Coxa 1 small or absent, mostly Ridden ray a Ralllowmne coxa , one of following
coxae longer than wide) (occasionally gnathopod 1 absent or vestigial).
Section D
Coxa 1 small or large but usually subequal to coxa 2 and never hidden by
following coxae (occasionally coxa 1 partially hidden but all following

coxae wider than long), (gnathopod 1 always fully developed) . . . . 4
Uropod 3 uniramous, or lacking rami, or absent. . . .. . . . Section E
Uropod 3 biramous, inner ramus often reduced .... . oy STEM eo:
Mandibular palp absent (use alternate if Section F Tea HaACtORy, because

of uncertain loss of palp) ... . cA gest Cpa SeCtONmn
Mandibular palp present (with very rare , exesoviion cesent) 2. of Sa eciee ete inan

Mandibular molar absent or if present not triturative, lacking numerous
ridges and teeth (occasionally with articulate spines), occasionally large
or immense and dominating mandible (fig. 166d) . . . . . . Section G

1 Gurjanova (1962) also has a modern key to the families.

10.

11.

12.

MARINH GAMMARIDEAN AMPHIPODA 107

Mandibular molar well developed, triturative, bearing ridges and teeth,
never extremely large or dominating mandible... . . SUMATRAN Lrg,
Peduncle of uropod 3 elongate, as long as or longer than rami of uropods 1-2
(fig. 40) and more than twice as long as telson and peduncle of uropod 2,
uropod 3 also greatly exceeding apices of either uropods 1 or 2 and rami
elongate, subequal and nearly as long as rami of uropods 1-2 (uropod 3 is
usually missing on individuals of this category but melphidippids are also
recognized by the presence of dorsal teeth and serrations on the pleon, in
combination with short, subequal and evenly quadrate anterior coxae,
plus strong hemispherical lateral ocular bulges on head). ...... 82
If peduncle of uropod 3 elongate as above then uropod 3 not strongly
exceeding apices of uropods 1—2; if peduncle of uropod 3 twice as long as
telson then uropod 3 not exceeding apices of uropods 1-2 or peduncle not
as long as rami of uropods 1-2; if uropod 3 greatly exceeding apices of
uropods 1—2 then inner ramus short and scale-like or peduncle not elongate
and not more than 1.2 times as long as peduncle of uropod 2, or rami much
shorter than peduncle; thus uropod 3 not like that of first part of this
couplet... . 2 9
Coxae 2-3 longer ben. pron Dleon Hoteally aroun mercoued I very
conspicuously longer than 3 or 4, head without lateral ocular bulges
(head often strongly rostrate) . . (exceptional) OEDICEROTIDAE (p. 373)
Coxae 2-3 very short and broader than long, pleon dorsally toothed and
serrate, pereopods 3-5 very long but subequal in length to each other,
head with strong hemispherical ocular bulges on sides.
MELPHIDIPPIDAE (p. 368)
Coxae 2-3 about as long as broad, pleonite 4 with one dorsal tooth, pereopods
3-5 long but subequal in length, head without lateral ocular bulges.
Casco (Gammaridae) (p. 238)
Coxae 1-3 successively and very distinctly shortened (fig. 43), (and see mimic
Megaluropus in Gammaridae and Casco with anteriorly acuminate coxa

lin Gammaridae) ... . . . . ARGISSIDAE (p. 157)
Coxae 2 or 3 always as long as or Neceer hen c Coxaelen ss Ory Renee ss tl
Telson entire, or emarginate, very short, coxa 4 not excavate posteriorly

(pereopods often glandular) . .. . Hae aaa SCION: le

Telson entire or cleft, short or long, coxa 4 ignellie excavate posteriorly or
acuminate (coxa 4 always excavate posteriorly or acuminate when telson
entire, but see Parapherusa in Section F and in Gammaridae), (pereopods
not glandular) ...... Bi atone red

Head massive (see dismaliion § in rlgssay he or eh Eronels downturned
rostrum, or “‘shark-nose’’ rostral projection ie 3l1c), [gnathopods weak
GSI aciico anneal: PARRY HDD

Head not massive or rose cf estan) on massive rend Hot downturned
(Hyperiopsidae have massive head, no rostrum; Pleustidae and some
Phoxocephalidae have downturned rostrum on small head)... . 13

Pereopod 5 conspicuously longer than pereopods 3-4, at least 1.5 times as
long as pereopod 4, accessory flagellum absent, or l-articulate and short if
present, peduncle of uropod 3 always elongate and telson always half or less
as long as that peduncle and usually entire or rarely emarginate.

OEDICEROTIDAE (p. 373)

Pereopod 5 subequal to pereopod 4 in length, though both often elongate,
accessory flagellum always present and usually multiarticulate; when 1-
articulate, accessory flagellum elongate; peduncle of uropod 3 rarely

2 See also the calliopiid Metaleptamphopus.

285-135 O - 69 - 7

108 U.S. NATIONAL MUSEUM BULLETIN 271

elongate but telson always (except Synopia with large accessory flagellum)
longer than and as twice as long as peduncle of uropod 38, cleft or

entine se ariens . . . SYNOPIIDAE (p. 454)

13. Pereopods 3-5 atronele spinose or 2 aeiOse ond rahi Gea setae (‘‘fossorial,”’
see glossary) (figs. 97-99) . aa it Ea at Mat elie gunn ab
Pereopods 3-5 poorly unos ¢ or SOU or noe Toscana ee epnet aia meter: 1)

14. Pereopod 5 conspicuously elongate, at least 1.5 times longer than pereopods
SAGs Nsw te + » . OEDICEROTIDAE (p. 373)
Pereopod 5 not pongoreonsley Smeets, subequal to, shorter than or slightly
longer than pereopod4.... . RES PR eer! (a Lis)

15. Telson elongate, nearly twice as long as epednmcle Of hinonod 3 or urosomite
Sey asus eed . . . (exceptional) sYNoPIIDAE (p. 454)
Telson not Soni, Rearcel: eased length of peduncle on uropod 3 or
urosomite3..... BM eriti ed 0)

16. Base of primary de eee a tena 1 camnictint, ane article 1 of flagellum
longer than peduncle (fig. 34a) . (exceptional) VITJAZIANIDAE (p. 476)
Base of primary flagellum of antenna 1 multiarticulate, article 1 of flagellum
less than half as long as peduncle... . . echapaeeh aeaenalledt
17. Rostrum usually evanescent, pereopod 5 ennai Solan ig Herconnd 4 in
structure; when pereopod 5 shorter than and of different structure than
pereopod 4 then rostrum absent (fig. 45).
(See Sancho in Calliopiidae) 18
Head with distinct, and depressed or downturned, rostrum, pereopod 5
always shorter and of different structure than pereopod 4 (fig. 44).
PHOXOCEPHALIDAE (p. 412)
18. Check the individual through the family key of Haustoriidae. Most haus-
toriids will be detected in the first 15 couplets. If one of the following
characters does not apply to the species, proceed to Gammaridae: (1)
pereopod 5 much shorter than 4 and article 2 lamellar (Pontoporeia and
Urothoides) ; (2) mandible immense, triturating surface large and smooth,
distal, palp relatively small and thin, and nearly asetose (Carangolia); (3)
article 2 of maxilla 1 palp shorter than article 1 (Phozocephalopsis); (4)
mandibular incisor unproduced and teeth absent or obsolescent and article
6 of pereopods 1-2 with stout spines (Urothoe and Urothoides).
HAUSTORIIDAE (p. 248) and GAMMARIDAE (p. 231)
19. Some anterior coxae acuminate midventrally (fig. 36), and accessory flagel-
lum less than 3-articulate.
PARAMPHITHOIDAE (p. 389) and ACANTHONOTOZOMATIDAR (p. 117) 3
Anterior coxae not acuminate midventrally, or if acuminate then accessory
flagellum more than 2-articulate . ... . Biveea ings X0)
20. Article 4 of pereopods 1-2 extremely elongate Polntive fo ened articles, palp
of one member of first maxillae distinctly geniculate and scaled.
(fig. 33) HYPERIOPSIDAE (p. 261)
Article 4 of pereopods 1-2 not ee palp of maxilla 1 neither strongly

geniculate nor scaled . ... . Pepe pret s(n val:
21. Telson entire (often with minute notch or Spoilers auindiantion) - ican 4
Melsonnelefty seek yi os in Aouce: bay hele die gh Men hey (elie cate Ian ocean ek Seem, OF

3 See also Halirages stebbingi Schellenberg (1931) (=H. huzleyanus Stebbing, 1888, not Bate) in family
Calliopiidae. Note that almost all acanthonotozomatids have a nontriturative mandibular molar or no
molar whereas almost all Paramphithoidae have triturative molars. Acanthonotozomatidae usually have
conically grouped mouth parts from lateral view whereas most Paramphithoidae have mouth parts grouped
in a quadrate bundle. See text for further discussion.

22.

23.

24.

25.

26.

27.

28.

29.

30.

ol.

32.

33.

34.

MARINE GAMMARIDEAN AMPHIPODA 109

Atecessory flagellum 2-articulate’. 29.059. 5 2 63 8 28
Accessory flagellum l-articulate or absent ... . ROU cra

Rami of uropod 3 cylindroconical, much shorter anon Sones peduncle.
Bathyphotis (Ischyroceridae) (p. 275)

Rami of uropod 3 lanceolate or flabellate, much longer than peduncle.

(exceptional) GAMMARIDAE (p. 231)
Lower lip with unpointed, tilted, oval lobes astride partially coalesced inner
lobes (fig. 37) .... . . . PLEUSTIDAE (p. 421)
Lower lip with tilted or mntlted putes lobes! but with distinct mandibular
extensions, inner lobes when present not coalesced ae 38), (examine also

key to Eusiridae if telsonic condition dubious) .. . Sumeineumetnin ssa7 4)
Telson longer than wide ... . ee CALLTOPITDAE = (DLO)
Telson as wide as or wider than lone . . . (exceptional) GAMMARIDAE

Article 1 of primary flagellum on antenna | as long as peduncle.
VITJAZIANIDAE (p. 476) and (exceptional) male SYNOPIIDAE (p. 454)
Article 1 of primary flagellum on antenna 1 not longer than half of pe-

clummcley rane oe sao cee ie Parl
Telson elongate (fig. 39), vice as s long : as peduncle at Groped Sue ee
Telson rarely longer than peduncle of uropod 3, never twice as long as
peduncle of uropod3 ..... een ee se COU,

Urosomite 3 twice as long as masonic 2, jhendl oa lenterodorcal margin
extended as blunt plow (fig. 66d) . Pseudotiron (Synopiidae) (p. 461)
Urosomite 3 less than 1.5 times as long as urosomite 2, head of normal

dimensions as in basic gammaridean . . . sy Seelass caste NAS)
Accessory flagellum 3-+--articulate . (xccutionel) SYNOPIIDAE (p. 454)
Accessory flagellum O-2-articulate Gane also key to Calliopiidae if
telsonic cleft dubious) ...... . . . BUSIRIDAE (p. 213)

Inner ramus of uropod 3 short and puale: ie (Ge. 96c).
(exceptional) GAMMARIDAE (p. 231)

Innerramustof unopod’3 elongate 4. vs 2)". fa 5 2 el. 3 Jol
Rami of uropod 3 foliaceous ... . coos GAMMARIDAE (p. 231)
Rami of uropod 3 lanceolate . .. . Loatiis SA Sh ele ees tl ue dea

Mandibular palp article 2 shorter than onttialts ie
(exceptional) GAMMARIDAE (p. 231)

Mandibular palp article 2 longer than article 1 . .... . dd

Gnathopod 1 simple ..... . . (exceptional) GAMMARIDAE ©. 231)

Gnathopod 1 subchelate . .. . EAA SUS As MVC aie SD Rc oe Ye

Accessory flagellum 2+ - aceite ee i oO GAM MAR ED Amys 2 oi)

Accessory flagellum 1-articulate or absent . . . . . BUSIRIDAE (p. 213)
SECTION B

Head and body strongly depressed, rugose, coxae splayed (fig. 10).

PHLIANTIDAE (p. 405)

Head subglobular, body cylindrical (fig. 82), coxae short, not splayed.
(exceptional) EOPHLIANTIDAE (p. 209)
Head compressed or subglobular, ae compressed laterally, rarely rugose,
coxae never splayed . ... . Reese
Article 2 of antenna 1 shorter han or sapell to article iL nronod 3 usually
biramous but when uniramous then article 2 of antenna 1 not elongate or
gnathopod 1 not chelate .. . 3
Article 2 of antenna 1 much longer ahem, or Secniell fa onfalle 1 (Ges 15), nropods
always uniramous, gnathopods strongly chelate . . . SEBIDAE (p. 435)

110 U.S. NATIONAL MUSEUM BULLETIN 271

10.

Lis

12.

13.

. Article 4 of pereopods 1—2 enormously elongate relative to other articles and

often inflated (fig. 338f) ..... Ree Aaa
Article 4 of pereopods 1—2 of normal lenedh wala to Saher aatiialles - eee)

. Palp of maxilla 1 claviform, slightly geniculate (figs. 33a, b.).

HYPERIOPSIDAE (p. 261)

Palp of maxilla 1 not geniculate .. . . . proceed to couplet 5
. Gnathopod 2 with form of typical lestamaseicl saittiom (fig-13) ee
Gnathopod 2 not typical of lysianassids .. . . SAS AEE ea

. Accessory flagellum absent and pleonites 1-3 wails han dowel processes,

articles 5 and 6 of gnathopod 2 lacking scales or minute coarse setules.
(exceptional) ACANTHONOTOZOMATIDAE (p. 117)

If accessory flagellum absent then pleonites 1—3 dorsally smooth; accessory
flagellum otherwise present, articles 5 and 6 of gnathopod 2 bearing scales

or minute coarse setules . .... . . . (fig. 3) LYSIANASSIDAE (p. 294)

. Urosomiteszally coalescedg ne 2) iic50 ieee obs eer oe es ea a ee
Wrosomites Separate i056 Gs sepia tec (ee aeo ule pels weary ikea te Won ea

. Uropod 3 biramous. ...... . . . (fig. 22) PROPHLIANTIDAE (p. 432)
Uropod 3 uniramous .. . See oo 6 (aalegs 113) aqenenaanyeus, (>, 2EO))

. Uropod 3 formed only of Simmel, Agelle- like peduncle.

Didymocheila (incertae sedis) (p. 478)

Uropod 3 bearing one or two rami (inner often reduced) . . avai uo
Uropod 3 with one ramus . . . .. . . Microprotopus Gsacidse) (p. 274)
Uropod 3 with tworami .. . dace seem hl

All mouthparts except emilline dt repliced Be sentra (buecell reel and bulbs.
(fig. 20) ANAMIXIDAE (p. 145)

Mandibles, lower lip and maxillae present. . .. . shaven bt ede
Accessory flagellum 1-2 articulate or absent, moaminaaNts om lateral view
usually grouped in conical bundle, pereopods not strongly spinose. . 13

Accessory flagellum always 1-+-articulate, mouthparts from lateral view
usually. quadrately grouped, pereopods strongly spinose and setose (‘‘fos-
sonial’y)m (filet 4 5) se . . (exceptional) HAUSTORIIDAE (p. 248)

Mandibular palp present, Picon snoaesstieenous (fig. 7).

(exceptional) ACANTHONOTOZOMATIDAE (p. 117)

Mandibular palp absent, pleon never processiferous (fig. 4).

STEGOCEPHALIDAE (p. 436)

SECTION C
Body plan cylindrical or depressed (tanaid-like, or cuit or extremely
broad and flat)... . Pe aN a: 2
Body plan compressed laterally. @ocmal cemmmenidlennn lia) 6
Mandible lacking palp and molar degraded .... . Bee mene Gul (Re
Mandible with palp of one to three articles, molar ftomattire 4

Body cylindrical, tanaid-like, smooth (fig. 8), coxae very small.
COLOMASTIGIDAE and EOPHLIANTIDAE (pp. 182 and 209)
Body strongly depressed, very broad and rugose (fig. 10), coxae large and
splayed or body, if not rugose, with pereonites laterally discontiguous.
PHLIANTIDABR (p. 405)
Urosomites 1-3 coalesced . . .. . Meal sea 25)
Urosomites 2—3 coalesced and urosoraies Ls very leloneate (Ge 52).
PODOCERIDAE (p. 426)
Uropods 1-2 of very diverse structure (fig. 53) . . . CHELURIDAE (p. 180)
Uropods 1-2 similar to each other and of normal gammaridean structure.
(exceptional) COROPHIIDAE (p. 184)

10.

Wik

12.

13.

14.

15.

16.

MARINE GAMMARIDEAN AMPHIPODA 111

WropodkescuniramOuUsice sta vst keeenel MEA CtoN. ay ths Dns OTN Seale 07
Uropod 3 biramous. . . . aa Cerne Ulu aeraganaay SETAC)
Coxa 1 visible, not covered By folloeane | CORE) iii. 5, BUR Ce i ee ee tS
Coxa 1 small and hidden by large following coxae .. . Bing a ah Sons
Mandible lacking palp, bearing molar, inner plates of mamillipeds obsolescent,

telson cleft. .... ... . (fig. 13) KURIIDAE (p. 280)
Mandible bearing palp, lacking! molar inner plates of maxillipeds fully

developed, telson entire. . . ... . . (fig. 14) PAGETINIDAE (p. 387)

Telson and pleonite 6 coalesced even though telson recognizable and un-
thickened, article 2 of pereopod 3 expanded. (fig. 16) CRESSIDAE (p. 198)
Telson thickened dorsoventrally and distinct from urosome, but urosomites

partially coalesced, article 2 of pereopod 3 linear.
(fig. 17) THAUMATELSONIDAE (p. 473)

Mandibularspalpyabsentba each) eae oe ene 28 UL
Mandibular palp present. . . Se asa, WEEE Se
Only urosomites 2-3 coalesced (fig. 23) . . . . . DEXAMINIDAE (p. 200)
Urosomites 1-3 coalesced (fig. 22)... . . . PROPHLIANTIDAE (p. 432)
Pereopod 5 of different structure and almost always shorter than pereopod 4

(igs i226) kis ae . . . . AMPELISCIDAE (p. 128)
Pereopod 5 similar nl pabedual oF or elonger than pereopod 4 (or these

pereopods' often brokenjorsmissing) ioe Pe Se ae es 1S
Telson entire orslightly emarginate ................ 14
Telson cleft or deeply emarginate ........ mee Was LG
Urosomite 1 elongate (fig.52) .... . . . . PODOCERIDAE (p. 426)
Urosomite 1 not elongate, or all eroeorntce ponlesecd Sr SERA ERY aay os oe RRR) EE

Uropods 1 and 2 of extremely diverse structure (fig. 53).
CHELURIDAE (p. 180)
Uropods 1 and 2 similar to each other and of normal gammaridean structure
(nike 399) Dae uae ne . . . COROPHIIDAE (p. 184)

Coxae 1-4 rounded, fuadente or Eplunt Penirallen

(fig. 25) ATYLIDAE (p. 161)
One or more of coxae 1-4 acuminate ventrally and often bifid (fig. 24).
LEPECHINELLIDAE (p. 286)

1 See Paracalliope in supplement.

SECTION D
Gnathopod 1 reduced to one or two articles or absent (figs. 20,30) . . . 2
Gnathopod 1 present, comprising six or seven articles... . 3
Mouthparts all present and normal . . . . . (fig. 30) BATEIDAE “3 (Ge 163)

Mandibles and maxillae absent or coalesced into sharp keel (fig. 20)
ANAMIXIDAE (p.145)
Pereopods 3-5 fossorial, article 4.0f antenna 2 strongly expanded, antennae
with plumose setae longer than any antennal article.
(exceptional) HAUSTORIIDAE (p. 248)
Pereopods 3-5 usually not fossorial but if fossorial then article 4 of antenna
2 not expanded, setae of antennae not elongate plumes. ...... 4
Gnathopod 1 strongly carpochelate, carpus larger than propodus (or ap-
pearing as propodochelate because of loss of dactyl) (fig. 20)... .. 5
Gnathopod 1 subchelate or simple, or carpus smaller than propodus (oc-
casionally carpus with long lobe but not distinctly chelate) . . ... 6
Mouthparts except maxilliped replaced by a ventral keel and bulbs.
(fig. 20) ANAMIXIDAE (p. 145)

112

10.

11.

12.

U.S. NATIONAL MUSEUM BULLETIN 271

Mandibles, lower lip and maxillae present.
(exceptional) LEUCOTHOIDAE (p. 289)

Uropod 3 biramous:: 12)) ihe eanreeh Gras tly Si sea ese Ucar enact aerial NU armG|
Uropod 3 uniramous . . aU as Sree ree ae eyes ih (0)
Coxa 4 not excavate aire. poreapods ie 2 eflesmdtcllon GET LOR Ose
Coxa 4 excavate posteriorly, pereopods 1-2 not glandular ...... 9

Rami of uropod 3 shorter than peduncle, outer ramus with distal hook or
distolateral denticles, lacking setae.
(exceptional) ISCHYROCERIDAE (p. 275)
Rami of uropod 3 longer than peduncle, outer ramus setose or spinose but
lacking special hooks or denticles.
(exceptional) ISAEIDAE (= Photidae) (p. 264)
Article 4 of pereopods 1-2 exceptionally elongate (fig. 33f), base of primary
flagellum on antenna 1 conjoint and forming an article longer than pe-
duncle, palp of maxilla 1 geniculate and scaled (figs. 102a, 6).
Parargissa (Hyperiopsidae) (p. 264)
Article 4 of pereopods 1—2 not elongate, base of primary flagellum on antenna
1 not conjoint, palp of maxilla 1 not geniculate or scaled.
AMPHILOCHIDAE (p. 132)
Gnathopods simple and slender, with dense setae as long as fifth and sixth
articles, all coxae at least as broad as long.
(exceptional) ISAEIDAE (=Photidae) (p. 264)
Gnathopods usually subchelate and one pair usually stout, if not then their
setae very short and sparse, some coxae very much longer than broad . 11
Article 2 of pereopod 3 slender and linear (like fig. 162 v) . . ..... 12
Article 2 of pereopod 3 expanded like pereopods 4—5 (like fig. 162s).
CRESSIDAE (p. 198)
Urosomal segments coalesced fully or partially, telson thickened dorso-
ventrally (fig. 17), or gnathopod 2 carpochelate.
(exceptional) THAUMATELSONIDAE (p. 473)
Urosomal segments separate, telson depressed (fig. 18), gnathopod 2 never
carpochelate ............. . . . STENOTHOIDAE (p. 444)

SECTION E

Uropod 3 essentially uniramous, lacking rami or occasionally absent; when
present inner ramus scalelike and outer ramus cylindrical; Cheluridae partially
included herein although outer ramus flat; observer urged to verify that one ramus
of uropod 3 not accidentally broken off.

1.

Mandibular molar bearing grinding oe (triturative) or if not triturative

then molar cupshaped. ... . CN rea ete
Mandibular molar smooth or pearing a fou erticulate snes, or molar

absent) or not:cUpshbaped va voy ae ae) Leen eee eee)
Mandibular palp' presenti oo hey 2 elena eae ee tes ae a
Mandibular palp absent. ... . aetna opie 20 (0)
Urosomite 1 elongate, longer than nelccomnlis 3 and more nan twice as

long as urosomite 2 (fig. 52)... . . . . . . PODOCERIDAE (p. 426)
Urosomite I notelongate as’aboves..- <1. 6 ee ee ee
All urosomites‘coalesced) Neco eee astve usin oe een ee rar
Urosomites separate . . HHMI aN ene mien 1 8S

Uropods 1-2 of strongly diverse perucitine Ge 53) . CHELURIDAE (p. 180)
Uropods 1-2 normally styliform, although occasionally with rami or ped-
uncles reducedin size . ...... . . (fig. 51) coROPHIIDAE (p. 184)

MARINE GAMMARIDEAN AMPHIPODA 113

6. All urosomites coalesced (fig. 13)... ... =... . KURIIDAE (p. 280)
Urosomites separate .... : SA bee mE

7. Antennae and pereopods strongly splines ¢ or ecioee spines anal setae long,
epistome nasiform (fig. 11) .... . . . .DOGIELINOTIDAE! (p. 207)

Antennae and pereopods poorly spinose or setose (spines if numerous very
small), epistome rounded or truncate in front.
(fig. 12) TALITROIDEA (p. 463)
8. Lower lip lacking inner lobes, outer lobes obsolescent (fig. 173g), gnathopods
small, chelate, slender, equal, sixth article elongate (figs. 173d, e).
Didy mocheila (incertae sedis) (p. 478)
Lower lip with inner lobes and strong mandibular se ee not

taking special form of figs. 173d,e ....... Be Prams cy NO

9. Urosome depressed... .. . . . COROPHIIDAE (p. 184)
Urosome not depressed . . (Cxeeptional) AORIDAE (p. 147) and ISAEIDAE

(p. 264)

LOM Viandibularipalp present’ soc. ol Lees ele SOR hea ee es) LD
Mandibular palp absent ... . Bi ca ceo ead iy

11. Gnathopods subchelate, outer plates oF aeciiped very painall: inner obsolete.

(fig. 14) PAGETINIDAE 2 (p. 387)

Gnathopods chelate (fig. 15e) inner and outer plates of maxilliped well formed.

(exceptional) SEBIDAE (p. 435)

12. Body cylindrical (fig. 87a), coxae small . . (fig. 8) EOPHLIANTIDAE (p. 209)
Body compressed (normal), coxae large, not splayed.

Najna (Talitroidea) (p. 470)

Body depressed, rugose, coxae splayed laterally, or if body not rugose pereon-

ites laterally discontiguous . . . . . . . (fig. 10) PHLIANTIDAE (p. 405)

1 Metoediceros Schellenberg (1931) resembles this family but its epistome is unknown and its telson is
nearly a perfect, uncleft circle, whereas Dogielinotidae have a rectangular, cleft telson. Metoediceros is pres-
ently classified with the Oedicerotidae but it probably should form the type of a new family and is therefore
removed to ‘‘incertae sedis.”’

2 Urosomites 2-3 are presumed to be coalesced.

SECTION F
le bodvyaplan cylindrical (fig. Sia). ie teenies Mae eee Tees aie oe 2,
Body plan compressed or depressed . . . . . 5)

2. Palp of maxilla 1 larger than outer plate, inner etnies of Peeained! dermaded:
(fig. 9) COLOMASTIGIDAE (p. 182)
Palp of maxilla 1 oa inner plates of maxilliped well developed,
Separate .... . . . . (fig. 8) EOPHLIANTIDAE (p. 209)
3. Mandibular molar of ted sizes trituratives(ige2f yet ts Bowes A

Mandibular molar enormous, smooth (fig. 166d).
Jeddo (Synopiidae) (p. 461)
Mandibular molar evanescent or absent, lacking grinding ridges ... . 6

4. Uropod 3 with subequal rami (fig. 50).

Sunamphitoe (Ampithoidae) (p. 145)

Uropod 3 with inner ramus reduced or absent . . 5
5. Accessory flagellum absent. ... . . (fig. 12) TALITROIDEA (os 463)
Accessory flagellum present .... . . (fig. 42) BEAUDETTIIDAE (p. 166)

6. Mouthparts from lateral view conically grouped below head, uropod 3 bi-
ramous and well developed, coxae 1—4 compressed, elongate, forming lateral
shield, body compressed, smooth . . (fig. 4) STEGOCEPHALIDAE (p. 436)

114 U.S. NATIONAL MUSEUM BULLETIN 271

Mouthparts from lateral view not conically grouped below head, uropod 3
degraded, coxae 1-4 not Pas lateral shield, splayed laterally, body
depressed, rugose . . . .. . . (fig. 10) PHLIANTIDAE (p. 405)

Mouthparts from lateral view nor conically grouped below head, uropod 3
biramous and well developed, coxae 1-4 compressed, very short, not
forming lateral shield, body compressed, smooth.

Parahalice (Pardaliscidae) (p. 403)

SECTION G
1. Maxillipedal palp with fewer than four articles ........... 2
Miaxaillipedal\palpy with; foursarticles| iio nes ar ae ae eS

2. Maxillipedal palp 3-articulate.
(fig. 7) (exceptional) ACANTHONOTOZOMATIDAE (p. 117)
Maxillipedal palp 2-articulate (fig. 6)... . . . . LAFYSTIIDAE (p. 281)
Maxilliped lacking palp (fig. 5) .... . . . OCHLESIDAE (p. 371)
3. Mouthparts from lateral view conically prouned belowsheadi see
Mouthparts from lateral view not conically grouped below head . . . . 5
4. Anterior coxae long, some acuminate ventrally (fig. 7).
ACANTHONOTOZOMATIDAE (p. 117)
Coxae short, not acuminate ventrally (fig. 28).
Halicella (Pardaliscidae) (p. 402)

5. Gnathopod 1 carpochelate (fig. 21) . . . . . . . LEUCOTHOIDAE (p. 289)
Gnathopod 1 propodochelate, subchelate, or simple ......... 6
6. Mandibular molar present (though occasionally very minute) ..... 7
Mandibular molar absent . . . PA ere S22

7. Pereopod 5 shorter and of diferent sinew inert merconad 4, head with
strongly depressed large rostrum.

(fig. 44) (exceptional) PHOXOCEPHALIDAE (p. 412)

Pereopod 5 of structure similar to pereopod 4, rarely shorter than 4, rostrum

if present not depressed (not dorsoventrally flattened) ....... 8
8. Telson elongate, more than twice as long as urosomite 3 ....... 9
Telson short, subequal to or shorter than urosomite 3 (seen laterally) . . 10

9. Accessory flagellum l-articulate; antennae with large calceoli; gnathopods
large and strongly subchelate (fig. 92h).

Eusirella (p. 225) and Eusiropsis (Eusiridae) (p. 225)

Accessory flagellum multiarticulate; antennae lacking calceoli; gnathopods

feeble (fig. 167) .... . . . SYNOPIIDAE (p. 454)

10. Pereopods 3-5 fossorial Gone fees of tinoxe pecconods at least half as long

as longest article of any pereopod, fig. 45)... . . é oa e ele:

Pereopods 3-5 not fossorial (setae and spines very dhar, seancelly exceeding

article 3 of those pereopods) ... . Aire isk: Goel

11. Accessory flagellum l-articulate and oxtroinaly eral) or “alpsen: pereopod

5 very elongate, at least 1.5 times as long as pereopods 3-4 (uropod 3

always elongate, rami very slender and lanceolate, subequal to peduncle,

but uropod 3 often broken, telson always short, linguiform and entire or

emarginate). ........ . . (exceptional) OEDICEROTIDAE (p. 373)

Accessory flagellum rarely vestigial (if so, uropod 3 very short, flabellate,

rami highly unequal), pereopod 5 shorter than or usually scarcely longer

than pereopod 4 (uropod 3 usually with short peduncle, unequal rami

one of which longer than peduncle, if not then accessory flagellum multi-
articulate, telson cleft even minutely, usually elongate).

(exceptional) HAUSTORIIDAE (p. 248)

12.

13.

14.

15.

16.

I2fe

18.

19.

20.

21.

22.

MARINE GAMMARIDEAN AMPHIPODA 115

MelSONRCLELG Ama ean ie Magica? seen eer S NENA rie Cet h aan Hea deneeeh Mayr ath be 1S
Telson entire .. . Ses ic Tae BY
Gnathopods porreneeilly piticnelete (fig. Ae pat like pe 609) MEP eR LA:
Gnathopods feeble (fig. 32d)... . . 15
Accessory flagellum 2-+--articulate, Tes ina pbsolescent 4s cleft 75
percent of itslength ....... (fig. 41) LILJEBORGIIDAE (p. 291)
Accessory flagellum absent, rostrum neslly as long as article 1 of antenna 1,
telson cleft 40 percent of its length.
Pseudamphilochus (Amphilochidae) (p. 139)
Accessory flagellum 0-1 articulate, rostrum obsolescent, telson cleft 25
percent of itslength .. . . . . Austropleustes (Pleustidae) (p. 422)
Mandibular molar a conical or trapezoidal, setose lamina ..... . 16
Mandibular molar a large setulose tuberosity.
Synopia (Synopiidae) (p. 462)
Accessory flagellum present ...... . . (fig. 32) asTyRIDAE (p. 157)
Accessory flagellum absent.

Eclysis and Epimeriella (Paramphithoidae and Astyridae) (pp. 161, 389)
Coxa 3 with about 3 times surface area of coxa 4, maxillipeds foliaceous,
article 3 of mandibular palp vestigial, subequal to article 1 (fig. 165e).

Synopia (Synopiidae) (p. 462)
Coxa 4 usually slightly larger than, subequal to, or scarcely smaller than
coxa 3, maxillipeds not foliaceous, article 3 of mandibular palp much

longer than articlel .... Srey te a
Coxae 1-4 wider Garcnomnosterion) ian Tene (Gormaleyentrsl) (fig.
TIVST GAs <a ie Bh Alan heNan. Pee te AO Snead phere yO)
Coxae 1-4 longer diner mide Ge 37) aac ale RRs: CUR Te Shine, SEES 20

Coxae contiguous or slightly overlapping.
(exceptional) CALLIOPIIDAE (p. 167)
Coxae not touching serially (fig. 111a).
Laphystiopsis (p. 285) and Prolaphystiopsis ena (p. 285)

Mandibular molar bulbous, ovate or cylindrical. . . pees 2
Mandibular molar asetose conicallamella . . (fig. 32), ASTYRIDAE (p. 157)
Article 4 of pereopods 1-2 enormously elongate (fig. 33f), accessory flagellum

long and atleast 3-articulate . .. . . . . .HYPERIOPSIDAE (p. 261)

Article 4 of pereopods 1-2 of normal aimensions! accessory flagellum short or
absent and less than 3-articulate.

PLEUSTIDAE (p. 421) and CALLIOPIIDAE (p. 167)

Accessory flagellum usually (probably always) present and 3-+-articulate,

maxillae not foliaceous (occasionally palp of maxilla 1 slightly foliaceous).

(fig. 28) PARDALISCIDAE (p. 397)

Accessory flagellum absent, vestigial or 2-articulate, extremely small, max-

illae foliaceous. ....... =... . (fig. 29) stmrpEDIDAE (p. 451)

SECTION H

Pereopod 5 exceeding 1.5 times length of pereopods 3-4 ! (fig. 27).
OEDICEROTIDAE (p. 373)
Pereopod 5 scarcely longer than or shorter than pereopod4...... 2
Outer lobes of lower lip notched apically or medially excavate, outer ramus of
uropod 3 bearing one or two giant, hooked, articulate spines, rami flattened.
(fig. 50) AMPITHOIDAE (p. 141)

1 Consult Aoridae if telson fleshy and pereopods 1-2 glandular.

116 U.S. NATIONAL MUSEUM BULLETIN 271

oo

10.

11.

12.

Outer lobes of lower lip not notched nor medially excavate, outer ramus of
uropod 3 not bearing giant hooked spines (except Jassa), occasionally
bearing small distal hook, or bent and serrate spine, rami lanceolate or
styliform. ... aD aap. 3

. Peduncle of uropod 3 aien snail as Mlonere as or rlonee ‘hora podiums of uropod

2, rami of uropod 3 much shorter than peduncle, outer ramus of uropod

3 distally hooked (hook a giant spine in Jassa), or with minute distolateral
denticles, pectinae or serrations (and see Parapherusa in Gammaridae).

(fig. 49) ISCHYROCERIDAE (p. 275)

Peduncle of uropod 3 usually much shorter than peduncle of uropod 2, or

rami of uropod 3 subequal to or longer than peduncle, or outer ramus of

uropod 3 not distally hooked and lacking distolateral ornamentation . 4

- Urosome depressed) 3 jeoi. she) SOON Oe Ce) Share ea earat

Urosome not depressed . . 7

. All urosomites coalesced, nropede ie 3 OP fehl divers Aine (Gig 53).

CHELURIDAE (p. 180)
Urosomites rarely all coalesced, uropods 1-2 (but rarely 3) similar to each
other and of normal gammaridean form. ............. 6

. Urosomite 1 elongate (fig. 52) . . .... =... . PODOCERIDAE (p. 426)

Urosomite 1 not elongate (fig. 51) .... . . . COROPHIIDAE (p. 184)

. Gnathopods 1-2 feeble, subequal to each cer! simple or nearly so, article 3

of mandibular palp vestigial (fig. 165e), article 1 of primary flagellum of
antenna 1 elongate, plates of maxilliped foliaceous.

Synopia (Synopiidae) (p. 462)

Gnathopods 1-2 strong, usually dimorphic and subchelate, article 3 of man-

dibular palp strong, article 1 of primary flagellum of antenna 1 not elongate,

plates of: maxilliped not: foliaceous:isc 43 n) ee eS

. Gnathopod 1 larger than 2 (fig. 48) ...... SMA esi Gee oO)

Gnathopod 2 larger than 1 or gnathopods equalinsize. ....... 10

. Telson thick and fleshy, perepods glandular. . . . . . AORIDAE (p. 147)

Telson thin dorsoventrally, pereopods not glandular.
Falklandella 2? (Gammaridae) (p. 241)
Telson thick and fleshy, pereopods usually glandular.
(fig. 47) ISAEIDAE (p. 264)
Telson thin dorsoventrally, pereopods not glandular ......... Ii
Accessory flagellum exceeding three articles.
(exceptional) GAMMARIDAE (p. 231)
Accessory flagellum 2-articulate or less, often absent ......... 12
Coxae 1-4 strongly or weakly acuminate midventrally or coxae midventrally
excavate to form weak or strong anterior and posterior acuminations.
(exceptional) PARAMPHITHOIDAE (p. 389)
Coxae 1—4 rounded or quadrate ventrally.
(exceptional) CALLIOPIIDAE (p. 167)

2 Schellenberg (1931) writes that coxa 4 is distinctly excavate posteriorly but no figure is given and such
excavation must be deep and distinct to avoid placing a genus in section H.

MARINE GAMMARIDEAN AMPHIPODA 117

Acanthonotozomatidae

Ficures 54-56

Dracnosis.—Mouthparts grouped into cone-shaped bundle pro-
jecting from ventral surface of head; accessory flagellum absent or
vestigial. See Paramphithoidae, Ochlesidae, Pardaliscidae, La-
fystidae.

Description.—Accessory flagellum absent, 1- or 2-articulate
when present, very small and short; body usually with dorsal processes;
rostrum well-developed; mouthparts rather linear in shape (to fit
conical bundle and probably adapted for piercing and sucking);
mandible especially linear, usually lacking molar or having only small
molar protuberance, although triturative molar present in some genera;
palp of mandible always present; lower lip variable, with or without
inner lobes, apices of outer lobes occasionally incised; maxilliped with
well-developed plates but palp with three or four articles and long
or short; coxae 1-4 usually acuminate or subacuminate; gnathopod
1 and usually gnathopod 2 very feeble, often minutely chelate, article
3 of gnathopod 2 occasionally elongate; rami of uropod 3 longer than
peduncle, rami flattened-lanceolate; telson entire or slightly cleft;
article 2 of pereopods 3-5 usually with posterior teeth.

RELATIONSHIP.—The mandibular molar of Astyridae is larger than
but not morphologically distinct from the protuberances occurring
in some acanthonotozomatids; the accessory flagellum of astyrids is
1-articulate but long, and the mouthparts are not arranged in a conical
bundle.

The Lafystidae have a 2-articulate maxillipedal palp, but otherwise
the family is so similar to the Acanthonotozomatidae that the two
families might be joined together.

The Stilipedidae resemble Acanthonotozomatidae but apparently
the mouthparts are not grouped in a conical bundle; the maxillae
are strongly foliaceous but this is true also of Mazilliphimedia.

With one exception the mouthparts of Pardaliscidae are not grouped
in a conical bundle; usually pardaliscids have a multiarticulate
accessory flagellum and the inner plates of the maxillipeds are obsoles-
cent. One of the mandibles is especially flattened and heavily toothed.

The Stegocephalidae lack a mandibular palp.

Mouthparts of the Paramphithoidae are not arranged in a conical
bundle but this distinction is confused by the acanthonotozomatid
Bathypanoploea ausiralis (Chilton, 1912) (see Schellenberg, 1931).
The mouthparts of B. australis not only are grouped in a quadrate
bundle but the molar is absent and the gnathopods are styliform as in
most acanthonotozomatids. The type genera of the two families differ
in the following ways:

118 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 54.—Acanthonotozomatidae: a, Echiniphimedia hodgsont (Walker), original draw-
ing; b, Acanthonotozoma cristatum (Ross) (Sars, 1895, pl. 131).

MARINE GAMMARIDEAN AMPHIPODA 119

Paramphithoe Acanthonotozoma
Mouthparts bluntly bundled conically bundled
Mandible stout substyliform
Mandibular molar large, ridged obsolescent, conical
Maxillae stout substyliform
Gnathopods small, subchelate styliform

Intergradations of these conditions occur in the genera of both

families. If the sea yields more genera similar to Bathypanoploea, the
fusion of the Paramphithoidae and Acanthonotozomatidae may be
required.

Acanthonotozomella oates: K. H. Barnard (1930) is transferred to

Acanthonotozomoides.

10.

12.

13.

Key to the Genera of Acanthonotozomatidae

Ralproremaxillayl uniarticulate (fig! 56d)) 2. 2
Palp on maxilla biarticulates(fig.;56c)) - S24 Ses ee
Gnathopod 2 slender and chelate (fig. 560). . . ......2.2.~. 3
Gnathopod 2 stout and subchelate (fig. 56p) ... . is fa Odius
Palp article 2 of maxilliped strongly produced medially along article 3

(e562). Sas . . . . Pariphimedia
Palp article 2 of raeeilleed not mroduced (fig. 56h) ess Barichinhime dis
Palp of maxilla 1 very short, not reaching end of outer plate (fig. 56d) . . 5
Palp of maxilla 1 reaching end of outer plate (fig. 56c) ........7
Gnathopods 1 and 2 simple (figs. 56k,n). . . . . . . . Panoploeopsis

Gnathopod 1 minutely chelate (figs. 561, m), prachened 2 not simple. . 6
Palp article 2 of maxilliped produced along inside of third (fig. 562), lower

lip usually incised (fig. 56a) . . . . . . . Panoploea
Palp artile 2 of maxilliped not produecd plone: neide of third (fig. 56g),
lower lip not incised (fig. 566) . . . . . . . Anchiphimedia
Mandible tapering to a smooth, pramenecnicall ecoopeshaned apex (fig. 55g). 8
Mandible not as above (e.g., fig. 55k) . 5 9
Upper lip rounded, not incised (fig. 55c); owes ie fet saciaedl Ge, 560).
Labriphimedia

Upper lip quadrangular, incised (fig. 55d); lower lip incised (fig. 56a).
Maoriphimedia
Gnathopod simple: (figs 56/4)! sete ay oh ene ciacee e See not. enema ml \()
Gnathopod 1 chelate (fig. 561)... ....... carte eens LS)
Telson entire, rounded (fig. 56g) .. . se “Acanthonotozomoides
Telson distally notched or emarginate (figs. 56r YS) Meena iinet ka |

Gnathopod 1 feeble, very slender, gnathopod 2 stouter ‘fhom i

Acanthonotozoma
Gnathopods 1 and 2 similar to each other, well developed. . . . .. 12
Palp of maxilliped shorter than outer plate; mandible pointed (fig. 55l).
Bathypanoploea
Palp of maxilliped longer than outer plate; mandible distally broad (fig. 55k).
Acanthonotozomella

Cutting edge of mandible drawn out in a long, needle-like apex (fig. 557).
Parapanoploea

120 U.S. NATIONAL MUSEUM BULLETIN 271

Mandible not drawn out in a long, needle-like apex . . . . 14

14. Palp of maxilla 1 enormously expanded into an ovate leant Tne lip
asymmetrically incised (fig. 55e), lower lip not incised (fig. 56b).

Maxilliphimedia

These characters not combined . . . BMH oo 55

15. Palp article 2 of maxilliped distinctly anedtnced siarays medial adlge af article

3 (fig. 562), outer lobes of lower lip ae and strongly notched (fig.

56a) Pe twe te CLG

Palp article 2 8 menlliped andlithaclliy or ner anedingedl allo medial edge

of article 3 (figs. 56g,h), outer lobes of lower lip unnotched or very weakly

mnotcheds. 23 va) ae sone tay IL
16. Pereonite 1 as long as est ihtce sans combined Ge 55m), mandibular
incisor apparently absent... . . ... . . Cypsiphimedia
Pereonite 1 not longer than each of next ines sesTnemti (fig. 55a), mandibular
incisor strong . . . . . . Iphimedia!
17. Upper lip deeply merce iGo “BEO), Past wor articles of maxillipedal palp
extraordinarily broadened (fig. 56f) ... . . . . Pseudiphimediella
Upper lip entire or emarginate, first two articles of maxillipedal palp not
broadened (fig. 56g) . . . . : eutinatat teres sit Ra ea NLS,
18. Some or all coxae with salnamaehiell teeth RU aaa ‘Echiniphimedia
Coxae lacking submarginal teeth .. . Ree si Whe)

19. Primary cutting edge of mandible elndlentete Ges. ih, ®.
Pariphimediella
Primary cutting edge of mandible 2- or less dentate (fig. 551) . . . .. 20
20. Mandible short, stout, Pee cutting edge very broad, smooth, molar
conical (fig. 557). ... . . . . . . Gnathiphimedia
Mandible slender, long, primary arin ede tapering, subacute, molar if
present, not conical (fig. 55/) ........... .. Iphimediella

1See also Panoploea spinosa Thomson (Hurley, 1954c) which keys here because of its scarcely shortened
palp of maxilla 1.

Genera of Acanthonotozomatidae

Acanthonotozoma Boeck

Acanthonotus Ross, 1835 (homonym, Pisces).
Acanthonotozoma Boeck, 1876 (new name for Acanthonotus) —Stebbing, 1906.

Type-species: Acanthonotus cristatus Ross, 1835 (original designa-
tion). See Sars, 1895.

Upper lip incised; mandible with narrow, rounded apex; lobes of
lower lip not incised; palp of maxilla 1 biarticulate, reaching or exceed-
ing end of outer plate; maxillipedal palp exceeding outer plate,
4-articulate, article 2 not produced; gnathopods simple, gnathopod 1
very slender, feeble, gnathopod 2 stouter than 1; telson apically
incised. Species: 4, arctic-boreal, littoral (rarely to 700 m).

MARINE GAMMARIDEAN AMPHIPODA 121

Figure 55.—Acanthonotozomatidae: a, Odius carinatus (Bate) (Sars, 1895, pl. 133).
Accessory flagellum: b, Iphimedia haurakiensis Hurley (1954c). Upper lip: c, Labriphi-
media vespuccii K. H. Barnard (1932); d, Acanthonotozoma serratum (Fabricius) (Sars,
1895, pl. 131); e, Odius. Mandible: f, Acanthonotozoma; g, Labriphimedia, apex; h,
Pariphimedia integricauda Chevreux (1906c); i, Gnathiphimedia mandibularis K. H.
Barnard (1930); 7, Parapanoploea oxygnathia Nicholls (1938); k, Maoriphimedia hinemoa
Hurley (1954c); 7, Odius. Front of body: m, Cypsiphimedia gibba K. H. Barnard (1955).

122 U.S. NATIONAL MUSEUM BULLETIN 271

Acanthonotozomella Schellenberg
Acanthonotozomella Schellenberg, 1926a.

Type-species: A. alata Schellenberg, 1926a (monotypy).

Upper lip ?incised [‘‘eingekerbt’’]; mandible with narrow, rounded
apex, minutely serrate; lobes of lower lip not incised; palp of maxilla
1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding
outer plate, 4-articulate, article 2 not produced; gnathopods simple,
alike; telson apically incised. Species: 1, antarctic, littoral.

Acanthonotozomoides Schellenberg
Acanthonotozomoides Schellenberg, 1931.

Type-species: A. sublitoralis Schellenberg, 1931 (monotypy).

Upper lip incised; mandible with narrow, slightly toothed apex;
lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching
end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate,
article 2 slightly produced medially; gnathopods simple, alike; telson
apically rounded, entire. Species: 2, antiboreal, littoral.

Anchiphimedia K. H. Barnard
Anchiphimedia K. H. Barnard, 1930.

Type-species: A. dorsalis K. H. Barnard, 1930, 1932 (monotypy).

Upper lip incised; mandible with narrow, acute apex; lobes of lower
lip not incised; palp of maxilla 1 biarticulate, very short; maxillipedal
palp exceeding outer plate, 3-articulate, article 2 not produced;
genathopod 1 minutely chelate, gnathopod 2 not simple; telson slightly
incised. Species: 1, antarctic, bathyal (to 550 m).

Bathypanoploea Schellenberg

Epimeriopsis K. H. Barnard, 1931 (void ab znitzo).

Iphimediopsis Schellenberg, 1931 (homonym, not Della Valle, 1893).

Bathypanoploea Schellenberg, 1939 (footnote p. 137; new name for Iphimediopsis
Schellenberg) .

Pseudiphimediopsis Ruffo, 1949.

Type-species: Acanthonotozoma australis Chilton, 1912 (monotypy).
K. H. Barnard (1932, p. 182) noted that H’pimeriopsis is void ab initio
because of its basis on a misidentified specimen of the designated
type-species, A. australis Chilton.

Upper lip incised; mandible with broad, toothed apex; lobes of
lower lip not incised; palp of maxilla 1 biarticulate, reaching end of
outer plate; maxillipedal palp shorter than outer plate, 4-articulate,
article 2 not produced; gnathopods simple, alike; telson emarginate.
Species: 1, subantarctic, abyssal.

MARINE GAMMARIDEAN AMPHIPODA 123

Cypsiphimedia K. H. Barnard

Cypstphimedia K. H. Barnard, 1955.

Type-species: C. gibba K. H. Barnard, 1955 (original designation).

Upper lip incised; mandibular incisor feeble or ?absent; lobes of
lower lip notched; palp of maxilla 1 biarticulate, reaching end of outer
plate; maxillipedal palp exceeding outer plate, 3-articulate, article
2 medially produced along article 3; gnathopod 1 chelate, gnathopod 2
subchelate; pereonite 1 as long as next three segments combined; telson
slightly emarginate. Species: 1, South Africa, littoral.

Ficure 56.—Acanthonotozomatidae: Lower lip: a, Iphimedia obesa Rathke (Sars, 1895,
pl. 132); b, Acanthonotozoma serratum (Fabricius) (Sars, 1895, pl. 131). Maxilla 1: c,
Acanthonotozoma; d, Odius carinatus (Bate) (Sars, 1895, pl. 133). Maxilla 2: e, Acantho-
notozoma. Maxilliped: f, Pseudiphimediella nodosa (Dana) (Schellenberg, 1931); g,
Labriphimedia vespuccur K. H. Barnard (1932); h, Acanthonotozoma; 1, Iphimedia; j,
Pariphimedia integricauda Chevreux (1906c).° Gnathopod 1: k, Acanthonotozoma; 1,
Iphimedia; m, Odius. Gnathopod 2: n, Acanthonotozoma; 0, Iphimedia; p, Odius. Tel-
son: gq, Paritphimedia; r, Iphimedia; s, Acanthonotozoma; t, Odius. Uropod 3: u,
Acanthonotozoma.

285-135 © = 69 - 9

124 U.S. NATIONAL MUSEUM BULLETIN 271

Echiniphimedia K. H. Barnard
Echiniphimedia K. H. Barnard, 19380.

Type-species: Iphimedia hodgsont Walker (see 1907) (present
selection).

Upper lip entire or slightly emarginate; mandible with narrow,
rounded apex; lobes of lower lip not incised or minutely so; palp of
maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp
exceeding outer plate, 4-articulate, article 4 extremely minute, article
2 scarcely or not produced; gnathopods 1 and 2 chelate; telson with
shallow, broad emargination; some or all coxae with submarginal teeth.
Species: 3, antarctic, bathyal to littoral (to 824 m).

Gnathiphimedia K. H. Barnard
Gnathiphimedia K. H. Barnard, 1930.

Type-species: G. mandibularis K. H. Barnard, 1930 (present
selection).

Upper lip entire; mandible short, broad, apex smooth, molar
conical; lobes of lower lip not incised; palp of maxilla 1 biarticulate,
reaching end of outer plate; maxillipedal palp exceeding outer plate,
3-articulate, article 2 indistinctly or not produced; gnathopod 1
chelate; telson ?emarginate. Species: 3, antarctic, bathyal (littoral)
(to 824 m).

Iphimedia Rathke

Iphimedia Rathke, 1843.—Stebbing, 1906.
Microcheles Kréyer, 1846.

Type-species: J. obesa Rathke, 1843 (monotypy). See Sars, 1895.

Upper lip slightly emarginate or truncate; mandible with medium-
broad, slightly toothed apex; lobes of lower lip notched; palp of maxilla
1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding
outer plate, 3-articulate, article 2 produced along article 3; gnathopods
both chelate or gnathopod 2 subchelate; telson incised or emarginate.
Species: 8, tropical Pacific to antarctic, littoral (one bathyal).

Iphimediella Chevreux
Iphimediella Chevreux, 1911e, 1912a, 1912b.

Type-species: J. marguerite. Chevreux 1912a (designated by
Chevreux) ; 1912b.

Upper lip entire or weakly incised; mandible medium-broad, apex
scarcely toothed; lobes of lower lip not incised; palp of maxilla 1

MARINE GAMMARIDEAN AMPHIPODA 125

biarticulate, reaching end of outer plate; maxillipedal palp exceeding
outer plate, 3-articulate, article 2 scarcely or not produced; gnathopods
both chelate; telson cleft one third. Species: 4, antarctic, bathyal
(110-457 m).

Labriphimedia K. H. Barnard

Labriphimedia K. H. Barnard, 1931.

Type-species: JL. vespuccw K. H. Barnard, 1931 (original designa-
tion).

Upper lip entire; mandible with smooth, scoop-shaped, tapering
apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate,
reaching end of outer plate; maxillipedal palp exceeding outer plate,
composed of three articles tipped with a very minute fourth, article 2
not produced; gnathopods both ?chelate (type-species not described) ;
telson notched. Species: 2, antarctic, littoral.

Moaoriphimedia Hurley
Maortphimedia Hurley, 1954c.

Type-species: M. hinemoa Hurley, 1954c (original designation).

Upper lip slightly incised; mandibular apex broad, untoothed,
scoop-shaped; lobes of lower lip slightly incised apically; palp of
maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp
exceeding outer plate, 3-articulate, article 2 not produced; gnathopods
both chelate; telson apically notched. Species: 1, New Zealand,
littoral.

Maxilliphimedia K. H. Barnard

Mazilliphimedia K. H. Barnard, 1930.

Type-species: Iphimedia longipes Walker (see 1907) (monotypy).

Upper lip broad, incised; mandible with broad, toothed apex;
lobes of lower lip not incised; palp of maxilla 1 biarticulate, article 2
enormously expanded; maxillipedal palp exceeding outer plate,
3-articulate, article 2 not produced; gnathopod 1 chelate, (gnathopod 2
not described) ; telson deeply notched. Species: 1, antarctic, bathyal,
(183-379 m).

Odius Liljeborg

Otus Bate, 1862 (homonym, Lepidoptera).
Odius Liljeborg, 1865.—Stebbing, 1906.

Type-species: Otus carinatus Bate, 1862 (monotypy).
Upper lip narrow, incised; mandible with narrow, dentate apex;
lobes of lower lip not incised; palp of maxilla 1 uniarticulate, tiny,

126 U.S. NATIONAL MUSEUM BULLETIN 271

not reaching end of outer plate; maxillipedal palp exceeding outer
plate, 4-articulate, article 2 not produced; gnathopod 1 slender,
chelate, gnathopod 2 stout, subchelate; telson notched. Species: 2,
arctic-boreal, N. Atlantic, littoral.

Panoploea Thomson

Panoploea Thomson, 1880.—Stebbing, 1906.
Iphimediopsis Della Valle, 1893.

Type-species: P. spinosa Thomson, 1880 (present selection) ;
Chevreux and Fage (1925) erroneously cite as type-species, Iphimedia
eblanae Bate (1857d), a species not included originally in the genus by
Thomson.

Upper lip narrow, ?incised or not; mandible with very narrow,
scarcely dentate apex; lobes of lower lip incised or not; palp of maxilla
1 biarticulate, not reaching end of outer plate; maxillipedal palp
exceeding outer plate, 3-articulate, article 2 produced medially;
gnathopod 1 chelate, gnathopod 2 slightly chelate or subchelate; telson
broadly or minutely incised or cleft. Species: 8, amphiboreal, littoral
(one in bathyal antarctic to 550 m).

Panoploeopsis Kunkel
Panoploeopsis Kunkel, 1910.

Type-species: P. porta Kunkel, 1910 (monotypy).

Upper lip narrow, possibly not incised; mandible with narrow,
dentate apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate,
not reaching end of outer plate; maxillipedal palp barely exceeding
outer plate, 3-articulate, article 2 slightly produced medially; gnatho-
pods simple; telson bilobate. Species: 1, Bermuda, littoral.

Paranchiphimedia Ruffo
Paranchiphimedia Ruffo, 1949.

Type-species: P. monodi Ruffo, 1949 (original designation).

Upper lip deeply emarginate; mandible subpyramidal, incisor un-
toothed; [lobes of lower lip unknown]; palp of maxilla 1 uniarticulate,
short; maxillipedal palp exceeding outer plate, 3-articulate, article 2
not produced; gnathopods chelate; telson emarginate. Species: 1.
antarctic, ?littoral.

MARINE GAMMARIDEAN AMPHIPODA 127

Parapanoploea Nicholls

Parapanoploea Nicholls, 1938.

Type-species: P. orygnathia Nicholls, 1938 (original designation).

Upper lip broad, faintly emarginate; mandible with needle-like
apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate,
reaching end of outer plate; maxillipedal palp exceeding outer plate,
4-articulate, article 2 slightly produced medially; gnathopods chelate;
telson notched. Species: 1, antarctic (220 m).

Pariphimedia Chevreux

Pariphimedia Chevreux, 1906a.

Type-species: P. integricauda Chevreux, 1906a (original designa-
tion).

Upper lip broad, faintly emarginate; mandible with narrow, dentate
apex; lobes of lower lip not incised; palp of maxilla 1 uniarticulate, not
reaching end of outer plate; maxillipedal palp exceeding outer plate,
3-articulate or with minute article 4, article 2 strongly produced
medially; gnathopods chelate; telson entire. Species: 2, antarctic,
littoral.

Pariphimediella Schellenberg

Pariphimediella Schellenberg, 1931.

Type-species: Iphimedia serrata Schellenberg, 1926a (original
designation).

Upper lip broad, faintly emarginate; mandible with narrow, toothed
apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate,
reaching end of outer plate; maxillipedal palp exceeding outer plate,
3-articulate, article 2 faintly produced; gnathopods chelate; telson
broadly emarginate or cleft. Species: 5, antarctic, littoral to bathyal.

Pseudiphimediella Schellenberg
Pseudiphimediella Schellenberg, 1931.

Type-species: Amphitoé nodosa Dana, 1853 (original designation).

Upper lip broad, incised; mandible with broad, toothed apex; lobes
of lower lip not incised; palp of maxilla 1 biarticulate, reaching end
of outer plate; maxillipedal palp exceeding outer plate, 3-articulate,
article 2 not produced, articles 1-2 broadened; gnathopod 1 chelate,
enathopod 2 slightly chelate; telson entire or faintly emarginate.
Species: 1, subantarctic, littoral.

128 U.S. NATIONAL MUSEUM BULLETIN 271

Ampeliscidae

Ficures 57, 58

Diaenosis.—Accessory flagellum absent; pereopod 5 shorter than
and of different structure from 4; article 4 of pereopods 1—2 elongate;
head elongate; eyes when present bearing maximum of two pairs of
anterolateral cuticular lenses; urosomites 2—3 coalesced; pereopods 1-2
glandular. See Atylidae, Haustoriidae, Argissidae.

Description.—Accessory flagellum absent; head elongate, tall,
compressed; body lacking dorsal processes, except on pleonite 4 and
occasionally low carinae on pleonites 1-3; antennae and their pe-
duncles elongate, often with long setae, male primary flagellum conjoint
basally ; anterior coxae long, coxa 1 often broader than 2, 2 occasionally
tapering; mouthparts basic but mandibular palp article 3 of many
species much shorter than article 2 and mandibular lobes of lower lip
obsolescent; gnathopods feeble, subchelate or nearly simple; article 4 of
pereopods 1-2 elongate; pereopod 5 always shorter and of different
morphology from pereopod 4; pereopods with a few elongate setae,
especially on article 2 of pereopod 5 and article 4 of pereopods 1-2,
pereopods 3-4 often with strong submarginal spines on articles 4-5;
rami of uropod 3 lanceolate, usually elongate, occasionally foliaceous
or shortened; telson either elongate or very short, usually cleft, cleft
often short.

RELATIONSHIP.—The shiny cuticular lenses of oculate ampeliscids
are unique to this family, except for their rare occurrence in a few
lysianassids. Oculate ampeliscids also have bright red, brown, or black
pigmentary masses in the head towards which, perhaps, the cuticular
lenses direct light.

Pereopods of this family are definitively fossorial but presumably
they are used not in digging but in clinging to or rearranging the
insides of their tubes spun by pereopods 1-2 or in creating water
currents and feeding.

The Phoxocephalidae always bear a rostrum and a multiarticulate
accessory flagellum. The Haustoriidae bear a multiarticulate accessory
flagellum. Both of those families form burrows in sediments whereas
Ampeliscidae are provided with glands in the pereopods for the
construction of tubes lying on the substrate surface.

The Atylidae have ommatidial eyes and pereopod 5 is usually
longer than pereopod 4 and of similar structure; the pereopods are
not glandular.

MARINE GAMMARIDEAN AMPHIPODA

129

IN &

Sa
>

N
\

SNS

Figure 57.—Ampeliscidae: a, Ampelisca tenuicornis Liljeborg (Sars, 1895, pl. 58);
b, Byblisoides arcillis J. L. Barnard (1961).

130 U.S. NATIONAL MUSEUM BULLETIN 271

Key to the Genera of Ampeliscidae

1. Pereopod 5: anterior edge of posterior lobe on article 2 lacking setae near its

junction with article 3 (fig. 581)... . . ERP henge 102)
Pereopod 5: anterior edge of posterior lobe on amills 2 penne setae up to

its junction with article 3 (figs. 58m,n,o) . . 4

2. Flagella of antennae poorly developed, with 2 46 4 amiclles Ge. 57b) [eniene-
ventral corner of head produced (fig. 57b)] . .. . =... . Byblisoides
Flagella of antennae long, usually with more than 10 articles [anteroventral
cornerorheadirarelyjproduced| aes) sites e alee eS

3. Anteroventral corner of head produced. ... . Joon ee eee Oo dos
Anteroventral corner of head sloping, Anorodecdt | .... . . Ampelisca

4. Posterior lobe of article 2 on pereopod 5 strongly expanded distally, posterior
edge oblique (fig. 58m) ...... Ale By Dlis
Posterior lobe of article 2 on pereopod 5 noe etoamdledl dis! ally, posterior edge
vertical (igs. 58730) 2 2 iss a ee ee enloons

Genera of Ampeliscidae

Ampelisca Kr¢yer

Ampelisca Krgyer, 1842.—Stebbing, 1906.
Pseudophthalmus Stimpson, 1853.
Araneops Costa, 1853c, 1857.

Tetromatus Bate, 1856, 1857a, 1857c.

Type-species: A. eschrichtia Krgyer, 1842 (monotypy). See Sars,
1895.

Pereopod 5 with posterior lobe of article 2 greatly expanded distally,
posterior edge oblique, anterior edge of posterior lobe lacking setae
near its junction with article 3; palp article 3 of mandible variable in
length; antenna 2 with more than five flagellar articles; anteroventral
corner of head unproduced. Species: 94, primarily littoral cosmo-
politan, about 20 bathyal and one abyssal species.

Byblis Boeck
Byblis Boeck, 1871.—Stebbing, 1906.

Type-species: Ampelisia (lapsus) gaamardu Krgyer, 1846 (mono-
typy). See Sars, 1895.

Pereopod 5 with posterior lobe of article 2 greatly expanded distally,
posterior edge oblique, anterior edge of posterior lobe bearing setae
near its junction with article 3; palp article 3 of mandible shorter
than article 2; antenna 2 with more than five flagellar articles; antero-
ventral corner of head unproduced. Species: 21, cosmopolitan,
littoral to abyssal.

MARINE GAMMARIDEAN AMPHIPODA 131

Soa

Ficure 58.—Ampeliscidae: Upper lip: a, Ampelisca typica (Bate) (Sars, 1895, pl. 57):
Mandible: b, Ampelisca; c, Byblis gaimardi (Kr@yer) (Sars, 1895, pl. 64). Lower lip:
d, Ampelisca; e, Byblis. Maxillae 1-2: f,g, Ampelisca. Maxilliped: h, Ampelisca.
Gnathopods 1-2, Pereopods 1,5: 1,7,k,/, Ampelisca. Pereopod 5: m, Byblis; n, Haploops
setosa Boeck (Sars, 1895, pl. 68); 0, Haploops tubicola Liljeborg (Sars, 1895, pl. 67). Uro=
pod 3: p, Ampelisca. Telson: g, Ampelisca; r, Byblis.

132 U.S. NATIONAL MUSEUM BULLETIN 271

Byblisoides K. H. Barnard

Byblisoides K. H. Barnard, 1931, 1932.

Type-species: B. juaticorns K. H. Barnard, 1931 (original
designation).

Pereopod 5 with posterior lobe of article 2 greatly expanded distally,
posterior edge oblique, anterior edge of posterior lobe lacking setae
near its junction with article 3; palp article 3 of mandible shorter than
article 2; antenna 2 with less than five flagellar articles; anteroventral
corner of head produced. Species: 4, tropics to antarctic, bathyal.

Haploops Liljeborg
Haploops Liljeborg, 1856.—Stebbing, 1906.

Type-species: H. tubicola Liljeborg, 1856 (original designation).
See Sars, 1895.

Pereopod 5 with posterior lobe of article 2 not expanded distally,
usually narrow, posterior edge vertical, anterior edge of posterior lobe
bearing setae near its junction with article 3; palp article 3 of mandible
as long as article 2; antenna 2 with more than five flagellar articles;
anteroventral corner of head unproduced. Species: 13, cosmopolitan,
cold-water, littoral to abyssal, primarily bathyal.

Triodos K. H. Barnard
Triodos K. H. Barnard, 1916.

Type-species: 7. insignis K. H. Barnard, 1916 (monotypy).

Pereopod 5 with posterior lobe of article 2 greatly expanded distally,
posterior edge oblique, anterior edge of posterior lobe lacking setae
near its junction with article 3; palp article 3 of mandible as long as
article 2; antenna 2 with more than five flagellar articles; anteroventral
corner of head produced. Species: 1, S. Africa, littoral.

Amphilochidae

Fiaures 59, 60

Dracnosis.—Accessory flagellum absent; coxa 1 very small, par-
tially hidden by following coxae. See Pleustidae, Calliopiidae, Leuco-
thoidae, Anamixidae, Stenothoidae, Thaumatelsonidae, Cressidae,
Stegocephalidae.

Description.—Accessory flagellum absent; rostrum conspicuous;
coxa 1 very small in all but one genus and partially hidden by fol-

134 U.S. NATIONAL MUSEUM BULLETIN 271

lowing coxae, coxa 2 occasionally as small as coxa 1 but often as large
as coxa 3 and not hidden; coxae 3-4 enlarged, either overlapping or,
when immensely enlarged, with contiguous margins abutting; upper
lip usually deeply incised, rarely slightly excavate; mandible usually
with 3-articulate palp, rarely absent, molar well developed and trit-
urative or formed of a smooth protuberance or evanescent; lower
lips of two kinds (see figures); maxilla 1 with 1- or 2-articulate palp;
maxilla 2 rarely reduced in size, in one genus formed of a single plate;
maxillipeds normal; gnathopods of medium size or small, subchelate
or nearly simple, often incompletely carpochelate; uropod 2 shortened ;
uropod 3 biramous, peduncle elongate (except Pseudamphilochus) ;
telson entire in all but one genus, often elongate and triangular, also
short and linguiform.

RELATIONSHIP.—The Pleustidae and Calliopiidae have large first
coxae, not hidden by following coxae. Leucothoidae have completely
carpochelate first gnathopods because article 6 is narrow.

The Stegocephalidae always lack a mandibular molar and palp; no
amphilochid lacks both at the same time; the mouthparts of stego-
cephalids project in a conical bundle, an accessory flagellum is present
and the first coxa is never hidden by the following coxae.

The Stenothoidae, Thaumatelsonidae, and Cressidae have uni-
ramous third uropods.

The Anamixidae have a completely carpochelate first gnathopod or
the appendage is absent, and a ventral cephalic keel replaces the
mandibles and maxillae.

Pseudamphilochus Schellenberg is an aberrant amphilochid in its
normal coxa 1 and cleft telson. It tends to fall into the Eusiridae in
the various keys but a special place has been made for it also in
Gammaridea Family Key, Section G. It also appears to be a pleustid
with cleft telson even though its lower lip is not fully typical of
pleustids. It can also be confused with liljeborgiids even though the
accessory flagellum is absent in Pseudamphilochus.

Incomplete carpochelation of gnathopods is ignored in the keys
and diagnoses to follow and the terms “simple” and ‘‘subchelate”’
refer only to the condition of the propodus (article 6).

Neocyproidea peninsulae Hurley (1955) is removed to Peltopes.

Key to the Genera of Amphilochidae

1. Coxae 3-4 not immensely broadened, with contiguous margins overlapping,
not concealing coxa 2 (fig. 59a) . . . (Amphilochinae, new subfamily) 2
Coxae 3-4 immensely broadened, contiguous margins abutting, concealing
the vestigial first two coxae (fig. 59b) . (Cyproideinae, new subfamily) 10

2. Mandibular molar large, with ridged and toothed triturating surface (fig.
COG re ABER EE TN et CAN Ca area tea ab En GTS CO LE EARS ee a 3

10.

12.

13.

14.

15.

16.

MARINE GAMMARIDEAN AMPHIPODA 135

Mandibular molar small, or absent, unarmed or bearing one to three spines

ieGOd) ae is |: BOA Be teny te Se" Ab. SRD terete at 0
Palp of maxilla 1 ith ‘ares Pericles (fig. 60;) . ...... . . Gitanopsis
Palp of maxilla 1 with one article (fig. 60k) ...... ee ee A:
Outer plate of maxilliped strongly excavate medially, prticle 1 of palp much

longer than other palp articles (fig. 60p) ... . . . . Gitanogeiton
Outer plate of maxilliped straight or slightly oun medially, article 1 of

palpssubequal-to article!2 (figiGQo) 9) Vrs 259° IR) Garg foc s Rob
Gnathopod 2 large, subchelate (fig. 60v) . ... . .. . Amphilochopsis
Gnathopod 2 small, nearly simple (fig. 60w) . . . . 3 . . Gitana
Maxilla 2 composed of only one elongate plate (fig. 60m) . : Agauhiloehelia
Maxilla 2 composed of two plates (fig. 60/) .... ape N,
Maxilla 2 degraded, plates tiny, subequal in width Ge 60n) . Ry eS
Maxilla 2 normal, inner plate much broader than outer (fig. 60/1) . . . . 9
Posterolateral angles of pleonite 6 not produced. . . . . Amphilochoides
Posterolateral angles of pleonite 6 produced, reaching apex of telson.
Cyclotelson
Melsonventire) (fig. G0z) . -oegek 2 »sgease «>. ev.) Amphilochus
Telson.cleft (fig; 60y).... =»... ... .. . . Pseudamphilochus
Article 2 of pereopods 4-5 flac. Blender Shires: SA cep cai srige al
Article 2 of pereopod 5 and usually pereopod 4 sanded sa Rue Drentas Ss
Palm of gnathopod 2 transverse, urosomite 3 vaulting over telson, telson

Smallest ser hae ... . . Cyproidea
Palm of gnathopod 2 oblique TOSS 3 not ~sanlGing over telson, telson

NWS eR Ay ee: i de une ee ay IP aracyproiged'
Urosomite 1 wealeacled, toner or hast Soh NALCO OBEN ERENT a arrue Lem pa aeea NA ka)
Urosomite 1 dorsally keeled, elongate . . . eee eee
Gnathopod 2 simple; uropod 2 shortened, not senrahtaye end of uropod 3

(igse 590,0)) i) 2) fa ... . . Stegoplax
Gnathopod 2 subchelate, ogi ene alan mroparl 2 reaching end of

uropod 3 (figs. 60bb,cc) . . . Rie lege: Se ls SPeltocoxa
Article 2 of pereopod 3 slender, tere Bee Serna Par ca FO ae anes ae oe MON al 10)
Article 2 of pereopod 3 expanded .... . Hie dee tar mae Sal 6)

Palp of maxilla 1 biarticulate, outer plate of card lived nendatlag to end of
palp article 2, article 7 of gnathopod 1 apparently fitting oblique palm.

Hoplopleon

Palp of maxilla 1 uniarticulate, outer plate of maxilliped reaching to end of

palp article 1, article 7 of gnathopod 1 greatly overlapping short, trans-

verse palm ... . ee ew ww ee es .). . . Hoplopheonoides
Mandibular palp cen PR aete Ceotan Sonat ae tae See ger obese Pe LLODeS
Mandibular palp absent .............. ... Neocyproidea

Genera of Amphilochidae

Subfamily Amphilochinae, new subfamily

Type-genus: Amphilochus Bate.
Coxae 2-4 not immensely broadened, with contiguous margins

overlapping.

i36). U.S. NATIONAL MUSEUM BULLETIN 271

Amphilochella Schellenberg
Amphilochella Schellenberg, 1926a.

Type-species: <A. simplicarpus Schellenberg, 1926a (monotypy).

Mandibular molar absent; palp of maxilla 1 biarticulate; maxilla 2
composed of one elongate plate; outer plates of maxillipeds not
excavate; palp article 1 shorter than article 2; gnathopod 2 small,
slender, scarcely subchelate; telson apparently entire. Species: 1,
antarctic, littoral.

Amphilochoides Sars
Amphilochoides Sars, 1895.—Stebbing, 1906.

Type-species: Amphilochus odontonyx Boeck, 1871 (original desig-
nation). This species is figured by Sars erroneously under his Amphilo-
choides pusillus which becomes a synonym of A. odontonyx; Sars’
description and figures of A. odontonyx are proposed by him on p. 690
to have the name A. boeckiit which Stebbing (1906) finds is a junior
synonym of Probolium serratipes Norman (1869a).

Mandibular molar obsolete; palp of maxilla 1 biarticulate; lower lip
atypical for family, with tilted oval outer lobes astride distinct inner
lobes; maxilla 2 degraded but with two distinct subequal lobes; outer
plates of maxillipeds not excavate, palp article 1 subequal to article 2;
gnathopod 2 large, subchelate; telson entire. Species: 4, N.E.
Atlantic, littoral.

Amphilochopsis Stephensen
Amphilochopsis Stephensen, 1925a.

Type-species: A. hamatus Stephensen, 1925a (monotypy).

Mandibular molar large, triturative; palp of maxilla 1 uniarticulate;
maxilla 2 normal, outer plate narrower than inner; outer plates of
maxillipeds not excavate, palp article 1 subequal to article 2; gnathopod
2 large, subchelate; telson entire. Species: 1, subarctic, bathyal to
abyssal (825-2702 m).

Amphilochus Bate

Amphilochus Bate, 1862.—Stebbing, 1906.
Callimerus Stebbing, 1876.
Type-species: A.manudens Bate, 1862 (monotypy). See Sars, 1895.
Mandibular molar small, not triturative; palp of maxilla 1 biarticu-
late; maxilla 2 normal, outer plate narrower than inner; outer plates
of maxillipeds not excavate, palp article 1 subequal to article 2;
enathopod 2 large, subchelate; telson entire. Species: 11, cosmo-
politan, littoral (one species to 913 m).

MARINE GAMMARIDEAN AMPHIPODA ey

Ficure 60.—Amphilochidae: Upper lip: a, Amphilochus manudens Bate (Sars, 1895, pl.
74); b, Pseudamphilochus shoemakeri Schellenberg (1931). Mandible: c, Gitanopsis
bispinosa (Boeck) (Sars, 1895, pl. 76); d, Amphilochus; e, Neocyproidea otakensis Hurley
(1955); f, Pseudamphilochus. Lower lip: g, Gitanopsis; h, Amphilochoides odontonyx
(Boeck) (Sars, 1895, pl. 76); 1, Amphilochus. Maxilla 1:7, Amphilochus; k, Gitana sarsi
Boeck (Sars, 1895, pl. 78). Maxilla 2: 1, Amphilochus; m, Amphilochella simplicarpus
Schellenberg (1926a); », Amphilochoides. Maxilliped: 0, Amphilochus; p, Gitanogeiton
sarsi Stebbing (1910). Gnathopod 1: g, Amphilochus; r, Gitana; 5, Hoplopheonotdes
obesa Shoemaker (1956a); t, Peltopes productus K. H. Barnard (1930). Gnathopod 2:
u, Gitana; v, Amphilochus; w, Hoplophenonoides. Telson: x, Amphilochus; y, Pseudam-
philochus; x, Cyclotelson purpureum Potts (1915). Uropod 3: aa, Amphilochus. Pleon
and Telson, lateral: bb, Hoplopheonoides; cc, Peltocoxa marioni Catta (Chevreux and Fage,
1925).

138 U.S. NATIONAL MUSEUM BULLETIN 271

Cyclotelson Potts
Cyclotelson Potts, 1915.

Type-species: C. purpureum Potts, 1915 (monotypy).

Mandibular molar absent; palp of maxilla 1 biarticulate; maxilla 2
degraded but with two distinct subequal lobes; outer plates of maxilli-
peds not excavate, palp article 1 subequal to article 2; gnathopod 2
small, subchelate; telson entire; lateral angles of urosomite 3 strongly
produced (combining character). Species: 1, Gulf of Mexico, littoral.

Gitana Boeck
Gitana Boeck, 1871.—Stebbing, 1906.

Type-species: G. sarst Boeck, 1871 (designated by Sars, 1895,
p. 229, ‘‘Remarks’’).

Mandibular molar large, triturative; palp of maxilla 1 uniarticulate;
maxilla 2 normal, outer plate narrower than inner; outer plates ‘of
maxillipeds slightly excavate, palp article 1 subequal to article 2;
gnathopod 2 small, nearly simple; telson entire. Species: 4, boreal,
littoral to bathyal (475 m).

Gitanogeiton Stebbing
Gitanogeiton Stebbing, 1910.

Type-species: G. sarsi Stebbing, 1910 (monotypy).

Mandibular molar large, triturative; palp of maxilla 1 uniarticulate;
maxilla 2 normal, outer plate narrower than inner; outer plates of
maxillipeds strongly excavate, palp article 1 much longer than other
articles; gnathopod 2 small, subchelate [telson damaged]. Species: 1,
off Manning R., Australia, littoral.

Gitanopsis Sars
Gitanopsis Sars, 1895.—Stebbing, 1906.

Type-species: Amphilochus bispinosus Boeck, 1871 (original desig-
nation).

Mandibular molar large, triturative; palp of maxilla 1 biarticulate;
maxilla 2 normal, outer plate narrower than inner; outer plates of
maxilliped rarely excavate, palp article 1 subequal to article 2;
enathopod 2 small, subchelate; telson entire. Species: 12, cosmo-
politan, littoral (2 species in bathyal to 1096 m).

MARINE GAMMARIDEAN AMPHIPODA 139

Pseudamphilochus Schellenberg
Pseudamphilochus Schellenberg, 1931.

Type-species: P. shoemakert Schellenberg, 1931 (monotypy).

Mandibular molar obsolete; palp of maxilla 1 biarticulate; maxilla
2 normal, outer plate narrower than inner; outer plates of maxilliped
not excavate, palp article 1 subequal to article 2; coxa 1 not reduced;
enathopod 2 small, subchelate; telson deeply cleft. Species: 1,
S. Georgia Is., littoral.

Subfamily Cyproideinae, new subfamily

Type-genus: Cyproidea Haswell.
Coxae 3-4 immensely broadened, with contiguous margins abutting,
hiding coxae 1 and 2.

Cyproidea Haswell

Cyproidea Haswell, 1880a.—Stebbing, 1906.
Cyproidia [sic] Haswell, 1880c.
Gallea Walker, 1904.

Type-species: Cyproidia [sic] ornata Haswell, 1880c (present
selection). See Walker, 1904 (as Gallea tecticauda).

Article 2 of pereopods 3-5 linear, slender; gnathopod 2 subchelate,
palm transverse; uropod 2 long; urosomite 1 not elongate, not keeled;
urosomite 3 vaulting over telson; telson normal; mandibular molar
absent or present, palp 3-articulate. Species: 2, tropics to antiboreal,
EK. Hemisphere, littoral.

Hoplopheonoides Shoemaker
Hoplopheonoides Shoemaker, 1956a.

Type-species: H. obesa Shoemaker, 1956a (monotypy).

Article 2 only of pereopod 5 expanded, of pereopods 3-4 slender and
linear (Hoplopleon); gnathopod 2 subchelate, tending to be chelate;
uropod 2 long; urosomite 1 elongate, dorsally keeled; urosomite 3 not
vaulting over telson; telson normal; mandibular molar well developed,
triturative, palp absent; palp of mazilla 1 uniarticulate (Hoplopleon).
Species: 1, Florida, littoral. Hoplopleon similis Schellenberg is inter-
mediate between this and the next genus.

285-135 O - 69 - 10

140 U.S. NATIONAL MUSEUM BULLETIN 271

f Hoplopleon K. H. Barnard
Hoplopleon K. H. Barnard, 1932.

Type-species: Peltocoxa australis K. H. Barnard, 1916 (original
designation).

Article 2 of pereopods 4-5 expanded, of pereopod 3 slender (Hoplo-
pheonoides); gnathopod 2 subchelate, palm oblique or transverse;
[uropod 2 unknown]; urosomite 1 elongate, keeled; urosomite 3 not
vaulting over telson; telson normal; mandibular molar triturative,
palp absent; palp of maxilla 1 biarticulate (Hoplopheonoides). Species:
3, antarctic to S. W. Africa, littoral.

Neocyproidea Hurley
Neocyproidea Hurley, 1955.

Type-species: Cyproidea otakensis Chilton, 1900 (original
designation).

Article 2 of pereopods 3-5 somewhat expanded; gnathopod 2 nearly
simple; uropod 2 long; urosomite 1 elongate, dorsally keeled, slightly
vaulting to base of telson or less; urosomite 3 not vaulting over telson;
telson normal; mandibular molar strong, palp absent. Species: 2,
New Zealand, littoral.

Paracyproidea Stebbing
Paracyproidea Stebbing, 1899d, 1906.

Type-species: Cyproidia [sic] lineata Haswell, 1880c (original
designation).

Article 2 of pereopods 3-5 slender, linear; gnathopod 2 subchelate,
palm oblique; uropod 2 long; urosomite 1 not elongate, not keeled;
urosomite 3 not vaulting over telson; telson huge; mandibular molar
triturative, palp 3-articulate. Species: 1, E. Australia, littoral.

Peltocoxa Catta
Peltocoza Catta, 1875.—Stebbing, 1906.

Type-species: P. marioni Catta, 1875 (original designation). See
Chevreux and Fage, 1925.

Article 2 of pereopods 4-5 expanded, of pereopod 3 slender and
linear; gnathopod 2 subchelate, palm transverse; uropod 2 long;
urosomite 1 short, unkeeled; urosomite 3 not vaulting over telson;
telson huge; mandibular molar triturative, palp 3-articulate. Species:
2, E. Atlantic, littoral.

MARINE GAMMARIDEAN AMPHIPODA 141

Peltopes K. H. Barnard
Peltopes K. H. Barnard, 1930.

Type-species: P. productus K. H. Barnard, 1930 (monotypy).

Article 2 of pereopods 3-5 expanded; gnathopod 2 simple; uropod
2 long; urosomite 1 elongate, dorsally keeled, process vaulting over
following segments; urosomite 3 not vaulting over telson; telson
normal; mandibular molar [unknown in type] strong, palp 3-articulate.
Species: 2, New Zealand, littoral.

Stegoplax Sars
Stegoplax Sars, 1882.—Stebbing, 1906.

Type-species: SS. longirostris Sars, 1882 (original designation). See
Sars, 1895.

Article 2 of pereopods 3-5 expanded; gnathopod 2 simple; uropod 2
shortened; urosomite 1 short, unkeeled; urosomite 3 slightly vaulting
over telson; telson normal; mandibular molar triturative, palp 3-
articulate. Species: 1, N. Atlantic, bathyal.

Ampithoidae

FIGURE 61

Draenosis.—Accessory flagellum present or absent; telson entire,
short, fleshy ; lower lip with anterior lobes notched or medially excavate
(figs. 61e,f); coxa 4 not excavate posteriorly; pereopods glandular;
uropod 3 with very small, short rami, shorter than peduncle, rami
quadrate, blunt, outer armed with one or two hooks. See Isaeidae,
Ischyroceridae, Oedicerotidae, Calliopiidae, Pleustidae.

Description.—Accessory flagellum varying from absent to long and
multiarticulate; body smooth; rostrum absent; coxae medium in size,
quadrate or rounded, coxa 4 not excavate posteriorly; mouthparts
basic except for one genus lacking mandibular palp and another with
reduced molar; lower lip with lateral (outer) lobes projecting, notched
(bilobed) or medially excavate; gnathopods usually powerful, sub-
chelate, in one genus feeble and chelate, gnathopod 1 usually smaller
than 2, occasionally larger than 2; uropod 3 with very short quadrate
rami, shorter than peduncle, outer ramus armed with one or two
hooks; telson short, entire, nearly circular, square or triangular, fleshy.

ReELATIONSHIP.—The notched outer lobes of the lower lip distinguish
this family from its relatives, the Isaeidae, Aoridae, Ischyroceridae,
and Corophiidae. The very short quadrate rami with a hook or hooks
on the outer are distinctive (but see Jassa in Ischyroceridae).

142 U.S. NATIONAL MUSEUM BULLETIN 271

The Ischyroceridae also have short rami but the peduncle is more
elongate, the rami are superficially lanceolate and poorly setose, and
the outer is armed with denticles, not large hooks (except Jassa).

The genus Amphitholina Ruffo was transferred to Eophliantidae by
Gurjanova (1938) ; it bridges these two families in its intersimilar third

Ficure 61.—Ampithoidae: a, Ampithoe rubricata (Montagu) (Sars, 1895, pl. 206). Upper
lip: b, Ampithoe. Mandible: c, Ampithoe; d, Sunamphitoe pelagica (Milne Edwards)
(Sars, 1895, pl. 208, as S. conformata). Lower lip: e, Ampithoe; f, Pleonexes gammaroides
(Bate) (Sars, 1895, pl. 207). Maxillae 1-2, Maxilliped: g,h,i, Ampithoe. Gnathopod
1: 3, Ampithoe; k, Macropisthopus stebbingi K. H. Barnard (1916). Gnathopod 2: J,
Ampithoe. Pereopod 5: m, Ampithoe; n, Pleonexes. Uropod 3: 0, Ampithoe. ‘Telson:
p, Ampithoe; q, Pleonexes.

MARINE GAMMARIDEAN AMPHIPODA 143

uropods, but the degeneration of mouthparts and coxae and the sub-
cylindrical body mark it as an eophliantid.

The shape of uropod 3 distinguishes the Ampithoidae from the
Calliopiidae, Pleustidae, and Oedicerotidae.

Nomenclatural Changes in Ampithoidae

Amphitholina Ruffo (1953) is removed to Kophliantidae.
Amphithoides patrizi. Maccagno (1936) is removed to Cymadusa.

Key to the Genera of Ampithoidae

1. Mandible lacking palp (fig. 61d) .......... .. . . Sunamphitoe
Mandibleypearimp=palp (fig. Gillie): 2k on SA aoe, Se a ARN
ZepAntenna,l lacking accessory flagellum: 64 os si. hee. ). Bole e. 8
Antenna | bearing accessory flagellum. . . pla har ene eens arse eee 0)
3. Gnathopod 1 larger and stouter than apetingned oy. . . . . . Exampithoe
Gnathopod 1 smaller and more slender than gnathopod2 ....... 4
4. Gnathopods feeble, chelate (fig. 61k). . . ... =. +... . .Macropisthopus
Gnathopods large, subchelate (figs. 617,1). . . . 5

5. Article 6 of pereopods 3-5 widened apically, siripemalieastlle, Gis. 61n), ‘iin
bearing large striated spines, telson with apical pair of reverted, elongate,
cornified processes (fig. PUD) Aiea etre . .. . . Pleonexes

Article 6 of pereopods 3-5 widened or mee Pidened rarely subprehensile
(fig. 61m), apical cornified processes of telson if present, obsolescent (fig.

Gil) te Panis Si nane me ages Jom eon AIM DIthOe

6. Outer ramus of ameyoudl 3 Beane one Thaliels ..... . . . Amphithoides
Outer ramus of uropod 3 bearing two hooks (fig. 6lo) ......... 7

7. Gnathopod 1 larger than gnathopod 2 ......... .. . Paragrubia
Gnathopod 1 smaller than gnathopod. ........... . Cymadusa

Genera of Ampithoidae

Amphithoides Kossmann
Amphithoides Kossmann, 1880.—Stebbing, 1906.

Type-species: A. longicornis Kossmann, 1880 (monotypy).

Antenna 1 with accessory flagellum; mandible with palp; gnatho-
pods large, subchelate, gnathopod 2 equal to or larger than 1; article 6
of pereopods 3-5 not apically widened; outer ramus of uropod 3 with
one hook. Species: 1, Red Sea, littoral.

Ampithoe Leach

Ampithoe Leach, 1814a.—Stebbing, 1906.

Type-species: Cancer (Gammarus) rubricatus Montagu, 1808
(monotypy). See Sars, 1895.

144 U.S. NATIONAL MUSEUM BULLETIN 271

Antenna 1 lacking accessory flagellum; mandible with palp; gnatho-
pods large, subchelate, gnathopod 2 equal to or larger than 1; article 6
of pereopods 3-5 scarcely widened apically, rarely prehensile (Pleo-
nexes); outer ramus of uropod 3 with two hooks; apical cornified
processes of telson, wf present, obsolescent (Pleonexes). Species: 35,
cosmopolitan, littoral.

Cymadusa Savigny

Cymadusa Savigny, 1816.
Grubia Czerniavski, 1868.
Acanthogrubia Stout, 1912.

Type-species: OC. filosa Savigny, 1816 (monotypy). See Chevreux
and Fage, 1925 (as Grubia hirsuta), Shoemaker, 1935 (as Grubia
filosa).

Antenna 1 with 1- or 2-articulate accessory flagellum; mandible
with palp; gnathopods large, subchelate, gnathopod 2 equal to or
larger than 1; article 6 of pereopods 3-5 not apically widened; outer
ramus of uropod 3 with two hooks. Species: ca. 10, generally tropical-
amphiboreal, littoral.

Exampithoe K. H. Barnard
Exampithoe K. H. Barnard, 1925.

Type-species: HH. natalensis K. H. Barnard, 1925 (monotypy).

Antenna 1 lacking accessory flagellum; mandible with palp, molar
reduced; gnathopods large, subchelate, gnathopod 1 stouter than 2;
article 6 of pereopods 3-5 apically widened; outer ramus of uropod 3
with two hooks. Species: 1, 8. Africa, littoral.

Macropisthopus K. H. Barnard
Macropisthopus K. H. Barnard, 1916.

Type-species: MM. stebbingi K. H. Barnard, 1916 (monotypy).

Antenna 1 lacking accessory flagellum; mandible with palp; gnatho-
pods feeble, chelate; article 6 of pereopods 3-5 scarcely widened
distally; outer ramus of uropod 3 with two hooks. Species: 1, S.
Africa, littoral.

Paragrubia Chevreux
Paragrubia Chevreux, 1901a.

Type-species: P. vorax Chevreux, 1901a (monotypy).

Antenna 1 with accessory flagellum; mandible with palp; gnathopods
large, subchelate, gnathopod 1 larger than 2; article 6 of pereopods
3-5 not apically widened; outer ramus of uropod 3 with two hooks.
Species: 1, tropical Pacific, littoral.

MARINE GAMMARIDEAN AMPHIPODA 145

Pleonexes Bate

Anisopus Templeton, 1836 (homonym, Diptera).
Pleonexes Bate, 1857a.

Type-species: P. gammaroides Bate, 1857a (monotypy). See Sars,
1895.

Antenna 1 lacking accessory flagellum; mandible with palp;
enathopods large, subchelate, gnathoped 2 equal to or larger than 1;
article 6 of pereopods 3-5 apically widened and subprehensile (Ampithoe) ;
outer ramus of uropod 3 with two hooks; telson with apical pair of
reverted, elongate, cornified . processes (Ampithoe). Species: 3, N.E.
Atlantic, New Zealand, littoral.

Sunamphitoe Bate
Sunamphitoe Bate, 1857a.

Type-species: Amphithoe pelagica Milne Edwards, 1830 (desig-
nated by Chevreux and Fage, 1925). See Sars, 1895 (as S. conformata).

Antenna 1 lacking accessory flagellum; mandible lacking palp;
enathopods large, subchelate, gnathopod 2 equal to or larger than
1; article 6 of pereopods 3-5 not apically widened; outer ramus of
uropod 3 with two hooks. Species: 2, N.K. Atlantic, epipelagic.

Anamixidae

FIGURE 62

Diaenosis.—Mandible, lower lip, and maxillae absent; buccal
regions with ventrally projecting keel; outer plates of maxillipeds
absent, inner plates small, fused together; coxa 1, when present,
hidden by the shield-like coxae 2—4; telson entire. See Leucothoidae,
Amphilochidae, Cressidae, Sebidae, Thaumatelsonidae.

Dzscrietion.—Body smooth, lacking dorsal processes; mouth-
parts aberrant as in diagnosis; accessory flagellum absent; gnathopod
1, when present, small and complexly chelate, having a chelate
process on article 5, articles 6 and 7 (when present) together forming
dactyl; gnathopod 2 immense, article 5 forming a false chela but
articles 6 and 7 large, and article 7 alone forming the true dactyl;
uropod 3 with styliform rami as long as the elongate peduncle; telson
entire.

RELATIONSHIP.—The Leucothoidae bear mandibles and maxillae.

Key to the Genera of Anamixidae

Pee EGmathopodlepresemti cies 8 fr eels ved ok eA Ce a ese inca a ein hae Anamixis
Cnathopodwltalosemt yeu nee ila mien eo ee ae aia ele iy aun Paranamixis

146 U.S. NATIONAL MUSEUM BULLETIN 271
Genera of Anamixidae

Anamixis Stebbing
Anamixis Stebbing, 1897; 1906.

Type-species: A. hanseni Stebbing, 1897 (monotypy).
Species: 4, Indo-Pacific tropics and subtropics, littoral (sponges).

Paranamivis Schellenberg

Paranamixis Schellenberg, 1938.

Type-species: P. bocki Schellenberg, 1938 (monotypy).
Species: 1, Gilbert Islands, littoral.

Figure 62.—Anamixidae: a, Anamixis hanseni Stebbing (1897). Anamixis linsleyi J. L.
Barnard (1955b): 6, oblique ventral view of head showing keel presumably formed of
mandibles and maxillae: c, maxilliped; d, e, gnathopods 1, 2; f, uropod 3; g, telson.

MARINE GAMMARIDEAN AMPHIPODA 147

Aoridae

FIGURES 638, 64

Draenosis.—Accessory flagellum variable, multiarticulate, or ab-
sent; telson entire, short, fleshy; coxa 4 not excavate posteriorly;
pereopods glandular; wropod 3 rarely projecting beyond uropods 1 and 2,
at least one of rami as long as or longer than peduncle; gnathopod 1
larger than gnathopod 2, in rare cases only as large as gnathopod 2
but with male secondary sexual modifications. See Isaeidae, Coro-
phiidae, Ischyroceridae.

Description.—Accessory flagellum varying from absent to long
and multiarticulate; body smooth; rostrum vestigial; coxae rounded
or quadrate below, varying from long to short, fourth not excavate
posteriorly; mouthparts basic; gnathopods powerful, subchelate or
complexly subchelate, occasionally nearly simple, but then often
extremely setose, first always larger or more complex than second in
both sexes and bearing most of male secondary sexual modifications;
pereopod 5 usually conspicuously elongate; uropod 3 short, rami
usually as long as or longer than peduncle, occasionally inner ramus
reduced in size or both rami reduced; telson short, nearly circular or
square, entire, occasionally falsely (?secondarily) cleft.

RELATIONSHIP.—Like the Isaeidae but having an enlarged first
gnathopod; otherwise Aoridae are related to other families in the
same way as are the Isaeidae. Keys of both families and the Coro-
phiidae should be checked when examining any genera with abnormally
enlarged or complexly subchelate gnathopods. One may understand
the difficulties of segregating Aoridae and Corophiidae when examining
the small differences between Grandidierella (Corophiidae) and
Neomicrodeutopus (Aoridae).

Nomenclatural Changes in Aoridae

The female of Dryopoides was first assigned to the Aoridae by
Stebbing (1888). Its first gnathopod is larger than the second. In 1910,
Stebbing discovered the male, with gnathopod 2 larger than 1, but the
species was transferred to the Corophiidae. Because the genus may yet
prove to be an aorid, the female is retained in the key to Aoridae as
if it were a male.

Paradryope and Dryopoides might be placed in the family Ischyro-
ceridae, especially Paradryope because of the short rami of its uropod
3 and the slightly elongate peduncle. The uropodal peduncle of
Dryopoides is not greatly elongate. Ischyroceridae would thus receive
genera with gnathopod 1 larger than 2, of which Bonnierella is an

U.S. NATIONAL MUSEUM BULLETIN 271

148

AQ
S
NS

S35 S
S535
SScasen

See.
oF a
>
=>
= a

J
A “le
{/'
H

Aora:

Figure 63.—Aoridae: a, Aora typica Krgyer (Sars, 1895, pl. 193, as 4. gracilis).
b, upper lip; c, mandible; d, lower lip; e, f, maxillae 1-2; g, maxilliped. Gnathopod 1:

h, female Aora; 1, male Aora; 7, Xenochetra fasciata Haswell (Pirlot, 1938); k, Lembos

websteri Bate (Sars, 1895, pl. 194); 2, Rudilemboides stenopropodus J. L. Barnard (1959a);

m, Lembopsis spinicarpus Pearse (1912). Uropod 3: n, Aora; 0, Dryopoides westwood

Stebbing (1888); p, Paraoroides unistilus Stebbing (1910); 9, Acuminodeutopus heteruropus
Telson: r, Aora.

J. L. Barnard (1959a).

MARINE GAMMARIDEAN AMPHIPODA 149

intermediate genus. This procedure would also justify combining the
remaining Aoridae with the Isaeidae (Photidae) and might eventually
result in the Ischyroceridae also being joined to the Isaeidae, perhaps
as a subfamily.

10.

Key to Genera of Aoridae

[MALES AND MALE-LIKE FEMALES]

Article 4 of gnathopod 1 produced into a long tooth (merochelate) (fig. 632)! 2
Article 4 of gnathopod 1 not produced into a one tooth Oe onay with

ashort tooth). ... : Die CeO
Accessory flagellum long, compeced of dine 6 or more amides Gee AOra
Accessory flagellum absent . . . j . .. . . Aoroides
Article 5 of gnathopod 1 with one or more icone distal or subposterior teeth

(carpochelate) (fig. 64j) .... Niece saga
Article 5 of gnathopod 1 lacking a enone dicen Gemn (Ge. 631) Geen tata eel lg
Uropod 3 uniramous (fig. 63p). .... =. . =... . Neomicrodeutopus
Uropodrs biramousy. (figs; GSq)) se Ne i ya eg BO)
Gnathopod I of both sexes alike. ............. . Hansenella
Gnathopod 1 differing in each sex . . . » 6

Gnathopod 2 heavily setose on full anterior adee of percle 5 Ge 649) . 7
Gnathopod 2 sparsely setose on anterior edge of article 5 or setae compacted
into a distal bundle only . . a eas 10
Article 6 of gnathopod 1 as long ond: as Tprondl ¢ as pide 5 Ge. 63m).
Lembopsis
Article 6 of gnathopod 1 shorter and/or narrower than article 5 (fig. 631) . 8
Article 5 of gnathopod 2 longest (fig. 64g), female gnathopod 1 simple (like
fi G4f) os oo: ees Neue peules
Article 6 of paathopod 2 lowes female eaathopod 1 subchelate (fig. 63h) .
Head elongate, antenna 2 attached halfway back on ventral cephalic margin,
gnathopod 2 of male as stout as 1, of female weakly setose anteriorly.
Amphideutopus
Head not elongate, antenna 2 attached to anteroventral corner of head in
front of eye, gnathopod 2 of male much more slender than 1, of female

densely setose anteriorly . .. . Be ROR Goremanas
Inner ramus of uropod 3 less than half as iene as OHS: ramus (fig. 63q).
Acuminodeutopus

Rami of uropod 3 subequal ne 63n)....... . .. . Microdeutopus
Uropod 3 lacking rami .. . Cea On mince en ee ne ee OM CInOtALSUS
Uropod 3 with at least oneramus .. . BIER etary NEGA LY
Rami of uropod 3 minute, less than half as fone as erredinclc (ae 630) ..13 2
Rami of uropod 3 not minute, as long as or longer than peduncle .. 14
Pleonite 6 evanescent dorsally, artiels 3 of antenna 1 shorter than article 1,
accessory flagellum vestigial . . . ... . . .. .. . Dryopoides, female

1 Note that juvenile males of Aora and Aoroides have a very short tooth and thus may key to Lembos but
gnathopod 1 of Lembos is always strongly subchelate, whereas gnathopod 1 of Aora and Aoroides is essentially
simple.

2 These genera may belong with the Ischyroceridae.

150 U.S. NATIONAL MUSEUM BULLETIN 271

Pleonite 6 visible dorsally, article 3 of antenna 1 ened than article 1, acces-

sory flagellum well developed. . . Berea Eavadtvone

14. Gnathopod 1 with article 6 equal to or 2 EEE 6 in Jength and breadth than
article SiGich O37) eae eee ba tasers ALG
Gnathopod 1 with article 6 shorten amcor narrower ‘hem price 5 (fig.
GSE) yk a Sone PDs seni mame etn ites OS ILS)

15. Uropod 3 uniramous, _ aoeeREny feeeliacn albsents .. . . . . Paraoroides
Uropod 3 biramous, accessory flagellum present. . . . . Wheto eer)

16. Article 3 of antenna | subequal to article 1, gnathopods 1-2 eouel: in size and
morphology, subchelate ... . ... . . Aorcho

Article 3 of antenna 1 shorter than ariel i aamoped 1 larger than gnatho-

pod 2 or gnathopod 2 extremely setose, occasionally gnathopod 2 simple. 17

iver Gnathepod 2:subchelate swan Aeon ay eee | ieee ee eNOS

Gnathopod 2 simple (fig. 1061) ..... . been tees) Itc)

18. Coxae 1 and 2 short, wider than long, pabequall in size, _gulsomadbnite coxa 2
not concealing coxa 1, article 5 of gnathopod 2 very bulbous (fig. 64d).

: Xenocheira

Coxa | smaller than and partially to fully hidden by coxa 2, latter longer than
wide, article 5 of gnathopod 2 not bulbous. ... . . . Leptocheirus

19. Gnathopods fully subchelate . .. . ... .. . . Lemboides
Gnathopods scarcely subchelate (figs. 631, 64/) . . . . . Rudilemboides

Genera of Aoridae

Acuminodeutopus J. L. Barnard
Acuminodeutopus J. L. Barnard, 1959a.

Type-species: A. heteruropus J. L. Barnard, 1959a (original
designation).

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 with large distal tooth on article 5, article
6 shorter and narrower than article 5, simple, female gnathopod 1 sub-
chelate; male gnathopod 2 nearly simple, poorly setose, article 5
longest; outer ramus of uropod 3 longer than peduncle, inner ramus less
than half as long as outer. Species: 1, California, littoral.

Amphideutopus J. L. Barnard
Amphideutopus J. L. Barnard, 1959a.

Type-species: A. oculatus J. L. Barnard, 1959a (original
designation).

[Antenna 1 unknown]; male gnathopod 1 with distal tooth on article
5, article 6 shorter and narrower than 5, simple, female gnathopod 1
subchelate; male gnathopod 2 with moderately setose anterior edge
on article 5, article 6 slightly the longest, subchelate; male gnathopod 2
as stout as gnathopod 1; head elongate, antenna 2 attached halfway back
on ventral cephalic margin; female gnathopod 2 weakly setose anteriorly
(Coremapus) ; rami of uropod 3 equal, longer than peduncle. Species:
1, California, littoral.

MARINE GAMMARIDEAN AMPHIPODA IVS

Ficure 64.—Aoridae: a, Microdeutopus gryllotalpa Costa (Sars, 1895, pl. 192); b, Aorcho
delgadus J. L. Barnard (1961). Gnathopod 2: c, Aora typica Krgyer female (Sars, 1895,
pl. 193, as 4. gracilis); d, Xenocheira fasciata Haswell (Pirlot, 1938); e, dora male; f,
Rudilemboides stenopropodus J. L. Barnard (1959a); g, Neomegamphopus roosevelt: Shoe-
maker (1942); h, Lembopsis spinicarpus Pearse (1912); 1, Lembos websteri Bate (Sars, 1895,
pl. 194). Gnathopod 1: 7, Microdeutopus.

152 U.S. NATIONAL MUSEUM BULLETIN 271

Aora Kr¢éyer

Aora Krgyer, 1845.—Stebbing, 1906.
Lalaria Nicolet, 1849.
Lonchomerus Bate, 1857a.
Type-species: A. typica Krgyer, 1845 (monotypy). See Sars, 1895.
Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 complexly subchelate, with long tooth on
article 4, articles 5-6 lacking teeth, article 6 slightly shorter and some-
times narrower than article 5, simple, female gnathopod 1 scarely
subchelate; male gnathopod 2 subchelate, not heavily setose on
anterior edge of article 5, articles 5 and 6 subequal; rami of uropod 3
subequal, longer than peduncle. Species: 2, Atlantic, amphiboreal,
littoral, bathyal.

Aorcho J. L. Barnard
Aorcho J. L. Barnard, 1961.

Type-species: A. delgadus J. L. Barnard, 1961 (original designa-
tion).

Article 3 of antenna 1 as long as article 1, accessory flagellum
present; male gnathopod 1 lacking teeth, scarcely larger than and
similar to gnathopod 2, [female unknown]; gnathopod 2 subchelate,
poorly setose, articles 5-6 subequal; rami of uropod 3 subequal, longer
than peduncle. Species: 1, Tasman Sea (610 m).

Aoroides Walker
Aoroides Walker, 1898.—Stebbing, 1906.

Type-species: A. columbiae Walker, 1898 (monotypy). See J. L.
Barnard, 1954.

Article 3 of antenna 1 shorter than article 1, accessory flagellum
absent; male gnathopod 1 complexly subchelate, with long tooth on
article 4, articles 5-6 lacking teeth, article 6 shorter and narrower
than 5, simple, female gnathopod 1 scarcely subchelate; male gnatho-
pod 2 not heavily setose, articles 5-6 subequal, subchelate; rami of
uropod 3 subequal, as long as peduncle. Species: 2, Pacific boreal,
littoral.

Coremapus Norman
Coremapus Norman, 1905.

Type-species: Lembos versiculatus Bate, 1857a (original designa-
tion). See Chevreux and Fage, 1925.

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 with distal tooth on article 5, article 6

MARINE GAMMARIDEAN AMPHIPODA 153

shorter and narrower than 5, simple, female gnathopod 1 subchelate;
male and female gnathopod 2 with heavily setose anterior edge of
article 5, article 6 longest, subchelate; male gnathopod 2 much more
slender than gnathopod 1; head not elongate, antenna 2 attached at antero-
ventral cephalic corner in front of eye (Amphideutopus); rami of uropod
3 equal, longer than peduncle. Species: 1, N. Atlantic boreal,
littoral.
Dryopoides Stebbing

Dryopoides Stebbing, 1888; 1906.

Type-species: D. westwoodi Stebbing, 1888 (monotypy). See
Stebbing, 1910.

Article 3 of antenna 1 shorter than 1, accessory flagellum minute;
male gnathopod 1 subchelate, articles 4-6 lacking teeth; female
enathopod 1 subchelate; male gnathopod 2 subchelate, not heavily
setose, article 6 longest; rami of uropod 3 minute, equal, much shorter
than peduncle; pleonal segment 6 evanescent dorsally (Paradryope).
Species: 1, Australia, littoral. Also assigned to Corophiidae.

Hansenella Chevreux
Hansenella Chevreux, 1909.

Type-species: H. longicornis Chevreux, 1909 (original designation).

[Male unknown, but probably like female]; article 3 of antenna 1
shorter than article 1, accessory flagellum present; gnathopod 1 like
that of male aorid, with distal tooth on article 5, article 6 shorter
and narrower than 5, poorly subchelate; gnathopod 2 subchelate,
article 6 longest, article 5 poorly setose anteriorly; rami of uropod 3
equal, slightly longer than peduncle. Possibly synonymous with
Microdeutopus and representing aberrant female. Species: 1, N.
Atlantic (1,360 m).

Lemboides Stebbing

Lemboides Stebbing, 1895; 1906.

Type-species: JL. afer Stebbing, 1895 (original designation).

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 subchelate, articles 4-5 lacking teeth, 6
occasionally with chela, article 5 longer and broader than 6, female
gnathopod 1 subchelate, male gnathopod 2 subchelate, article 5
longest, poorly setose; rami of uropod 3 equal, longer than peduncle.
Species: 4, antiboreal, littoral.

Lembopsis Pearse
Lembopsis Pearse, 1912.

Type-species: L. spinicarpus Pearse, 1912 (original designation).
Article 3 of antenna 1 shorter than article 1, accessory flagellum

154 U.S. NATIONAL MUSEUM BULLETIN 271

present; male gnathopod 1 with long posterior tooth on article 5,
article 6 longer and as broad as 5, subchelate; male gnathopod 2
heavily setose on anterior edge of article 5, articles 5-6 subequal;
rami of uropod 3 equal, longer than peduncle. Species: 1, Gulf of
Mexico, littoral.

Lembos Bate

Lembos Bate, 1857a.—Stebbing, 1906.
Autonoe Bruzelius, 1859.
Bemlos Shoemaker, 1925.

Type-species: JL. webstert Bate, 1857a (designated by Chevreux
and Fage, 1925). See Sars, 1895.

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 subchelate, with articles 4—5 lacking teeth,
article 6 not narrower but usually longer than 5, female gnathopod 1
subchelate; male gnathopod 2 subchelate, with article 5 longest, poorly
setose on anterior edge; rami of uropod 3 equal, longer than peduncle.
Species: 31, cosmopolitan, primarily tropical, littoral (2 bathyal, 1
abyssal).

Leptocheirus Zaddach

Leptocheirus Zaddach, 1844.—Stebbing, 1906.
Ptilocheirus Stimpson, 1853.
Boeckia Malm, 1871.

Type-species: JL. pilosus Zaddach, 1844 (original designation). See
Sexton, 1911; Chevreux and Fage, 1925.

Article 3 of antenna 1 shorter than article 1, accessory flagellum
3+-articulate; male gnathopod 1 subchelate, articles 4-5 lacking
teeth; female gnathopod 1 subchelate and extremely setose; gnathopod
2 simple in both sexes and extremely setose on articles 2 and 5 but
article 5 not especially bulbous; rami of uropod 3 equal or unequal,
one or both rami longer than peduncle; coza 1 often very small and
occasionally large but always slightly smaller than coxa 2 and often tri-
angular, hidden fully or partially by enlarged coxa 2, latter longer than
wide (Xenocheira). Species: 10, boreal N. Atlantic, littoral.

Microdeutopus Costa

Microdeutopus Costa, 1853c.—Stebbing, 1906.
Stimpsonia Bate and Westwood, 1863 (homonym, Nemertea).
Stimpsonella Della Valle, 1893.

Type-species: MM. gryllotalpa Costa, 1853c (monotypy). See Sars,
1895.

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 with long distal tooth on article 5, article

MARINE GAMMARIDEAN AMPHIPODA 155

6 shorter and narrower than article 5, simple, female gnathopod 1
subchelate; male gnathopod 2 subchelate, articles 5-6 variable in
length, article 5 not heavily setose anteriorly; rami of uropod 3 sub-
equal, as long as or longer than peduncle. Species: 10, cosmopolitan,
littoral.

Neomegamphopus Shoemaker
Neomegamphopus Shoemaker, 1942.

Type-species: WN. rooseveltt Shoemaker, 1942 (original designation).

Article 3 of antenna 1 as long as article 1, accessory flagellum small;
male gnathopod 1 with long distal tooth on article 5, article 6 simple, as
long but narrower than article 5, female gnathopod 1 simple; male
enathopod 2 with article 5 longer than 6 and heavily setose on anterior
edge; rami of uropod 3 subequal, slightly longer than peduncle.
Species: 1, E. Pacific tropical, littoral.

Neomicrodeutopus Schellenberg
Neomicrodeutopus Schellenberg, 1925b.

Type-species: N. cabindae Schellenberg, 1925b (monotypy).

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 with long distal tooth on article 5, article
6 shorter and narrower than article 5, simple, [female unknown]; male
enathopod 2 subchelate, stout, article 5 longer than 6, article 5 poorly
setose anteriorly; uropod 3 with a single ramus twice as long as pe-
duncle, latter with a broad distomedial lobe. Species: 2, W. Africa,
littoral. Differs from Grandidierella in Corophiidae by normally short
article 1 of mandibular palp.

Paradryope Stebbing
Paradryope Stebbing, 1888; 1906.

Type-species: P. orguion Stebbing, 1888 (monotypy).

Article 3 of antenna 1 longer than article 1, accessory flagellum
present; male gnathopod 1 subchelate, articles 4—5 lacking teeth,
article 6 longer and broader than 5; male gnathopod 2 subchelate, not
heavily setose, articles 5-6 equal; rami of uropod 3 minute, equal,
much shorter than peduncle. Species: 1, N. Pacific, abyssal (4,200
m).

Paraoroides Stebbing
Paraoroides Stebbing, 1910.

Type-species: P. unastilus Stebbing, 1910 (monotypy).
Article 3 of antenna 1 shorter than article 1, accessory flagellum

285-135 O-69-11

156 U.S. NATIONAL MUSEUM BULLETIN 271

minute, vestigial; male gnathopod 1 subchelate, articles 4—5 lacking
teeth, articles 5-6 subequal, female gnathopod 1 scarcely subchelate;
male gnathopod 2 subchelate, poorly setose, article 5 slightly longer
than 6; uropod 3 uniramous, ramus equal to peduncle. Species: 1,
S.E. Australia, littoral.

Rudilemboides J. L. Barnard
Rudilemboides J. L. Barnard, 1959a.

Type-species: . stenopropodus J. L. Barnard, 1959a (original
designation).

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 simple, lacking teeth, article 5 much
broader and longer than article 6, female gnathopod 1 subchelate;
male gnathopod 2 nearly simple, poorly setose, article 5 longest;
rami of uropod 3 equal, longer than peduncle. Species: 1, California,
littoral.

Uncinotarsus L’Hardy and Truchot
Uncinotarsus L’ Hardy and Truchot, 1964.

Type-species: U. pellucidus L’ Hardy and Truchot, 1964 (original
designation).

Article 3 of antenna 1 shorter than article 1, accessory flagellum
2-articulate; male gnathopod 1 subchelate, articles 4—5 lacking teeth,
article 6 broader and longer than article 5, female gnathopod 1 sub-
chelate; male gnathopod 2 subchelate, not heavily setose, article 6
longer than 5; uropod 2 with one ramus; rami of uropod 3 absent.
Species: 1, Atlantic France, littoral. Differs from Concholestes in
Corophiidae by slender antenna 2 being shorter than antenna 1, by
enathopod 1 being larger than 2, by the short article 3 of antenna 1,
acuminate coxae 5-7, slender pereopods 1—2, and elongate pereopods
o-9.

Xenocheira Haswell
Xenocheira Haswell, 1880b.—Stebbing, 1906.

Type-species: X. fasciata Haswell, 1880b (monotypy). See Pirlot,
1938.

Article 3 of antenna 1 shorter than article 1, accessory flagellum
present; male gnathopod 1 subchelate, articles 4—5 lacking teeth;
female gnathopod 1 subchelate; male gnathopod 2 simple, female
minutely subchelate, article 6 longest, article 5 strongly setose
anteriorly and very bulbous; gnathopod 2 extremely setose; rami of
uropod 3 equal, longer than peduncle; corae 1-2 subequal i size,
short, wider than long, coxa 1 not concealed by coxa 2 (Lep tocheirus).
Species: 2, tropical Pacific, littoral.

MARINE GAMMARIDEAN AMPHIPODA 157
Argissidae
FIGURE 65

Draenosis.—Accessory flagellum 2-articulate; coxae 1 and 4 long,
coxae 1-3 successwely smaller, coxa 4 larger than coxa 1; gnathopods
feeble, simple or poorly subchelate. Monogeneric. See Gammaridae,
Haustoriidae, Synopiidae, Ampeliscidae, Vitjazianidae.

Description.—Accessory flagellum 2-articulate; when present,
primary flagellum basally conjoint in male; dorsal teeth only on
urosome, rostrum obsolescent; coxa 1 large, coxae 2 and 3 successively
smaller, coxa 4 larger than 1; mouthparts basic, upper lip slightly
incised; mandibular palp with article 3 longer than article 2; gnatho-
pods feeble, simple, except one species having gnathopod 1 poorly
subchelate; uropod 3 biramous; telson of medium length, cleft.

RELATIONSHIP.—Argissids are distinguished by the consecutive
reduction in size of coxae 1-3. They closely resemble Gammaridae,
especially the genus Megaluropus in which coxae 1-4 also are irregular
but in which coxa 2 is larger than 1 and coxa 3 smaller than 2.

Despite their definitive similarity to Gammaridae, argissids bear
external resemblance to Haustoriidae and were at one time included
in that family. They call attention to the close relationship of haus-
toriids to gammarids. Again the unusual coxae distinguish them from
haustoriids. The peculiar eyes of Argissa hamatipes, four bigeminous
lenticular bodies imbedded at the periphery of a common pigmentary
mass, seem unique to the family but the other species of Argissa, A.
stebbingi, lacks eyes, so that they are not diagnostic.

Sars (1895) has noted the similarity of the genus Argissa to the
Ampeliscidae (at that time Argissa was placed in the Haustoriidae).
Resemblance occurs in the antennae, pereopods 1-2 and 5, the uropods,
and the telson. Argissa nevertheless has some primitive characters
which place it between the Ampeliscidae and the Gammaridae: the
accessory flagellum, the intermediately modified pereopods 3 and 4,
and a weakly 2-articulate outer ramus of uropod 3. The eyes of Argissa
may represent a stage in the development of the ampeliscid corneal
lens. Of course, Argissa cannot stand on the direct line between
Ampeliscidae and Gammaridae because of its specializations in coxae
but it may well be a close relative of the organisms that occurred
directly on the evolutionary pathway. The most important specializa-
tions of the Ampeliscidae have been the development of tube-spinning
glands in pereopods 1-2, a function apparently not developed in
Argissa, and the amalgamation of the last two pleonal segments.

Article 2 of pereopods 3-5 in Argissa does have conspicuous glands.

Vitjazianids have many characters in common with argissids but
Argissa may be recognized by the special configuration of coxae.

158 U.S. NATIONAL MUSEUM BULLETIN 271

Nomenclatural Changes in the Argissidae

Parargissa Chevreux (1908c; Pirlot, 1934) is removed to the
Hyperiopsidae.

Phylluropus K. H. Barnard (1932) is a synonym of Megaluropus
(Gammaridae).

Phylluropus capensis K. H. Barnard (1932) is a synonym of Megalu-
ropus agilis (fide K. H. Barnard, 1940).

\

|
Si
HN

\\

/ AK
LN

N

Ficure 65.—Argissidae: Argissa hamatipes (Norman) (Sars, 1895, pl. 48, as 4. typica):
a, upper lip; b, lateral; c, mandible; ¢, lower lip; f,g, maxillae 1, 2; h, maxilliped; 7, uropod
3; 7,k, gnathopods 1, 2; /, uropod 3. Argissa stebbingi Bonnier (1896): d, mandible; m,
gnathopod 1.

MARINE GAMMARIDEAN AMPHIPODA 159
Genera of Argissidae

Argissa Boeck

Argissa Boeck, 1871.—Stebbing, 1906.
Chimaeropsis Meinert, 1890 (homonym, Pisces).

Type-species: Syrrhoe hamatipes Norman, 1869a (monotypy and
subsequent synonymy). See Sars, 1895.

Species: 2 (possibly synonymous), N. Hemisphere and possibly
cosmopolitan, eurybathic (30-1919 m).

Astyridae

FIGURE 66

Diaenosis.—Accessory flagellum 1- or 2-articulate, slender (or
absent in dubious genera); mandibular molar nontriturative; lower
lip with outer lobes very widely separated; gnathopods feeble. Mono-
generic. See Eusiridae, Calliopiidae, Pleustidae, Liljeborgiidae,
Haustoriidae, Synopiidae, Vitjazianidae, Hyperiopsidae, Paramphi-
thoidae, Stilipedidae.

Description.-—Accessory flagellum a single, medium-sized article
or 2-articulate; body smooth; rostrum inconspicuous, head usually
normal in appearance but occasionally massive and with “‘shark nose”
appearance (as in some Synopiidae [=Tironidae] and some Par-
daliscidae) ; coxae of medium size or long, rounded or quadrate below
but 2 and 4 occasionally subacuminate; mandibular molar a conical,
setose, nontriturative lamina; outer lobes of lower lip very widely
separated, with or without small erect inner lobes; remainder of
mouthparts basic, occasionally maxillae subfoliaceous; gnathopods
feeble, barely subchelate; rami of uropod 3 very elongate; telson
short, entire or cleft.

ReEwatTionsuHip.— Usually, the Synopiidae strongly differ from the
Astyridae only by their multiarticulate accessory flagella (except the
genus Jeddo). The combination of very broadly separated lobes of
the lower lip, the fact that coxa 4 is usually larger than coxa 3 and
the peculiar, sharp, conical, nontriturative molar serve to distinguish
Astyridae.

The short coxae and the absence of a mandibular molar distinguish
the Pardaliscidae.

Oedicerotidae have strongly setose pereopods, lack an accessory
flagellum, have relatively short rami and therefore a relatively long
peduncle of uropod 3 and a non-astyrid lower lip.

Most Eusiridae have either a ridged mandibular molar, do not bear

160 U.S. NATIONAL MUSEUM BULLETIN 271

the astyrid lower lip or have an elongate telson. Most Calliopiidae
have triturative molars and the nontriturative molars of Pleustidae
are bulbous, not conical.

In terms of accessory flagellum, mandibular molar, maxillae,
maxillipeds, gnathopods, uropods, and telson the Astyridae resemble
Paramphithoidae closely, especially to the genera Helysis and Epi-
meriella, which have the same mandibular molar and gnathopods,
maxillae and maxillipeds. Astyridae differ from the Paramphithoidae

spp howe
x ON

\
i
i
i
iy
.
RS
Ns

Ficure 66.—Astyridae: a, Astyra abyssi Boeck (Sars, 1895, pl. 73); 6, upper lip; c, mandible;
e, lower lip; g,h, maxillae 1, 2; 7, maxilliped; j,k, gnathopods 1, 2; m, telson; 0, uropod 3;
d, head of Astyra gardinert (Walker) (Pirlot, 1934, as Parastyra longidactyla); f, lower lip
of A. bogorovi Birstein and Vinogradov (1955); /, telson of A. zenkevitchi Birstein and
Vinogradov (1955); n, uropod 3 of A. gardinert.

MARINE GAMMARIDEAN AMPHIPODA 161

only by the peculiar lower lip. That this character is sufficient to
remove Astyridae from Paramphithoidae is doubtful, especially
because of the subacuminate condition of coxae 2 and 4 in astyrids
and the distally expanded and rounded-quadrate coxa 1 of Eelysis.

The Laphystiopsidae lack an accessory flagellum, have normal
lower lips, and entire telsons.

The Haustoriidae do not have the characteristic astyrid lower lip
and have fossorial (spiny and setose) pereopods 3-5.

Astyroides Birstein and Vinogradova is synonymized with Alez-
andrella in the Stilipedidae. Stilipedids differ from Astyra in the absence
of a mandibular molar, the broadening of the mandibular body,
the greater foliaceousness of the maxillae, but the two families are
strongly related by the condition of coxae 1-2, of which coxa 1 is
broader than 2 and 2 tends to taper distally. Stedipes stands between
Astyra and Alexandrella in the condition of its upper lip.

Key to the Genera of Astyridae

(PE CoxalesubaGUMINALG inh coisas uel elena ase) SOR SESS [Epimeriella] !
Coxaylsrounded-subquadrate distally 2) ie 2 ann ee eee Ag 2,
Pee Ne cessorysiagellumepnresemt ..9.. 25) 3 Ge eae ee ies cl pel Astyra
NCcCeSsonyehacellummalbsenmtim.) coin nie pies cea ore iyi Me les [Eclysis] !

1See these genera in the Paramphithoidae (pp. 394, 395).

Genera of Astyridae

Astyra Boeck

Astyra Boeck, 1871.—Stebbing, 1906.
Chagosia Walker, 1909.
Parastyra Pirlot, 1934.
Type-species: A. abyss1 Boeck, 1871 (monotypy). See Sars,
1895.
Species: 5, cosmopolitan, bathypelagic (100-2000 m).

Atylidae

FIGURES 67, 68

Driacnosis.—Accessory flagellum vestigial or absent; urosomites
2-3 coalesced. Monogeneric. See Ampeliscidae, Dexaminidae, Pro-
phliantidae, Lepechinellidae.

Description.—Accessory flagellum vestigial, 1-articulate or absent;
body with dorsal carinae or teeth; rostrum prominent; urosomites

162 U.S. NATIONAL MUSEUM BULLETIN 271

2 and 3 coalesced; mouthparts basic, lower lip lacking distinct inner
lobes; gnathopods small, subchelate; uropod 3 biramous; telson of
medium length, cleft.

RELATIONSHIP.—The Ampeliscidae have cuticular visual lenses
and pereopod 5 is shorter and of different structure than pereopod 4.
Other nonisaeid families with coalesced urosomites 2-3 are the Dexa-

Ficure 67.—Atylidae: a, Atylus falcatus Metzger (Sars, 1895, pl. 164); b, Atylus carinatus,
(Fabricius) (Sars, 1895, pl. 166).

MARINE GAMMARIDEAN AMPHIPODA 163

minidae, Lepechinellidae, and members of the Eophliantidae, Pro-
phliantidae, and Kuridae.

The Dexaminidae lack a mandibular palp.

The Lepechinellidae have acuminate or very short coxae and a
spine-like accessory flagellum.

The Eophliantidae, Prophliantidae, and Kuriidae all lack mandib-
ular palps and have other mouthparts departing from the basic
configuration.

Atylus aberrantis J. L. Barnard (1962d) is removed to

Lepechinella.
Genera of Atylidae

Atylus Leach

Atylus Leach, 1815.—Stebbing, 1906.—J. L. Barnard, 1956.
Nototropis Costa, 1853c, 1857.

Epidesura Boeck, 1861.

Paratylus Sars, 1895.

Type-species: Gammarus carinatus J. C. Fabricius, 1793 (mono-
typy). See Sars, 1895.
Species: 22, cosmopolitan, littoral (2 boreal bathyal).

Ficure 68.—Atylidae: Atylus carinatus (Fabricius) (Sars, 1895, pl. 166); a, upper lip; d
lower lip; c,d, maxillae 1, 2; e, gnathopod 1; f, maxilliped; g, mandible; h, telson; 7, uropod
3; 7, gnathopod 2. Atylus swammerdami (Milne Edwards) (Sars, 1895, pl. 163): k,
mandible.

164 U.S. NATIONAL MUSEUM BULLETIN 271

Bateidae

FIGURE 69

DraGnosis.—Coxa 1 vestigial or absent, remainder of gnathopod
1 degraded to a single article; accessory flagellum absent. See Gam-
maridae, Kusiridae.

Description.—Accessory flagellum absent; rostrum conspicuous;
back often with carinae; mouthparts basic, lower lip with or without
inner lobes; gnathopod 1 and its coxa degraded; gnathopod 2 feeble,
subchelate; others of first 3 coxae rounded ventrally; uropod 3 with
elongate, lanceolate rami; telson of medium length, longer than
peduncle of uropod 3, cleft.

RELATIONSHIP.—This is the only family with a degraded first
gnathopod except the genus Paranamizis in the Anamixidae but
that family is far removed from the Bateidae. Except for gnathopod
1, bateids resemble the Eusiridae.

Bateidae have a grade of structure similar to the Amphilochidae
and the two families casually resemble one another in overall
appearance.

Key to the Genera of Bateidae

1. Back with carinae, palp article 1 of maxilla 1 elongate . . . Carinobatea
Back smooth, palp article 1 of maxilla 1 of normal dimensions. . . Batea

Genera of Bateidae

Batea Muller
Batea Miller, 1865.—Stebbing, 1906.

Type-species: B. catharinensis Miiller, 1865 (monotypy). See
Shoemaker, 1926.
Species: 4, Caribbean, tropical eastern Pacific, littoral.

Carinobatea Shoemaker
Carinobatea Shoemaker, 1926.

Type-species: C. cuspidata Shoemaker, 1926 (original designation).
Species: 2, Caribbean, littoral.

MARINE GAMMARIDEAN AMPHIPODA 165

Ficure 69.—Bateidae (all figures after Shoemaker, 1926): a, Carinobatea cuspidata Shoe-
maker. Head: b, Batea catharinensis Muller. Mandible: c, Batea catharinensis. Lower
lip: d, Carinobatea; e, Batea. Mouthparts, Batea: f, maxilla 1; g, maxilla 2; f, maxilliped.
Gnathopod 1: 1, Batea. Telson: 7, Batea transversa; k, Batea rectangulata; 1, Batea
catharinensis. Uropod 3: m, Batea catharinensis. Gnathopod 2: n, Batea catharinensis.

166 U.S. NATIONAL MUSEUM BULLETIN 271

Beaudettiidae

Figure 70

Draenosis.—Accessory flagellum 2-articulate; mandible lacking
palp; inner ramus of uropod 3 reduced; coxa 4 not excavate posteri-
orly; telson very short, entire (but emarginate). Monogeneric. See
Gammaridae.

Description.—Body like that of Gammaridae, dorsally unorna-
mented; rostrum obsolete; antennal peduncles elongate, accessory
flagellum 2-articulate; mandible with molar but lacking palp; mouth-
parts otherwise basic; gnathopod 2 larger than 1; coxa 4 unexcavate
posteriorly; uropod 3 short, inner ramus half as long and one third
as broad as outer ramus; telson very short, entire (but emarginate).

RELATIONSHIP.—This monotypic family apparently is derived from
the Gammaridae. It is like the genus Elasmopus but has lost the man-
dibular palp, has a reduced inner ramus of uropod 3, and a short,
emarginate telson.

Ficure 70.—Beaudettiidae: Beaudettia palmeri J. L. Barnard (1965): a, lateral aspect;
b, mandible; c, lower lip; d,e, maxillae 1, 2; f, maxilliped; g, telson; A, upper lip; 7, uropod 3.

MARINE GAMMARIDEAN AMPHIPODA 167
Genera of Beaudettiidae

Beaudettia J. L. Barnard
Beaudettia J. L. Barnard, 1965.

Type-species: B. palmera J. L. Barnard, 1965 (original designation).
Species: 1, Micronesia, littoral.

Calliopiidae
Figures 71-73, 146 7-k

Diacnosis.—Accessory flagellum absent or composed of a tiny
article or scale; telson entire, apically emarginate or apically notched,
never deeply split. See Eusiridae, Pleustidae, Isaeidae, Laphystiopsi-
dae, Oedicerotidae, Amphilochidae, Ampithoidae, Paramphithoidae.

Description.—Accessory flagellum vestigial or absent; rostrum
usually inconspicuous; upper lip usually poorly incised or not;
mouthparts basic, but mandibular molar occasionally nontriturative;
lower lip with or without inner lobes, never taking the form of Pleu-
stidae (see p. 421); coxae of medium length or short; occasionally
coxa 4 not posteriorly excavate; gnathopods powerful or feeble,
usually subchelate, occasionally simple; uropod 3 with elongate
lanceolate rami, outer occasionally shorter than inner; telson of
medium length, entire, acuminate, emarginate, or notched.

RELATIONSHIP.—EKusiridae are close to this family but have a
distinctly cleft telson; when only notched apically the telson is elongate
in eusirids. Interfamilial gradation is seen by comparing telsons of
Chosroes, Atylopsis, and Halirages in Calliopiidae and Pontogeneoides
in EKusiridae.

That the Callopiidae are simply Eusiridae-Pontogenelidae with
fused telsonic lobes is well demonstrated in the fact that some callio-
pid genera are paired with some eusirid genera. Djerboa (Eusiridae)
and Metaleptamphopus (Calliopiidae) share pectinate dactyls of pere-
pods; Sancho (Calliopiidae) has eusirid gnathopods; Harpinioides
(Calliopiidae) and Harpinioidella (Eusiridae) have similar gnathopods;
Stenopleura (Calliopiidae) and Metewsiroides (Eusiridae) are very
similar in all characters except telsons.

Calliopiids no longer differ from other families by their incon-
spicuous rostra because several have been discovered lately that have
a longer rostrum than do some eusirids and pleustids.

Pleustidae differ from Calliopiidae only by the special configuration
of their lower lips. The genera Mesopleustes (Pleustidae) and Harpini-
oides (Calliopiidae) have lower lips which are very difficult to classify

168

U.S. NATIONAL MUSEUM BULLETIN 271

SEQ
2
ee:

co
Sas
SS

iS
ry
H
A
ry
i\
Kt
a
aN
\\
A
nt
K
Nt
\\
\\
Ny
i\
i
\
i

Ficure 71.—Calliopiidae: a, Calliopius laeviusculus (Kréyer) (Sars, 1895, pl. 158);
b, Cleippides quadricuspis Heller (Sars, 1885). Upper lip: ¢, Calliopius. Lower lip:
d, Calliopius; e, Halirages fulvocinctus (M. Sars) (Sars, 1895, pl. 154). Mandible and
palp: f, Calliopius; g, Leptamphopus sarsi Vanhoffen (Sars, 1895, pl. 162, as L. longimanus);
h, Oradarea walkeri Shoemaker (Walker, 1903).

Maxilla 1: 7, Calliopius; j, Laothoes
meinerti Boeck (Sars, 1895, pl. 160).

Maxilla 2: k, Calliopius.

MARINE GAMMARIDEAN AMPHIPODA 169

Ficure 72.—Calliopiidae: a, Apherusa tridentata (Bruzelius) (Sars, 1895, pl. 156). Pectinate
dactyl of pereopod: b, Metaleptamphopus pectinatus Chevreux (1912b). Maxilliped:
c, Calliopius laeviusculus (Krgyer) (Sars, 1895, pl. 158). Telson: d, Calliopius; e, Har-
pinioides drepanocheir Stebbing (1888); f, Stenopleura atlantica Stebbing (1888); g, Atylopsis
emarginata Stebbing (1888). Gnathopod 1: h, Leptamphopus sarsi VanhOffen (Sars, 1895,
pl. 162, as L. longimanus); 1, Regalia fascicularis K. H. Barnard (1930); 7, Sancho platynotus
Stebbing (1897). Gnathopod 2: k, Leptamphopus; l, Laothoes meinerti Boeck (Sars, 1895,
pl. 160); m, Haliragoides inermis Sars (1895, pl. 153); n, Calliopius; 0, Harpinioides.
Uropod 3: p, Calliopius.

170 U.S. NATIONAL MUSEUM BULLETIN 271

because they are almost perfect intergrades between the lower lips
of pleustids and calliopiids.

Calliopiids (and pleustids) may be confused with isaeids, ischyroce-
rids, and ampithoids unless one notes the absence of pereopodal glands
in Calliopiidae. The isaeid-like families have a fleshy telson which is
usually much shorter than broad, and generally have unexcavate
posterior margins on the fourth coxae.

The Laphystiopsidae, very closely related to the Calliopiidae,
differ primarily by the poorly developed mandibular molar. One
laphystiopsid genus, Prolaphystius, however is included in the key
to the Calliopiidae because of its uncertain position in the Laphystiop-
sidae. It has a moderately well developed molar.

The Vitjazianidae are distinguished by their distinctly simple
first gnathopods. Pleustidae have their characteristic lower lips.

The Paramphithoidae are characterized by cuspidate or pointed
anterior coxae. Occasionally these acuminations are extremely weak
and coxae 1-4 are scarcely excavate ventrally.

One calliopiid, Halirages regis (Stebbing, 1914) (=H. stebbingi
Schellenberg, 1931; =H. huzleyanus Stebbing, not Bate, 1862) has
subacuminate anterior coxae and thus keys to Paramphithoe, type-
genus of Paramphithoidae. Halirages regis has strongly developed
enathopods with ovate sixth articles and thus differs from most of its
congeners and Paramphithoe. Superficially, this species forms the
perfect intergrade between Calliopiidae and Paramphithoidae and
distinctions between the two families are completely confounded;
further study of the problem is warranted. Some species of Cleippides
also have subacuminate coxae.

Some genera of the Gammaridae, such as Falklandella, might be
placed in the Calliopiidae, except that Falklandella has short rami
of uropod 3, not characteristic of Calliopiidae. Gammarellus and
Weyprechtia with entire telson, and Parapherusa are not assignable to
Calliopiidae because of their multiarticulate accessory flagella.

The Oedicerotidae have disproportionately long fifth pereopods and
densely setose coxae and pereopods.

Most synopiids have a multiarticulate accessory flagellum but
a few have an accessory flagellum similar to that of eusirids. The
massive head of synopiids is characteristic, especially because of its
strongly deflexed rostrum. Synopiid gnathopods are feeble, whereas
those of eusirids are usually powerful.

Gnathopods within a genus are highly variable as to proportions
of articles 5 and 6; departures from a ‘‘normal theme” are seen
in Oradarea and Leptamphopus, which have exceptionally elongate
sixth articles of gnathopod 2. Cleippides has a relatively elongate
second gnathopod. Harpinioides has peculiar, bent sixth articles.

MARINE GAMMARIDEAN AMPHIPODA 171

Even the type-genus, Calliopius, has enlarged gnathopods, not
typical of the remaining members of the family. Extremes of variation
are shown in a group of gnathopodal figures presented herein.

Apherusa, Bouvierella, Callioprella, Halirages, and Haliragoides
are often very difficult to separate. Their type-species are easily dis-
tinguished by conditions of lower lip, anteroventral cephalic corners,
coxa 1 and accessory flagella, but those characters are highly variable
among other species of the five genera.

Gurjanova (1962) has pointed out the unusual character of Sancho
and Chosroes. They have broadly vaulted, extremely depressed
bodies and heads and splayed coxae like those of Phliantidae but
they also have subfossorial pereopods and gnathopod 1 like that
of fossorial families Phoxocephalidae, Haustoriidae, and Oedicerotidae.

a

Figure 73.—Calliopiidae: Gnathopod 2: a, Sancho platynotus Stebbing (1897); b, Clarencia
chelata K. H. Barnard (1932); c, Stenopleura atlantica Stebbing (1888); d, Cleippides
bicuspis Stephensen (Gurjanova, 1951); ¢, Atylopsis dentatus Stebbing (1888).

The mouthparts of both genera have a distinct look of the Oedi-
cerotidae. Neither genus has the fully developed pereopod 5 nor
elongate peduncle of uropod 3 found in the Oedicerotidae and Chosroes
has an elongate telson. Gnathopod 2 of Sancho distantly resembles
that of the phoxocephalid Joubinella (and various eusirids) ; and since
Phoxocephalidae and Haustoriidae seem to have strong affinities one
might consider Sancho to be an aberrant member of the Haustoriidae,
were it not for the uncharacteristic short peduncle of antenna 1, the
large male gnathopod 2, and the shortened outer rami of the uropods.
The conditions of the telson and pereopod 5 exclude the genera from
the Phoxocephalidae. But the short outer rami of the uropods of
Sancho and Chosroes like those of the Calliopiidae and Eusiridae
are not characteristic of the fossorial families. Probably a new family

285-135 O - 69 - 12

172 U.S. NATIONAL MUSEUM BULLETIN 271

should be created for these two genera which combine in a remarkable
way the characteristics of phliantids, oedicerotids, and eusirids.
This problem is also discussed under the Phliantidae (p. 405).

Remarks on the key to genera: Differences among genera are
minor, for example: couplets 5, 6, 9, 16. Stebbing’s (1906) key to
the genera utilized such quantitative features as vaulting of the
dorsum of the body, the dentition of the segments, and ornaments
of the telson, of which the present key resorts only to the latter.
Nevertheless, Halirages and Bouwerella are not clearly separated by
qualitative characters and the validity of the genera Calliopiella,
Atylopsis, or Amphithopsis is not convincing, the latter being related
closely to Halirages.

Clarencia is not firmly assignable to the Calliopiidae because the
uropods and telson were missing on the unique type-specimen.

- Metaleptamphopus appears to be a primitive melphidippid in its
elongate uropod 3 and especially resembles Melphisana in the uncleft
telson, obsolete accessory flagellum, slightly enlarged coxae and
well-developed rostrum.

Almost all of the species, except types, need refiguring of mouth-
parts and accessory flagella.

Nomenclatural Changes in Calliopiidae

Atylopsis latupalpus Walker and Scott (1903) should be removed to
a eusirid genus. It appears to resemble some species of Pontogeneia.

Pseudopontogeneia Oldevig (1959) is removed to the Eusiridae.

Leptamphopus litoralis Gurjanova (1938) is removed to Calliopiella.

Key to the Genera of Calliopiidae

(Paracalliope and Paraleptamphopus occupy freshwater and are not
included in the key.)

1.) -Gnathopod: 2’ chelate (figs 73b) ais Vea) Pe Se es Clarencia
Gnathopod 2 subchelate or simple ... . 2 ie Caley,

2. Articles 5-6 of gnathopod 2 together amen cl Solenente Sender rectan-
gular, sixth article, 75%, as long as article 2) (fig. W2k) 4) 2) Seu
Hither article 5 or 6 of gnathopod 2 elongate and slender, article 6 never more
than 60% aslongasarticle2 ... . ras el sie 5 747
Neither articles 5 or 6 elongate and Slondles, net foceanes ecently elongate,
usually slightly inflated oroval ..... SR ae ieee)

3. Article 3 of mandibular palp shortened, Afout about ae nid as long as
article 2 (fig. 71h), accessory flagellum present, lower lip with inner lobes
Oradarea !

Article 3 of mandibular palp subequal to or about four fifths as long as article

2, relatively slender, faleate or pointed terminally (fig. 71g), accessory
flagellum absent, lowerliplackinginnerlobes ........... 4

! See footnote 1 on p. 173.

15.

16.

Wie

18.

MARINE GAMMARIDEAN AMPHIPODA 173

Outer ramus of uropod 3 half or less as long asinnerramus . Leptamphopus !
Outer ramus of uropod 3 three fourths or more as long as inner ramus.
Bouvierella !

Mowerlip withanner lobes! (igs (id) Want ee he: Fe eae AG
Lower lip lacking inner lobes (fig. 7le). . . . Soden lA
Article 5 of gnathopods with distinct posterior lobe Ge SRY Ae et Senders
Article 5 of gnathopods without distinct posterior lobe (fig. 727) . . . 10
Article 6 of gnathopods slender, bent distally (fig. 720), essentially simple,
mandibular molar nontriturative .. . . . . . . Harpinioides
Article 6 of gnathopods stout, not bent, spandtiballen molar triturative. . 8
Body dorsally processiferous, pereopod 5 greatly elongate, article 5 as long as
articles 2, 3,4, and 6 together ... . . . . Stenopleuroides
Body dorsally smooth or without sharp arTeRSeS, pereopod 5 slightly longer
than 4, article 5 not disproportionately elongate. . .. . a or Mees)
Antenna 1 longer than or equal to antenna 2, telson srovteallls notched or
pointed (fig. 72f).. . . . . . . . Stenopleura
Antenna 2 slightly longer ‘Pham 1, telson poically somndad (fig. 72d).
Calliopius
Gnathopods bearing grossly distinct palms (fig. 72m) . . Seep eanedeala|
Gnathopods not bearing distinct palms, essentially simple Ge C20) tee els,
Articulate accessory flagellum present. ... . . . . . Atylopsis 3
Accessory flagellum absent, occasionally senneasatatl ibys OROCESSiieys Ah eae U2
Coxa 4 slightly longer than coxa 1 (fig. 71a) . . Halirages and Apherusa ?
Coxa 4 not longer than coxa 1 (fig. 71b) . . . ... .. . Haliragoides
Article 5 of gnathopods much longer than 6 (fig. 73d) . . . . Cleippides
Articles 5 and 6 of gnathopods subequal in length (fig. 727). . . Regalia
Palp of maxilla 1 reduced, reaching only halfway along outer plate (fig. 71)).
Laothoes
Palp of maxilla 1 of normal length, Nea to or ae apex of outer
platen Giese) 3) cages Ai hah em oa Be tay: a Ee)
Dactyls of pereopods mechinate Ge 72b) . Ae aah Metaleptamphonus
Dactyls of pereopods weakly serrate or smooth but not pectinate . . . 16

Gnathopod 2 of the EHusirus kind, a large suboval sixth article attached to a
slender fifth article (fig. 73a), (body and head extremely flattened).

Sancho

Gnathopod 2 not of the Husirus kind .... . ys ine eer ate Wh
Accessory flagellum as long as article 1 of ane “leeallenn (body mod-
erately depressed) .... ....... . . Amphithopsis
Accessory flagellum scale-like or “sibson pa Has ni Sean aR ecw rene
Article 2 of pereopods 1 and 2 smooth anteriorly, body comprcesed lat-
erallyaeeie eee BA eS)
Article 2 of mer opods 1 and 2 sft paterion mroceet! body premely flattened.
Chosroes

1 See also Atylopsis megalops Nicholls (1938); and a few species of Apherusa, Halirages, and Laothoes. Except
for A. megalops, none of these species has article 6 of the gnathopods more than 60% as long as article 2. Lao-
thoes is characterized by the condition of maxilla 1 palp.

2 Pontogeneoides (Eusiridae) resembles Atylopsis but has an especially elongate telson; it also keys to
Stenopleura but has powerfully subchelate gnathopods like Eusiroides, large coxae and elongate telson.

3A few species of Rhachotropis (Eusiridae) may be keyed to this point because of obsolescent telsonic
clefts but they may be distinguished by their extremely elongate telsons being more than twice as long as
the peduncle of uropod 3.

174 U.S. NATIONAL MUSEUM BULLETIN 271

19. Accessory flagellum absent or represented by a process on article 3 of

antennal ... ENE RER: sn AAD)
Accessory flagellum fence of a | eeaneall artmoulleis seals aaiigs Calliopiella

20. Pereopods 1-5 minutely subcheliform ...... .... . Bouvierella
Pereopods: 1-5) simple teas votive Nine! psa Gee die ie Waa AONE DED RANE Renae ates So aL Mie en

21. Male antennae with caleeoli ........ 2.2... +... . Halirages
Male antennae without calceoli ............ . . Apherusa

22. aeNccessony flagellum presenti. dee erenae tne eal ate Aone Sh fn ieee enna)
Accessory flagellum absent. . . . ATER ae

23. Article 5 of gnathopods elongate, aaron faneen and diene thicker than
emo G64) bc . . . (exceptional) Cleippides
Articles 5 and 6 hoeethen slamainie andl of similar thickness. . . Oradarea

24. Lower lip withsmallinnerlobes . . somespecies of Apherusa and Halirages
Lower lip lacking inner lobes. . . . : Div. As eee 74)

25. Outer ramus of uropod 3 half as fone as inner ramus. . iiveprariphonie
Outer ramus of uropod 3 three fourths or more aslong asinnerramus . . 26

26. Palp of maxilla 1 reduced, not reaching end of outer plate . . Laothoes
Palp of maxilla 1 normal, reaching or exceeding outer plate .... . 27

27. Gnathopod 2 perfectly simple, inner plate of maxilla 1 without setae, palp
of maxilliped not as long as outer plate and its parent article combined.
Prolaphystius (Laphystiopsidae)

Gnathopod 2 subchelate, inner plate of maxilla 1 setose, palp of maxilliped

nearly twice as long as article and its lobe of outer plate... . . . 28
284e<Body= dorsally smooth: 2.) (005%. 30. 2 we Cae aoe a Bouvierella
Body dorsally processiferous . ... . . . . some species of Halirages

Genera of Calliopiidae

Amphithopsis Boeck
Amphithopsis Boeck, 1861.—Stebbing, 1906.

Type-species: A. longicaudata Boeck, 1861 (selected by Boeck,
1876). See Sars, 1895.

Antennae equal in length; accessory flagellum as long as article 1
of primary flagellum; lower lip without inner lobes; gnathopodal
palms distinct, article 5 slightly longer than article 6, articles together
not elongate, article 5 of gnathopod 1 with slight posterior lobe; telson
rounded apically, outer ramus of uropod 3 less than half as long as
inner, body moderately depressed. Species: 1, subarctic Atlantic
(40-1,505 m).

Apherusa Walker

Gossea Bate and Westwood, 1863 (homonym, Coelenterata).

Apherusa Walker, 1891b.—Stebbing, 1906.

not Phaedra Bate, 1858b (possibly type-species a composite of two specimens,
head and body).

Type-species: Amphithoe jurinei Milne Edwards, 1830 (monotypy
and subsequent synonomy). See Sars, 1895.

MARINE GAMMARIDEAN AMPHIPODA 175

Antenna 1 shorter than antenna 2; accessory flagellum absent;
lower lip with slight inner lobes; gnathopodal palms distinct, article 5
slightly longer than article 6 and occasionally elongate, articles 5-6
together not elongate, article 5 of gnathopod 1 lacking posterior lobe;
telson pointed, toothed, emarginate or rounded; coza 4 slightly longer
than coxa 1 (Haliragordes) ; calceolr absent on male antennae (Halirages).
Species: 16, biboreal, littoral.

Atylopsis Stebbing
Atylopsis Stebbing, 1888; 1906.

Type-species: A. emarginatus Stebbing, 1888 (present selection).

Antenna 1 shorter than antenna 2 in type but otherwise variable in
length; accessory flagellum shorter than article 1 of primary flagellum ;
lower lip with inner lobes; gnathopodal palms distinct, articles 5 and 6
together not elongate, article 5 on gnathopod 2 shorter than 6, article
5 of gnathopod 1 as long as 6 or shorter than 6, lacking posterior lobe;
enathopod 2 occasionally slightly elongate; telson with deep notch,
broad or narrow. Species: 3, antiboreal, subantarctic, littoral
(one species to 567 m).

Bouvierella Chevreux
Bouvierella Chevreux, 1900.

Type-species: Paramphithoe carcinophilus Chevreux, 1889 (mono-
typy).

Antenna 1 slightly longer than antenna 2; accessory flagellum
absent; lower lip without inner lobes; gnathopodal palms distinct,
article 5 of gnathopod 1 slightly shorter than article 6, with weak
posterior lobe, articles 5 and 6 together on gnathopods slightly
elongate; telson emarginate; pereopods 1-5 minutely subcheliform
(Apherusa, Calliopiella, and Halirages). Very close to Leptamphopus
but gnathopod 2 not fully elongate and outer ramus of uropod 3
nearly as long as inner. Species: 1, Atlantic bathyal (620-1,386 m).

Calliopiella Schellenberg
Calliopiella Schellenberg, 1925b.

Type-species: Calliopiella michaelsent Schellenberg, 1925b (mono-
typy).

Antenna 1| slightly shorter than antenna 2; accessory flagellum scale-
like (type) or absent; lower lip without inner lobes; gnathopodal palms
distinct, article 5 shorter than article 6, together not elongate, article
5 with slight posterior lobe; telson rounded. Species: 3, W. Africa,
N. Pacific temperate, littoral.

176 U.S. NATIONAL MUSEUM BULLETIN 271

Calliopius Liljeborg

Calliope Bate, 1857a (homonym, Aves).
Calliopius Liljeborg, 1865 (new name).—Stebbing, 1906.

Type-species: Amphithoe laeviuscula Krgyer, 1838 (monotypy and
subsequent synonymy). See Sars, 1895.

Antennae subequal; accessory flagellum absent or article 3 of
antenna 1 with a process; lower lip with inner lobes; gnathopods stout,
palms distinct, article 5 shorter than article 6, together not elongate,
article 5 with distinct posterior lobe; telson rounded. Species: 2,
boreal, littoral.

Chosroes Stebbing

Chosroes Stebbing, 1888; 1906.

Type-species: C. incisus Stebbing, 1888 (monotypy).

Body and head extraordinarily depressed; antenna 1 slightly shorter
than antenna 2; accessory flagellum absent; lower lip lacking inner
lobes; gnathopodal palms distinct, articles 5 and 6 subequal, together
not elongate, article 5 lacking posterior lobe; telson notched or cleft;
article 2 of pereopods 1 and 2 with large anterior process. Species: 2,
antarctic, littoral (1 to 342 m).

Clarencia K. H. Barnard

Clarencia K. H. Barnard, 1931; 1932.

Type-species: C.chelata K. H. Barnard, 1931 (original designation) ;
1932.

Antenna 1 very short, stout, with few articles; accessory flagellum
absent; lower lip with tiny fused inner lobes; gnathopod 1 with distinct
palm, articles 5 and 6 subequal; gnathopod 2 large, chelate, article 5
tiny and masked by articles 4 and 6, article 6 very large, bearing a thumb;
pleonite 4 elongate; [telson unknown]. Species: 1, antarctic (342 m).

Cleippides Boeck
Cleippides Boeck, 1871.—Stebbing, 1906.

Type-species: Acanthonotus tricuspis Krgyer, 1846 (monotypy). See
Bate, 1862; Gurjanova, 1951.

Antenna 1 much longer than antenna 2; accessory flagellum about
half as Jong as article 1 of primary flagellum; inner lobes of lower lip
obsolescent; gnathopodal palms indistinct, article 5 much longer than
6, together slightly elongate, article 5 with weak or no posterior lobe;
telson rounded. Close to Leptamphopus but inner lobes of lower lip
slightly developed and accessory flagellum present. Species: 3,
subarctic, littoral to bathyal (1,960 m).

MARINE GAMMARIDEAN AMPHIPODA eee

Halirages Boeck
Halirages Boeck, 1871.—Stebbing, 1906.

Type-species: Amphithoe fulvocuncta M. Sars, 1858 (selected by
Boeck, 1876). See Sars, 1895.

Antennae subequal or antenna 1 short; accessory flagellum absent
or article 3 of antenna 1 with process; lower lip with small or large
inner lobes; gnathopodal palms distinct, articles 5 and 6 subequal,
together not elongate or article 5 elongate, article 5 lacking distinct
posterior lobe; telson emarginate, toothed or pointed; telson and
peduncle of uropod 3 elongate, body usually dorsally processiferous
(Bounmerella); calceol present on male antennae (Apherusa). Species:
10, bipolar, littoral to abyssal (2,500 m). See note on H. regis (Stebbing,
1914) in “Relationship” of Calliopiidae.

Haliragoides Sars
Haliragoides Sars, 1895.—Stebbing, 1906.

Type-species: Halirages vnermis Sars, 1882 (monotypy). See Sars,
1895.

Antenna 1 shorter than antenna 2; accessory flagellum absent;
lower lip with inner lobes; gnathopodal palms distinct, article 5
longer than article 6, together not elongate, article 5 lacking posterior
lobe; telson pointed or toothed; cora 4 not longer than cora 1 (Apherusa).
Species: 3, bipolar, littoral to abyssal (2,450 m).

Harpinioides Stebbing
Harpinioides Stebbing, 1888; 1906.

Type-species: H. drepanocheir Stebbing, 1888 (monotypy).

Antenna 1 longer than antenna 2; accessory flagellum more than
half as long as article 1 of primary flagellum; mandibular molar tumid,
ovate, not triturative; lower lip with slight inner lobes; sizth articles of
gnathopods slender, simple, apically bent, tapering, article 7 half as long
as article 6, article 5 much shorter than article 6, with distinct posterior
lobe; telson notched. This genus also assigned to Pleustidae. Species:
1, Indonesia, Kerguelen (108-496 m).

Laothoes Boeck

Laothoes Boeck, 1871.—Stebbing, 1906.

Thoelaos Della Valle, 1893.
Type-species: L. meinerti Boeck, 1871 (monotypy). See Sars 1895.
Antenna 1 shorter than antenna 2; accessory flagellum absent; lower

lip lacking inner lobes; gnathopodal palms distinct, articles 5 and 6

equal, together not elongate or slightly elongate, article 5 lacking

178 U.S. NATIONAL MUSEUM BULLETIN 271

posterior lobe; telson slightly emarginate; palp of maxilla 1 reduced,
reaching halfway along outer plate. Species: 4, subarctic, littoral to
bathyal (1,500 m).

Leptamphopus Sars

Leptamphopus Sars, 1895.—Stebbing, 1906.

Type-species: JL. sars1 Vanhéffen, 1897 (monotypy; L. sarsi is a
new name for Sars’ misidentification of Amphithopsis longimana Boeck).
For stability, this arrangement should be preserved.

Antenna 1 slightly longer than antenna 2; accessory flagellum absent;
lower lip without inner lobes; gnathopodal palms distinct, articles 5 and
6 together elongate; telson notched; article 3 of mandibular palp sub-
equal to article 2 or very slightly shorter, relatwely slender, falcate or
pointed (Oradarea); outer ramus of uropod 3 half as long as inner
(Bouvierella). Species: 2, ?amphiboreal, bipolar, littoral to bathyal
(1,505 m).

Metaleptamphopus Chevreux

Metaleptamphopus Chevreux 1911c; 1912a; 1912b.

Type-species: M. pectinatus Chevreux, 1912a (designated by
Chevreux) ; 1912b.

Antenna 1 longer than antenna 2; accessory flagellum absent (type)
or uniarticulate; lower lip without inner lobes; gnathopodal palms
scarcely distinct, articles 5 and 6 equal, together not elongate; article 5
lacking posterior lobe; telson rounded; dactyls of pereopods strongly
pectinate. Species: 2, littoral subantarctic to S. Atlantic bathyal
(GRY ian).

Oradarea Walker
Oradarea Walker, 1903.—Shoemaker, 1930.

Type-species.—O. walkert Shoemaker, 1930 (monotypy; O. walkeri
is new name for Walker’s 0. longimana which is a homonym of Boeck’s
Amplhithopsis longimana later transferred to Oradarea).

Antenna 2 probably longer than antenna 1; accessory flagellum
cylindrical but shorter than article 1 of primary flagellum; lower
lip with inner lobes; gnathopodal palms distinct, articles 5 and 6
together elongate; telson truncate, rounded but with minute notch
or apically pyramidal; article 3 of mandibular palp short, stout, two
thirds as long as article 2 (Leptamphopus). Species: 9, primarily
antarctic (6), one bathyal Indonesia, one boreal, littoral to bathyal
(15165 mm);

Regalia K. H. Barnard

Regalia K. H. Barnard, 1930.
Type-species: fF. fascicularis K. H. Barnard, 1930 (monotypy).

MARINE GAMMARIDEAN AMPHIPODA 179

[Antennae unknown; accessory flagellum unknown]; lower lip with
slight inner lobes; gnathopodal palms indistinct, articles 5 and 6
equal, together not elongate, article 5 lacking posterior lobe; telson
short, broad, slightly emarginate. Species: 2, one pelagic at New
Zealand and one at 304 m in Indonesia.

Sancho Stebbing
Sancho Stebbing, 1897; 1906.

Type-species: WS. platynotus Stebbing, 1897 (monotypy).

Body and head extremely depressed; antennae subequal, peduncle of
antenna 1 very short; accessory flagellum as long as article 1 of primary
flagellum; lower lip without inner lobes; gnathopods not elongate,
male gnathopod 1 with transverse palm, article 5 longer than article 6,
gnathopod 2 much larger, Eusirus-like; female gnathopod 2 like male
gnathopod 1; pereopods spinose as in Haustorvidae; telson tapering
but apically truncate. Species: 1, E. Australia, littoral.

Stenopleura Stebbing
Stenopleura Stebbing, 1888; 1906.

Type-species: S. atlantica Stebbing, 1888 (monotypy). See
Stephensen, 1915; Birstein and Vinogradov, 1958; J. L. Barnard,
1962d.

Antenna 1 slightly longer than or equal to antenna 2; accessory
flagellum absent; lower lip with inner lobes; gnathopods moderately
stout, palms indistinct, article 5 much shorter than 6 and with distinct
posterior lobe, articles 5 and 6 together not elongate, article 7 as long as
article 6, latter not bent (Harpinioides); telson apically toothed.
Species: 1, probably cosmopolitan epipelagic in tropics. Compare
diagnosis of Stenopleuroides.

-

Stenopleuroides Birstein and Vinogradov
Stenopleuroides Birstein and Vinogradov, 1964.

Type-species: S. macrops Birstein and Vinogradov, 1964 (mono-
typy).

Antenna 1 much longer than antenna 2; accessory flagellum absent;
[lower lip unknown]; gnathopods moderately stout, palms indistinct,
article 5 much shorter than 6 and with distinct posterior lobe, articles
5-6 together not elongate, article 7 nearly as long as article 6, latter
not bent (Harpinioides); telson entire, wnornamented; body dorsally
cuspidate (pereonite 7 and pleonites 1-4); pereopod 5 very elongate,
article 5 as long as articles 2, 3, 4, and 6 together (Stenopleura). Species:
1, Indian Ocean, probably epipelagic.

180 U.S. NATIONAL MUSEUM BULLETIN 271

Cheluridae

FIGURE 74

DraGcnosis.—Body depressed; segments of urosome immovably
fused together, marked ventrally by sutures, segment 3 very large;
uropods 1, 2, and 8 radically dissimilar to each other in structure and
size; telson foliate, entire; accessory flagellum present; flagellum
of antenna 2 clavate, composed largely of one article in the adult.
See Corophiidae.

Description.—Accessory flagellum 1l-articulate, stout; flagellum
of antenna 2 largely composed of one clavate article; head either
bulbous and lacking supra-antennal line, or quadrate and bearing
such line; mouthparts basic; coxae short, not overlapping; gnathopod
1 either feeble and slightly chelate, or large and subchelate; gnathopod
2 feeble and slightly chelate; uropods dissimilar, uropod 1 composed
of normal peduncle and two short rami; uropod 2 composed of foli-
aceous peduncle, with or without rami; uropod 3 composed of short
peduncle, a greatly enlarged outer ramus and a scale-like or no
inner ramus; telson entire.

ReLatronsuip.— Unlike the Cheluridae, uropods 1 and 2 of the
Corophiidae are similar to each other, the second antennal flagellum
is distinctly multiarticulate, the third urosomal segment, although
occasionally fused to the others, is never greatly enlarged, and the
third uropods are never immensely enlarged as in the Cheluridae.

Key to the Genera of Cheluridae

1. Uropod 3 bearing a scale-like inner ramus (fig. 741) ....... Chelura
Wropod)3 Jacking an immeriramUs:) 9 is fons eee rte ee ene 2

2. Uropod 2 with two rami (fig. 74k), gnathopod 1 large, subchelate (fig. 747).
Tropichelura

Uropod 2 lacking rami (fig. 740), gnathopod 1 small, slightly chelate (fig. 74h).
Nippochelura

Genera of Cheluridae

Chelura Philippi
Chelura Philippi, 1839—Stebbing, 1906.

Type-species: (. terebrans Philippi, 1839 (monotypy). See Sars,
1895.

Supra-antennal line absent except in defining the lateral ocular
lobes; gnathopods small; uropod 2 with two rami; uropod 3 with
small inner ramus. Species: 1, cireumamphiboreal, littoral, bores
wood.

MARINE GAMMARIDEAN AMPHIPODA 181

Nippochelura J. L. Barnard
Nippochelura J. L. Barnard, 1959c.

Type-species: Chelura brevicauda Shiino, 1948 (original designa-
tion), 1957.

Supra-antennal line absent, anterodorsal part of head supporting
a large process; gnathopods small; uropod 2 lacking rami; uropod 3
lacking inner ramus. Species: 1, Japan, littoral, bores wood.

Ficure 74.—Cheluridae: Chelura terebrans Philippi (Sars, 1895, pl. 225): a, lateral; b, upper
lip; c, mandible; d, lower lip; e,f, maxillae 2, 1; g, maxilliped; h,i, gnathopods 1, 2; 7,k,1,
uropods 1, 2, 3; m, telson. Tropichelura insulae (Calman) (J. L. Barnard, 1959c): n, male
gnathopod 1. Nippochelura brevicauda (Shiino) (J. L. Barnard, 1959c): 0, uropod 2.

182 U.S. NATIONAL MUSEUM BULLETIN 271

Tropichelura J. L. Barnard
Tropichelura J. L. Barnard, 1959c.

Type-species: Chelura insulae Calman, 1910 (original designation).

Supra-antennal line present; gnathopod 1 large; uropod 2 with
two rami; uropod 3 lacking inner ramus. Species: 1, cireumtropical,
littoral, bores wood.

Colomastigidae

FIGguRE 75

Draenosis.—Accessory flagellum absent; body subcylindrical,
urosomites 2 and 3 coalesced; coxae very short but overlapping;
mandibular palp absent, primary incisor cut into a series of long
spine-like teeth; molar problematical, large but nontriturative in
some species, absent or unknown in others; inner plates of maxillipeds
very small, fully or partially coalesced, unarmed or acuminate;
uropod 8 biramous; telson entire. Monogeneric. See Eophliantidae,
Phliantidae, Prophliantidae.

Description.—Accessory flagellum absent; body subcylindrical;
coxae short but overlapping; epistome projecting acutely in front;
upper lip bilobed; mandible lacking palp, its body cut into several
long spine-teeth, molar large and unridged, small or ‘‘absent’’; lower
lip poorly known; maxillae stunted, maxilla 1 with 1-articulate palp
larger than outer plate; lobes of maxilla 2 partially coalesced; inner
plates of maxilliped very small, fused, or split but forming a sub-
conical piece, outer lobes of medium size but appearing large because
the articles they are attached to are elongate; gnathopod 1 simple,
usually armed apically with spines or setae forming a brush, some
males lacking these setae; gnathopod 2 in male enlarged, subchelate,
in female small and subchelate; uropod 3 biramous, occasionally
outer ramus considerably reduced in size; telson short to medium in
length, entire, often apically ornamented.

ReLatTionsuip.—The Eophliantidae also have cylindrical bodies
but differ from colomastigids by the well-developed inner lobes of the
maxillipeds and the cleft telson; the mandible is not cut into a series
of long projecting teeth and the palp of maxilla 1 is degraded.

Phliantidae and Prophliantidae have compressed, or depressed, not
cylindrical bodies. The latter have cleft telsons. Probably all phliantids
have a reduced inner ramus of uropod 3 and prophliantids have all
urosomal segments coalesced.

MARINE GAMMARIDEAN AMPHIPODA 183

Possibly the primary cutting edge of the colomastigid mandible
is missing and the digitations represent enlarged and basally fused
spines of the normal gammaridean spine-row.

Corophiidae and Podoceridae may resemble Colomastigidae be-
cause of depressed and subcylindrical bodies but the former two
families have mandibular palps and normal maxillipeds.

Figure 75.—Colomastigidae: a, Colomastix pusilla Grube (J. L. Barnard, 1955a). Telson:
b, C. pusilla; c, Colomastix subcastellata Hurley (1954d). Mandible: d, C. pusilla.
Maxilla 1: e, C. subcastellata; f, C. pusilla. Maxilla 2: g, C. subcastellata; h, C. pusilla.
Maxillipeds: i, C. subcastellata; 7, C. pusilla. Uropod 3: k, C. pusilla; l, C. magnirama
Hurley (1954d). Gnathopod 1, C. pusilla: m, female; n, male. Gnathopod 2, C. pusilla:

o, male; p, female.

184 U.S. NATIONAL MUSEUM BULLETIN 271

Genera of Colomastigidae

Colomastix Grube

Colomastix Grube, 1861; 1864a.—Stebbing, 1906.
Cratippus Bate, 1862.
Exunguia Norman, 1869b.
Type-species: Colomastiz pusilla Grube, 1861 (monotypy). See
J. L. Barnard, 1955a.
Species: 9, cosmopolitan, littoral.

Corophiidae

FigurEs 76-80

Diacnosis.—Accessory flagellum variable, often absent; urosome
visibly depressed; wrosomal segment 2 subequal to segment 1 in length;
telson entire; pereopods often glandular. See Isaeidae, Aoridae,
Podoceridae, Ischyroceridae, Cheluridae.

DescriptTion.—Accessory flagellum multiarticulate, 1-articulate or
absent; body often subcylindrical, generally depressed, especially
urosome, some or all of urosomites often coalesced; body rarely
carinate and only on urosome; coxae long, of medium length, or short,
often not touching each other, occasionally acuminate; mouthparts
basic, except mandibular palp occasionally reduced to one or two
articles and upper lip occasionally bilobed; gnathopods variable,
either pair the larger, often complexly subchelate; uropods 1 and 2
usually normal, rarely with inner rami reduced, rarely with inner
ramus on uropod 2 absent; uropod 3 variable: (1) bearing two rami,
the inner reduced in size; (2) bearing one ramus, either longer than
peduncle or very short; or (3) rami absent; telson entire, fleshy, circular
or symmetrically trapezoidal, or very broad and short, occasionally
armed with hooks; pereopod 5 occasionally very elongate and bearing
long setae like fossorial families.

ReELATIONSHIP.—The Corophiidae are a mixture of isaeid, aorid,
and ischyrocerid kinds having the urosome especially depressed and
usually losing the inner ramus of the third uropods, although this
condition is found also in some isaeids. It is very difficult to distinguish
these three families, as exemplified by the genus Aorcho, originally
assigned to Corophiidae but now provisionally assigned to Aoridae
(see discussion under the Isaeidae). Because of its tall urosome,
Unciolella is another genus of doubtful assignment to the Corophiidae.
Neomicrodeutopus (Aoridae) and Grandidierella (Corophiidae) are very
closely related.

MARINE GAMMARIDEAN AMPHIPODA 185

Ficure 76.—Corophiidae: a, Cerapus crassicornis (Bate) (Sars, 1895, pl. 217); b, Corophium
affine Bruzelius (Sars, 1895, pl. 221); c, Gaviota podophthalma J. L. Barnard (1958b);
d, Kamaka kuthae Derjavin (Gurjanova, 1951).

186 U.S. NATIONAL MUSEUM BULLETIN 271

Because of its peculiar mandibular palp and immensely thickened
male antenna 2, Corophium, the type-genus, is not typical of most
other genera assigned to the family. Dr. E. L. Bousfield (in litt.) has
pointed out to me the probability that Hricthonius, with its elongate
peduncle of uropod 3, belongs with the Ischyroceridae.

The Podoceridae differ from the Corophiidae by their elongate
urosomal segment 1.

Corophiidae may resemble Eophliantidae and Colomastigidae be-
cause of their depressed, subcylindrical bodies and short, serially
gaping coxae but the latter two families have degraded mandibles.

Nomenclatural Changes in Corophiidae

Aorcho J. L.. Barnard (1961) is removed to the Aoridae.

Bogenfelsia J. LL. Barnard (1962d) is removed to the Ischyroceridae.

Dryopoides Stebbing (1888) was in Aoridae in Stebbing (1906) and
transferred to Corophiidae by Stebbing (1910). Female keys out
to the Aoridae.

Runanga J. L. Barnard (1961) removed from Podoceridae.

Key to the Genera of Corophiidae

(All species having gnathopod 1 larger than 2 should also be keyed out
in the Aoridae.)

1. Uropod 3 lacking rami (fig. 80n) . . ....... .. . . Concholestes
Uropod:3) bearingal—2 rami (figs 80b—2)) 3 a. 5 a 2 ee

2. Mandibular palp 3-articulate (fig. 789). . . . . Lig 2a a2 ee ne
Mandibular palp less than 3-articulate (figs. 78e, f). Peer del xh Pall

3. Inner ramus of uropod 8 distinct (fig. 80h)... ... REN Stine
Inner ramus of uropod 3 absent or indistinct (uropod 3 Stee one ramus only)
(higstiSObS gu renieiie SON plead voi ahh Wate. ineepeee a aaa

4. Coxae not touching erally, Ges, 76b 7a) OR aC)
Coxae touching serially (figs. 76c,d) .... . SY Sahay take 1

5. Gnathopod 1 scarcely larger than and similar to pnethonod 2, waalice smooth,
inner ramus of uropod 3 distinct but much shorter than outer ramus (fig.
80h), third pleonal epimeron smoothly quadrate ... .. . Camacho

Gnathopod 1 much larger than 2, palm sculptured, inner ramus of uropod 3
usually represented by a large peduncular process (fig. 80z), rarely distinct
or marked off, when distinct scarcely shorter than outer ramus, third

pleonal epimeron bearing posteroventral tooth . . ... . . . Unciola
6. Inner ramus of uropod 3 much shorter than outer ramus, gnathopod 2 mero-
chelate... . . . Paracorophium

Rami of uropod 3 sapmally lone, poathonedl 2 not meroehelare:
male of Dryopoides

10.

Jhaik,

12.

13.

14.

15.

16.

IT

18.

I).

20.

21.

MARINE GAMMARIDEAN AMPHIPODA 187

Wropodyawithioneramusx(i gS 0) vee ter eee Se ee ne a8
Uropod 2 with tworami (fig. 80h) ...... 5 1@
Pleopod 3 with one ramus, [article 3 of antenna 1 loner chan antici 1].
Runanga
Pleopod 3 with two rami, [article 3 of antenna 1 rarely longer than article
Laan ceo 9
Male peetnonad 2, mate paethonod) 1, pampleniy, Baenelere Ge 791), head
with thorn-like rostrum andeyes. . . . . . Cerapus
Male gnathopod 1, not gnathopod 2, complexly Sapenelits head with ob-
solescent rostrum and no eyes . . . .. . . . Chevreuxius
Inner rami of uropods 1—2 less than half as Tener as outer rami (fig. 807).
Pseudericthonius
Inner rami of uropods 1—2 usually subequaltoouterrami ....... Ii
PMILenmagelackine accessory tagellum™ 9. 9, foe eee ees MD
Antenna 1 bearing accessory flagellum .. . is 1)
Male gnathopod 2 complexly subchelate Gaol) i 79k). Ericenonine
Male gnathopod 2 not complexly subchelate ... . ; ficial o gaelic}
Ficonitest4—5srcoalescedeise othe Cota oye o.oo a. a 7s Kamaka
Pleonites 4-5 separate... . Btacor lah Ue eect mantat shenmiienl Ul
Male gnathopod 2 chelate Gugpododnens) - . . .. . . . Cerapopsis
Male gnathopod 2 subchelate (fig. 793) . .......... .. Gaviota
PNCEESSOLYAlagellUmMeMNUltiantiCulate pala ie eee eee ha os ae iG
PNCCeSSOlveHacellum: dearticulater, ws enue smi eae een ations 1G
Coxae angular .. . 2 SURED ROA E PR SAR eR COLE ORME ANG:
Coxae with softly semadled Cornersi aye Ck. mapasgedles,

Article 6 of gnathopod 2 an elongate nagiemele. Prarvenclae or palm trans-
verse, antennae usually much shorter than body and flagellum of antenna
2 scarcely longer than peduncular article 5, peduncle of uropod 3 often
expanded medially very eran antenna 2 generally considered stout,
often strongly so... . . . Unciola

Article 6 of gnathopod 2 ouate or ene ordeal eepanded palm oblique,
antennae usually as long as or longer than body, flagellum of antenna 2
nearly as long as all articles of peduncle, peduncle of uropod 3 rarely
expanded medially and usually poorly so, antenna 2 very slender.

Neohela

Inner plate of maxilla 1 with one seta, ramus of uropod 3 about 1.5 times
aslongas peduncle. ... . . .. . . Unciolella
Inner plate of maxilla 1 with about 10 seine lesadiays medial edge, ramus of
uropod 3 about twice as long as peduncle. . . ... . . . Parunciola
Male gnathopod 1 complexly subchelate. . . .. . . . . Grandidierella
Male gnathopod 1 simply subchelate .. . eee 20

Male antenna 2: articles 3-5 stout (fig. 76b), loneen Paani uropod 3
with asymmetrically prolonged peduncle (fig. 802), or uropod 2 with
vestigial inner ramus . . . . . Unciola

Male antenna 2: articles 3-5 Blender Ges. 76a, a, 77a), monedl 2 with sym-
metrical peduncle (fig. 80f), rami of uropod 2 subequal.

Unciolella (juvenile)

Mandibular palp l-articulate (fig. 78e). . .... . . . . Siphonoecetes

Mandibular palp 2-articulate (fig. 78f) . .... .. .. . . Corophium

285-135 O - 69 - 13

188 U.S. NATIONAL MUSEUM BULLETIN 271
Genera of Corophiidae

Camacho Stebbing

Camacho Stebbing 1888; 1906.

Type-species: C. bathyplous Stebbing, 1888 (monotypy). See
J. L. Barnard, 1961.

Accessory flagellum multiarticulate; article 3 of antenna 1 much
shorter than article 1, flagellum very long; antenna 2 much shorter
than antenna 1, slender, flagellum shorter than article 5 of peduncle;
mandibular palp 3-articulate; coxae short, not touching serially;
enathopods normally subchelate; uropod 2 biramous; uropod 3
with outer ramus longer than peduncle, inner ramus one third as
long as outer. Species: 1, antiboreal, eurybathic, littoral to abyssal.

Cerapopsis Della Valle
Cerapopsis Della Valle, 1893.—Stebbing, 1906.

Type-species: C. longipes Della Valle, 1893 (monotypy).

Accessory flagellum absent; article 3 of antenna 1 equal to article
1; flagellum slightly longer than peduncle; antenna 2 slender, slightly
longer than antenna 1, flagellum slightly longer than peduncle;
mandibular palp 3-articulate; coxae large, touching serially; gnatho-
pods normally subchelate in female, gnathopod 2 chelate in male,
enlarged, with digitate thumb; uropod 2 biramous; uropod 3 with
its single ramus as long as peduncle; head with long lateral, ocular
lobes. Species: 1, Mediterranean, littoral.

Cerapus Say

Cerapus Say, 1817.—Stebbing, 1906.
Cerapodina Milne Edwards, 1840.

Type-species: C. tubularis Say, 1817 (monotypy). See J. L.
Barnard, 1962a.

Accessory flagellum absent; article 3 of antenna 1 shorter than
or 120% as long as article 1, flagellum as long as article 3; antenna 2
slender, equal to antenna 1, flagellum as long as article 5; mandibular
palp 3-articulate; coxae short, not touching serially; gnathopod 1
normally subchelate, gnathopod 2 in female simple even though article
6 slightly inflated, in male very large, essentially carpochelate, with
teeth on article 5, article 6 simple and more slender than 5, dactyl
very large and folding back to middle of article 5; uropod 2 with one
ramus; uropod 3 with one vestigial, hooked ramus; head with thorn-like
rostrum. Species: 5, (possibly less), cosmopolitan, littoral (one

bathyal).

MARINE GAMMARIDEAN AMPHIPODA 189

Figure 77.—Corophiidae: a, Unciolella lunata Chevreux (191la); 6, Paracorophium
excavatum (Thomson, 1884).

190 U.S. NATIONAL MUSEUM BULLETIN 271

Chevreuxius Bonnier
Chevreuxius Bonnier, 1896.—Stebbing, 1906.

Type-species: C. grandimanus Bonnier, 1896 (monotypy).

Accessory flagellum 2-articulate; article 3 of antenna 1 slightly
shorter than article 1, flagellum slightly longer than article 3; antenna
2 slender, equal to antenna 1, flagellum shorter than article 5; mandib-
ular palp 3-articulate; coxae short, not touching serially; male
gnathopod 1 much larger than gnathopod 2, carpochelate, article
6 stout and subchelate but narrower than article 5; gnathopod 2 sub-
chelate; uropod 2 with one ramus; uropod 3 with its single ramus
as long as peduncle. Species: 1, N. Atlant-c, bathyal (950 m).

Concholestes Giles
Concholestes Giles, 1888.—Stebbing, 1906.

Type-species: C. dental Giles, 1888 (monotypy).

Accessory flagellum absent; article 3 of antenna 1 subequal to article
1, flagellum equal to article 3; antenna 2 of medium stoutness, longer
than antenna 1, flagellum much shorter than article 5; [mandible not
described]; coxae not touching serially; gnathopod 2 normally sub-
chelate, gnathopod 1 nearly simple; uropod 2 with ?one ramus; uropod
3 lacking rami. Species: 1, Bay of Bengal, littoral, in shell of
Dentalium. See Uncinotarsus in Aoridae.

Corophium Latreille

Corophium Latreille, 1806.—Stebbing, 1906.
Audouinia Costa, 1851b (nom. nud.).

Type-species: Oniscus volutator Pallas, 1776 (monotypy and sub-
sequent synonymy). See Sars, 1895 (as C. grossipes).

Accessory flagellum absent; article 3 of antenna 1 shorter than
article 1, flagellum longer than article 3; antenna 2 equal to or longer
than antenna 1, stout, especially in male flagellum much shorter than
article 5, male article 4 usually with a distal tooth; mandibular palp
2-articulate; coxae short, not touching serially; gnathopods small,
gnathopod 1 subchelate, gnathopod 2 simple, extremely setose, articles
4 and 5 elongate and anteroposteriorly apposed; uropod 2 biramous;
uropod 3 flattened, its single ramus equal to peduncle; urosomites
coalesced or free. Species: 47, cosmopolitan, open-sea littoral, brackish,
fresh-water and Caspian Sea.

Dryopoides Stebbing
Dryopoides Stebbing, 1888.—Stebbing, 1906.

Type-species: D. westwood: Stebbing, 1888 (monotypy). See
Stebbing, 1910.

MARINE GAMMARIDEAN AMPHIPODA 191

u
f

=

eX
AR
?

e/a mr

Figure 78.—Corophiidae: a, Bogenfelsia incisa J. L. Barnard (1962d); b, Camacho bathy-
plous Stebbing (J. L. Barnard, 1961). Upper lip: c, Neohela monstrosa (Boeck) (Sars,
1895, pl. 224); d, Corophiwm volutator (Pallas) (Sars, 1895, pl. 219 as C. grossipes). Man-
dible: e, Siphonoecetes colletti Boeck (Sars, 1895, pl. 218); f, Corophium; g, Ericthonius
brasiliensis (Dana) (Sars, 1895, pl. 215, as E. abditus). Mouthparts, Corophium: h,
lower lip, 2,7, maxillae 1, 2, &, maxilliped. Maxilliped: /, Ericthonius.

192 U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum absent; article 3 of antenna 1 much shorter
than article 1, flagellum longer than peduncle; antenna 2 slender,
slightly shorter than antenna 1, flagellum shorter than article 5;
mandibular palp 3-articulate; coxae of medium size, touching serially;
enathopods normally subchelate, gnathopod 2 larger than 1 in male,
smaller than 1 in female; uropod 2 biramous; uropod 3 biramous, rami
shorter than peduncle. Female assigned to Aoridae. Species: 1, E.
Australia, littoral.

Ericthonius Milne Edwards

Ericthonius Milne Edwards, 1830.—Stebbing, 1906.
Pyctilus Dana, 1852b.

Type-species: HH. difformis Milne Edwards, 1830 (selected by
Stebbing, 1888, p. 142!). See Sars, 1895.

Accessory flagellum absent; article 3 of antenna 1 longer than
article 1, flagellum subequal to peduncle; antenna 2 slender, equal to
antenna 1, flagellum as long as peduncle; mandibular palp 3-articulate ;
coxae short, scarcely touching serially; male gnathopod 2 larger than
1, carpochelate, article 6 simple, female gnathopods normally sub-
chelate; uropod 2 biramous; uropod 3 with its single, medium length,
uncinate ramus shorter than peduncle. Species: 8, mainly N.
Hemisphere and tropics (one species), littoral (one extends to 1,134 m).

Gaviota J. L. Barnard
Gaviota J. L. Barnard, 1958b.

Type-species: G. podophthalma J. L. Barnard, 1958b (original
designation).

Accessory flagellum absent; article 3 of antenna 1 equal to article 1,
flagellum longer than article 3; antenna 2 slender, equal to antenna 1,
flagellum longer than article 5; mandibular palp 3-articulate; coxae
long, touching serially; gnathopods normally subchelate; uropod 2
biramous; uropod 3 with its single ramus shorter than peduncle; head
with vmmense, pedunculate lateral ocular lobes. Species: 1, California,
littoral.

Grandidierella Coutiére
Grandidierella Coutiére, 1904.

Type-species: G. mahafalensis Coutiére, 1904 (monotypy). See
Ruffo, 1958.

Accessory flagellum 1l-articulate, usually elongate; article 3 of
antenna | shorter than article 1, flagellum nearly as long as peduncle;
antenna 2 slender (rarely stout in male), nearly equaling antenna 1,
flagellum often shorter than article 5; mandibular palp 3-articulate;

MARINE GAMMARIDEAN AMPHIPODA 193

——=—

—
a eS=
Ss

Ficure 79.—Corophiidae: a, Neohela monstrosa (Boeck) (Sars, 1895, pl. 224). Gnathopod
1: b, Ericthonius brasiliensis (Dana) (Sars, 1895, pl. 215, as FE. abditus); c, Unciola leucopis
(Krgyer) (Sars, 1895, pl. 222); d, Siphonoecetes colletti Boeck (Sars, 1895, pl. 218); e,
Corophium volutator (Pallas) (Sars, 1895, pl. 219, as C. grossipes); f, male Cerapopsis
longipes Della Valle (1893). Gnathopod 2: g, Cerapopsis; h, Corophium; 1, Unciola; 7,
Siphonoecetes; k, Ericthonius; 1, Cerapus crassicornis (Bate) (Sars, 1895, pl. 217).

194 U.S. NATIONAL MUSEUM BULLETIN 271

coxae short, scarcely touching serially; male gnathopod 1 large,
carpochelate, article 5 bearing a tooth (or two), article 6 simple,
enathopod 2 smaller, normally subchelate or slightly chelate, and
articles 4-5 sometimes like Corophium; uropod 2 biramous; uropod 3
with its single ramus longer than peduncle. Species: 15, circum-
tropical and warm-temperate, littoral, especially brackish water.
Differs from Neomicrodeutopus in Aoridae by elongate article 1 of
mandibular palp, that article as long as article 2.

Kamaka Derjavin
Kamaka Derjavin, 1923.—Gurjanova, 1951.

Type-species: K. kuthae Derjavin, 1923 (monotypy).

Accessory flagellum absent; antenna 1 short, article 3 slightly
shorter than article 1, flagellum equal to peduncle; antenna 2 much
longer than antenna 1, slender, flagellum equal to or shorter than
article 5, often about 2-articulate; mandibular palp 3-articulate; coxae
of medium size, touching serially ; gnathopods normally subchelate and
male gnathopod 2 with large palmar tooth; uropod 2 biramous; uropod
3 with its single ramus slightly shorter than (or subequal to) peduncle;
pleonites 4-5 coalesced. Species: 3, boreal N. W. Pacific, littoral.

Neohela Smith

Hela Boeck, 1861 (homonym, Decapoda).
Neohela Smith, 1881 (new name).—Stebbing, 1906.
Helella Sars, 1882.

Type-species: Hela monstrosa Boeck, 1861 (monotypy). See Sars,
1895.

Accessory flagellum multiarticulate; article 3 of antenna 1 longer
than article 1, flagellum longer than peduncle; antenna 2 slender,
slightly longer than 1, flagellum longer than peduncle; mandibular
palp 3-articulate; coxae short, some sharp, not touching serially;
enathopods normally subchelate, gnathopod 1 slightly larger than
enathopod 2; uropod 2 biramous; uropod 3 with its single ramus equal
to or twice as long as peduncle; telson slightly enlarged and apparently
coalesced with urosomite 3; inner plate of maxilla 1 unth about five setae
mostly terminal (Parunciola). Species: 3, subarctic-boreal, littoral to
abyssal (2,288 m).

Paracorophium Stebbing

Paracorophium Stebbing, 1899b, c; 1906.

Type-species: Corophium excavatum Thomson, 1884 (monotypy).
See Hurley, 1954b.

MARINE GAMMARIDEAN AMPHIPODA 195

Figure 80.—Corophiidae: a, Runanga coxalis J. L. Barnard (1961). Uropod 2: b, Cerapus
crassicornis (Bate) (Sars, 1895, pl. 217). Uropod 3: c, Cerapus; d, Ericthonius brasili-
ensis (Dana) (Sars, 1895, pl. 215, as E. abditus); e, Corophium crassicorne Bruzelius (Sars,
1895, pl. 220); f, Siphonoecetes colletti (Boeck) (Sars, 1895, pl. 218); g, Neohela monstrosa
(Boeck) (Sars, 1895, pl. 224); h, Camacho bathyplous Stebbing (J. L. Barnard, 1961); 2,
Unciola leucopis (Krdyer) (Sars, 1895, pl. 222). Urosome, dorsal: 7, Pseudericthonius
gausst Schellenberg (1926a); k, Corophium volutator (Pallas) (Sars, 1895, pl. 219, as C.
grossipes); 1, Cerapus; m, Corophium crassicorne; n, Concholestes dentalii Giles (1888).
Telson: 0, Cerapus; p, Ericthonius difformis Milne Edwards (Sars, 1895, pl. 216); gq,
Neohela; r, Unciola; s, Corophium.

196 U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum absent; article 3 of antenna 1 shorter than
article 1, flagellum longer than peduncle; antenna 2 stout, shorter
than antenna 1, flagellum slightly longer than article 5; mandibular
palp 3-articulate; coxae of medium size, touching serially; gnathopod
1 normally subchelate; gnathopod 2 subchelate or chelate, article 4
greatly prolonged as in Corophium but diverging from article 5 and strongly
setose; uropod 2 biramous; uropod 3 with outer ramus slightly shorter
than peduncle, inner ramus half as long as outer. Species: 2, New
Zealand, fresh-water.

Parunciola Chevreux

Parunciola Chevreux, 1911a.

Type-species: P. seuratt Chevreux, 191la (monotypy).

Accessory flagellum multiarticulate; article 3 of antenna 1 much
longer than article 1 [flagellum broken]; antenna 2 slender, equaling
antenna 1 [flagellum broken]; mandibular palp 3-articulate; coxae
small, with softly rounded corners, but touching serially; gnathopods
normally subchelate; uropod 2 biramous; uropod 3 with its single
ramus twice as long as peduncle; inner plate of maxilla 1 with about 10
medial setae (Neohela, Unciolella). Species: 1, Algeria, littoral.

Pseudericthonius Schellenberg
Pseudericthonius Schellenberg, 1926a.

Type-species: P. gaussi Schellenberg, 1926a (monotypy).

Antenna | not described but apparently as in Hricthonius; antenna 2
slender; mandibular palp 3-articulate; coxae short, not touching;
enathopods normally subchelate; uwropods 1 and 2 with inner rami
much shorter than outer; uropod 3 with its single elongate uncinate
ramus shorter than peduncle. Species: 1, antarctic, littoral (to
342 m).

Runanga J. L. Barnard

Runanga J. L. Barnard, 1961.

Type-species: R&. coxalis J. L. Barnard, 1961 (original designation).

Accessory flagellum scale-like; article 3 of antenna 1 much longer
than article 1, flagellum longer than article 3; antenna 2 slender, equal
to antenna 1, flagellum longer than article 5; mandibular palp 3-
articulate; coxae short, not touching serially; gnathopods normally
subchelate; uropod 2 with one ramus; uropod 3 with two vestigial
rami, outer ramus hooked; pleopod 3 with one ramus. Species:
1, Tasman Sea, bathyal (610 m).

MARINE GAMMARIDEAN AMPHIPODA 197

Siphonoecetes Kréyer
Stphonoecetes Kréyer, 1845.—Stebbing, 1906.

Type-species: S. typicus Krgyer, 1845 (monotypy). See Stephensen,
1944a.

Accessory flagellum absent; article 3 of antenna 1 subequal to article
1, flagellum subequal to or longer than article 3; antenna 2 slender or
slightly thickened, longer than antenna 1, flagellum shorter than
article 5; mandibular palp 1l-articulate; coxae short, not touching
serially; gnathopods weakly or normally subchelate; uropod 2 with
1-2 rami; uropod 3 with medially expanded pedunele, its single ramus
shorter than peduncle. Species: 9, arctic-boreal to ?antisubtropics,
littoral to bathyal (565 m).

Unciola Say

Unciola Say, 1818.—Stebbing, 1906.
Glauconeme Krgyer, 1845 (homonym, Bryozoa).
Dryope and Driope Bate, 1862 (Dryope=homonym, Diptera).

Type-species: U. irrorata Say, 1818 (monotypy). See Shoemaker,
1945b.

Accessory flagellum elongate, multiarticulate or 1-articulate; article
3 of antenna 1 equal to article 1, flagellum subequal to peduncle;
antenna 2 of medium to strong stoutness, slightly shorter than antenna
1, flagellum subequal to article 5; mandibular palp 3-articulate; coxae
short, not touching serially; gnathopod 1 larger than gnathopod 2,
both normally subchelate or gnathopod 2 simple or parachelate;
uropod 2 biramous; uropod 3 often with medially expanded pedunele,
its single ramus slightly shorter than peduncle. Species: 15, arctic-
boreal, littoral to abyssal.

Unciolella Chevreux
Unciolella Chevreux, 1911la.

Type-species: U. lunata Chevreux, 191la (monotypy).

Accessory flagellum 1-articulate, elongate or multiarticulate; article
3 of antenna 1 equal to or shorter than article 1, flagellum equal to
peduncle; antenna 2 slender, shorter than antenna 1, flagellum shorter
than article 5; mandibular palp 3-articulate; coxae short, touching or
not touching serially, with softly rounded corners; gnathopods nor-
mally subchelate, gnathopod 1 shghtly the larger; uropod 2 biramous;
uropod 3 with its single ramus about 1.5 times longer than peduncle;
inner plate of maxilla 1 with one terminal seta (Neohela, Parunciola).
Species: 2, Mediterranean and S. Atlantic, littoral (to 392 m).

198 U.S. NATIONAL MUSEUM BULLETIN 271

Cressidae
Ficures 81, 82

Diacnosis.—Accessory flagellum absent; mandibular molar evanes-
cent; coxa 1 very small, always partially covered by following coxae;
coxa 4 large, either excavate posteriorly or shield-like and not excavate;
uropod 3 uniramous; outer lobes of maxilliped vestigial; telson entire
and fused with pleonite 6, latter often coalesced with pleonite 5;
article 2 of pereopod 3 expanded. See Stenothoidae, Amphilochidae,
Thaumatelsonidae, Pagetinidae, Leucothoidae, Anamixidae, Phliantidae.

if g y

Ficure 81.—Cressidae: Cressa dubia (Bate) (Sars, 1895, pls. 98-99): a, lateral; b, upper
lip; c, lower lip; d, telson; ¢,f, gnathopods 1, 2; g, maxilliped; h,7, maxillae 1, 2; 7, mandible.

MARINE GAMMARIDEAN AMPHIPODA 199

DescripTion.—Rostrum inconspicuous; accessory flagellum absent;
body smooth or with slight dorsa] teeth; coxa 1 small, hidden by follow-
ing coxae; coxa 4 large, either shield-like or excavate posteriorly;
upper lip incised; mandible with weak, sparsely spinose molar, not
triturative, palp long, 3-articulate; lower lip with coalesced inner
lobes, outer lobes with blunt extremities; palp of maxilla 1 uniartic-
ulate; maxilla 2 small, lobes blunt; maxillipeds slender, outer lobes
vestigial; gnathopods small, first simple, second subchelate; uropod 3
uniramous, the single ramus 2-articulate; telson of medium length,
entire, fused with pleonite 6; article 2 of pereopod 3 expanded.

a

SS
ase
ae
4

LEE

\

Figure 82.—Cressidae: a, Cressina monocuspis Stephensen (1931); b, uropod 3 of Cressa
minuta Boeck (Sars, 1895, pl. 99); c, pleonite 3 and urosome, left to right, of Cressina,
showing absence of urosomite 3.

RELATIONSHIP.—The Stenothoidae have all segments of the urosome
and the telson distinct, coxa 4 is always shield-like, not excavate
posteriorly (like Cressina, however) and article 2 of pereopod 3 is
always linear.

All Thaumatelsonidae have coxa 4 shield-like, not excavate pos-
teriorly. Although the segments of the urosome may be coalesced the
telson is not clearly fused with them and is usually thickened dorso-
ventrally; article 2 of pereopod 3 is always linear.

The Phliantidae have depressed, rugose bodies, and relatively equal
coxae.

200 U.S. NATIONAL MUSEUM BULLETIN 271

Key to the Genera of Cressidae

1. Coxa 4 rectangular, excavate posteriorly (fig. 8la) ........ Cressa
Coxa 4 shield-like, not excavate posteriorly (fig. 82a). ..... Cressina

Genera of Cressidae

Cressa Boeck

Danaia Bate, 1857a (homonym, Anthozoa).
Cressa Boeck, 1871.—Stebbing, 1906.

Type-species: Danaia dubia Bate, 1857a (selected by Boeck, 1876,
as Cressa schiodte. Boeck, 1871). See Sars, 1895.

Species: 6, arctic-boreal, N. Atlantic, littoral to bathyal.

Cressina Stephensen
Cressina Stephensen, 1931.

Type-species: C. monocuspis Stephensen, 1931 (monotypy).
Species: 1, subarctic, bathyal (682—1,096 m).

Dexaminidae

[including Anatylidae Bulycheva, 1955]
Figures 83-85

Draenosis.—Accessory flagellum absent; urosomites 2 and 3
coalesced; mandible lacking palp, molar well developed or vestigial;
uropod 3 biramous; telson deeply cleft; body laterally compressed;
gnathopods feeble but subchelate. See Atylidae, Prophliantidae, Phlian-
tidae. See Sphaerophthalmus in ‘‘Incertae Sedis.”’

Description.—Accessory flagellum absent; body laterally com-
pressed and often dorsally carinate or toothed; urosomites 2 and 3
coalesced; mandible lacking palp, molar well developed, triturative
(Dexamininae) or evanescent (Anatylinae) ; lower lip with or without
mandibular processes, inner lobes well developed or vestigial, occa-
sionally fused; palp of maxilla 1 uni- or biarticulate; maxillipeds with
3- or 4-articulate palp, inner plates occasionally vestigial; gnathopods
feeble, subchelate; pereopods occasionally cheliform; uropod 3
biramous; telson long or short, deeply cleft.

Rexationsuie.—The Talitroidea have fully separated urosomal
segments.

Phliantidae and Prophliantidae have the mandibular molar evanes-
cent. The Phliantidae, furthermore, have the palp of maxilla 1 de-
graded or absent. The Prophliantidae have all urosomal segments
coalesced.

MARINE GAMMARIDEAN AMPHIPODA 201

Ficure 83.-—Dexaminidae: a, Polycheria antarctica (Stebbing, 1888); b, Dexamonica redun -
cans J. L. Barnard (1958c).

202 U.S. NATIONAL MUSEUM BULLETIN 271

Probably Sphaerophthalmus Spandl is assignable to the Dexamin-

idae. See its diagnosis under ‘“‘Incertae Sedis.”’

co

(oy)

“I

Key to Subfamilies of Dexaminidae
Mandibular molar large, triturative (fig.847) ... . . . . Dexamininae
Mandibular molar evanescent (fig. 85e) ...... . . . .Anatylinae

Key to the Genera of Dexaminidae

(expanded after Sheard, 1938)

Palp of maxilliped with three articles (fig. 84d). ........... 2.2
Palp of maxilliped with four articles (fig. 84e) ............ .4
Inner lobes of lower lip well developed (fig. 84h) . . . . . . Dexaminella
Inner lobes of lower lip vestigial (fig. 847) . . . . BY ke

Pereopods 1—5: article 4 shorter than articles 5-6 combined Ge 85n).
Dexamine

Pereopods 1-5: article 4 longer than articles 5-6 combined (fig. 850).
Tritaeta
Pereopods 1-5 cheliform (fig. 85p) ........... .... .Polycheria
Pereopods 1-5 simple. . . . Sane ACME INEM CCE Gy AG ison) 6
Palp of maxilla 1 uniarticulate Ge. 84m) PURER. VR OI UU ara
Palp of maxilla 1 biarticulate (fig. 847) .... Manes itiay nats)
Inner plates of maxillipeds vestigial, short and bud-like Gee, 84c) . Pee n ats 276
Inner plates of maxillipeds of medium size (fig. 84d) ......... .8
Inner lobes and mandibular lobes of lower lip distinct; palp article 4 of
maxilliped claw-shaped (fig. 84e) ... . . . . . .Dexaminoides
Inner lobes of lower lip coalesced, remelirale lobes obsolescent (fig. 84f);
palp article 4 of maxilliped stout, blunt (fig. 84c). . . . . Dexamonica
Mandibular processes of lower lip absent (fig. 84g). . . . . Syndexamine
Mandibular processes of lower lip present (fig. 847). . . . . Paradexamine

Palp article 2 of maxilla 1 large (fig. 841); inner plates of maxillipeds well
developed (fig. 857); coxa 5 small, coxa 4 not posteriorly excavate.

Anatylus

Palp article 2 of maxilla 1 small (fig. 84k) or large; inner plates of maxillipeds

vestigial, short and bud-like (fig. 84c); coxa 5 larger than anterior coxae,

coxa) 4) posteriorly excavate 7Ms 099525 6 a0 4s ees Gennes

Genera of Dexaminidae

Anatylus Bulycheva

Anatylus Bulycheva, 1955.

Type-species: A. pavlovskii Bulycheva, 1955 (original designation).
Mandibular molar evanescent; lower lip bearing inner lobes, man-

dibular processes distinct, palp of maxilla 1 biarticulate, artrcle 2 long
(Guernea) ; maxillipedal palp 4-articulate, inner plates well developed;
coxa 4 not excavate posteriorly. Species: 1, boreal N. W. Pacific,
littoral.

285-135 O - 69 - 14

MARINE GAMMARIDEAN AMPHIPODA 203

Dexamine Leach

Dexamine Leach, 1814b.—Stebbing, 1906.
Amphithonotus Costa, 1851b.

Type-species: Cancer spinosus Montagu, 1813 (monotypy). See
Sars, 1895.

Lower lip with poorly developed inner lobes, mandibular processes
distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 3-articulate,
inner plates well developed; article 4 of pereopods shorter than articles
5-6 combined (Tritaeta). Species: 5, boreal N. E. Atlantic, Mediter-
ranean, Ceylon, littoral.

Dexaminella Schellenberg
Dexaminella Schellenberg, 1928b.

Type-species: D. aegyptiaca Schellenberg, 1928b (monotypy).

Lower lip with well-developed inner lobes, mandibular processes
distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 3-articulate,
inner plates absent. Species: 1, Red Sea, littoral.

Dexaminoides Spandl
Dexaminoides Spandl, 1923.

Type-species: D. orientalis Spandl, 1923 (monotypy). See J. L.
Barnard, 1965.

Lower lip with well-developed inner lobes, mandibular processes
distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 4-articulate,
article 4 claw-like (Dexamonica), inner plates vestigial, bud-like.
Species: 1, Indo-Pacific, tropical, littoral.

Dexamonica J. L. Barnard
Dexamonica J. L. Barnard, 1958c.

Type-species: D. reduncans J. L. Barnard, 1958c (original
designation).

Lower lip with distinct, fused inner lobes, mandibular processes
obsolescent; palp of maxilla 1 uniarticulate; maxillipedal palp 4-
articulate, article 4 short, not claw-like (Dexaminoides), inner plates
vestigial, bud-like. Species: 1, California, littoral.

Guernea Chevreux

Helleria Norman, 1868 (homonym, Isopoda).
Guernea Chevreux, 1887 (new name).—Stebbing, 1906.
Prinassus Hansen, 1887.

Type-species: Helleria coalita Norman, 1868 (monotypy). See
Chevreux and Fage, 1925.

204

U.S. NATIONAL MUSEUM BULLETIN 271

B

{\

LOM) Zi al

TT. AR EN
Th 7U
Le rk
Da \
HY “nN =

iN IN
4 \ N

\\\
Bi NN Xs
1 \
a) \
i \
a
ey
\\
V
Th
\
\)

a=

p
Figure 84.—Dexaminidae: a, Dexamine spinosa (Montagu)(Sars, 1895, pl. 167).
lip: b, Dexamine.

Upper
Maxillipeds: c, Guernea coalita (Norman) (Chevreux and Fage, 1925);
d, Dexamine; e, Polycheria antarctica (Stebbing, 1888). Lower lip: f, Dexamonica redun-
cans J. L. Barnard (1958c); g, Syndexamine carinata Chilton (1914); 4, Dexaminella
aegyptica Schellenberg (1928); 7, Dexamine. Mandible: 7, Dexamine. Maxilla 1: k,
Guernea; 1, Polycheria; m, Dexamine. Maxilla 2: n, Dexamine. Gnathopods 1, 2: 0, 9,
Dexamine.

MARINE GAMMARIDEAN AMPHIPODA 205

t

Figure 85.—Dexaminidae: Anatylus pavlovskii Bulycheva (1955), parts as follows:
a, body without appendages; b,c, antennae 1, 2; d, upper lip; ¢, mandible; f, lower lip;
g,h, maxillae 1, 2; 7, maxilliped; 7, uropod 3; k, telson; /,m, gnathopods 1, 2. Pereopod
4: n, Dexamine spinosa (Montagu) (Sars, 1895, pl. 167); 0, Tritaeta gibbosa (Bate) (Sars,
1895, pl. 168). Pereopod 3: p, Polycheria antarctica (Stebbing, 1888). Uropod 3: gq,
Dexamine. Telson: 7, Guernea coalita (Norman) (Chevreux and Fage, 1925); 5, Dexamine.

Mandible: #, Guernea.

206 U.S. NATIONAL MUSEUM BULLETIN 271

Lower lip with well-developed inner lobes, mandibular processes
absent; palp of maxilla 1 biarticulate, artecle 2 short (Anatylus) except
in G. petalocera Ruffo; maxillipedal palp 4-articulate, inner plates
vestigial, bud-like; coza 5 larger than anterior coxae (Anatylus).
Species: 3, boreal-tropical in N. and E. hemispheres, littoral.

Paradexamine Stebbing
Paradexamine Stebbing, 1899d; 1906.

Type-species: Dexamine pacifica Thomson, 1879 (original designa-
tion). See Stephensen, 1927.

Lower lip with well-developed inner lobes, mandibular processes
distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 4-articulate,
inner plates of moderate size. Species: 9, antarctic-antiboreal, littoral.

Polycheria Haswell
Polycheria Haswell, 1880c.—Stebbing, 1906.

Type-species: Dexamine antarctica Stebbing, 1875 (present selec-
tion, denoted by synonymy of Chilton, 1912). See Stebbing, 1888 (as
Tritaeta antarctica and T. kerguelent).

Lower lip with well-developed inner lobes, mandibular processes
absent; palp of maxilla 1 ?biarticulate (uniarticulate=auct. other
than Stebbing); maxillipedal palp 4-articulate, inner plates well
developed; pereopods cheliform. Species: 4, Antarctic to N. Pacific,
littoral.

Syndexamine Chilton

Syndexamine Chilton, 1914.

Type-species: SS. carinata Chilton, 1914 (monotypy).

Lower lip with small but distinct inner lobes, mandibular processes
absent; palp of maxilla 1 uniarticulate; maxillipedal palp 4-articulate,
inner plates small. Species: 1, New Zealand, littoral.

Tritaeta Boeck

Lampra Boeck, 1871 (homonym, Lepidoptera).
Tritaeta Boeck, 1876 (new name).—Stebbing, 1906.

Type-species: Atylus gibbosus Bate and Westwood, 1863 (mono-
typy). See Sars, 1895.

Lower lip with tiny imner lobes, mandibular processes distinct;
palp of maxilla 1 uniarticulate; maxillipedal palp 3-articulate, imner
plates well developed; article 4 of pereopod 5 longer than articles 5-6
combined (Dexamine). Species: 2, EK. Atlantic, littoral.

MARINE GAMMARIDEAN AMPHIPODA 207

Dogielinotidae

FIGURE 86

Diacnosis.—Accessory flagellum absent; epistome with a pendant
lobe; mandible lacking palp; palp of maxilla 1 vestigial; uropod 3
uniramous or lacking a ramus; telson broad, cleft shortly; antennae
and pereopods densely spinose and setose (fossorial). See Talitroidea,
Haustoriidae, Kurtidae, Phliantidae.

Description.—Accessory flagellum absent; body not carinate or
toothed; rostrum small; coxae of medium length; mandible lacking
palp, molar large, triturative; palp of maxilla 1 vestigial, 1-articulate;
enathopods of medium size, subchelate; uropod 3 uniramous or ramus
absent, ramus small, bearing apical setae; telson short, broad, shortly
cleft; antennae and pereopods densely spinose as in phoxocephalids
and haustoriids.

RELATIONSHIP.—This remarkable family combines characters of
Talitroidea and Phoxocephalidae-Haustoriidae. Its two species have
these characters of Talitroidea: no accessory flagellum; uniramous
[or aramous] third uropod; the telson of a Hyale; no mandibular palp;
and a vestigial first maxillary palp. Dogielinotids have these characters
of the Phoxocephalidae-Haustoriidae: shape of head and rostrum;
short, very spinose antennae; similar gnathopods and pereopods; long
peduncular setae of the first and second uropods; and the strong tooth
at the posteroventral corner of the third pleonal epimeron.

Metoediceros Schellenberg (1931), now assigned erroneously to
Oedicerotidae probably should form the type of a new family with
resemblance to Dogielinotidae (see Incertae Sedis). Metoediceros
differs from dogielinotids in its circular, uncleft telson. Its epistome
is unknown.

Dogielinotids are distinguished from Talitroidea by their phoxo-
cephalid characters and vice-versa, and from the Haustoriidae by the
uniramous uropod 3 and the absence of a mandibular palp.

The Kurtidae have coalesced urosomal segments.

The Prophliantidae and Phliantidae have evanescent mandibular
molars.

Key to the Genera of Dogielinotidae

i ee Oropodisiwithtonewramusys ca5 45 yas estes ee ae see Dogielinotus
Wropodis lacking, rami un mus eyy ices ate Pewter bea Neen § Haustorioides

208 U.S. NATIONAL MUSEUM BULLETIN 271
Genera of Dogielinotidae

Dogielinotus Gurjanova

Dogielinotus Gurjanova, 1953.

Type-species: Allorchestes moskvitina Dershavin, 1930 (original
designation).
Species: 1, boreal Pacific, littoral.

Ficure 86.—Dogielinotidae: Dogielinotus sp., California, original figures: a, lateral aspect;
b, mandible; c, maxilla 1; d, maxilliped; ¢, telson; f, uropod 3; g, lower lip; h, anterior view
of epistome pendant in front of upper lip; 7, maxilla 2.

MARINE GAMMARIDEAN AMPHIPODA 209

Haustorioides Oldevig
Haustorioides Oldevig, 1958.

Type-species: H. munsterhjelmi Oldevig, 1958 (monotypy).
Species: 1, boreal Pacific, littoral.

Eophliantidae

(Sheard, 1936a, elevated by Gurjanova, 1958]
FicursEs 87, 88

Diacnosis.—Accessory flagellum absent; body cylindrical; all
coxae short, generally not touching; mandible lacking palp, molar
vestigial or absent; palp of maxilla 1 degraded or absent; gnathopods
usually feeble; uropod 3 becoming degenerate; telson short, weakly
to fully cleft. See Prophliantidae, Phliantidae, Talitroidea, Kuriidae,
Colomastigidae.

Description.—Body vermiform, head globular, lacking rostrum,
urosome tending to be reduced and its segments coalesced, uropod 3
becoming degenerate, with two, one or no rami; accessory flagellum
absent; mandible lacking palp and molar obsolescent; palp of maxilla
1 small, 1-articulate or absent; remaining mouthparts basic; gnatho-
pods feeble, simple or weakly subchelate or parachelate, article 3 of
enathopods occasionally elongate; telson short, weakly to fully cleft;
pleopods biramous or uniramous, requiring further study in the
several genera.

Uropod 3 and its relationship to the urosome need careful study;
it is necessary to distinguish clearly pleonite 6, the peduncle of
uropod 3, and its ramus. Sometimes pleonite 6 is fused to 5 but a
peduncle of uropod 3 may be confused with the segment if one mistakes
a ramus for a peduncle.

ReLationsuip.—The Phliantidae differ from the Eophliantidae by
their depressed or compressed bodies with long coxae.

The Talitroidea differ by their strongly triturative mandibular
molars (except Najna) and compressed bodies.

The Colomastigidae have cylindrical bodies but the inner plates
of the maxillipeds are very small, the telson is cleft, and the mandib-
ular cutting edge is divided into large teeth.

Eophliantidae may resemble Corophiidae and Podoceridae because
of their depressed, subcylindrical bodies but the latter two families
have mandibular palps and molars.

210 U.S. NATIONAL MUSEUM BULLETIN 271

f g

Figure 87.—Eophliantidae: a, Wandelia crassipes Chevreux (1906c); b, Ceinina japonica
Stephensen (Gurjanova, 1951). Upper lip: c, Wandelia. Head: d, Bircenna fulva
Chilton (Nicholls, 1939) (arrow marks ventral flange). Mandible: e, Amphitholina
cuniculus (Stebbing) (Chevreux and Fage, 1925); f, Wandelia; g, Wandelia japonensis
(Nicholls, 1939). Lower lip: h, Bircenna ignea Nicholls (1939).

MARINE GAMMARIDEAN AMPHIPODA P11

Key to the Genera of Kophliantidae

1. Uropod 3 biramous (figs. 88l,0,q) 2
Uropod 3 not biramous (figs. 88m,n,p) . 4

2. Maxilla 1 lacking palp (fig. 88a) . Apes Senne Oiy eT REN ¢ 5 3
Maxilla 1 with small palp (fig. 88c) .. . Se frophiianticn

3. Uropod 3 with short, broad rami less than half as longs as peduncle (fig. 880),
gnathopod 2 powerful (fig. 887). . . . . . Amphitholina
Uropod 3 with slender rami as long as s pedunele (iw 88), gnathopod 2 feeble
HeAS Sa) iiss ta) é . . . Biancolina

4. Pleonites 5-6 small emt Shatin. tas alae Nar LR a ania as ee OLN RN eM
Pleonites 5-6 coalesced. . . . . . BV US HMO gd

5. Antennae half as long as head, avila, 1 lackine alo: mena lacking
mn@lene 55) 6) 6 . . . . (theoretical position) Ceinina
Antennae as tones as heel rts les 1 with minute palp, mandible bearing
small molar (one member at least). . . . . . . =... . . Eophliantis*

6. Pleopods uniramous (fig. 88k) ........... . .. Cylindryllioides
Pleopods biramous. . . aaica hey See MT t canaene east Mes SRSA

7. Pereonite 1 with a ential fiance Ge. 87d). ieee eee ee ea TRE ene
Rereonitemilackineayventraliflamgel cis ais) ye) aS ee 8

8. Coxae tiny, serially noncontiguous, flagellum of antenna 2 biarticulate,
mandibular palp absent... . .... .+. Ceinina
Coxae small, serially contiguous, Abyscllhonn of POA 2 about 6-articulate,
mandible with vestigal palp. . ........... =... Wandelia

1 Sheard (1936a) and Nicholls (1939) have different concepts of uropod 3 on Eophliantis.

Genera of Eophliantidae

Amphitholina Ruffo

Amphitholina Ruffo, 1953.
Type-species: Amphithoe cuniculus Stebbing, 1874 (original des-
ignation). See Chevreux and Fage, 1925 (as Biancolina cuniculus).
Maxilla 1 lacking palp; pleonites 5-6 apparently separate; uropod
3 with two short broad rami, half as long as peduncle, gnathopod 2
powerful (Biancolina). Species: 1, Mediterranean, littoral.

Biancolina Della Valle
Biancolina Della Valle, 1893.—Stebbing, 1906 (in part).

Type-species: B. algicola Della Valle, 1893 (monotypy).

Maxilla 1 lacking palp; pleonites 5-6 separate; uropod 3 with
slender rami as long as peduncle, gnathopod 2 feeble (Amphitholina).
Species: 2, Mediterranean, Australia, littoral.

212 U.S. NATIONAL MUSEUM BULLETIN 271

Bircenna Chilton
Bircenna Chilton, 1884.—Stebbing, 1906.

Type-species: B. fulvuus Chilton, 1884 (monotypy). See Sheard,
1936a; Nicholls, 1939.

Maxilla 1 lacking palp; pleonites 5-6 coalesced; uropod 3 with a
single vestigial ramus or none; pereonite 1 with a ventral flange.
Species: 4, 5S. Australia and New Zealand, littoral.

r

a

Figure 88.—Eophliantidae: Maxilla 1: a, Wandelia crassipes Chevreux (1906c); b, Am-
phitholina cuniculus (Stebbing) (Chevreux and Fage, 1925); c, Eophliantis tindale: Nicholls
(1939). Maxilliped: d, Wandelia. Gnathopod 1: e, Wandelia; f, Amphitholina; g,
Biancolina algicola Della Valle (1893). Gnathogod 2: h, Wandelia; 1, Amphitholina;
j, Biancolina algicola. Pleopod: k, Cylindryllioides mawsont Nicholls (1938). Uropod 3:
1, Biancolina australis Nicholls (1939); m, Eophliantis; n, Bircenna nichollst Sheard
(Nicholls, 1939). Telson, uropod 3 and dorsum of urosome: 0, Amphitholina; p, Wan-
delia; q, Biancolina algicola. Urosome, lateral: r, Eophiiantts.

MARINE GAMMARIDEAN AMPHIPODA 213

Ceinina Stephensen
Ceinina Stephensen, 1933.

Type-species: C. japonica Stephensen, 1933 (monotypy). See
Gurjanova, 1951; Nicholls, 1939 (as Wandelia gaponensis).

Maxilla 1 lacking palp; pleonites apparently coalesced; uropod 3
with one ramus; antennae half as long as head, mandible lacking molar
(Hophliantis) ; coxae tiny, serially noncontiguous (Wandelia). Species:
1, Japan, littoral.

Cylindryllioides Nicholls

Cylindryllioides Nicholls, 1938.

Type-species: C. mawsoni Nicholls, 1938 (original designation).
Maxilla 1 lacking palp; pleonites 5-6 coalesced; uropod 3 lacking
rami; pleopods uniramous (combining character). Species: 1, antarctic,
littoral.
Eophliantis Sheard

Eophliantis Sheard, 1936a.

Type-species: . tindalei Sheard, 1936a (original designation).

Maxilla 1 with small palp; pleonites 5-6 separate; uropod 3 with
two blunt rami (Sheard), no rami (Nicholls, 1939) ; antennae as long as
head, mandible bearing small molar (Ceinina). Species: 1, Tasmania,
littoral.

Wandelia Chevreux
Wandelia Chevreux, 1906a; 1906b.

Type-species: W. crassipes Chevreux, 1906a (original designation) ;
1906b.

Maxilla 1 lacking palp; pleonites 5-6 coalesced; uropod 3 lacking
rami; coxae small, serially contiguous (Ceinina). Species: ?2, antarctic
to N. Pacific, littoral.

Eusiridae
including Pontogeneiidae]

Fiaures 89-93

Diacnosts.—Accessory flagellum either a single styliform article
or two short articles, a scale or absent; telson cleft. See Calliopiudae,
Pleustidae, Liljeborgiidae, Gammaridae, Vitjazianidae, Astyridae,
Lepechinellidae, Paramphithoidae.

Descriptron.—Accessory flagellum absent, or composed of a scale
or one long article or two short articles; rostrum present or obsolete;
upper lip not incised (bilobed in one genus); mandibles bearing 3-

214 U.S. NATIONAL MUSEUM BULLETIN 271

—

SESE TASS BEISDERIBPAIIES N

a oe

Ficure 89.—Eusiridae: a, Rhachotropis aculeata (Lepechin) (Sars, 1895, pl. 149); b, Eustrus
cuspidatus Krdéyer (Sars, 1895, pl. 146). Kinds of accessory flagella: c, Liouvillea oculata
Chevreux (1912b); d, Rhachotropis; e, fused process, Prostebbingia gracilis (Chevreux,
1912b); f, Pontogeneoides abyssi Nicholls (1938); g, no accessory flagellum on Pontogeneia
inermis (Krdyer) (Sars, 1895, pl. 159); h, Djerboa furcipes Chevreux (1906c); 2, Eusirus
propinquus Sars (1895, pl. 147). 7, Calceolus.

MARINE GAMMARIDEAN AMPHIPODA 215

articulate palp, molar triturative (except in a few genera); lower lip
with or without inner lobes; other mouthparts basic, except one genus
with shortened palp of maxilla 1; gnathopods either powerful and
subchelate, occasionally small, or feeble, occasionally nearly simple
(Harpinioidella); coxae of medium length or short; uropod 3 with
elongate, lanceolate rami; telson often elongate, much longer than
peduncle of uropod 3, or increasingly shortened, cleft deeply or slightly
cleft only apically, occasionally appearing to be emarginate; in
Eusirogenes telson shorter than peduncle of uropod 3.

Revationsuip.—The Astyridae differ from the Eusiridae by the
nontriturative mandibular molar and the characteristic shape of the
lower lip. Some astyrids have a massive head and all have a very
short telson.

The Vitjazianidae are distinguished by their distinctly simple first
enathopods and the elongate article 1 of the flagellum on antenna 1
which equals the peduncle in length. Pleustidae have their character-
istic lower lips.

Since Calliopiidae appear to be Eusiridae-Pontogeneiidae with fused
telsonic lobes and some Eusiridae have nearly entire telsons, it must
be pointed out that Sancho in the Calliopidae, with entire telson and
with 1-articulate accessory flagellum as long as article 1 of the primary
flagellum has gnathopod 2 similar to the “‘Kusirus” gnathopods and
might be placed more logically in Eusiridae than in Calliopiidae. See
the latter family for further discussion.

Most synopiids have a multiarticulate accessory flagellum but a few
have an accessory flagellum similar to that of eusirids. The massive,
galeate head of most synopiids is characteristic, especially because of
its strongly deflexed rostrum. Synopiid gnathopods are feeble whereas
those of eusirids are usually powerful.

Rhachotropis aculeata bears resemblance to Paramphithoidae in its
slightly acute first four coxae but differs by its powerful gnathopods.
Gnathopods of Paramphithoidae are feeble and one or more of the
first four coxae are acuminate.

All Liljeborgiidae have a nontriturative mandibluar molar, but
eusirid resemblance to this family is close because some liljeborgiids
have a 2-articulate accessory flagellum and a few eusirid genera have a
poorly triturative mandibular molar.

Most Gammaridae differ from Eusiridae by their multiarticulate
accessory flagella but a number of genera have 1-2-articulate accessory
flagella (see couplets 6-17 in Gammaridae key). Eusiridae differ from
those genera of Gammaridae by their elongate telsons and the slight
elongation of the metasome. Nearly half of those gammarid genera
have short inner rami on uropod 3, and most of the remainder have
either foliaceous rami on uropod 3, and elongate article 2 of the

216 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 90.—Eusiridae: a, Eurymera monticulosa Pfeffer (Chevreux, 1906c). Upper lips:
b, Eusirus propinquus Sars (1895, pl. 147); c, Pontogeneia inermis (Krgyer) (Sars, 1895,
pl. 159). Mandible: d, Eusirus; e, Pontogeneia; f, Liouvillea oculata Chevreux (1912b).
Lower lip: g, Pontogeneia; h, Eusirus; 1, Bovallia gigantea Pfeffer (Chevreux, 1906c); 7,
Pontogeneoides abyssi Nicholls (1938); k, Eurymera; 1, Meteusiroides keyensis Pirlot (1934).

MARINE GAMMARIDEAN AMPHIPODA Dilvy,

mandibular palp or a simple gnathopod 1. Only Elasmopus, Liagocera-
docus, and Anelasmopus do not fit these statements.

Though originally assigned to Amphilochidae, Pseudamphilochus
Schellenberg falls to the Eusiridae, especially through the illustrated
family key. That genus has the head of an amphilochid or pleustid,
enathopods of an amphilochid, a lower lip faintly resembling that of
Pleustidae, but coxa 1 is not reduced as it is in Amphilochidae and the
cleft telson is unlike either pleustids or amphilochids. Pseudamphilo-
chus has been given a special place in Gammaridea Family Key G
(p. 114) and is mentioned also under the Amphilochidae.

Lepechinellids have acuminate coxae and apparently always have
pleonites 5-6 coalesced.

Key to the Genera of Eusiridae !

1. Article 6 of gnathopods attached to the strongly produced or extended apex
of article 5 (‘‘Eusirus gnathopods’’), article 6 as broad as long (figs.

COP TY Ost) Vi ance aber ae TSA Me eae,
Article 6 of Pahonods with nerraal deehment. Bericle 6 longer than
broad (figs. 92a,c,d,i,m) .. . 5

2. Articles 4 and 6 of gnathopods saucehiars e. 92h), ariniennaes sla fcalecol
(fig. 897) .. . . . Eusiropsis
eis 4 and 6 aie pontnepadst not Vouching| pCparated ie a lobe of article

5 (figs. 92b,7), antennae lacking calceoli. . . 3

3. Nettie 6 of gnathopod 1 larger than article 6°of pannaped 2, alto amielle 3 of
mandible shorter or scarcely shorter than article 2 (fig. 90f) . Eusirogenes
Article 6 of gnathopod 1 equal to or smaller than article 6 of gnathopod 2,
palp article 3 of mandible equal to or longer than article 2 (fig.90d). . 4

4. Posterior border of article 5 of gnathopods forming a very narrow projecting
lobe, much narrower than length of tapered distal end of article 5 (fig.

Q2b) Ree eB ee SUSInUS
Posterior border of price 5 of Pee honoda foamed of a » NONE shallow setose
lobe, its length greater than the tapered distal end of article 5 (fig. 92e).

Pareusirogenes
5. Palp of maxilla 1 short, article 1 longer than 2 (fig. 91f). . . . . Eusirella
Palp of maxilla 1 long, article 2 longer than 1 (fig. 91g)... . 6
6. Articles 4, 5, 6 of pereopods 3-5 each 1.5 times as long as omits 2 (if lees
broken proceed to couplet 9) (fig. 89a)... . dl
Articles 4, 5, 6 of pereopods 3-5 each as long as aheie poco preteen Gt
SOO) ee ee Ea SSA NMRA ENE US IanC Rr Si li OU ae OR an SU se nee)
7. Gnathopods not pubenerire RO en ie aN pence mh aga ey ALCL GO
Gnathopods subchelate .. . sea CES

8. Body carinate (often weakly), coxa 4 Dodie’ Aarawnendl Ge 89a).
Rhachotropis
Body not carinate, coxa 1 not producedforward. .... . . . Cleonardo
9. Epistome with large anterior process (seen from lateral view) . . . . . 102

1 Pseudopontogencia Oldevig (1959) and Pontogenevides Nicholls (1938) are not sufficiently described for
inclusion in the following key.

2 See also small process of Afetewsiroides and broadly flabelliform (dorsal-ventral) but shallow process of
Chosroes and footnote 3 to couplet 31.

218

13.

14.

15.

16.

Mas

28.

U.S. NATIONAL MUSEUM BULLETIN 271

Hpistome lacking largelanterion processie. vei) co bike eu ened
Postantennal cephalic angle acutely produced. . ... .. . .. Atyloella
Postantennal cephalic angle presumably rounded. . . . . . Liouvillea
Both articles 5 and 6 of gnathopod 2 long and sublinear (fig. 92k) Se Miele
Either article 5 or 6 of gnathopod 2notlongandsublinear. ...... 14
Convex side of dactyli on pereopods 3-5 with accessory spine or spines
(fie OSC) Maw ig : Reese Nasa... Ue}
Dactyli of soreenodls 3 cl facie Dontenion EOSIN sine . . Schraderia
Pereopods 1 and 2 with dactylar spine, article 2 of first antennal peduncle
shorter than article]... .. . . . . . Djerboa
Pereopods 1 and 2 lacking dectlen g spine, _ amelie 2 of first antennal peduncle
longer than article]... . . . . Bathyschraderia
Gnathopods not subchelate (ike Pleoeneneoniies tt in Calhionide) (figs. 92d,p).
15

Gnathopods distinctly subchelate. . . . «ee hG
Probably each of articles 4-6 of syarsonronls au 5 nor fonees neva article 2.
Harpinioidella

Each of articles 4-6 of pereopods 3-5 longer than article 2. . . Harcledo
Accessory flagellum distinctly articulate, usually cylindrical, often as long
as article 1 of primary flagellum (figs. 89h,z) . . . . BER al) esi dy ALT
Accessory flagellum usually absent, when present foomed on a fused scale or
process, not articulate (figs. 89e,f,g) .... . ; . 26

Inner plate of maxilla 1 with more than four ne. seine penal lining
medial edge and apex (figs. 91d,e).
Paramoera (and Atylopsis in Calliopiidae)

Inner plate of maxilla 1 with four or fewer terminal setae (figs. 91c,g). . 18
Article 6 of gnathopod 2 twice as long as article5. ......2.2.2. «19
Article 6 of gnathopod 2 subequal to article 5inlength ....... 24
Article 6 of pereopods 3-5 subequal to or shorter than article 2... . 20
Article 6 of pereopods 3-5 longer than article2........... =. 23
Accessory flagellum as broad as long (fig. 89f) .......2.2.=.. 2i1
Accessory flagellum long, rectangular (fig. 89h) ..... . 22
Melsonedeephy, cleft) (igs Ol) eerie ne dooce (nie wee nnn ees air Necederioenn

Telson apically notched (fig. 91m).
Paramoera (and Atylopsis in Calliopiidae)

Upper lip bilobed .. . wi nachauny visti an crip a A Unocal NeMtOR Ag Ses ee OTE CO:
Upper lip rounded outa y eA ie he Hoe usss ae) UeStroides
Body ecarinate, accessory flagellum 2 darticulate . . . . . . Rhachotropis
Body smooth, accessory flagellum l-articulate . . ... . . . Cleonardo
Body. canimate Wan le Woah clas voles) Soleo seul eC LEONG ODSIS
Body smooth. . . . BINA Han eee cane Meee Gy ys) PAD)
Article 6 of Paainonods narrower 3 dhvam fifth . ..... . . Pseudomoera
Article 6 of gnathopods as broad as fifth . . . . . Accedomoera (in part)
Anteroventral cephalic angle with large tooth, coxa 1 produced forward to
anterior edge of eye (fig. 93b) ... . . . . . . Rozinante
Anteroventral cephalic angle with small ‘root or none, coxa 1 not pro-
ducediats once WU es ane oiten POU
Inner plate of eile 1 it tour or fear avast sone Mieaiicrb eo" cs Ze:
Inner plate of maxilla 1 with more than four setae and some setae lining
medial edge (fig. 9le) . . . BENET GE OY sig EAE oR

Articles 4, 5, and 6 of nrorasoods 3- 5 cera 15 ines as long as article 2
Harcledo

Articles 4, 5, and 6 of pereopods 3-5 not longer than article2. .. . . 29

MARINE GAMMARIDEAN AMPHIPODA P19

29. Gnathopods simple (like Harpinioides in Calliopiidae) (figs. 92d,p).

Harpinioidella

Gnathopods subchelate .. . so BW

30. Outer lobes of lower lip broadly conamtcd: inner Nlobes farce (Ge 901), telson
extending three fourths along rami of uropod 3... . . . Meteusiroides

Outer lobes of lower lip with small medial gap, inner lobes obsolescent or
absent (fig. 90g), telson extending one third or less along rami of uropod

31. Inner lobes of lower lip small but distinct, inner plate of maxilla 2 with
poiae lining medial edge, upper lip rounded or truncate below, gnathopod
astavalliliman ie a sie ... . . Pontogeneia 3

Inner lobes of lower iD Apeent inner yale on jedi & with one medioter-
minal seta, upper lip with small incision on ventral margin, gnathopod 2

very large (like Rhachotropis) Ane . . . . . Dautzenbergia

32. Dactyls of pereopods 3 and 4 as long as erncle 6, pereopods very setose and
oedicerotid-like (fig. 93g) ... . .. . . . Zaramilla
Dactyls of pereopods 3 and 4 very shorts nereopode anon setose, slender . 33

SORE loweripnwivasimmer lObesiey kl ke he Rn eA ela nie
Wowerliplackingimner lobes) 394 4%) = 2 =. 2) ee 3 enone

34. Coxa 4 not excavate posteriorly. ............ prostebbineia
Coxamtvexcavate POSteLlOnlyamea mae iis (chee is til, alien BT vy re MGURE oD

Se bodyaumibbonate (ie 90G)ea.h feswer es eee oc on. ae. Burnymera
Body not umbonate. . . . Pera Seles 0)

36. Inner lobes of lower lip very ene inner alate a soneallle 1 oath setae mostly
eT MMII Aall ess ey eens 5 .. . . . Pontogeneia

Inner lobes of lower Ep tere: inner plate of oceaill 1 with setae lining full
medialedge .... ... . . Pontogeneiella

37. Coxa 4 deeply ae agosto, upper 2 iti rounded below, telson cleft
halfway, maxillipedal palp artiele 4 shorter than article3 . . . . Bovallia

Coxa 4 scarcely excavate posteriorly, upper lip slightly incised, telson cleft
one fifth, maxillipedal palp article 4 longer than article 3.
(theoretical possibility) Dautzenbergia

3 Also see Chosroes decoratus K. H. Barnard (1932), which is related to the type, C. incisus (Calliopiidae),
but which has a cleft telson and so inadvertently keys to Pontogeneia.

Genera of Eusiridae

Nore: ‘“‘eustrip” gnathopods have article 6 as broad as long and
attached to the produced apex of article 5.

Accedomoera J. L. Barnard
Accedomoera J. L. Barnard, 1964c.

Type-species: Pontogeneia tricuspidata Gurjanova, 1938 (original
designation).

Accessory flagellum articulate, short; lower lip with small inner
lobes; inner plate of maxilla 1 with four or fewer terminal setae;
enathopods not eusirid, subchelate, article 5 not lobate, shorter than
6, 6 as broad as 5 (Pseudomoera); pereopods 3-5 with each of articles
4-6 not distinctly longer than article 2. Species: 3, subantarctic,
boreal, N.W. Pacific, littoral.

285-135 O - 69 - 15

NATIONAL MUSEUM BULLETIN 271

220 U.S.

a (|
LY
a
=,
ea *y
R 7h, F f
f 4
i \ fill 7 of
Ww # qf Ae
1i\ D MEY
Z\\’ fh ‘
_—, —A

{ ff :
(AU =
Cal) Re

» i
oA : \ \
: »

=
SSI L/
ome ee

BZ

Y
(\
\\
ih
i

k
m

Maxilla 2: h, Eusirus; 1,

Ficure 91.—Eusiridae: a, Pontogeneia inermis (Krdyer) (Sars, 1895, pl. 159); b, Cleonardo
appendiculatus (Sars, 1885). Maxilla 1: c, Eusirus propinquus Sars (Sars, 1895, pl. 147);
d, Eurymera monticulosa Pfeffer (Chevreux, 1906c); ¢, Pontogeneia; f, Eusirella elegans
Telson: 1, Pontogeneia; m, Ponto-

Chevreux (1908b); g, Pseudomoera gabrieli (Sayce, 1901).

Maxillipeds: 7, Pontogeneia; k, Eusirus.
geneoides abyssi Nicholls (1938); n, Eusirus minutus Sars (1895, pl. 147); 0, Eusirus

Pontogeneia.
propinquus; p, Rozinante fragilis Goés (Gurjanova, 1951); g, Liouvillea oculata Chevreux

(1912b).

MARINE GAMMARIDEAN AMPHIPODA 221

n

Ficure 92.—Eusiridae: Gnathopod 1: a, Bovallia gigantea Pfeffer (Chevreux 1906c); b,
Eusirus propinquus Sars (1895, pl. 147); c, Djerboa furcipes Chevreux (1906c); d, Harcledo
plumipes (Birstein and Vinogradov, 1955); e, Pareusirogenes carinatus Birstein and
Vinogradov (1955); f, Rhachotropis aculeata (Lepechin) (Sars, 1895, pl. 149); g, Pontogeneia
inermis (Kroyer) (Sars, 1895, pl. 159); h, Eusiropsis riisei Stebbing (1897). Gnathopod
2: 1, Bovallia; j, Eusirus; k, Djerboa; 1, Rozinante fragilis (Goés) (Gurjanova, 1951); m,
Rhachotropis; n, Pontogeneoides abyssi Nicholls (1938); 0, Pontogeneia; p, Harcledo.

222 U.S. NATIONAL MUSEUM BULLETIN 271

Atyloella Schellenberg
Atyloella Schellenberg, 1929a.

Type-species: Atylopsis magellanicus Stebbing, 1888 (monotypy).

Accessory flagellum articulate, short; lower lip with inner lobes;
inner plate of maxilla 1 with four or five terminal and medial setae;
gnathopods not eusirid, subchelate, article 5 not lobate, shorter than
6; pereopods 3-5 with each of articles 4-6 not longer than article 2;
postantennal cephalic angle acutely produced (Laouvillea) ; epistome unth
anterior process.* Species: 3, antarctic, littoral (to 441 m).

*See footnote with Meteusiroides (p. 227).

Bathyschraderia Dahl
Bathyschraderia Dahl, 1959.

Type-species: B. magnifica Dahl, 1959 (monotypy).

Accessory flagellum very short but articulate; lower lip lacking
inner lobes; inner plate of maxilla 1 densely furnished with terminal
and medial setae; gnathopods not eusirid, scarcely subchelate, both
articles 5 and 6 of both gnathopods elongate; [pereopods 3-5 not
described for lengths of articles]; pereopods 3-5 with a row of spines
on convex side of dactyli, not present on pereopods 1-2 (combining
character and Djerboa). Species: 1, Kermadec Trench, hadal.

Bovallia Pfeffer
Bovallia Pfeffer, 1888.—Stebbing, 1906.

Type-species: B. gigantea Pfeffer, 1888 (monotypy). See Chev-
reux, 1906b.

Accessory flagellum absent; lower lip lacking inner lobes; inner
plate of maxilla 1 with numerous terminal setae and setae partially —
lining medial edge; gnathopods not eusirid, subchelate, article 5
strongly lobate, shorter than article 6; pereopods 3-5 with articles
4-6 each not longer than article 2; coxa 4 deeply excavate posteriorly,
upper lip rounded ventrally; telson cleft halfway; mazillipedal palp
article 4 shorter than article 3 (Dautzenbergia). Species: 1, antarctic,
littoral.

Cleonardo Stebbing
Cleonardo Stebbing, 1888; 1906.

Type-species: C. longipes Stebbing, 1888 (monotypy).

Accessory flagellum articulate, short; lower lip with inner lobes;
inner plate of maxilla 1 with less than four terminal setae; gnathopods
not eusirid, scarcely subchelate, article 5 strongly lobate, shorter than

MARINE GAMMARIDEAN AMPHIPODA Dae

article 6; pereopods 3-5 with articles 4-6 each much longer than
article 2; body dorsally smooth (Rhachotropis). Species: 7, N. Hemi-
sphere largely, bathyal-abyssal, pelagic or ?demersal.

Cleonardopsis K. H. Barnard

Cleonardopsis K. H. Barnard, 1916.
Amathillopleustes Pirlot, 1934.

Type-species: C. carinata K. H. Barnard, 1916 (monotypy).

Accessory flagellum articulate and long; lower lip with broad
semifused inner lobes; maxilla 1 with four or fewer terminal setae;
gnathopods not eusirid, subchelate, article 5 lobate, as long as article
6; pereopods 3-5 with articles 4-6 each not longer than article 2;
body carinate (Pseudomoera). Possibly belongs in the Pleustidae
because of the lower lip. Species: 1, probably cosmopolitan,
bathypelagic.

Dautzenbergia Chevreux
Dautzenbergia Chevreux, 1900.

Type-species: Amphithopsis grandimana _ Chevreux, 1887
(monotypy).

Accessory flagellum absent; lower lip lacking inner lobes; inner
plate of maxilla 1 with four or fewer terminal setae; gnathopods
not eusirid, subchelate, article 5 of gnathopod 2 with slender pos-
terior lobe, article 6 in male very large, about 5 times as long as
article 5; pereopods 3-5 with articles 4-6 each not longer than article
2; coxa 4 scarcely excavate posteriorly, upper lip slightly incised, telson
cleft one fifth, maxillipedal palp article 4 longer than article 3 (Bovallia).
Species: 3, subarctic, N. Atlantic to ?Indian Ocean, bathyal to abyssal.

Djerboa Chevreux
Djerboa Chevreux, 1906b.

Type-species: D. furcipes Chevreux, 1906b (monotypy).

Accessory flagellum articulate, long; lower lip lacking inner lobes;
inner plate of maxilla 1 densely lined with setae on medial edge;
genathopods not eusirid, subchelate, article 5 not lobate, on gnathopod
2 especially, both articles 5 and 6 greatly elongated and sublinear;
pereopods 3-5 with articles 4-6 each not longer than article 2;
pereopods 1-5 with accessory spine on conver side of dactyli (combining
character and Bathyschraderia). Species: 1, antarctic, littoral.

Eurymera Pfeffer
Eurymera Pfeffer, 1888.—Stebbing, 1906.

Type-species: E. monticulosa Pfeffer, 1888 (monotypy). See
Chevreux, 1906b.

DIA U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum absent; lower lip with inner lobes; inner plate
of maxilla 1 densely lined with setae on medial edge; gnathopods
not eusirid, subchelate, article 4 not lobate, slightly shorter than 6;
pereopods 3-5 with articles 4-6 each not longer than article 2; body
umbonate. Species: 1, antarctic, littoral.

Ficure 93.—Eusiridae: a, Djerboa furcipes Chevreux (1906c); b, Rozinante fragilis Goés
(Shoemaker, 1930). Uropod 3: c, Eusirus propinquus Sars (1895, pl. 147); d, Eusirus

minutus Sars (1895, pl. 147). End of pereopod 3: e, Djerboa. Right coxa 4: f, Prosteb-
bingia gracilis (Chevreux, 1906c). Pereopod 5: g, Zaramulla kergueleni Stebbing (1888).

MARINE GAMMARIDEAN AMPHIPODA 225

Eusirella Chevreux

Eusirella Chevreux, 1908b.

Type-species: LE. elegans Chevreux, 1908b (original designation).

Accessory flagellum absent; mandibular molar not triturative; [lower
lip unknown]; palp of mazilla 1 short, article 1 much longer than article
2; gnathopods not eusirid, scarcely subchelate but article 6 not
fully linear, article 5 not or scarcely lobate, shorter than article 6;
pereopods 1 and 2 with article 6 bearing long plumose posterior setae;
pereopods 3-5 with articles 4-6 each longer than article 2. Species:
4, cosmopolitan, bathypelagic.

Eusirogenes Stebbing
Eustrogenes Stebbing, 1904.

Type-species: £. dolichocarpus Stebbing, 1904 (monotypy).

Accessory flagellum articulate, very long; lower lip with inner
lobes; inner plate of maxilla 1 with four or fewer terminal setae;
gnathopods eusirid, subchelate, articles 4-6 each not longer than
article 2; palp article 3 of mandible shorter than article 2 (Eusirus,
Pareusirogenes); antennae lacking calceoli (Eusiropsis); article 6 of
gnathopod 1 larger than that of gnathopod 2. Species: 5, cosmopolitan
(N. Hemisphere), bathypelagic.

Eusiroides Stebbing
Eusiroides Stebbing, 1888; 1906.

Type-species: Atylus monoculoides Haswell, 1880c (indicated by
Della Valle, 1893, through erroneous synonymy and thus by synonymy
of Stebbing, 1906, but firmly selected by Chevreux and Fage, 1925).

Accessory flagellum articulate, of medium size; lower lip with small
inner lobes; inner plate of maxilla 1 with four or fewer terminal setae;
gnathopods not eusirid, subchelate, article 5 lobate, shorter than
article 6, palms armed with very stout spines; pereopods 3-5 with
articles 4-6 each not longer than article 2. Species: ca. 6, ?cosmo-
politan, littoral to ?bathyal.

Eusiropsis Stebbing
Eustropsis Stebbing, 1897; 1906.

Type-species: LE. riisei Stebbing, 1897 (monotypy).

Accessory flagellum articulate, minute; mandibular molar not trit-
urative; lower lip with inner lobes; inner plate of maxilla 1 with four
or fewer terminal setae; gnathopods eusirid, articles 4-6 touching;
pereopods 3-5 with articles 4-6 each much longer than article 2;
antennae with calceoli (Eusirus, Eusirogenes, Pareusirogenes). Species:
1,S. Hemisphere (?cosmopolitan), bathypelagic.

226 U.S. NATIONAL MUSEUM BULLETIN 271

Eusirus Kréyer
Eusirus Kr¢yer, 1845.—Stebbing, 1906.

Type-species: EE. cuspidatus Kroyer, 1845 (monotypy). See Sars,
1895.

Accessory flagellum articulate, of medium size; lower lip with
inner lobes; inner plate of maxilla 1 with four or fewer terminal
setae; gnathopods eusirid, articles 4-6 not touching; pereopods 3-5
with articles 4-6 each not longer than article 2; antennal calceoh
absent (Eusiropsis); posterior border of gnathopod article 5 forming a
projecting lobe much narrower than length of tapered distal end of article
5 (Pareusirogenes). Species: 17, bipolar and tropical submergent,
primarily bathyal to abyssal.

Harcledo J. L. Barnard
Harcledo J. L. Barnard, 1964c.

Type-species: Meteusiroides plumipes Birstein and Vinogradov,
1955 (original designation). Birstein and Vinogradov (1964) synony-
mized M. plumipes with M. curvidactyla (Pirlot, 1934).

Accessory flagellum presumably absent; lower lip unknown, but
presumably with inner lobes; inner plate of maxilla 1 with four or
fewer terminal setae; gnathopods not eusirid, not distinctly sub-
chelate, article 6 tapering and with article 7 folded back on its setose
posterior edge, article 5 slightly lobate and shorter than 6; pereopods
3-5 with articles 4-6 each much longer than article 2. Species: 1,
N. W. Pacific, ?3500 m (has eyes), possibly epipelagic.

Harpinioidella Schellenberg
Harpinioidella Schellenberg, 1926a.

Type-species: H. fissicauda Schellenberg, 1926a (monotypy).

[Accessory flagellum unknown]; lower lip with inner lobes; inner
plate of maxilla 1 with four or fewer terminal setae; gnathopods not
eusirid, simple, article 6 long, bent distally, article 5 much shorter
than 6, apparently lobate; [pereopods 3-5 unknown]. Species: 1,
antarctic to S. Atlantic, bathyal to ?abyssal (4,898 m), possibly
bathypelagic.

Liouvillea Chevreux
Liouvillea Chevreux, 1911c; 1912a; 1912b.

Type-species: L. oculata Chevreux, 1912a (designated by Chevreux) ;
1912b.

Accessory flagellum articulate, short; lower lip with inner lobes;
inner plate of maxilla 1 densely lined with setae on medial edge;

MARINE GAMMARIDEAN AMPHIPODA 237

gnathopods not eusirid, subchelate, article 5 nearly as long as 6, not
lobate; pereopods 3-5 with articles 4-6 each not longer than article
2: postantennal cephalic angle presumably rounded (Atyloella); rostrum
reaching end of article 1 of antenna 1 (Paramoera). Species: 1, antarctic,
littoral.

Meteusiroides Pirlot

Meteusiroides Pirlot, 1934.

Type-species: MM. keyensis Pirlot, 1934 (original designation).

Accessory flagellum absent; lower lip with large inner lobes, outer
lobes very widely spread (Pontogeneia)*; inner plate of maxilla 1 with
four or fewer terminal setae; gnathopods not eusirid, subchelate,
article 5 lobate, shorter than article 6; pereopods 3-5 with articles
4-6 each not longer than article 2. Species: 2, Indonesia, N. Atlantic,
bathyal to abyssal.

*Apparently inner lobes shown by Pirlot (1934), are horizontal basal processes
unnaturally flattened and outer lobes therefore excessively spread; small inner
lobes present as in Pontogeneia; information from antarctic individual of Mete-
usiroides (possible new species in colls. of El Tanin expeds.). Meteusiroides has
telson extending 3/4 along rami of uropod 3, whereas telson of Pontcgeneza extends
only 1/3 along rami of uropod 3. Also Meteusiriodes has small adze-shaped process
on epistome and might be confused with Atyloella.

Paramoera Miers

Paramoera Miers, 1875.—Stebbing, 1906 (in part, not Atylozdes Stebbing, 1888).—
Schellenberg, 1929a.—K. H. Barnard, 1932.

Stebbingza Pfeffer, 1888.

Aucklandia Walker, 1908.

Type-species: Amphithoe fissicauda Dana, 1852a (monotypy and
subsequent synonymy). See Schellenberg, 1931; Atyloides australis
and A. assimilis Stebbing, 1888.

Accessory flagellum articulate, of medium length; lower lip with
inner lobes indistinct or absent; inner plate of maxilla 1 densely
setose along medial edge; gnathopods not eusirid, subchelate, article 5
not lobate, shorter than article 6; pereopods 3-5 with articles 4-6 each
not longer than article 2; rostrum inconspicuous (Liouvillea). Species:
28, bipolar, coldwater, littoral.

Pareusirogenes Birstein and Vinogradov
Pareusirogenes Birstein and Vinogradov, 1955.
~Type-species: P. carinatus Birstein and Vinogradov, 1955 (original
designation).

Accessory flagellum absent; lower lip with inner lobes; inner plate
of maxilla 1 with four or fewer terminal setae, hairy along medial edge;

228 U.S. NATIONAL MUSEUM BULLETIN 271

enathopods eusirid, articles 4-6 not touching; pereopods 3-5 with
articles 4-6 each much longer than article 2; antennal calceoli absent
(Eusiropsis); posterior border of gnathopodal article 5 formed of a broad,
setose lobe, its width greater than tapered distal end of article 5 (Eusirus).
Species: 1, N.W. Pacific, abyssopelagic.

Pontogeneia Boeck
Pontogeneia Boeck, 1871.—Stebbing, 1906.

Type-species: Amphithoe inermis Krgyer, 1838 (monotypy). See
Sars, 1895.

Accessory flagellum absent; lower lp with small inner lobes;
inner plate of maxilla 1 with four or more terminal or subterminal
setae; gnathopods not eusirid, article 5 not lobate, sightly longer
than article 6; pereopods 3-5 with articles 4-6 each not longer than
article 2, article 4 slender (Zaramilla); outer lobes of lower lip not
broadly separated (Meteusiroides). Species: 25, cosmopolitan, littoral.

Pontogeneiella Schellenberg
Pontogeneiella Schellenberg, 1929a.

Type-species: Atyloides brevicornis Chevreux, 1906a, 1906b (pres-
ent selection).

Accessory flagellum absent; lower lip with inner lobes; inner
plate of maxilla 1 densely lined with setae on medial edge; gnathopods
not eusirid, subchelate, article 5 not lobate, nearly as long as article
6; pereopods 3-5 with articles 4-6 each not longer than article 2,
article 4 one third as wide as article 2 (Zaramilla); antennae subequal,
coxa 4 strongly excavate posteriorly (Prostebbingia). Species: 2,
antarctic, littoral.

Pontogeneoides Nicholls
Pontogeneoides Nicholls, 1938.

Type-species: P. abyssi Nicholls, 1938 (original designation).

Accessory flagellum articulate, short; lower lip with small inner
lobes; inner plate of maxilla 1 with four or fewer terminal setae;
gnathopods not eusirid, subchelate, article 5 lobate on gnathopod
2, shorter than article 6, pereopods unknown; telson minutely notched.
Species: 2, antarctic, bathyal (480—-1,590 m).

Prostebbingia Schellenberg

Prostebbingia Schellenberg, 1926a.

Type-species: Stebbingia gracilis Chevreux, 1912a, 1912b (present
selection).

MARINE GAMMARIDEAN AMPHIPODA 929

Accessory flagellum absent; lower lip with inner lobes; inner
plate of maxilla 1 densely lined with setae on medial edge; gnathopods
not eusirid, subchelate, article 5 not lobate, slightly shorter than
article 6; pereopods 3-5 with articles 4-6 each not longer than
article 2; antenna 1 longer than antenna 2, coxa 4 scarcely excavate
posteriorly (combining character). Species: 2, antarctic, littoral (to
400 m).

Pseudomoera Schellenberg

Pseudomoera Schellenberg, 1929a.

Type-species: Atyloides gabrieli Sayce, 1901 (monotypy).

Accessory flagellum articulate, of medium length; lower lip with
inner lobes; inner plate of maxilla 1 with four or fewer terminal setae;
gnathopods not eusirid, subchelate, article 5 lobate, equal to article
6 in length, 5 slightly broader than 6 (Accedomoera); pereopods 3-5
with articles 4-6 each not longer than article 2; body smooth (Cleo-
nardopsis). Species: 1, Australia, freshwater.

Pseudopontogeneia Oldevig
Pseudopontogeneia Oldevig, 1959.

Type-species: P. intermedia Oldevig, 1959 (monotypy).

[Accessory flagellum and lower lip not described]; inner plate of
maxilla 1 with four or fewer terminal setae; gnathopods [not distinctly
described]; article 6 longer than 5; pereopods 3-5 with articles 4-6
each not longer than article 2. Species: 1, Bering Island, littoral.

Rhachotropis Smith

Tritropis Boeck, 1871 (homonym, Reptilia).
Rhachotropis Smith, 1883 (new name).—Stebbing, 1906.—Shoemaker, 1930.
Gracilipes Holmes, 1908.

Type-species: Oniscus aculeatus Lepechin, 1780 (monotypy). See
Sars, 1895.

Accessory flagellum 2-articulate, short; lower lip with small inner
lobes; inner plate of maxilla 1 with four or fewer terminal setae;
gnathopods not eusirid, subchelate, article 5 lobate, shorter than
article 6; pereopods 3-5 with articles 4-6 each longer than article 2;
body carinate or toothed (Cleonardo). Species: 30, bipolar, tropical
submergents, primarily bathyal-abyssal.

Ronco J. L. Barnard
Ronco J. L. Barnard, 1965.
Type-species: R. sosa J. L. Barnard, 1965 (original designation).

230 U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum articulate, of medium length; upper lip bilobate;
lower lip with inner lobes; inner plate of maxilla 1 with four or fewer
terminal setae; gnathopods not eusirid, subchelate, article 5 lobate,
shorter than article 6; pereopods 3-5 with articles 4-6 each not longer
than article 2. Species: 1, Micronesia, littoral.

Rozinante Stebbing
Rozinante Stebbing, 1894; 1906.

Type-species: Paramphithoé fragilis Goés, 1866 (original desig-
nation). See Shoemaker, 1930.

Accessory flagellum absent; lower lip with inner lobes obsolete
(examined by me); inner plate of maxilla 1 with four or fewer terminal
setae; gnathopods not eusirid, subchelate, article 5 not lobate, equal
to or slightly longer than article 6; pereopods 3-5 with articles 5-6
(not article 4) each longer than article 2 (examined by me); antero-
ventral cephalic corner with large tooth projecting forward to end of small
rostrum, coxa 1 produced forward to anterior edge of eye. Species: 1,
arctic-boreal, N. Atlantic, littoral (to 372 m).

Schraderia Pfeffer

Schraderia Pfeffer, 1888.—Stebbing, 1906.—K. H. Barnard, 1932.
Atyloides Stebbing, 1888.

Type-species: SS. gracilis Pfeffer, 1888 (monotypy).

Accessory flagellum articulate, of medium length; lower lip lacking
inner lobes; inner plate of maxilla 1 densely lined with setae along
medial edge; gnathopods not eusirid, subchelate, article 5 not lobate,
equal to article 6 in length, in gnathopod 2 especially articles 5-6
greatly elongate and sublinear; pereopods 3-5 with articles 4-6 each
not longer than article 2. Species: 2, antarctic, littoral.

Zaramilla Stebbing
Zaramilla Stebbing, 1888; 1906.

Type-species: Z. kergueleni Stebbing, 1888 (monotypy).

Accessory flagellum absent; lower lip with scarcely discrete inner
lobes; inner plate of maxilla 1 densely lined with setae on medial edge;
enathopods not eusirid, subchelate, article 5 not lobate, slightly
longer than article 6; pereopods 3-5 with articles 4—6 each not longer
than article 2, article 4 broadly expanded (especially Bovallia and
Pontogeneia), article 4 two thirds as wide as article 2 (Pontogenevella) ;
antennae subequal in length, coxa 4 strongly excavate posteriorly (Pros-
tebbingia) ; dactyli of pereopods 3-4 as long as article 6, pereopods re-
sembling those of oedicerotids (combining character). Species: 1,
Kerguelen, littoral. Probably referrable to Haustoriidae and included
in that family key (p. 252); absence of accessory flagellum in contrast
to Haustoriidae.

MARINE GAMMARIDEAN AMPHIPODA 231

Marine Gammaridae
Figures 94-96

Dracnosis.—Like the basic gammaridean, from which all other
diagnoses in this paper are used to distinguish families, but accessory
flagellum occasionally reduced to one long article, maxillipedal palp
occasionally 3-articulate, gnathopod 1 rarely larger than 2, telson
occasionally uncleft, inner ramus of uropod 3 reduced, coxae 1-4
rarely of nonuniform shape, coxa 4 occasionally not posteriorly
excavate and pereopod 5 rarely elongate.

Description.—The Gammaridae are variable and many genera
form links to other families. Accessory flagellum always present but
varying from one long article to more than 20 short articles; peduncle of
antenna 1 elongate; rostrum present or absent; upper lip not incised;
mandible always bearing strong triturative molar and 3-articulate
palp; lower lips variable, never ampithoid or pleustid in structure;
maxillae fully developed, not strikingly foliaceous; palp of first maxilla
2-articulate; maxillipeds with well-developed plates, palp 3- or 4-
articulate; gnathopods usually powerful and subchelate, occasionally
slender and simple, gnathopod 1 rarely larger than 2; coxae of medium
length, occasionally shortened; coxa 4 occasionally not excavate
posteriorly; uropod 3 highly variable but rami never shorter than
peduncle (except Parapherusa, Falklandella, and Gammarella) and
usually flattened, not cylindrical, generally lanceolate, foliaceous or
subquadrate, inner ramus occasionally very short; telson not elongate
(except Parapherusa), usually deeply cleft but occasionally broader
than long, uncleft or emarginate.

Re.ationsHips.—Gammaridae are difficult to define precisely be-
cause so many genera have exceptional morphology. Four of the
genera have uncleft telsons and thus resemble Calliopiidae but all have
well-developed accessory flagella. They may also resemble some of the
isaeid-like families but their telsons are not fleshy (not thickened
dorsoventrally). The most difficult member of this quartet is Para-
pherusa Stebbing which confounds all possible simplifications made
in the keys and diagnoses of families and genera. The unique species
of this genus has a third uropod resemblnig that of ischyrocerids or
ampithoids (fig. 96d), unexcavate coxa 4, and its telson is entire but
elongate, uropod 1 has a ventrodistal peduncular tooth like that of
various ischyrocerids and isaeids. The absence of pereopodal glands
apparently demonstrates its correct assignment to the Gammaridae,
rather than to the glandular families. The other marine gammarid
genera with uncleft telson have deep posterior excavations on coxa 4
and thus cannot be confused with isaeid-like genera. One ischyrocerid,
Bathyphotis, however, has a strongly excavate coxa 4 and has a special

250, U.S. NATIONAL MUSEUM BULLETIN 271

Poy] |/K » K
j YH
—A/ i)
YZ H|= ff

Ficure 94.—Gammaridae: a, Maera lovent (Bruzelius) (Sars, 1895, pl. 182); b, Megaluropus
tlongimerus Schellenberg (J. L. Barnard, 1962b). Accessory flagella: c, Marinogam-
marus marinus (Leach) (Sars, 1895, pl. 175); d, Elasmopus rapax Costa (Sars, 1895, pl.
183). Coxa 4: e, Maera othonis (Milne Edwards) (Sars, 1895, pl. 182); f, Gammarellus

homari (Fabricius) (Sars, 1895, pl. 172).

MARINE GAMMARIDEAN AMPHIPODA 233

place in Gammaridea Family Key A (p. 109). Its uropod 3 is even more
typically that of an ischyrocerid than is uropod 3 of Parapherusa.
Another 13 genera of Gammaridae (in couplets 6-18) have vestigial
accessory flagella and cleft telsons and thus resemble Eusiridae. All
Eusiridae except Eusirogenes Stebbing have elongate telsons and that
genus is easily recognizable as a eusirid in its gnathopodal morphology.
Eusiridae also have slightly enlarged metasomal segments, the meta-
some having the length of 6 pereonal segments, whereas gammarids
have a metasome the length of only 5 pereonal segments. This is a

Ficure 95.—Gammaridae: a, Melita dentata (Krdyer) (Sars, 1895, pl. 181). Mouthparts
of Elasmopus rapax Costa (Sars, 1895, pl. 183): b, upper lip; c, lower lip; ¢, mandible;
f,g, maxillae 1, 2; h, maxilliped. Lower lip: d, Gammarellus homari (Fabricius) (Sars,
1895, pl. 172). Gnathopod 2: 1, Elasmopus; j, Cheirocratus sundevalli (Rathke) (Sars,
1895, pl. 184). Gnathopod 1: k, Elasmopus; 1, Cheirocratus.

234 U.S. NATIONAL MUSEUM BULLETIN 271

difficult character analysis to practice however, and one must use
simply a process of elimination by checking generic keys and diagnoses
in both families for genera which are not obviously gammarid or
eusirid. Five of the genera in couplets 6-18 have reduced inner rami
of uropod 3 (melita-like uropod 3) and thus are easily eliminated from
consideration as eusirids, for in the latter family, the inner ramus of
uropod 3 is always as long as or longer than the outer.

Key to the Genera of Marine Gammaridae

Only those genera of Gammaridae with marine representatives are
included in this key (except Falklandella). Some generic pairs, such as
Maeropsis-Paraceradocus and Ceradocus-Ceradocoides have not been
separated by firm distinctions. Other genera are cited twice in the
key because of their intergrading or variable structures.

One must pursue both pathways in couplet 6 if the species at hand
has three accessory flagellar articles. Since other characters are as
variable as accessory flagella and more often poorly known, couplet 6
has been found to be more satisfactory than conceivable substitutes
until some of the poorly described genera are clarified.

1.) Belson entire (fig. 96g) but often emarginate. ©7250 2 2 2°. 2.025 292
Telson cleft (fig. 96h)... . . BRENDAN NEMS" (6)

2. Urosomal segments with puonlles of Glove eines paoesommies with deep
serrations on posterodorsal margins . . . . .. . . Mesogammarus
Urosomal segments lacking dorsal bundles of spines, metasome lacking
serrations but often with carinae. .. . Le) oh Sale ea

3. Rami of uropod 3 at least 1.5 times as long as 2 pede - wos eho ee
Rami of uropod 3 equal to or shorter than peduncle ......... 5

4. Body dorsally carinate (even weakly), article 3 of mandibular palp shorter
than article 2, lower lip without inner lobes . . . . . . Gammarellus !

Body dorsally smooth, article 3 of mandibular palp longer than article 2,
lower lip with inner lobes. . .. . . . . . . Weyprechtia

5. Accessory flagellum 1-articulate, telson Beendes near long, emarginate, lower
lip without inner lobes, coxa 4 posteriorly excavate. . . Falklandella

Accessory flagellum 8-articulate or more, telson longer than broad, not
emarginate, lower e with strong inner lobes, coxa 4 not excavate pos-

teniorlye yuan a). . . . . . Parapherusa

6. Accessory flagellum a or D artrenlate! “eoeionallly Bi EGE En gy
Accessory flagellum 4- or more articulate, occasionally 3. ..... . 19

7. Inner ramus of uropod 8 scale-like and much smaller than outer (fig. 96f). 8
Inner ramus of uropod 3 nearly as large as outer . . . Mea ca ts LP

8. Apex of uropod 3 not exceeding extent of uropods 1-2 (fig. 96a) . Melitoides
Apex of uropod 3 greatly exceeding extent of uropods 1-2. ...... 9

9. One or more of coxae 1-4 very a longer (almost GO) than coxae
Mp oviiae Cony renee Aca taal: KAS gine cui VEC Ital
All coxae selocamell 3 in Tort very Short Ae i A Gay gL Ea ected a ame LO)

1 Gammaracanthus has a telson broader than long but so deeply emarginate as to be considered cleft.

10.

be

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

28.

MARINE GAMMARIDEAN AMPHIPODA 235

Maxilla 2, medial edge of inner plate lined with setae (fig. 95g), article 2 of
outer ramus of uropod 3 subequal to article 1 in length (fig. 96f) . Eriopisa
Maxilla 2, medial edge of inner plate unarmed, article 2 of outer ramus of

uropod 3 much shorter than article 1 or absent. ........ . Il
Outer ramus of uropod 3 biarticulate. ....... .... . Eriopisella
Outer ramus of uropod 3 uniarticulate. .. . . . . . Netamelita
Rami of uropod 3 foliaceous, oval, apically Poured (Gig. 96e), either coxa 2

or 3 much shorter thancoxal..... . . . . . Megaluropus
Rami of uropod 3 lanceolate, or apices Aanoete es 96b,d), not foliaceous,

coxae 2-3 aslongascoxal.. . RS ie eat aN Ne ee NAIR ee U)
Mandibular palp article 2 shorter ‘them penele ioe . . . . Parelasmopus
Mandibular palp article 2 equal to orlongerthanarticlel....... 14
Cuathopodwlysubchelatemuh maieweePikts wheel hier el Rey sees 2 LO
Gnathopod 1 simple (fig. 957). . . . DEV us wie Wie: ENE iL aie
Inner plate of maxilla 1 poorly seen (2- 5 ‘gaind) . . . . Elasmopus
Inner plate of maxilla 1 densely setose (11 setae) ... Bee OU LG
Uropod 3 strongly exceeding uropod 1, outer ramus Dearticulate:

Liagoceradocus
Uropod 3 not exceeding uropod 1, outer ramus l-articulate. Anelasmopus ”
Female gnathopod 2 simple (fig. 951). . . ... =.=. +... . Cheirocratus
Hemaleynathopod 2; sulbehelate ims 5) san yi Bem ele We ee ue GA sla Ween We)
Antennal nearly as long as;antennaiy2, 2 5... . .). 4. Gheirgecatella
Antenna 1 reaching only to end of peduncle of antenna2... . . Casco
All pereonites carinate. ... . . . . . . Gammaracanthus
Pereonites not carinate, or pecanionally aumealboe 6-7 are carinate . . . 20
Urosome with two or three segments panape bundles or rows of articulate

Spines). . ; ea urea varie
Urosome saansitonellhy fli seniierad syne nec on fll {pene and not ar-

ranged in bundles, spines usually absent . ... . spe Wausi er Geis aoe

Some segments anterior to pleonite 4 bearing dorsal prcalerte spines.
“echinogammarus” members of Anisogammarus
No segments anterior to pleonite 4 bearing dorsal articulate spines (oc-

casionally bearing teeth or serrations) .. . is er aR Cea le at 274
Metasome with posterodorsal teeth and/or someone, La MALL oc adlves De
Metasome lacking posterodorsal teeth and serrations . . bate 5 ills AD)
Telsonic apices quadrate, lobes closely appréssed and peoerted by a slit.

Maerella
Telsonic apices subacute, lobes gaping. . . MOV io niet iO

Metasomites with a single dorsoposterior feothe
‘“Marine Carinate Gammarus”

Metasomites multiserrate dorsoposteriorly . . .. . . . Mesogammarus
Gnathopod 1 slightly larger than 2, both gnathopodal palms bearing two or

more chisel-shaped spines in rows. . . . . . . . Anisogammarus
Gnathopod 1 smaller than or oe to 2, spalizes bearing one or no chisel-

shaped spines. ... . . . . . Gammarus
Inner ramus of uropod 3 sone, very “ sirocill Ge 96f) IM aed aa Set,
[nner ramus.of ureped 3 nearly,as long as outer. . 6... ...,.,. 28
Uropod 3 strongly exceeding apex of uropod1 ....... ... Melita
Uropod 3 not exceeding apex of uropod1........ .. .. Melitoides
Rami of uropod 3 foliaceous, oval, apices rounded (fig. 96e). . . . . . 29
Rami of uropod 3 lanceolate, square, or apices truncate or pointed . . . 30

2 Not clearly described. See diagnoses. Hornellia Walker resembles Anelasmopus but the inner plate of
maxilla 1 has not been described.

285-135 O - 69 - 16

236 U.S. NATIONAL MUSEUM BULLETIN 271

29. Kyes four in tae inner plate of maxilla 1 with medial edge fully lined

Walthesetaer nies cl: .. . . . . Quadrivisio

Eyes two in number; inner wallet oi snes, 1 Ruin only terminal setae.
Paraceradocus
30. Maxilla 2, medial edge of inner plate not lined with setae ...... 383i
Maxilla 2, medial edge of inner plate lined with setae ........ 388
31. Palp of maxilliped with three articles ......... =... Maerella
Palp of maxilliped with four articles. . . . LH as ED MEET, RED eR Ny
32. Mandibular palp article 3 stout, falcate (fig. 95e) . .. . . . Elasmopus
Mandibular palp article 3 slender, not faleate. . . . . . . . . Maera

33. Palm and posterior margin of article 6 on gnathopod 2 nee distinct from one
another (fig. 951), article 7 of gnathopod 2 nearly as long as article 6,

rami of uropod 3 equal to peduncle in length. . . . . . Gammarella
Mheseicharacters not: combined aia cee ey yer eee kee Oe A See

34. Lower lip lacking inner lobes... . BS chil oa)
Lower lip bearing inner lobes (presumptive in @oradeeories and Maera-
cunha). . Aiba eho)

35. Mandibular alo cntiiale 3 naed on meal eee! Saw of Peeailined about
75% as long as article 3, claw-shaped, coxa 4 not excavate posteriorly.

Ceradocopsis
Mandibular palp article 3 setose medially, dactyl of maxilliped 25% as long
as article 3, setose, coxa 4 excavate posteriorly ... . Bathyceradocus
36. Maxilla 1, medial edge of inner lobe not lined with setae. . ..... 37
Maxilla 1, medial edge of inner lobe lined with setae. . ...... . 38
37. Outer ramus of uropod 3 bearing minute article2 ... . . Maeracunha
Outer ramus of uropod 3 lacking article2....... . .. . Maeropsis
38:3) | Uropod 3) with’ 2-articulate outer ramus si) an eee
Uropod 3 with l-articulate outer ramus .. . OAS aan A)

39. Mandibular palp article 3 subequal to article 1, haaing only two setae.
Maeracunha

Mandibular palp article 3 slightly longer than article 1, multisetose.

Metaceradocoides
40. Male gnathopod 2 very small and slender. . ... . . . Metaceradocus
Male gnathopod 2 enlarged (fig. 95a). . . . Sian ASL
41. Mandibular palp article 3 as long as article 2, elietiily digdeshoned, article 1
simple. . . . . . Elasmopoides

Mandibular niin pcticle 3 les chen 7 5% andl penerally half or less as long
as article 2, article 1 with distomedial process or spines.
Ceradocus and Ceradocoides 4

3 Not clearly described.

(Genera of Marine Gammaridae

The term ‘‘NORMAL GNATHOPODS” indicates that both pairs of
gnathopods are subchelate and gnathopod 2 is larger than 1 in the
male.

Anelasmopus Oliveira
Anelasmopus Oliveira, 1953.

Type-species: A. kraui Oliveira, 1953 (monotypy).

MARINE GAMMARIDEAN AMPHIPODA 237

Accessory flagellum 2-articulate; lower lip [unclear], apparently
with inner lobes; inner plate of maxilla 1 densely setose (11 setae),
inner plate of maxilla 2 setose partially on medial edge; gnathopods
normal; uropod 3 short, not exceeding uropod 1, rami short, rec-
tangular, inner slightly shorter than outer, outer 1-articulate; telson
deeply cleft; body lacking dorsal carinae, teeth or spines. Species:
1, Brazil, littoral.

Anisogammarus Dershavin

Antsogammarus Dershavin, 1927.—Gurjanova, 1951.
Eogammarus Birstein, 1933 [a valid subgenus].

as

h

Figure 96.—Gammaridae: a, Gammaracanthus loricatus (Sabine) (Sars, 1895, pl. 174, as
G. relictus). Uropod 3: b, Cheirocratus sundevalli (Rathke) (Sars, 1895, pl. 184); ¢,
Melita dentata (Krgyer) (Sars, 1895, pl. 181); d, Elasmopus rapax Costa (Sars, 1895,
pl. 183); e, Gammaracanthus; f, Eriopisa elongata (Bruzelius) (Sars, 1895, pl. 181). Tel-
son: g, Gammarellus homari (Fabricius) (Sars, 1895, pl. 172); h, Elasmopus; 1,
Gammaracanthus.

238 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: Gammarus pugettensis Dana, 1853 (monotypy and
subsequent synonymy). See J. L. Barnard, 1954.

Accessory flagellum more than 3-articulate; lower lip with small
or obsolete inner lobes; inner plates of maxillae both densely setose
medially; gnathopod 1 slightly larger than 2, palms bearing many
chisel-shaped spines; uropod 3 exceeding uropod 1, outer ramus long,
2-articulate (? rarely 1-articulate), inner ramus long or very short;
telson deeply cleft; urosome and occasionally metasome armed with
dorsal bundles of spines, except one species [type of Anzsogammarus]
having a tooth on urosomite 2 instead of spines. Species: 18, boreal
Pacific, littoral and freshwater. Includes former marine members
of Echinogammarus. Possibly this genus divisible into 4 subgenera.

Bathyceradocus Pirlot

Bathyceradocus Pirlot, 1934

Type-species: B. stephenseni Pirlot, 1934 (original designation).

Accessory flagellum multiarticulate; lower lip lacking inner lobes;
inner plates of maxillae 1-2 densely setose medially; gnathopods
small, subchelate, slender; uropod 3 exceeding uropod 1, rami of me-
dium length, subequal, broadly lanceolate, outer 1-articulate; telson
deeply cleft; metasome and urosome toothed and carinate, urosome
with scattered dorsal spines; palp article 4 of maxilliped short, blunt,
apically setose (Metaceradocoides). Species: 1, Indo-Pacific, bathyal-
abyssal (1,165-4,930 m).

Casco Shoemaker

Casco Shoemaker, 1930.

Type-species: Cheirocratus bigelowi Blake, 1929 (original desig-
nation).

Accessory flagellum 2-articulate; lower lip with inner lobes; inner
plates of maxillae 1-2 densely setose medially; gnathopod 1 essentially
simple, but dactyl large and folded back on article 6, gnathopod 2
slightly larger than 1 and slightly subchelate; uropod 3 exceeding
uropod 1, rami subequal, long, lanceolate, outer 1-articulate; telson
deeply cleft; urosome with a tooth and scattered spines; antenna 1
reaching only to end of peduncle of antenna 2 (Cheirocratella); antero-
ventral corner of head strongly produced. Species: 1, boreal W. Atlantic,
littoral.

Ceradocoides Nicholls

Ceradocoides Nicholls, 1938.
Type-species: ©. chiltoni Nicholls, 1938 (original designation).

MARINE GAMMARIDEAN AMPHIPODA 239

Accessory flagellum more than 4-articulate; [lower lip unknown];
inner plates of maxillae 1-2 densely setose medially; gnathopods
apparently normal; uropod 3 not exceeding uropod 1, rami equal,
lanceolate [outer ramus articles not described]; telson cleft shortly;
hody lacking dorsal teeth or spines (distinctions from Ceradocus by
Nicholls, 1938, unclear). Species: 1, antarctic (420 m).

Ceradocopsis Schellenberg
Ceradocopsis Schellenberg, 1926a.

Type-species: C. kergueleni Schellenberg, 1926a (monotypy).

Accessory flagellum more than 4-articulate; lower lip lacking inner
lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods
normal; uropod 3 not exceeding uropod 1, rami equal, short, outer
2-articulate; telson deeply cleft; body lacking dorsal teeth or spines.
Species: 1, Kerguelen Island, littoral.

Ceradocus Costa
Ceradocus Costa, 1853c, 1857.—Stebbing, 1906.

Type-species: C. orchestiipes Costa, 1853c (monotypy). See
Chevreux and Fage, 1925.

Accessory flagellum 3- or more articulate; lower lip with inner
lobes; inner plate of maxilla 1 densely setose medially, of maxilla
2 moderately to strongly setose medially; gnathopods normal; uropod
3 exceeding uropod 1, rami equal, lanceolate, broad or slender, outer
l-articulate; telson deeply cleft; palp article 3 of mandible half as
long as article 2 (Metaceradocus, Elasmopoides), article 1 of palp
with medial process or spines or cusp. Species: 18, cosmopolitan,
littoral (and 3 bathyal).

Cheirocratella Stephensen
Chetrocratella Stephensen, 1940.

Type-species: C. thori Stephensen, 1940 (original designation).

Accessory flagellum 2-articulate; [lower lip unknown]; inner plates
of maxillae 1-2 apparently densely setose medially; gnathopod 1
simple; gnathopod 2 larger and subchelate (only female is known);
[uropod 3 unknown]; telson deeply cleft; metasomal and urosomal
segments dorsally toothed, urosome with scattered spines; antennae
subequal in length (Casco). Species: 1, Iceland, littoral.

Cheirocratus Norman
Chetrocratus Norman, 1867a.—Stebbing, 1906.

Type-species: Gammarus assimilis Liljeborg, 1851 (monotypy
and subsequent synonymy). See Sars, 1895.

240 U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum 2-, occasionally 3-articulate; lower lip with
inner lobes; inner plates of maxillae 1-2 densely setose medially;
female gnathopods simple, slender, male gnathopod 1 simple, male
gnathopod 2 powerfully subchelate; uropod 3 exceeding uropod 1,
rami equal, lanceolate, outer 1l-articulate; telson deeply cleft; uro-
some with dorsal teeth and setae. Species: 4, boreal N.E. Atlantic,
littoral.

Elasmopoides Stebbing

Elasmopoides Stebbing, 1908b.

Type-species: E. chevreuxi Stebbing, 1908b (monotypy).

Accessory flagellum exceeding 20 articles; lower lip with inner lobes;
inner plate of maxilla 1 densely setose medially, of maxilla 2 moder-
ately to strongly setose medially; gnathopods normal; uropod 3 not
exceeding uropod 1, rami equal, broad, outer 1-articulate; telson
deeply cleft, lobes separated basally; palp article 3 of mandible as
long as article 2 and slightly sickle-shaped; article 1 simple (Ceradocus).
Species: 1, S. Africa, sublittoral.

Elasmopus Costa

Elasmopus Costa, 18538c, 1857.—Stebbing, 1906.
Neogammaropsis Stout, 1913.

Type-species: EH. rapax Costa, 1853c (monotypy). See Sars;
1895; Chevreux and Fage, 1925; J. L. Barnard, 1962b.

Accessory flagellum 3- or more, occasionally 2-articulate; lower
lip with inner lobes; inner plates of maxillae 1-2 with only terminal
setae; gnathopods normal; uropod 3 variable in length, rami equal,
rectangular, outer l1-articulate; telson deeply cleft; urosome occasionally
with dorsal teeth; palp article 3 of mandible falcate, stout (Maera).
Species: 35, tropico-warm-temperate, cosmopolitan, littoral (one
bathyal record).

Eriopisa Stebbing

Eriopis Bruzelius, 1859 (homonym, Coleoptera).
Eriopisa Stebbing, 1890 (new name); 1906.

Type-species: riopis elongata Bruzelius, 1859 (monotypy). See
Sars, 1895.

Accessory flagellum 2-articulate; lower lip with slight inner lobes;
inner plates of maxillae 1-2 densely setose medially; gnathopods
normal; uropod 8 greatly exceeding uropod 1, immensely elongate,
outer ramus with two very long, subequal articles, or article 2 oc-
casionally short, inner ramus very short, scale-like; telson deeply
cleft; body lacking dorsal teeth or spines. Species: 6, cosmopolitan,
littoral to bathyal (752 m).

MARINE GAMMARIDEAN AMPHIPODA 241

Eriopisella Chevreux

Eriopisella Chevreux, 1920.

Type-species: EH. pusilla Chevreux, 1920 (monotypy).

Accessory flagellum 2- articulate or less; lower lip with scarcely
defined inner lobes fused with outer lobes; inner plates of maxillae
1-2 setose only terminally; gnathopods small, subchelate, equal in
size; uropod 3 greatly exceeding uropod 1, immensely elongate, outer
ramus 2-articulate, article 2 much shorter than article 1, inner ramus
very short, scale-like; telson deeply cleft; body lacking dorsal teeth
or spines. Species: 3, France, Seychelles, India, littoral.

Falklandella Schellenberg

Falklandella Schellenberg, 1930, 1931.

Type-species: FF. obtusa Schellenberg, 1930, 1931 (monotypy).

Accessory flagellum 1-articulate; lower lip lacking inner lobes;
inner plates of maxillae 1-2 densely setose medially; gnathopod 1
larger than gnathopod 2, subchelate; uropod 3 short, not exceeding
uropod 1, rami semirectangular, outer l-articulate, inner half as long
as outer; telson short, emarginate; body lacking dorsal teeth or
spines. Species: 2, Falkland Islands, freshwater close to sea.

Gammaracanthus Bate

Gammaracanthus Bate, 1862.—Stebbing, 1906.

Type-species: Gammarus loricatus Sabine, 1821 (original designa-
tion!, Bate, 1862, p. 203). See Gurjanova, 1951.

Accessory flagellum 3- or more articulate; lower lip with inner
lobes obsolescent; inner plate of maxilla 1 setose only terminally, of
maxilla 2 densely setose medially; gnathopods large, nearly equal in
size, nearly eusirid in structure (fig. 96a); uropod 3 slightly exceeding
uropod 1, rami subequal, foliaceous or subfoliaceous; outer 1-artic-
ulate; telson very short, cleft, lobes gaping; body carinate through-
out. Species: 1, arctic and Caspian Sea, littoral.

Gammarella Bate, new synonymy

Pherusa Leach, 1814a.—Stebbing, 1906 (homonym, Polychaeta).
Gammarella Bate, 1857a.
Pherusana J. L. Barnard, 1964c (new name for Pherusa).

Type-species: Pherusa fucicola Leach, 1814a (monotypy and
subsequent synonymy). See Walker, 1891la, b; Chevreux and
Fage, 1925.

Accessory flagellum 4- or more articulate; lower lip with inner
lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods

242 U.S. NATIONAL MUSEUM BULLETIN 271

normal; uropod 3 short, not exceeding uropod 1, rami not longer
than peduncle, outer l-articulate, inner slightly shortened; telson
cleft; urosome dorsally carinate; palm and posterior margin of article
6 on gnathopod 2 not separate, article 7 nearly as long as article 6 (com-
bination of characters). Species: 1, subtropical E. Atlantic and
Mediterranean, littoral.

Gammarellus Herbst

Gammarellus Herbst, 1793.—Stebbing, 1906.

Amathia Rathke, 1837 (homonvm, Hydrozoa).

Grayia Bate, 1862.—Bate and Westwood, 1863.

Amathilla Bate and Westwood, 1863 (new name for Amathza).

Type-species: <Astacus homari J. C. Fabricius, 1779 (type indicated
by Stebbing, 1899a, but firmly selected by Chevreux and Fage,
1925). See Sars, 1895.

Accessory flagellum more than 3-articulate; lower lip lacking inner
lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods
normal; uropod 3 short, not exceeding uropod 1, rami lanceolate,
outer l-articulate; telson entire or emarginate; body dorsally carinate.
Species: 2, arctic and high boreal, littoral.

Gammarus Fabricius

Gammarus Fabricius, 1775.—Stebbing, 1906.

Marinogammarus Schellenberg, 1937a—Sexton and Spooner, 1940 [subgenus,
often considered a full genus].

Pectenogammarus Reid, 1940 [valid subgenus].

Type-species: Cancer locusta Linné, 1758 (selected by Boeck, 1876).
See Sars, 1895.

Accessory flagellum 3- or more articulate; lower lip with inner
lobes obsolescent or absent; inner plates of maxillae 1-2 densely
setose medially; gnathopods normal or gnathopod 1 slightly larger
than 2; uropod 3 exceeding uropod 1, outer ramus 2-articulate, inner
variable, long or short; telson deeply cleft; no dorsal teeth or carinae,
urosome with dorsal spine-bundles. Marine and brackish species:
13-15, arctic (boreal in Atlantic), littoral. See ‘“Marine Carinate
Gammarus” (p. 244).

Hornellia Walker

Hornellia Walker, 1904.

Type-species: H. incerta Walker, 1904 (monotypy).

Accessory flagellum 2-articulate; inner lobes of lower lip obsoles-
cent; [inner plate of maxilla 1 undescribed], inner plate of maxilla 2
densely setose on medial edge; gnathopods normal; uropod 3 not
exceeding uropod 1, rami equal, broadly lanceolate, outer 1-articulate;

MARINE GAMMARIDEAN AMPHIPODA 243

telson long and deeply cleft; metasome and urosome dorsally serrate.
Species: 1, Ceylon, littoral.

Liagoceradocus J. L. Barnard
Liagoceradocus J. L. Barnard, 1965.

Type-species: L. pusillus J. L. Barnard, 1965 (original designation)

Accessory flagellum 2-articulate; lower lip lacking inner lobes;
inner plates of maxillae 1-2 densely setose medially, gnathopods
feeble, subchelate, uropod 3 exceeding uropod 1, rami long, equal,
broadly lanceolate, outer 2-articulate; telson deeply cleft; body lack-
ing dorsal teeth or spines. Species: 1, Micronesia, littoral.

Maera Leach

Maera Leach, 1814a.—Stebbing, 1906.—Schellenberg, 1938.
Mulleria Leach, 1814a (nomen nudum).

Leptothoe Stimpson, 1853.

Linguimaera Pirlot, 1936.

Type-species: Cancer grossimanus Montagu, 1808 (monotypy).
See Chevreux and Fage, 1925.

Accessory flagellum 3- or more articulate; lower lip with inner
lobes; inner plates of maxillae setose only terminally; gnathopods
normal; uropod 3 long or short, rami equal, lanceolate or slightly
rectangular, outer 1-articulate; telson deeply cleft; urosome occa-
sionally toothed; mandibular palp article 3 slender, not falcate (Elas-
mopus). Species: 35, cosmopolitan, primarily tropical, littoral to
bathyal (1,238 m).

Maeracunha Stephensen
Maeracunha Stephensen, 1949.

Type-species: MM. tristanensis Stephensen, 1949 (monotypy).

Accessory flagellum more than 4-articulate; lower lip with small
inner lobes; inner plate of maxilla 1 with seven partially medial setae,
inner lobe of maxilla 2 densely setose medially; gnathopods normal;
uropod 3 short, rami subequal, sublanceolate, outer 2-articulate;
telson deeply cleft; metasome and urosome untoothed; mandibular
palp article 3 subequal to article 1, bearing only two setae (combining
character). Species: 1, Tristan da Cunha, littoral.

Maerella Chevreux

Maerella Chevreux, 191 1a.

Type-species: Gammarus tenuimanus Bate, 1862 (monotypy). See
Chevreux and Fage, 1925.

244 U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum 4- or more articulate; lower lip with inner lobes;
inner plates of maxillae setose only terminally; gnathopods normal;
uropod 3 exceeding uropod 1, rami equal, outer 1-articulate; telson
cleft shortly; metasome and urosome dorsally toothed, urosome with
scattered spines; mazillipedal palp with only three articles. Species:
1, temperate N. E. Atlantic, littoral.

Maeropsis Chevreux

Maeropsis Chevreux 1919; 1927.

Type-species: MM. perrieri Chevreux, 1919, 1927 (monotypy).

Accessory flagellum more than 4-articulate; lower lip with inner
lobes; inner plate of maxilla 1 setose only terminally, of maxilla 2
densely setose medially; gnathopods normal; uropod 3 not exceeding
uropod 1, rami equal, rectangular, outer l-articulate; telson deeply
cleft; body lacking dorsal teeth or spines. Species: 1, N. Atlantic,
bathyal (698 m).

**Marine Carinate Gammarus”’

Species: Gammarus mucronatus Say, 1818.

Accessory flagellum 3- or more articulate; lower lip with inner lobes;
inner plates of maxillae 1-2 densely setose medially; gnathopods
normal, subequal in size; uropod 3 exceeding uropod 1, rami long,
outer 2-articulate, inner slightly shorter than outer; telson deeply
cleft; each metasomite with one dorsal tooth; urosomites with dorsal
bundles of spines. Species: 1, Atlantic and Gulf United States,
littoral lagoonal. Formerly included with Baikalian Carinogammarus
but having affinities with freshwater Rivulogammarus. See Meso-
gammarus.

Megaluropus Hoek

Megalonoura Herdman, 1889 (nomen nudum).
Megaluropus Hoek, 1889.—Stebbing, 1906.—K. H. Barnard, 1940.
Phylluropus K. H. Barnard, 1932.

Type-species: M. agilis Hoek, 1889 (monotypy). See Chevreux
and Fage, 1925.

Accessory flagellum 2-articulate; lower lip with inner lobes; inner
plates of maxillae 1-2 densely setose medially; gnathopods feeble,
poorly subchelate; uropod 3 exceeding uropod 1 or not, rami equal,
foliaceous, outer 1-articulate; telson deeply cleft; metasome and
urosome often dorsally serrate. Species: 3, bi-boreal, littoral (pri-
marily neritic).

MARINE GAMMARIDEAN AMPHIPODA 9A5

Melita Leach, new synonymy

Melita Leach, 1814a.—Stebbing, 1906.—Shoemaker, 1941b.

Boscia Leach, 1814a (nomen nudum).

Megamoera Bate, 1862 (type by present selection, Gammarus dentatus Krgyer,
1842).

Ps Stout, 1913.

Dulichiella Stout, 1918.

Type-species: Cancer palmatus Montagu, 1804 (monotypy). See
Sars, 1895.

Accessory flagellum 2- or more articulate; lower lip with slight
inner lobes; inner plate of maxilla 1 with largely terminal setae, of
maxilla 2 densely setose medially; gnathopods normal; uropod 3
greatly exceeding uropod 1, outer ramus very long, 1-articulate, inner
short and scale-like; telson deeply cleft; metasome often toothed,
urosome often toothed and bearing scattered dorsal spines. Species:
45, cosmopolitan, littoral to abyssal.

Melitoides Gurjanova

Melitoides Gurjanova, 1934.

Type-species: MM. makarovt Gurjanova, 1934 (monotypy). See
Gurjanova, 1951.

Accessory flagellum 4-articulate; lower lip with inner lobes; inner
plates of maxillae densely setose medially; gnathopods normal;
uropod 3 not exceeding uropod 1, outer ramus long, 1-articulate,
inner ramus short, scale-like; telson deeply cleft; urosome dorsally
toothed. Species: 1, subarctic, littoral.

Mesogammarus Tzvetkova

Mesogammarus Tzvetkova, 1965.

Type-species: MM. melitoides Tzvetkova, 1965 (original desig-
nation).

Accessory flagellum more than 3-articulate; lower lip with scarce
indication of inner lobes; inner plates of maxillae 1-2 lined medially
with setae; gnathopod 1 slightly larger than 2; uropod 3 apparently
exceeding uropod 1, rami long, lanceolate, subequal in length, outer
ramus with short article 2; telson short, broad, cleft halfway, lobes
gaping [possibly interpreted as deeply emarginate]; pleonites 1-3
with posterodorsal margins cut into several medium-sized teeth;
urosome armed dorsally with rows of spines. Species: 1, Japan Sea,
near Vladivostok, littoral.

Metaceradocoides Birstein and Vinogradova

Metaceradocoides Birstein and Vinogradova, 1960.

246 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: MM. vitjazi Birstein and Vinogradova, 1960 (original
designation).

Accessory flagellum greatly more than 4-articulate; lower lip
with inner lobes; inner plates of maxillae 1-2 densely setose medially;
gnathopod 2 longer and more slender than gnathopod 1, both sub-
chelate; uropod 3 not exceeding uropod 1, rami equal, lanceolate,
outer 2-articulate; body lacking dorsal teeth or spines; palp article
4 of maxilliped long, claw-like (Bathyceradocus). Species: 1, N.W.
Pacific, hadal (7,210 m).

Metaceradocus Chevreux

Metaceradocus Chevreux, 1925.

Type-species: M. perdentatus Chevreux, 1925 (monotypy).

Accessory flagellum 3- or more articulate; lower lip with inner
lobes; inner plate of maxilla 1 mostly setose terminally, of maxilla
2 densely setose medially; gnathopods feeble, slender, subchelate;
uropod 3 exceeding uropod 1, rami long, equal, lanceolate, outer
l-articulate; telson deeply cleft; pleonites dorsally serrate and with
scattered spines; mandibular palp articles 2 and 3 subequal (Ceradocus).
Species: 2, E. Atlantic, E. Pacific, littoral.

Netamelita J. L. Barnard

Netamelita J. L. Barnard, 1962b.

Type-species: N. cortada J. L. Barnard, 1962b (original desig-
nation).

Accessory flagellum 1-articulate; lower lip with inner lobes; inner
plates of maxillae 1-2 setose only terminally; gnathopods small,
subchelate, equal in size; uropod 3 exceeding uropod 1, outer ramus
long, 1-articulate, inner short, scale-like; telson deeply cleft; body
lacking dorsal teeth or spines. Like Eriopisella but outer ramus of
uropod 3 uniarticulate. Species: 1, California, littoral.

Paraceradocus Stebbing

Paraceradocus Stebbing, 1899a; 1906.

Type-species: Megamoera miersii Pfeffer, 1888 (original designa-
tion). See K. H. Barnard, 1932.

Accessory flagellum 4- or more articulate; lower lip with imner
lobes; inner plate of maxilla 1 only terminally setose, of maxilla 2
densely setose medially; gnathopods normal; uropod 3 exceeding
uropod 1, rami foliaceous, equal, long, outer 1-articulate; telson
deeply cleft; some urosomal segments dorsally toothed. Note: P.
micramphopus Stebbing, 1910, is probably referrable to Metaceradocus.
Species: 2, subantarctic, Australia, littoral (to 310 m).

MARINE GAMMARIDEAN AMPHIPODA 247

Parapherusa Stebbing

Harmomia [lapsus for Harmonia] Haswell, 1880c (Harmonia =homonym,

Coleoptera).

Chloris Haswell, 1880a (homonym, Aves).
Parapherusa Stebbing, 1906 (new name for Harmomia).

Type-species: Harmonia crassipes Haswell, 1880c (monotypy).
See Stephensen, 1949.

Accessory flagellum 4- or more articulate; lower lip with large
inner lobes; inner plates of maxillae 1-2 densely setose medially;
gnathopods normal; uropod 3 short, not exceeding uropod 1, rami
equal, shorter than peduncle, outer 1-articulate and bearing enlarged
serrate spine; telson simple, elongate; body lacking dorsal teeth or
spines. Species: 1, antiboreal, Australia-New Zealand, littoral.

Parelasmopus Stebbing

Parelasmopus Stebbing, 1888; 1906.

Type-species: Gammarus suluensis Dana, 1852a (monotypy).
See Chevreux, 1901b (as P. setiger).

Accessory flagellum 2- or 3- articulate; lower lip with inner lobes;
inner plate of maxilla 1 setose only terminally, of maxilla 2 generally
terminally setose; gnathopods normal; uropod 3 not exceeding uropod
1, rami short, broad, lanceolate, outer l-articulate; telson deeply
cleft; metasome and urosome dorsally carinate, no spines; mandibular
palp article 2 shorter than article 1. Species: 3, tropical Pacific,
littoral (to 522 m).

Quadrivisio Stebbing

Quadrivisio Stebbing, 1907.
Pseudoceradocus Shoemaker, 1933b.
Type-species: Q. bengalensis Stebbing, 1907 (monotypy).
Accessory flagellum greatly more than 4-articulate; lower lip
with inner lobes; inner plates of maxillae 1-2 densely setose medially ;
enathopods normal; uropod 3 exceeding uropod 1, rami equal, foli-
aceous, outer l-articulate; telson deeply cleft; body lacking dorsal
teeth or spines; eyes four in number. Species: 2, circumtropical,
littoral.

Weyprechtia Stuxberg

Weyprechtia Stuxberg, 1880.—Stebbing, 1906.

Type-species: Amathilla heuglini Buchholz, 1874 (monotypy and
subsequent synonymy). See Gurjanova, 1951.

Accessory flagellum 4- or more articulate; inner lobes of lower lip
obsolescent (in drawings they appear distinct); inner plates of maxillae

248 U.S. NATIONAL MUSEUM BULLETIN 271

1-2 densely setose medially; gnathopods slender, feeble, poorly sub-
chelate; uropod 3 not exceeding uropod 1, rami equal, lanceolate,
outer l-articulate; telson entire; body lacking dorsal teeth or spines.
Species: 2, arctic, littoral.

Haustoriidae

Figures 97-100

Dracnosis.—Antenna 1 usually with accessory flagellum; rostrum
usually absent but when present then pereopod 5 longer than 4; otherwise
pereopod 5 shorter and of different structure than pereopod 4; pereo-
pods strongly spinose and setose (fossorial); gnathopods feeble. See
Gammaridae, Phoxocephalidae, Liljeborgiidae, Ampeliscidae, Param-
phithoidae, Astyridae, Argissidae.

Description.—Accessory flagellum absent (rare) or 1- to multi-
articulate; peduncles of antennae elongate or not; antenna 2 peduncle
occasionally spinose, often with elongate plumose setae; rostrum
usually inconspicuous, occasionally present; upper lip not strongly
incised; mandible always with 3-articulate palp and molar rarely
absent but often nontriturative; lower lip with inner lobes; maxillae
basic; maxilliped with well-developed plates, palp 3-4 articulate, occa-
sionally of peculiar shape; gnathopods feeble, varying from minutely
chelate, through simple to lacking article 7; coxae of medium length
or long, highly variable, occasionally acuminate; uropod 3 variable
but one or both rami longer than or equal to peduncle, lanceolate,
rectangular, or subcylindrical, rarely flabellate; telson short or medium
in length, deeply cleft or subentire.

RELATIONSHIP.—The family is so neatly linked to the Gammaridae
by the genera Pontoporeia and Bathyporeia that on gross morpho-
logical grounds there seems little reason to maintain it; nevertheless
taxonomists seem to have been able to recognize its members (ex-
cepting Hurinia Tucolesco). Of particular significance are the feeble
enathopods of haustoriids and the strong development of spines and
setae for these digging animals. Certainly the fossorial habit dis-
tinguishes them from most Gammaridae.

Liljeboriids have a weak mandibular molar, poorly spinose append-
ages and very powerful gnathopods.

Priscillina and Haustorius resemble the Paramphithoidae through
their subacute first four coxae but the excessive spination and seta-
tion of appendages and multiarticulate accessory flagella are distinc-
tive of Haustoriidae.

The Phoxocephalidae are closely related and merge into Haus-
torlidae through genera such as Platyischnopus and Zobracho. Most

MARINE GAMMARIDEAN AMPHIPODA QAO

haustortids lack a distinct rostrum. All phoxocephalids have pereopod
5 shorter than 4 and of different structure, and a large visor-like
rostrum. Some haustoriids have a small rostrum and pereopod 5
shorter than 4 and of different structure. With some genera, one
must flip a coin to choose the proper family.

g

Ficure 97.—Haustoriidae: a, Priscillina armata (Boeck) (Sars, 1895, pl. 42); b, Urothoe
elegans Bate (Sars, 1895, pl. 47, as U. norvegica). Upper lip: c, Haustorius arenarius
(Slabber) (Sars, 1895, pl. 46). Mandible: d, Urothoe; e, Haustorius; f, Carangolia mandt-
bularis J. L. Barnard (1962d). Lower lip: g, Haustorius; h, Pontoporeia femorata Krgyer
(Sars, 1895, pl. 41).

250 U.S. NATIONAL MUSEUM BULLETIN 271

re p

HN P09

Ficure 98.—Haustoriidae: a, Pontoporeia femorata Krdyer (Sars, 1895, pl. 41); b, Zobracho
canguro J. L. Barnard (1961). Maxilla 1: c, Haustorius arenarius (Slabber) (Sars, 1895,
pl. 46). Maxilla 2: d, Pontoporeia; e, Haustorius. Maxilliped: f, Haustorius; g, Prisctl-
lina armata (Boeck) (Sars, 1895, pl. 42). Urosome: h, Urohaustorius halei Sheard (1936b).

MARINE GAMMARIDEAN AMPHIPODA 251

All Ampeliscidae have pereopod 5 shorter than 4 and of different
structure, similar to all Phoxocephalidae and some Haustoriidae.
Unlike the burrowing Phoxocephalidae and Haustoriidae, the Am-
peliscidae have glands in pereopods 1-2 that secrete a substance
used for the formation of tubes. Ampeliscidae never have multiple
ommatidia in the eyes as in Phoxocephalidae and Haustoriidae but
form cuticular Jenses. They have elongate, laterally compressed
heads. Unlike phoxocephalids and haustoriids, Ampeliscidae lack
any trace of an accessory flagellum. The accessory flagellum of
Carangolia is vestigial but the animal is otherwise clearly an
haustoriid.

Oedicerotidae, with fossorial pereopods and always with vestigial
or absent accessory flagella resemble a few haustoriids, but Oedicero-
tidae always have pereopod 5 very elongate, uropod 3 with elongate
peduncle, elongate subequal rami and linguiform, entire (rarely
emareginate) telson.

A major division of the key to genera concerns the absence of
pereopodal dactyls. The ends of those legs often are highly spinose
and if one of the ‘‘spines” is not elongate nor obviously distinct from
the others then dactyls are considered to be absent.

Nomenclatural Changes for Haustoriidae

See Bousfield (1965) for specific nomenclatural changes.

Haustoriopsis Schellenberg (1938) is removed to the Prophliantidae.

Urothoe simplignathia J. L. Barnard (1962d) is removed to Par-
daliscella, Pardaliscidae.

Note the comments in the Calliopiidae on the genus Sancho.
Haustoriids are perhaps the most interesting group of amphipods

and so diversified that by necessity many genera have been illus-

trated herein.

Key to the Genera of Haustoriidae

ieeeAlleotepereopods, lo lacking dactyls (ig. 990) 2 2 ae i) eee) ee
Some of pereopods 1-5 bearing dactyls (fig. 98a) ! . ........ aa)

2. Pereopod 2 smaller than and morphologically distinct from pereopod 1
(its orientation apparently reversed)... .. . . . . .EHohaustorius
Pereopods 1-2 subequal in size and morphologically intersimilar. . . . 3

See Vera pedal pal) with four anticless 49 =. eee) 0s ne celica uel ran
Maxillipedalspalp: wathathreevarticles 3s Sens G. ce se se 5

1 See explanation in text (p. 251).

285-135 O- 69-17

252

17.

18.

9:

20.

U.S. NATIONAL MUSEUM BULLETIN 271

Head with large rostrum (fig. 986), telson longer than broad and deeply

cleft (fig. 99f) . . . . . . . Zobracho 2
Head lacking rostrum, ialaon BE andes bem lone amd seam cleft (fig. 99q).
Carangolia

Inner ramus of uropod2absent. ........ ... . .. Neohaustorius
Inner ramus of uropod 2 present. ... . AME aN pul (0)
Maxillipedal palp article 3 weakly penientate! or clavate, anion plate of
maxilla 2 scarcely enlarged (fig. 98d) .... . 7
Maxillipedal palp article 3 strongly geniculate (Gee 98f), autier mate of
maxilla 2 greatly enlarged (fig. 98e) ... . 8
Rami of uropod 1 expanded distally, sostanadansall ‘bowls: of dleoatke 3
forming a lobe overhanging urosome. .. . . . . Pseudohaustorius
Rami of uropod 1 not expanded distally, posterodorsal border of pleonite 3
not forming lobe overhanging urosome. ... .. . Protohaustorius
Pleonal epimeron 3 with large posteroventral tooth . . . Acanthohaustorius

Pleonal epimeron 3 rounded posteroventrally.
Haustorius and Parahaustorius

Antenna 1 geniculate between articles 1 and 2 (fig. 99c) ...... . 10
Antenna 1 not geniculate, or occasionally between articles 2 and3. . 11
Gnathopod 2 lacking dactyl (fig. 1002)... ... .. . .. Bathyporeia
Gnathopod 2 bearing dactyl. .... . . . . . Amphiporeia
Coxa 1 vestigial, less than one fourth as lone. AS /COKA Gi en ernie
Coxa’ l not) vestigial’ at least half asilong as coxa: 3)... eee ie
Coxa 2 vestigial, less than one fourth as long as coxa 3. . Urohaustorius
Coxa 2 not vestigial, almost as large as coxa 3... . .. =. +. . Cardenio
Coxai 2 obtusely, pointed below, (fies Oia). | 5 nas oe ee
Coxa: 2 rounded or, quadrate below, (igs 100g) 593-7.) 9 | eo
Telson/split nearly its fulllength) 97> 5 3. 2 ee Zobracho
Telson split: one third its length: 3°00. <2 =. i.e. aye Briscillina
Gnathopods chelate (fig. 100d) ............ . Platyischnopus
Gnathopods subchelate or simple . ... . Pee ses ae La)
Inner ramus of uropod 3 scale-like, half as fonee as een ramus (fig. 100m).
Carangolia
Inner ramus of uropod 3 subequal to outer. . . . Rime hi Lah MIU
Mandibular palp large, borne on basal process (like ie. 1509), article 3
clavate, molar of medium size, ridged ... . . .. . Phoxocephalopsis
Mandibular palp small, directly attached to mandibular body, article 3
linearsmolar large or, medium; smooth) 0) 2) a a eS
Outersramus of uropodia umiarticulatem 0) een oe 6)
Outer ramus of uropod 3 biarticulate . .. . Ae O)
Accessory flagellum present, usually 2- cnouliaite: aanaerarl 5 shorter and of
different structure than pereopod 4... . . . . . . . Pontoporeia

Accessory flagellum absent; pereopod 5 similar to Dereon 4,
Zaramilla (Eusiridae)

Article 4 of pereopod 4 not expanded, article 2 of pereopod 5 not greatly

produced downward .. . . . . Urothoe
Article 4 of pereopod 4 coronmledl Aericle 2 of neieapod 5 prem produced
downward) suey Aieiek slot ica a, Wenner Mian wee ne gO ROUMOLLES

2 Zobracho J. L. Barnard (1961). Note that the basal article of the maxillipedal palp was not inked on the
original drawing; the palp has four articles.

MARINE GAMMARIDEAN AMPHIPODA 253

Ayo

Rw
[K
H

A JU oe
€ i g h

Ficure 99.—Haustoriidae: a, Carangolia puliciformis J. L. Barnard (1961); b, Haustorius
arenarius (Slabber) (Sars, 1895, pl. 46). Antenna 1:c, Bathyporeia guilliamsoniana (Bate)
(Sars, 1895, pl. 43, as B. norvegica). Telson: d, Bathyporeia: e, Priscillina armata (Boeck)
(Sars, 1895, pl. 42); f, Pontoporeia femorata Krgyer (Sars, 1895, pl. 41); g, Carangolia;
h, Haustorius.

254 U.S. NATIONAL MUSEUM BULLETIN 271

Genera of Haustoriidae

Acanthohaustorius Bousfield

Acanthohaustorius Bousfield, 1965.

Type-species: A. millsi Bousfield, 1965 (original designation).

Antenna 1 not geniculate; mandibular palp not on basal process,
molar ridged; outer plate of maxilla 2 greatly enlarged; maxillipedal
palp 3-articulate, article 3 geniculate; coxae 1-2 of normal size, coxae
1-3 obtusely pointed below; gnathopod 1 simple, article 5 longer than
6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod 3
slightly elongate, rami subequal; telson short, cleft nearly to base;
pleonal epimeron 3 with spine-like posteroventral process (Parahaus-
torius, Haustorius); lower lip lacking mandibular lobes. Species: 4,
N.W. Atlantic, littoral.

Amphiporeia Shoemaker

Amphiporeia Shoemaker, 1929.

Type-species: A. lawrenciana Shoemaker, 1929 (original desig-
nation).

Antenna 1 geniculate between articles 1 and 2; mandibular palp
attached to basal process, molar ridged; outer plates of maxilla 2 not
enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size,
rounded-quadrate below; gnathopods subchelate, article 5 of gnath-
opod 1 slightly longer than article 6; gnathopod 2 bearing article 7
(Bathyporeia); pereopods dactylate; inner ramus of uropod 3 half as
long as outer; telson fully cleft. Species: 2, N.W. Atlantic, littoral.

Bathyporeia Lindstrom

Bathyporeia Lindstrom, 1855.—Stebbing, 1906.
Thersites Bate, 1857a, 1857b (homonym, Mollusca).

Type-species: B. pilosa Lindstrom, 1855 (monotypy). See Sars,
1895.

Antenna 1 geniculate between articles 1 and 2; mandibular palp
attached to basal process, molar ridged; outer plate of maxilla 2 not
enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size,
rounded-quadrate below; gnathopods simple, gnathopod 2 lacking
article 7 (Amphiporeia), article 5 of gnathopod 1 longer than article 6;
at least pereopods 1-2 dactylate; inner ramus of uropod 3 short,
scale-like; telson fully cleft. Species: 11, boreal N. Atlantic, littoral.

MARINE GAMMARIDEAN AMPHIPODA 955

Ficure 100.—Haustoriidae: a, Platyischnopus herdmani Walker (Pillai, 1957). Gnathopod
1: b, Bathyporeia guilliamsoniana (Bate) (Sars, 1895, pl. 43, as B. norvegica); c, Pontoporeia
femorata Krgyer (Sars, 1895, pl. 41); d, Platyischnopus mirabilis Stebbing (1888); ¢,
Priscillina armata (Boeck) (Sars, 1895, pl. 42); f, Urothoe; g, Haustorius arenarius (Slabber)
(Sars, 1895, pl. 46); h, Carangolia puliciformis J. L. Barnard (1961). Gnathopod 2:
i, Bathyporeia; 7, Haustorius. Uropod 3: k, Carangolia; 1, Haustorius; m, Priscillina;
n, Pontoporeia; 0, Urothoe.

256 U.S. NATIONAL MUSEUM BULLETIN 271

Carangolia J. L. Barnard
Carangolia J. L. Barnard, 1961.

Type-species: C. mandibularis J. L. Barnard, 1961 (original
designation).

Antenna 1 not geniculate; mandibular palp not on process, molar
large, smooth, terminal in position; outer plate of maxilla 2 not en-
larged, maxillipedal palp 4-articulate; gnathopods nearly simple,
article 7 immensely elongate, articles 5 and 6 of gnathopod 1 subequal
in length; pereopods dactylate (except ?pereopod 1); uropod 3 greatly
reduced in size, rami very short, subcircular, inner shorter than outer;
telson short, broad, minutely cleft. Species: 2, antiboreal, bathyal
(610-1,861 m).

Cardenio Stebbing

Cardenio Stebbing, 1888; 1906.

Type-species: (C. paurodactylus Stebbing, 1888 (monotypy).

Antenna 1 not geniculate; mandibular palp not on basal process,
molar ridged or toothed; outer plates of maxilla 2 not enlarged;
maxillipedal palp 3-articulate; coxa 1 very small, coxae rounded-
quadrate below; gnathopods simple, article 5 of gnathopod 1 longer
than article 6; pereopods 3-5 dactylate, pereopods 1—2 apparently not
dactylate; uropod 3 short, rami equal in length; telson long, deeply
cleft; lower lip apparently lacking mandibular processes. Species: 1,
subantarctic, littoral.

Eohaustorius J. L. Barnard
Eohaustorius J. L. Barnard, 1957b.

Type-species: Haustorius washingtonianus Thorsteinson, 1941
(original designation).

Antenna 1 not geniculate; mandibular palp not on basal process,
molar smooth; outer plates of maxilla 2 not enlarged; maxillipedal
palp 3-articulate, article 3 not geniculate; coxae 1 and 2 much smaller
than and partially hidden by coxae 3-4; gnathopod 1 simple, article 5
longer than article 6, gnathopod 2 minutely chelate; pereopod 2
similar to pereopod 3 in orientation, and smaller than pereopod 1;
uropod 3 short, rami equal in length; telsonic lobes completely detached
basally; outer lobes of lower lip lacking mandibular processes. Species:
4, boreal Pacific, littoral.

Haustorius Muller

Haustorius Muller, 1775.—Stebbing, 1906.
?Lepidactylis Say, 1818 (see Bousfield, 1965).
Pterygocerus Latreille, 1825.

MARINE GAMMARIDEAN AMPHIPODA 257

Bellia Bate, 1851 (homonym, Decapoda).
Sulcator Bate, 1854.

Type-species: Oniscus arenarius Slabber, 1769 (monotypy). See
Sars, 1895.

Antenna 1 not geniculate; mandibular palp not on basal process,
molar ridged; outer plate of maxilla 2 greatly enlarged; maxillipedal
palp 3-articulate, article 3 geniculate; coxae 1—2 of normal size, coxae
1-3 obtusely pointed below; gnathopod 1 simple, article 5 longer than
article 6, gnathopod 2 minutely chelate; pereopods lacking dactyls;
uropod 3 short, rami equal in length; telson short, partially cleft;
pereopods 4-5 urth articles 4-5 three fourths as broad as article 2 of
pereopod 5 (Zobracho and Priscillina); pleonal epimeron 3 rounded
posteroventrally (Acanthohaustorius); outer lobes of lower lip lacking
mandibular processes. Species: 2, N. Atlantic, littoral.

Neohaustorius Bousfield

Neohaustorius Bousfield, 1965.

Type-species: N. schmitzi Bousfield, 1965 (original designation) .

Antenna 1 not geniculate; mandibular palp not on basal process,
molar triturative; outer plate of maxilla 2 greatly enlarged; maxil-
lipedal palp 3-articulate, article 3 geniculate; coxae 1-2 of normal
size, to some extent coxae 1-2 obtusely pointed below; gnathopod 1
simple, article 5 longer than 6, gnathopod 2 minutely chelate; pereo-
pods lacking dactyls; uropod 3 short, rami subequal; telson short,
uncleft; inner ramus of uropod 2 absent; lower lip lacking mandibular
lobes. Species: 2, N.W. Atlantic, littoral.

Parahaustorius Bousfield

Parahaustorius Bousfield, 1965.

Type-species: P. longimerus Bousfield, 1965 (original designation).

Antenna 1 not geniculate; mandibular palp not on basal process,
molar triturative; outer plate of maxilla 2 greatly enlarged; maxil-
lipedal palp 3-articulate, article 3 geniculate; coxae 1-2 of normal
size, coxae 1-2 obtusely pointed below; gnathopod 1 simple, article
5 longer than 6, gnathopod 2 minutely chelate; pereopods lacking
dactyls; uropod 3 slightly elongate, rami subequal; telson short,
cleft about halfway; pleonal epimeron 3 rounded posteroventrally
(Acanthohaustorius); lower lip lacking mandibular lobes. Species: 3,
N.W. Atlantic, littoral.

Phoxocephalopsis Schellenberg

Phoxocephalopsis Schellenberg, 1931.
Haustoriella K. H. Barnard, 1931.

258 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: P. zvmmeri Schellenberg, 1931 (monotypy).

Antenna 1 not geniculate; mandibular palp on basal process, large,
article 3 clavate (Urothoe, Urothoides), molar ridged; outer plates of
maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxa 1 nearly
obtusely pointed below, others rounded-quadrate; gnathopods sub-
chelate or nearly simple, article 5 of gnathopod 1 longer than article
6; pereopods dactylate; uropod 3 short, rami subfoliaceous, subequal
in length; telson short, deeply cleft. Species: 2, subantarctic-anti-
boreal, littoral.

Platyischnopus Stebbing

Platyischnopus Stebbing, 1888; 1906.

Type-species: P. mirabilis Stebbing, 1888 (monotypy).

Antenna '1 not geniculate; mandibular palp not on basal process,
molar “ridged,’”’ smooth or absent; outer plates of maxilla 2 not
enlarged; maxillipedal palp 4-articulate; coxae of medium length,
rounded-quadrate below; gnathopods chelate, article 5 of gnathopod
1 longer or shorter than article 6; pereopods dactylate; uropod 3
(outer ramus only) immensely elongate, inner ramus short, scale-like;
telson partially cleft or emarginate; head with “‘shark-nose’’ rostrum.
Species: 6, warm N. temperate through tropics to antiboreal, littoral.

Pontoporeia Kr¢yer

Pontoporeia Kr¢yer, 1842.—Stebbing, 1906.

Type-species: P. femorata Kroyer, 1842 (monotypy). See Sars,
1895.

Antenna 1 not geniculate; mandibular palp not on basal process,
molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal
palp 4-articulate; coxae 1-2 of normal size, rounded-quadrate below;
gnathopod 1 subchelate, articles 5-6 subequal in length, gnathopod
2 slightly chelate; at least pereopods 1, 2, 5 dactylate; rami of uropod
3 equal in length; telson cleft halfway or more. Species: 5; 1
Caspian, 4 subarctic-boreal and glacial relicts, littoral.

Priscillina Stebbing

Priscilla Boeck, 1871 (homonym, Coleoptera).
Priscillina Stebbing, 1888 (new name); 1906.

Type-species: Pontoporeia armata Boeck, 1861 (monotypy). See
Sars, 1895.

Antenna 1 not geniculate; mandibular palp not on basal process,
molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal
palp 4-articulate; coxae 1-2 of normal size, coxae 1-4 obtusely pointed
below; gnathopods subchelate, articles 5-6 of gnathopod 1 subequal

MARINE GAMMARIDEAN AMPHIPODA 259

in length; some pereopods dactylate; uropod 3 short, inner ramus
scale-like; telson cleft one third of its length; articles 4-5 of pereopods
4-6 one third as broad as article 2 of pereopod 5 (Haustorvus). Species:
1, arctic-subarctic, littoral.

Protohaustorius Bousfield

Protohaustorius Bousfield, 1965.

Type-species: P. deichmannae Bousfield, 1965 (original designa-
tion).

Antenna 1 semigeniculate; mandibular palp not on basal process,
molar triturative; outer plate of maxilla 2 not enlarged; maxillipedal
palp 3-articulate, article 3 not geniculate; coxae 1-2 of normal size,
not obtusely pointed below; gnathopod 1 simple, article 5 longer than
6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod
3 short, rami subequal; telson short, slightly emarginate; rami of uro-
pod 1 not distally expanded; posterodorsal border of pleonite 3 not form-
ing lobe overhanging urosome (Pseudohaustorius); lower lip. lacking
mandibular lobes. Species: 2, N.W. Atlantic, littoral.

Pseudohaustorius Bousfield
Pseudohaustorius Bousfield, 1965.

Type-species: P. caroliniensis Bousfield, 1965 (original designa-
tion).

Antenna 1 not geniculate; mandibular palp not on basal process,
molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal
palp 3-articulate, article 3 clavate; coxae 1-2 of normal size, 1-3
subobtusely pointed below; gnathopod 1 simple, article 5 longer
than 6, gnathopod 2 minutely chelate; pereopods lacking dactyls;
uropod 3 short, rami subequal; telson short, slightly emarginate;
rami of uropod 1 distally expanded; posterodorsal border of pleonite
3 forming lobe overhanging urosome (Protohaustorius); lower lvp lack-
ing mandibular lobes. Species: 3, N.W. Atlantic and Gulf of Mexico,
littoral.

Urohaustorius Sheard
Urohaustorius Sheard, 1936b.

Type-species: U. halet Sheard, 1936b (original designation).

Antenna 1 not geniculate; mandibular palp apparently not on
basal process, molar large and projecting but apparently lacking
ridges or teeth; outer plates of maxilla 2 not enlarged; maxillipedal
palp 4-articulate; coxae 1 and 2 vestigial; gnathopod 1 simple, article
5 longer than article 6, gnathopod 2 minutely chelate; pereopods
dactylate but some dactyls resembling spines; rami of uropod 3 sub-
equal in length; telson short, entire. Species: 2, S. Australia, littoral.

260 U.S. NATIONAL MUSEUM BULLETIN 271

Urothoe Dana

Urothoe Dana, 1852b.—Stebbing, 1906.
Egidia Costa, 1853c, 1857.

Type-species: Urothoe irrostratus Dana, 1853 (indicated by elimi-
nation and ipso-facto monotypy of Boeck, 1876, but firmly selected
by Stebbing, 1891). Type-species remains dubious and genus herein
based provisionally on U. elegans Bate, 1857a (Sars, 1895, as U.
norvegica Boeck, 1861).

Antenna 1 not geniculate; mandibular palp not on basal process,
small, article 3 linear (Phoxocephalopsis), molar large, poorly ridged;
outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate;
coxae 1-2 of normal size, rounded-quadrate below; gnathopods sub-
chelate or simple, article 5 of gnathopod 1 longer than article 6;
pereopods dactylate; rami of uropod 3 equal in length; telson fully
cleft; article 4 of pereopod 4 not strongly expanded, article 2 of pereopod
5 not greatly produced downward (Urothoides). Species: 22, cosmo-
politan, littoral to abyssal.

Urothoides Stebbing

Urothoides Stebbing, 1891; 1906.

Type-species: Urothoe lachneessa Stebbing, 1888 (monotypy).

Antenna 1 not geniculate; mandibular palp not on basal process,
small, article 3 linear (Phoxocephalopsis), molar large, poorly ridged;
outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate;
coxae 1-2 of normal size, rounded-quadrate below; gnathopods
subchelate, article 5 of gnathopod 1 longer than article 6; pereopods
dactylate; uropod 3 of medium length, inner ramus half as long as
outer; telson deeply cleft; article 4 of pereopod 4 expanded, article 2
of pereopod 5 greatly produced downward (Urothoe). See other remarks
by K. H. Barnard, 1932. Species: 1+ ?1, subantarctic, bathyal
(236-720 m).

Zobracho J. L. Barnard

Zobracho J. L. Barnard, 1961.

Type-species: Z. canguro J. U. Barnard, 1961 (original designation).

Antenna 1 not geniculate; mandibular palp not on basal process,
molar small and slightly toothed; outer plates of maxilla 2 not en-
larged; maxillipedal palp 4-articulate; coxa 1 strongly reduced in
size, coxae 1—2 obtusely pointed below; gnathopods subchelate, article
5 of gnathopod 1 longer than article 6; some pereopods presumably
dactylate but dactyls resembling spines; uropod 3 short, inner ramus
half as long as outer; telson long, nearly fully cleft; articles 4-5 of
pereopods 4-5 one third as broad as article 2 on pereopod 5 (Haustorius) ;
head with large rostrum. Species: 1, Australia, bathyal (875 m).

MARINE GAMMARIDEAN AMPHIPODA 261

Hyalellidae and Hyalidae

[see Superfamily Talitroidea]

Hyperiopsidae
Ficures 101, 102

Diacnosis.—Accessory flagellum well developed; article 4 of
pereopods 1-2 greatly elongate; palp of first maxilla claviform, slightly
geniculate; gnathopods feeble, simple. See Vitjazianidae, Astyridae,
Stilipedidae, Synopiidae.

Description.—Accessory flagellum well developed, its articles
very elongate; body variable, smooth or carinate, head globular
(resembling hyperiids), subglobular or nearly normal, but rarely
bearing distinct lateral lobes; coxae variable, first four small and
uniform or coxa 4 enlarged and covering anterior coxae; upper lip
with minute incision; mandibular body very short, poorly toothed,
molar very small, weakly triturative or not, palp immense; lower lip
without inner lobes, outer lobes often alate; palp of maxilla 1 biartic-
ulate, article 2 slightly or greatly bent medially (at least on one side) ;
maxilla 2 and maxilliped normal; gnathopods feeble or nearly simple;
uropod 3 biramous; telson cleft or entire, with apical ornamentation.

It is difficult, at first glance, to justify a familial association of the
two hyperiopsid genera. Hyperiopsis resembles an hyperiid amphipod.
It has a globular head, smooth body and short uniform coxae. Parar-
gissa has the normal appearance of a gammaridean and although the
head is subglobular it is rather small and may have lateral lobes; its
fourth coxa is enlarged and covers the smaller coxae 1-3. Nevertheless,
Hyperiopsis and Parargissa share some features that are unusual in
the Gammaridea: (1) the peculiar first maxillary palps of which the
second articles are claviform and slightly bent on at least one of the
maxillae; (2) the very short body of the mandible with a nearly
untoothed cutting edge and a very small, poorly developed molar; (3)
the great elongation of article 4 on pereopods 1 and 2, a condition
rarely occurring in other families (cf. Ampeliscidae); (4) the elonga-
tion of article 1 of the first antennal flagellum (not uncommon in some
other families, viz. Lysianassidae, Synopiidae).

RELATIONSHIP.—The combination of the elongate article 4 of pere-
opods 1 and 2 with the bent first maxillary palp separates this family
from the following similar families: Astyridae, Synopiidae, Pardalis-
cidae, and Vitjazianidae.

The Ampeliscidae resemble hyperiopsids because of the elongate
fourth articles of pereopods 1-2, but ampeliscids lack accessory
flagella and have well-developed mandibular molars.

U.S. NATIONAL MUSEUM BULLETIN 271

h 1

Figure 101.—Hyperiopsidae: a, Hyperiopsis voringt Sars (1885); b, Parargissa galatheae
J. L. Barnard (1961). Accessory flagellum: c, Hyperiopsis laticarpa Birstein and Vino-
gradov (1955). Lower lip: d, Parargissa arquata (Birstein and Vinogradov, 1955);
e, Hyperiopsis. Mandible: f, Hyperiopsis gibbosa Pirlot (1934); g, Parargissa arquata.

Telson: h, Parargissa arquata; 1, Hyperiopsis gibbosa.

MARINE GAMMARIDEAN AMPHIPODA 263
Alezandrella, in the Stilipedidae, has a bent first maxillary palp

but hyperiopsids have well-developed accessory flagella and mandi-
bular molars.

Key to the Genera of Hyperiopsidae

1. Coxae 1-4 small, similar in size and shape (fig. 101a) . . . . MHyperiopsis
Coxa 4 greatly enlarged, concealing the small first three coxae (fig. 1016)
Parargissa

Figure 102.—Hyperiopsidae: Maxilla 1: a,b, Parargissa arquata (Birstein and Vinogradov,
1955); c, Hyperiopsis gibbosa Pirlot (1934); d, other palp of Hyperiopsis. Maxilla 2:
e, Hyperiopsis. Pereopods 1 and 2: f, Hyperiopsis laticarpa Birstein and Vinogradov
(1955); g, Parargissa arquata. Maxilliped: h, Hypertopsis.

Genera of Hyperiop sidae

Hyperiopsis Sars
Hyperiopsis Sars, 1885.—Stebbing, 1906.

Type-species: H. voringvi Sars, 1885 (monotypy). See Stephensen ;
1934.
Species: 7, cosmopelagic, abyssal and bathyal.

264. U.S. NATIONAL MUSEUM BULLETIN 271

Parargissa Chevreux

Parargissa Chevreux, 1908¢e.—Birstein and Vinogradov, 1958.
Protohyperiopsis Birstein and Vinogradov, 1955.

Type-species: P. nasuta Chevreux, 1908c (original designation).
Species: 5, cosmopelagic, abyssal.

Isaeidae

[including Photidae]
Ficures 103-106

Diacnosis.—Accessory flagellum variable in length or absent; telson
entire, short, fleshy; coxa 5 not excavate posteriorly; pereopods 1-2
olandular; wropod 3 rarely projecting beyond uropods 1 and 2, rami (at
least one) as long as or longer than peduncle, occasionally uropod 3
uniramous. See Aoridae, Corophiidae, Ischyroceridae, Calliopidae,
Laphystiopsidae, Oedicerotidae, Pleustidae.

Description.—Accessory flagellum varying from absent to long
and multiarticulate; body smooth, rostrum absent; coxae rounded or
quadrate ventrally, long or short, fourth not excavate posteriorly;
mouthparts basic; gnathopods usually powerfully subchelate or
extremely setose; uropod 3 short, rami as long as or longer than pe-
duncle, not uncinate, occasionally inner ramus reduced or absent;
telson entire, short, fleshy, nearly circular or square, occasionally
falsely cleft.

ReELATIONSHIP.—The Isaeidae do not warrant distinction from the
Photidae on the basis of prehensile or partially subchelate pereopods
and so they must be joined, Isaeidae (Isaeinae Dana, 1853) taking
precedence. This move is unfortunate because of the long use of the
name Photidae for a diversity of genera and the relatively infrequent
use of the name Isaeidae.

The Laphystiopsidae have a nontriturative mandibular molar and
are otherwise like calliopiids and pleustids.

See Calliopiidae for calliopiid-pleustid differences.

The Gammaridae usually have a cleft telson; when entire, the telson
is not circular or subcircular and coxa 4 is strongly excavate posteriorly ;
some Gammaridae have a small, square, unexcavate coxa 4 but in
those species the telson is cleft; Gammaridae lack glandular pereopods,
do not build tubes, usually have uropod 3 projecting beyond uropods
1 and 2 and the rami are enlarged, either by dorsoventral depression,
elongation, or foliaceousness.

The Aoridae have gnathopod 1 larger than gnathopod 2. Linkage is
shown to that family by Aorcho, in the Aoridae, having gnathopods 1

MARINE GAMMARIDEAN AMPHIPODA 265

y SK

aE ey \\ \ >
Of fe IT

S\
Lo)
c) ye
rN
SI
w
Ke,

Ml
Y
s
ps
L>
S
‘Ss
aS

Ficure 103.—Isaeidae: a, Gammaropsis (=Eurystheus) maculata (Johnson) (Sars, 1895,
pl. 198, as G. erythrophthalma); 6, Photis reinhardi Krg¢yer (Sars, 1895, pl. 202);
c, Megamphopus cornutus Norman (Sars, 1895, pl. 200).

J

Ficure 104.—Isaeidae and Aoridae: a, Isaea montagui Milne Edwards (Chevreux and Fage,
1925); b, Leptocheirus pilosus Zaddach (Sars, 1895, pl. 197) [=Aoridae]. Pereopod 1:
c, Ampelisciphotis tridens Pirlot (1938). Telson: d, Microprotopus maculatus Norman
(Sars, 1895, pl. 201); ¢, Photis reinhardi Krdyer (Sars, 1895, pl. 202). Uropod 3: f, Micro-
protopus; g, Leptocheirus; h, Photis;1, Megamphopus cornutus Norman (Sars, 1895, pl. 200);
J, Cheiriphotis megacheles (Giles) (J. L. Barnard, 1962a).

MARINE GAMMARIDEAN AMPHIPODA 267

and 2 equal in size. One must also examine Neomegamphopus, now
assigned to Aoridae, as an intermediary between Isaeidae and Aoridae.
All Isaeidae with male gnathopod 1 having secondary sexual modi-
fications have been removed to the Aoridae.

The Ischyroceridae have the rami of uropod 3 shorter than the
elongate peduncle and usually have one or more distinct hooks or an
uncinate condition on the outer ramus.

The Ampithoidae have an even more stunted third uropod than do
isaeids, the outer ramus of which usually bears hooks; the ampithoid
lower lip has notched or medially excavate outer lobes.

The Corophiidae have a depressed urosome although this is not
clearly distinct in some intergrading genera; all but a few corophiids
have aberrant third uropods lacking one ramus, although a few
Isaeidae also demonstrate this; some corophiids and isaeids are
not clearly distinct and the taxonomist must use both family keys
for doubtful genera.

Camacho and Parunciola in the Corophiidae and Aorcho, now
assigned to the Aoridae, show the difficulty in clearly distinguishing
among isaeids, aorids, and corophiids. These genera, with their
moderately or fully depressed urosomes, equal-sized gnathopods,
biramous third uropod [or uniramous third uropod with long ramus],
and circular or quadrate telsons, lacking hooks, could be assigned
to any of the three families.

The loss of one ramus of uropod 3 in five isaeid genera confounds
technical distinction of Isaeidae from several other, distantly re-
lated families having uncleft telsons. Cressidae, Stenothoidae, and
Thaumatelsonidae are distinguished from isaeids by their small
first coxae. Eophliantidae have cylindrical bodies and Phliantidae,
Talitroidea, and Dogielinotidae lack mandibular palps. Sebidae
have chelate gnathopods and obsolescent mandibular molars and
Pagetinidae have evanescent mandibular molars and poorly developed
outer plates of the maxillipeds.

Nomenclatural Changes in Isaeidae

Amphideutopus J. L. Barnard (1959a) is removed to the Aoridae.

Audulla Chevreux (1901a) is assigned in the Isaeidae as subgenus
to Gammaropsis.

Bathyphotis Stephensen (1944b) is removed to the Ischyroceridae.

Bonnierella Chevreux (1900) is removed to the Ischyroceridae but
is also assigned in the Isaeidae to Megamphopus.

Eurystheus dentatus Holmes (1908) is removed to Protomedeia
(Cheirimedeia).

Leptocheirus Zaddach (1844) is removed to the Aoridae.

285-135 O - 69 - 18

268 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 105.—Isaeidae: a, Chetriphotis megacheles (Giles) (J. L. Barnard, 1962a); b, Chevalia
aviculae Walker (J. L. Barnard, 1962a). Head: c, Ampelisciphotis tridens Pirlot (1938).
Accessory flagella: d, Gammaropsis (= Eurystheus) maculata (Johnston) (Sars, 1895, pl.
199, as G. melanops); e, Microprotopus maculatus Norman (Sars, 1895, pl. 201).

MARINE GAMMARIDEAN AMPHIPODA 269

Neomegamphopus Shoemaker (1942) is removed to the Aoridae.
Pseudeurystheus Schellenberg (1931) is assigned in the Isaeidae as

10.

11.

18.

subgenus to Gammaropsis.

Key to the Genera of Isaeidae

Mropod auuminramousndienMO4p) ie eo ea en Me. 2
Uropod 3 biramous (figs. 104g-j). . . . EAS Ua ONG
Lateral cephalic lobes pedunculate, article 6 oi Beopode 1-2 elongate
(igss 1O5e 104) ine . . . . . Ampelisciphotis
Lateral cephalic lobes and Artiele 6 of percopods 1-2 notelongate. ... 3
Gnathopod 1 simple (fig. 106h). . ....... . . . . Kuphocheira
Cnathopod lisubbchelateq sks. see eu oa Ne cle 3) aa eae
Matbennamtewithvaccessony Hagellumy. 402). were ae eee eu yiueery =) iO
Antenna 1 lacking accessory flagellum... . . . . . . Microphotis
Coxae 1-4 similar to each other in size and shape (fe. 103a) . Microprotopus
Coxae 1—4 of varying shapes and sizes (fig. 105a) . . . . . Cheiriphotis
Uropod 3 with one ey shortened ramus (less than 60% as long as
outer) (fig 04h) 2 xo: Sa ME RS 1 Sc SU CO a
Uropod 3 with subequal rami Ge 104i) SES CNG sigh areal Sam Oa Taam gaa sk Sees AL)
Gnathopods 1-2 simple. . . . . . .. . . . Haplocheira
Gnathopod 1 subchelate, pneehoped 2 ‘simple . . Leptocheirus (Aoridae)
Gnathopods 1-2 subchelate. . .. . Sah Ace ta eat:
Antenna 1 lacking accessory dopelum or bearing a otra. anticls or scale.
Photis

Antenna 1 with multiarticulate accessory flagellum. ......... 9
Uropod 3 scale-like, peduncle plate-like (fig. 1047). . . . . . Cheiriphotis
Uropod 8 cylindrical (fig. 104g) . . . Protomedeia (subgenus Cheirimedeia)
Gnathopod 1 complexly subchelate (fig. 64a), gnathopod 2 with well-
developedpalm. .. . BN tener eae tenn os ee (Aoridae)
These characters not combined. Aslan eval BAe tae aie aa eens eal FL
Article 3 of antenna 1 as long as article 1 or lone: Cent spi vatantec! ioe reheat cue Leet
Article 3iof antenna 1) shorter than article) 2.0.0.4... 3 .-,-,2h6
Accessory flagellum of antenna 1 absent... . . . . . Podoceropsis
Accessory flagellum of antennal present. ........ ay eta eenet alte
Accessory flagellum composed of ascale ........ - Kermystheus
Accessory flagellum composed of one or more long articles. . ..... 14

Accessory flagellum composed of one or two articles.
Megamphopus (Segamphopus and Audulla), Bonnierella !

Accessory flagellum composed of three or more articles . . .... . .15
Pereopods 1-5 slightly subchelate (fig. 104a)... . Noa ea SAR
Pereopods 1-5simple .. . . . Gammaropsis (eubeeaue Pseudeurystheus)
Accessory flagellum absent. . ... . . . . . Dryopoides? and Goesia
Accessory flagellum present. . .. . ' aaa uch Li
Gnathopod 2 subchelate, coxae 1 and ‘fils aaa of Donraalle balanced

dimensions. . . AU semi lye gal lke)
Gnathopod 2 simple Ge 1061), « coxa aL very ral aad cov percd by enlarged

COXAw2 a iene HAY aie a cota at . . . Leptocheirus (Aoridae)
Pleonites 4 and 5 apace OH Sek va al eae Protomedeia
Pleonites 4 and 5 coalesced (fig. 1058) . Beet ss ae Te ecm a oe ee CHOVEIIB

1 See this genus in the Ischyroceridac.
2 See male, Stebbing (1910) and Aoridae (p. 153).

270 U.S. NATIONAL MUSEUM BULLETIN 271

n
ay

m

Ficure 106.—Isaeidae and Aoridae: Mouthparts, Photis reinhardi Kréyer (Sars, 1895,
pl. 202): a, upper lip; b, mandible; c, lower lip; d,e, maxillae 1, 2: f, maxilliped. Max-
illiped: g, Isaea montagui Milne Edwards (Chevreux and Fage, 1925). Gnathopod 1:
h, Kuphocheira setimanus K.. H. Barnard (1932); 7, Photis tenuicornis (Sars, 1895, pl. 203);
j, Leptocheirus pilosus Zaddach (Sars, 1895, pl. 197) [=Aoridae]; k, Megamphopus cornutus
Norman (Sars, 1895, pl. 200). Gnathopod 2: /, Leptocheirus; m, Megamphopus; n,

Photts tenuicornis, female; 0, Kuphocheira; p, Photis tenuicornis, male.

MARINE GAMMARIDEAN AMPHIPODA o71
Genera of Isaeidae

Ampelisciphotis Pirlot
Ampelisciphotis Pirlot, 1938.

Type-species: A. tridens Pirlot, 1938 (original designation).

Lateral cephalic lobes and article 6 of pereopods 1—2 elongate (com-
bining character); accessory flagellum absent; article 3 of antenna 1
equal to or slightly longer than article 1; gnathopods subchelate;
uropod 3 uniramous, ramus shorter than peduncle. Species: 1,
Indonesia, littoral.

Cheiriphotis Walker

Cheiriphotis Walker, 1904.

Type-species: Melita megacheles Giles, 1885 (monotypy). See J. L.
Barnard, 1962a.

Accessory flagellum 3- or more articulate; article 3 of antenna 1
shorter than article 1; gnathopods subchelate; wropod 3 scale-like,
peduncle plate-like (Cheirimedeia), outer ramus equal to or shorter
than peduncle, inner ramus half as long as outer or absent; cozrae
1-4 of varying sizes and shapes (Microprotopus). Species: 2, tropical
Indo-Pacific, littoral.

Chevalia Walker

Chevalia Walker, 1904.
Neophotis Stout, 1913.

Type-species: C. aviculae Walker, 1904 (monotypy). See J. L.
Barnard, 1962a.

Accessory flagellum 2-articulate; article 3 of antenna 1 slightly
shorter than article 1; gnathopods subchelate, gnathopod 1 poorly;
uropod 3 short, biramous, rami subequal to each other, equal to pe-
duncle in length; pleonites 4 and 5 coalesced. Species: 1, tropical
Indo-Pacific, littoral.

Gammaropsis Liljeborg, new synonymy

(Gammaropsis) Liljeborg, 1855 [subgenus].

Eurystheus Bate, 1857a.—Stebbing, 1906.

Paranaenia Chilton, 1884.

Maeroides Walker, 1898.

Audulla Chevreux, 1901a [a valid subgenus].
Fimbriella Stout, 1913.

Pseudeurystheus Schellenberg, 1931 [a valid subgenus].

Type-species: Gammarus maculatus Johnston, 1827 (selected by
Boeck, 1876, with subsequent synonymy). See Sars, 1895.

Accessory flagellum 3- or more articulate; article 3 of antenna 1
equal to or longer than article 1; gnathopods subchelate; uropod 3

DD U.S. NATIONAL MUSEUM BULLETIN 271

biramous, rami equal to each other, variable in length, generally
equal to or longer than peduncle. Species: 53, cosmopolitan, littoral

to bathyal.
Subgenus Audulla Chevreux (190la): Like Gammaropsis but
flagellum of antenna 2 stout. Species: 1, tropical Indian Ocean,

littoral.

Subgenus Pseudeurystheus Schellenberg (1931): Like Gammaropsis
but article 5 of male gnathopod 2 at least 1.6 times as long as article 6.
Species: 1, subantarctic, littoral to 310 m.

Goesia Boeck

Goesia Boeck, 1871.—Stebbing, 1906.

Type-species: Autonoe depressa Goés, 1866 (monotypy). See
Stephensen, 1942, 1944a.

Accessory flagellum absent; article 3 of antenna 1 shorter than ar-
ticle 1; gnathopods subchelate, gnathopod 2 poorly; uropod 3 short,
biramous, rami equal to each other, subequal to peduncle. Species:
1, arctic-subarctic, littoral.

Haplocheira Haswell

Haplocheira Haswell, 1880b.—Stebbing, 1906.

Type-species: Gammarus barbimanus Thomson, 1879 (monotypy
and subsequent synonymy). See Stebbing, 1888.

Accessory flagellum 2- or more articulate; article 3 of antenna 1
shorter than article 1; gnathopods simple; uropod 3 biramous, outer
ramus subequal to peduncle, inner ramus very short. Species: 3,
subantarctic-antiboreal, littoral to bathyal (457 m).

Isaea Milne Edwards

Isaea Milne Edwards, 1830.—Stebbing, 1906.

Type-species: J. montagui Milne Edwards, 1830 (monotypy). See
Chevreux and Fage, 1925.

Accessory flagellum 3- or more articulate; article 3 of antenna 1
equal to article 1 in length; gnathopods subchelate; uropod 3 biramous,
rami equal to each other, slightly longer than peduncle; pereopods
1-6 slightly prehensile or subchelate, or article 6 widened distally.
Species: 4, N. Atlantic, littoral, some species inhabit decapods.

Kermystheus J. L. Barnard

Kermystheus J. L. Barnard, 1962a.

Type-species: Podoceropsis kermadeci Stebbing, 1888 (original
designation).

MARINE GAMMARIDEAN AMPHIPODA DHS)

Accessory flagellum a short, 1-articulate scale; article 3 of antenna
1 equal to or longer than article 1; gnathopods subchelate, gnathopod
1 occasionally poorly so; uropod 3 apparently biramous, rami sub-
equal to each other and to peduncle. Species: 2, Kerguelen Island
and California, littoral to bathyal (1,152 m).

Kuphocheira K. H. Barnard
Kuphocheira K. H. Barnard, 1931; 19382.

Type-species: KK. setimanus K. H. Barnard, 1931 (original des-
ignation); 1932.

Accessory flagellum absent; [article 3 of antenna 1 undescribed];
enathopods simple, dactyl of gnathopod 2 reduced in size and setose;
uropod 3 uniramous, ramus shorter than peduncle. Species: 1, 5.
Orkney Islands, littoral.

Megamphopus Norman

Megamphopus Norman, 1869a.—Stebbing, 1906.

(Segamphopus) J. L. Barnard, 1962a [subgenus].

[Bonnierella Chevreux, 1900], as possible synonym, but also assigned to Ischyro-
ceridae.

Type-species: MM. cornutws Norman, 1869a (original designation).
See Sars, 1895.

Accessory flagellum 1- or 2-articulate, article 1 elongate; article 3
of antenna 1 equal to or longer than article 1; gnathopods subchelate,
occasionally poorly; uropod 3 biramous, rami subequal to each other,
longer than peduncle. Species: 15, biboreal, littoral to bathyal
(385 m).

Subgenus Bonnierella Chevreux (1900): Like Megamphopus but
article 2 of pereopods 3-5 with parallel edges. This subgenus also is
assigned as a genus to the Ischyroceridae. Species: 5, bathyal to
abyssal.

Subgenus Segamphopus J. L. Barnard (1962a): Like Megamphopus
but article 5 of gnathopod 2 at least 1.6 times as long as article 6.
Species: 1, S. Georgia Island, littoral.

Microphotis Ruffo
Microphotis Ruffo, 1952.

Type-species: MM. blachei Ruffo, 1952 (monotypy).

Accessory flagellum absent; article 3 of antenna 1 equal to article
1; gnathopods subchelate; uropod 3 uniramous, ramus shorter than
peduncle. Species: 1, S. China Sea, littoral.

274. U.S. NATIONAL MUSEUM BULLETIN 271

Microprotopus Norman

?Dercothoe Dana, 1852b.
Microprotopus Norman, 1867b.—Stebbing, 1906.
Orthopalame Hoek, 1879.

Type-species: MM. maculatus Norman, 1867b (monotypy). See Sars,
1895.

Accessory flagellum 2-articulate, article 1 elongate; article 3 of
antenna 1 shorter than article 1; gnathopods subchelate; uropod 3
uniramous, ramus slightly longer than peduncle. Species: 2, boreal
to subtropical E. Atlantic and Mediterranean, littoral.

Photis Kr¢éyer

Photis Kroyer, 1842.—Stebbing, 1906.
Eiscladus Bate and Westwood, 1863. Also Heiscladus, lapsus by Norman, 1869a.

Type-species: P. reinhardi Kroyer, 1842 (monotypy). See Sars,
1895.

Accessory flagellum absent or formed of a vestigial article or scale;
article 3 of antenna 1 equal to or longer than article 1; gnathopods
subchelate; uropod 3 biramous, outer ramus equal to or shorter than
peduncle, inner ramus much shorter than outer ramus. Species: 41,
cosmopolitan, littoral (rare to abyssal).

Podoceropsis Boeck

Podoceropsis Boeck, 1861.—Stebbing, 1906.
Naenia Bate, 1862 (homonym, Lepidoptera).
Xenoclea Boeck, 1871.
Type-species: P. sophia Boeck, 1861 (monotypy). See Sars, 1895.
Accessory flagellum absent; article 3 of antenna 1 equal to or longer
than article 1; gnathopods subchelate; uropod 3 biramous, rami equal
to each other, equal to peduncle in length. Species: 6, Atlantic,
littoral to abyssal.

Protomedeia Kr¢yer ©

Protomedeia Krgyer, 1842.—Stebbing, 1906.
Cheirimedeia J. L. Barnard, 1962a [valid subgenus].
Type-species: P. fasciata Kroyer, 1842 (monotypy). See Sars, 1895.
Accessory flagellum 3- or more articulate; article 3 of antenna 1
shorter than article 1; gnathopods subchelate; uropod 3 biramous,
rami subequal to each other, subequal to peduncle. Species: 14,
subarctic, boreal Pacific (one Atlantic), littoral (?to bathyal).
Subgenus Cheirimedeia J. L. Barnard (1962a): Like Protomedeia
but inner ramus of uropod 3 half as long as outer ramus. Species: 2,
boreal Pacific.

MARINE GAMMARIDEAN AMPHIPODA 275

Ischyroceridae

FicurEs 107, 108

Diacnosis.—Accessory flagellum variable; telson short, entire,
fleshy; coxa 4 not excavate posteriorly except in one genus; pereopods
glandular; uropod 3 rarely projecting beyond uropods 1 and 2, peduncle
elongate, rami styliform, shorter than peduncle, outer ramus usually
uncinate, either with hooked tip, or hooked spine at tip, or denticles.
See Isaeidae, Ampithoidae, Calliopiidae, Pleustidae, Corophiidae.

Description.—Accessory flagellum varying from uniarticulate to
multiarticulate; body smooth; rostrum absent; coxae rounded or
quadrate below, varying from long to short, fourth not excavate
posteriorly (with exception in Bathyphotis); mouthparts basic, possibly
epistome always acutely produced anteriorly; gnathopods usually
powerfully subchelate; structure of uropod 3 as in diagnosis but
merging with conditions typical of Isaeidae by tendencies to elongation
of rami and lack of uncination; telson entire, subcircular or nearly
square, thick.

RELATIONSHIP.—The Isaeidae (= Photidae) differ from the Ischy-
roceridae by the lack of uncination on the outer ramus of uropod 3,
the longer rami and shortened peduncles, but some species now
assigned to Isaeidae bear third uropods merging with the Ischyro-
ceridae.

Calliopiids and pleustids have a nonfleshy telson and a short
peduncle of uropod 3.

Ericthonius in the Corophiidae has an ischyrocerid uropod 3 without
an inner ramus. Dryopoides and Paradryope in the Aoridae have
third uropods which technically are ischyrocerid. Parapherusa in the
Gammaridae lacks pereopodal glands but has uropod 3 similar to
that of ischyrocerids.

Key to the Genera of Ischyroceridae

1. Coxae minute, not touching. .. . .. . . . Bogenfelsia

Coxae small or large, overlapping or banal foocite AS esis a Em Ie oa MESA hte
Gnathopod 1 distinctly larger than 2

Dryopoides (Aoridae) and Paradryope (Aoridae)

Gnathopod 1 smaller than 2 or gnathopods subequal to each other . . . 3

3. Second articles of pereopods 3-5 narrowly linear, palp article 4 of maxilliped

claw-like, longer than article3.... . . . . . . Bonnierella

Second articles of pereopods 3-5 suboval or br wudly Peoria ular: palp article 4

of maxilliped shorter than article 3, blunt,! or subconical,! (setose) (fig.

OMS) Mee ANE Cc RE RAR EMRE RES Mt MORON RN ERNIE cor MeN) jhe aN ENA A aa

i)

1 Not including terminal spine.

276 U.S. NATIONAL MUSEUM BULLETIN 271

4. Article 5 of gnathopod 1 much longer than article6.......... 5
Article 5 of gnathopod 1 shorter than or subequal to article6. ..... 6

5. Spines on outer plate of first maxilla nine or more, article 4 of pereopod 3 not
wider than on pereopods 4-5 .... . . . . . . Pseudischyrocerus
Spines on outer plate of maxilla 1 reduced i four, article 4 of pereopod 3
twice as wide as on pereopods 4-5 . ... . . . . . . Bathyphotis

6. Accessory flagellum a small scale, articulate or Hot (Gg 107c). . Parajassa
Accessory flagellum elongate (figs. 107d,e) . . . . se) ie Rie Sua,

7. Coxa 5 subequal to 6 in length, coxa 1 half as long as coxa 2. . Microjassa
Coxa 5 much longer than 6, coxa 1 three fourths as long as coxa 2. . . . 8

8. Pereopods subchelate (fig. 108d) ............ .. .. .Isaeopsis
Pereopods simple (fig. 107b) . ... . Reiner 8)

9. Outer ramus of uropod 3 with basally ranma, hooked femiaal spine,
distolateral margin of ramus with one to three large, reverted, sharp scales
or several denticles (fig. 108k); male gnathopod 2 usually with large tooth
on proximal portion of palm and with tooth on distal portion of palm.

Jassa ?

Outer ramus of uropod 3 with or without basally immersed terminal spine,
never hooked, distolateral margin of ramus with 0-7+ perpendicular, blunt,
usually extremely small denticles (fig. 1087); male gnathopod 2 lacking
proximal palmer tooth but often with distal palmar process. Ischyrocerus ”

2 These genera extremely difficult to distinguish except in adult males.

Genera of Ischyroceridae

Bathyphotis Stephensen
Bathyphotis Stephensen, 1944b.

Type-species: JB. tridentata Stephensen, 1944b (monotypy).

Accessory flagellum multiarticulate, elongate; article 5 of gnathopod
1 longer than article 6; coxa 1 more than three fourths as long as
coxa 2, coxa 4 subequal to coxa 6 in length, coxa 4 excavate posteriorly;
article 4 of pereopod 8 tuice as wide as on pereopods 4-5; outer plate of
maxilla 1 with four spines (Pseudischyrocerus). Species: 1, subarctic
Atlantic, bathyal (1,096 m).

Bogenfelsia J. L. Barnard
Bogenfelsia J. L. Barnard, 1962d.

Type-species: 8B. incisa J. L. Barnard, 1962d (original designation).

[Accessory flagellum unknown]; article 5 of gnathopod 1 longer than
article 6; coxae minute, not touching, equal in length. Species: 1,
antiboreal Atlantic, abyssal (4,893 m).

Bonnierella Chevreux
Bonnierella Chevreux, 1900.

Type-species: Podoceropsis abyssi Chevreux, 1887 (monotypy).

MARINE GAMMARIDEAN AMPHIPODA ORT

g Di \ OE

Ficure 107.—Ischyroceridae: a, Microjassa macrocoxa Shoemaker (1942); b, Ischyrocerus
anguipes Kréyer (Sars, 1895, pl. 209). Accessory flagella: c, Parajassa pelagica (Leach)
(Sars, 1895, pl. 214, as Janassa capillata); d, Jassa falcata (Montagu) (Sars, 1895, pl.
212); e, Ischyrocerus megacheir (Boeck) (Sars, 1895, pl. 211). Mouthparts, Ischyrocerus :
f, upper lip; g, mandible; h, lower lip; 7,7, maxillae 1, 2; &, maxilliped.

278 U.S. NATIONAL MUSEUM BULLETIN 271

Accessory flagellum 2-articulate, elongate; article 6 of gnathopod
1 longer than or equal to article 5; coxa 1 as long as coxa 2, coxa 5
longer than coxa 6; article 2 of pereopods 3-5 narrow, linear, palp
article 4 of marilliped claw-like, longer than article 3 (combining char-
acter). Species: 5, Atlantic, E. Pacific, bathyal to abyssal.

Isaeopsis K. H. Barnard
Tsaeopsis K. H. Barnard, 1916.

Type-species: J. tenaz K. H. Barnard, 1916 (monotypy).

Accessory flagellum 1-articulate; article 6 of gnathopod 1 longer than
article 5; coxa 1 “‘small,”’ coxa 5 longer than coxa 6; pereopods pre-
hensile. Species: 1, S. Africa, littoral.

Ficure 108.—Ischyroceridae: Gnathopod 1: a, Ischyrocerus anguipes Krgyer (Sars, 1895,
pl. 209); b, Jassa falcata (Montagu) (Sars, 1895, pl. 212); c, Pseudischyrocerus denticauda
Schellenberg (1931). Pereopod 1: d, Isaeopsis tenax K.. H. Barnard (1916). Gnathopod
2: ¢, Jassa, note sharp distal palmar tooth and poor development of proximal tooth as an
exception to couplet 9 of key; f, Ischyrocerus. Pereopods 3, 4, 5: g,h,1, Microjassa macro-
coxa Shoemaker (1942). Uropod 3:7, Ischyrocerus; k, Jassa. Telson: 1, [schyrocerus.

MARINE GAMMARIDEAN AMPHIPODA 279

Ischyrocerus Kr¢yer
Ischyrocerus Kr¢yer, 1838.—Stebbing, 1906.

Type-species: J. anguipes Krgyer, 1838 (monotypy). See Sars,
1895.

Accessory flagellum 2-articulate, elongate; article 6 of gnathopod 1
longer than article 5; coxa 1 more than three fourths as long as coxa 2,
coxa 5 longer than coxa 6; article 6 of male gnathopod 2 lacking large
posteroproximal tooth; outer ramus of uropod 3 with or without basally
immersed, unhooked terminal spine, distolateral margin usually with
perpendicular, blunt, mostly very minute denticles (Jassa). Species:
32, possibly cosmopolitan but primarily boreal, littoral to bathyal
(2,090 m).

Jassa Leach

Jassa Leach, 1814a.—Stebbing, 1906.
Lusyta Nardo, 1847.

Cratophium Dana, 1852b.

Macleayia Haswell, 1880a.

Wyvillea Haswell, 1880c.

Bruzeliella Norman, 1905.

Hemiyassa Walker, 1907.

Type-species: Cancer falcatus Montagu, 1808 (selected by
Chevreux and Fage, 1925; indicated by subsequent montoypy
through elimination by Stebbing, 1899a, and thereafter synonomy
of species). See Sars, 1895 (as Podocerus falcatus).

Accessory flagellum 2-articulate, elongate; article 6 of gnathopod
1 longer than article 5; coxa 1 more than three fourths as long as
coxa 2, coxa 5 longer than coxa 6; article 6 of male gnathopod 2 usually
bearing a large posteroproximal tooth; outer ramus of uropod 3 with
distal, basally immersed, hooked spine, distolateral margin with 1-3
large, sharp, reverted scales (Ischyrocerus). Species: 7, cosmopolitan,
littoral.

Microjassa Stebbing

Microjassa Stebbing, 1899b; 1906.

Type-species: Podocerus cumbrensis Stebbing and _ Robertson,
1891 (monotypy). See Chevreux and Fage, 1925.

Accessory flagellum 1-articulate, very small; article 6 of gnathopod
1 longer than article 5; coxa 1 less than half as long as coxa 2; coxa
5 not longer than coxa 6. Species: 2, E. Atlantic, E. Pacific, littoral.

Parajassa Stebbing

Janassa Boeck, 1871 (homonym, Pisces).
Parajassa Stebbing 1899a; 1906 (new name for Janassa).

280 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: Jassa pelagica Leach, 1814a (monotypy of Boeck
and subsequent synonymy). See Sars, 1895 (As Janassa capillata).
Accessory flagellum a small tubercle; article 6 of gnathopod 1
longer than article 5; coxae 1 and 2 equal in length, coxa 5 longer
than coxa 6. Species: 4, Atlantic, E. Pacific, amphiboreal, littoral.

Pseudischyrocerus Schellenberg
Pseudischyrocerus Schellenberg, 1931.

Type-species: P. denticauda Schellenberg, 1931 (present selection).

Accessory flagellum multiarticulate, elongate; article 5 of gnathopod
1 longer than article 6; coxae 1 and 2 equal in length; coxa 5 probably
longer than coxa 6; article 4 of pereopod 3 wider than on pereopods
4-5; outer plate of maxilla 1 with nine spines (Bathyphotis). Species:
1, antiboreal, littoral.

Kuriidae
[J. L. Barnard, 1964c]

FicureE 109

Driaenosis.—Accessory flagellum absent; body laterally compressed ;
urosomites all coalesced, mandible lacking palp, molar well developed,
triturative; gnathopods feeble, subchelate; uropod 3 uniramous; telson
deeply cleft. Monogeneric. See Phliantidae, Talitroidea, Dogielinotidae,
EKophlhiantidae, Prophliantidae, Pagetinidae.

DescripTion.—Body laterally compressed, all urosomal segments
coalesced; mandible lacking palp, molar well developed; [lower lip
and maxillae unknown]; maxilliped with outer lobes strongly reduced ;
egnathopods feeble, slender, subchelate; uropod 3 uniramous; telson
short, deeply cleft.

RELATIONSHIP.—The Talitroidea have discrete urosomal segments
and well-developed outer plates on the maxillipeds.

The molar of the mandible is evanescent in the Prophliantidae
and Pagetinidae.

Eophhantidae have cylindrical bodies.

The Phliantidae have evanescent mandibular molars and well-
developed outer plates of the maxillipeds. Probably the Kuriidae
should eventually be assigned as a subfamily to Phliantidae to demon-
strate their close relationship. The Kuriidae show a link between
Phliantidae and Talitroidea.

The Dexaminidae have only the last two urosomal segments co-
alesced and possess biramous third uropods but the Kuriidae other-
wise demonstrate a link between the Dexaminidae and the Talitroidea.

MARINE GAMMARIDEAN AMPHIPODA 281
Genera of Kuriidae

Kuria Walker and Scott

Kuria Walker and Scott, 1903.

Type-species: K. longimana Walker and Scott, 1903 (monotypy).
Species: 1, Abd-el-Kuri, littoral.

ye

Ficure 109.—Kuriidae: Kurita longimanus Walker and Scott (1903): a, lateral aspect,
note overlap of pereopods 3-5 probably erroneous; b, mandible; c, maxilliped; d, uropod 3;
e, telson; f,g, gnathopods 1, 2.

Lafystidae
FicurE 110

Diacnosis.—Accessory flagellum absent; mouthparts arranged into
a conical bundle; mandibular molar absent; palp of maxilla 1 reduced
to a single tiny article; maxilliped with 2-articulate palp; coxa 4 acu-

282 U.S. NATIONAL MUSEUM BULLETIN 271

minate, cozae 1-3 quadrate; gnathopod 1 simple; telson oval, short,
entire. Monogeneric. See Laphystiopsidae, Acanthonotozomatidae.

Description.—Accessory flagellum absent; body broad, not carin-
ate, urosomite 1 elongate; rostrum large, mouthparts arranged into a
conical bundle; mandible lacking molar and spine row; lower lip lack-
ing inner lobes; palp of maxilla 1 reduced to a single article; palp of
maxilliped reduced to two small articles; coxae 1-3 quadrate, coxa 4
acuminate; gnathopods feeble, 1 simple, 2 weakly subchelate; uro-
pod 3 biramous; telson small, oval, entire.

ReELATIONSHIP.—This family is so closely related to Acanthonoto-
zomatidae that they might be joined together as subfamilies.

The Laphystiopsidae bear a 4-articulate maxillipedal palp, a man-

Ficure 110.—Lafystiidae: Lafystius sturionis Krgyer (Sars, 1895, pl. 134): a, lateral;
b, dorsal view of head; c, upper lip; d, mandible; ¢, lower lip; f,g, maxillae 1, 2; 4,maxilliped;
1,7, gnathopods 1, 2; k, uropod 3; J, telson.

MARINE GAMMARIDEAN AMPHIPODA 283

dibular molar, a large palp on the first maxilla and an incised, broad-
ened upper lip.

Superficially, the Lafystiidae resemble several other families through
the Laphystiopsidae [which should be consulted]. As discussed therein
the Laphystiopsidae are still considered distinct from Lafystiidae,
necessitating the removal of Prolaphystiopsis to the Laphystiopsidae.

Genera of Laphystiidae

Lafystius Kr¢gyer

Lafystius Krgyer, 1842.—Stebbing, 1906.
Darwinia Bate, 1857a.
?Dermophilus Beneden and Bessels, 1870.
?Ichthyomyzocus Hesse, 1878.
Type-species: JL. sturionis Krgyer, 1842 (monotypy). See Sars,

1895.
Species: 1, boreal N.E. Atlantic, littoral, occurring on fishes.

Laphystiopsidae

FicurE 111

DiaGcnosis.—Accessory flagellum absent; mandibular molar non-
triturative; gnathopods feeble, simple; telson entire. See Isaeidae,
Stilipedidae, Oedicerotidae, Calliopiidae, Pleustidae, Lafystiidae.

Description.—Accessory flagellum absent; body broad, carinate
only on a few pleonites; urosomite 1 elongate or not; rostrum large
and spatulate or absent; upper lip incised; mandible bearing molar
process, simple or slightly triturative; lower lip with or without inner
lobes; palp of maxilla 1 large, 1- or 2-articulate; maxilliped with
small 4-articulate palp; coxae variable, either very short and rectan-
cular and serially disjunct or of medium length, quadrate or slightly
acuminate and overlapping; gnathopods feeble, simple; outer rami of
uropods 1-2 slightly shortened; uropod 3 biramous, peduncle short;
telson short or of medium length, oval, entire.

Reationsurp.—The Laphystiopsidae are simply Calliopiidae- and
Pleustidae-like organisms with feeble, simple gnathopods, feeble,
poorly setose maxillipedal palps and poorly developed mandibular
molars. Even the lower lip of the type-genus is similar to that of
Pleustidae, although Prolaphystius lacks inner lobes. Even Cleippides
and Regalia in the Calliopiidae have simple gnathopods.

The Lafystiidae faintly resemble Laphystiopsidae but the latter
have lost the mandibular molar and spine row, two articles of the

285-135 O - 69 - 19

284 U.S. NATIONAL MUSEUM BULLETIN 271
maxillipedal palp, and have the first maxillary palp reduced in size.
The unique species of lafystiid, however, has retained the subchelate
condition of its second gnathopod, although it is feeble.

The uniformity of the Laphystiopsidae is complicated by the loss
of the rostrum in Prolaphystvus and that genus warrants further in-
vestigation on its familial assignments. Prolaphystius has an elongate
urosomite 1 like Podoceridae but otherwise bears no resemblance to
that family. It has been included in the key to Calliopiidae where it

seems to have strong affinities.
The Astyridae bear a small accessory flagellum and broadly sepa-

rated outer lobes on the lower lip.
The Oedicerotidae warrant close inspection of their slightly sub-

chelate gnathopods as a feature distinguishing them from laphysti-

=
=

> ——
——
ee a oo ry

Cs
Laphystiopsis planifrons Sars (1895, pl. 135): a, lateral;

Figure 111.—Laphystiopsidae:
b, upper lip; c, mandible; d, lower lip; e,f, maxillae 1, 2; g, maxilliped; h,7, gnathopods 1, 2;
Prolaphystius tsopodops K. H. Barnard (1930): /, head, right side,

j, uropod 3; k, telson.
stippled; m, maxilla 1; n, mandible; 0, telson.

MARINE GAMMARIDEAN AMPHIPODA 285

opsids. Oedicerotids have an elongate peduncle of uropod 3, equal
rami of uropods 1-2, unnotched upper lips, strongly setose appendages
and disproportionately long fifth pereopods.

Stilipedidae have a broad, pardaliscid mandibular incisor, no molar
and foliaceous maxillae.

Isaeids and their relatives have strongly triturative mandibular
molars and usually have subchelate and/or large gnathopods.

Key to the Genera of Laphystiopsidae

(Ae AGlarcelrostrumaspresent, (liga vWblla) ior fa bo: fee meters) a Poa ol Waele eas 2
Rostrumyalosemte chica IWl)) newer eens ici ae nts lees Prolaphystius
Zealpiotimaxilla libiarticulate: ies Tila) yo one. Prolaphystiopsis
Halprommaxitia leumanticulate: (ig: ile). 29. 8 2 Laphystiopsis

Genera of Laphystiopsidae

Laphystiopsis Sars
Laphystiopsis Sars, 1895.—Stebbing, 1906.

Type-species: L. planifrons Sars, 1895 (monotypy).

Rostrum well developed; mandibular molar conical, unridged; palp
of maxilla 1 uniarticulate; telson short, oval; pleonite 3 not forming a
projecting shelf; coxae not touching serially. Species: 3, boreal to
China Sea, littoral to bathyal (900 m).

Prolaphystiopsis Schellenberg
Prolaphystiopsis Schellenberg, 1931.

Type-species: P. platyceras Schellenberg, 1931 (monotypy).

Rostrum well developed; mandibular molar truncate, poorly ridged;
palp of maxilla 1 biarticulate; telson short, oval; pleonite 3 not
forming a projecting shelf; coxae not touching serially. Species: 1,
subantarctic, littoral.

Prolaphystius K. H. Barnard
Prolaphystius K. H. Barnard, 1930.

Type-species: P. isopodops K. H. Barnard, 1930 (monotypy).

Rostrum absent; mandibular molar truncate, poorly ridged; palp
of maxilla 1 biarticulate; telson long, linguiform; pleonite 3 forming a
shelf-like projection; coxae of medium size, touching serially. Species:
1, antarctic, bathyal (406 m).

286 U.S. NATIONAL MUSEUM BULLETIN 271

Lepechinellidae

Ficure 112

DraGnosis.—One or more and usually all of coxae 1-4 acutely
pointed below or anteriorly, occasionally bifid; accessory flagellum
composed of one spine-like article; pleonites 5-6 ?coalesced. See
Paramphithoidae, Atylidae, Eusiridae, Pleustidae.

Description.—Body processiferous; accessory flagellum 1-articu-
late, spine-like; rostrum conspicuous; upper lip probably always
incised slightly; molar of mandible triturative, palp 3-articulate,
slender, short or long; lower lip apparently with coalesced inner lobes,
remaining mouthparts basic; gnathopods feeble, slender, subchelate;
coxae of medium length or shortened, usually acuminate, often bifid;
rami of uropod 3 greatly elongate, very slender, 3-6 times as long as
peduncle; telson short, cleft, lobes diverging.

ReLationsuip.—Originally this family was a part of the Param-
phithoidae, because the type-genus Lepechinella was placed there by
Stebbing (1908a); the junior synonym Dorbanella Chevreux (1914)
was placed in the Tironidae but Schellenberg (1925a) removed that to
a family Dorbanellidae, later to take the name of the senior synonym
of the type-genus. Lepechinellidae resemble Paramphithoidae but
presumably all species have pleonites 5-6 coalesced. The inner lobes
of the lower lip of lepechinellids apparently are always fused together
in contrast to the Paramphithoidae.

The coxae of Atylidae are elongate and not acuminate distally and
their gnathopods are shorter and stouter than those of Lepechinellidae.
Coxae 2-4 of Paralepechinella are scarcely, if at all acuminate.

No pleustid has coalesced urosomal segments nor a_ spine-like
accessory flagellum. That family has characteristic lower lips.

Dexaminidae, Kuriidae, Prophliantidae all lack mandibular palps.

Key to the Genera of Lepechinellidae

1. Mandibular palp article 3 much shorter than article 2 (fig. 112c).
Lepechinella
Mandibular palp article 3 immense, much longer than article 2 (fig. 112h).
Paralepechinella

Genera of Lepechinellidae

Lepechinella Stebbing

Lepechinella Stebbing, 1908a.
Dorbanella Chevreux, 1914.

Type-species: L. chrysotheras Stebbing, 1908a (monotypy).
Species: 13, cosmopolitan, bathyal and abyssal.

MARINE GAMMARIDEAN AMPHIPODA 287

Ficure 112.—Lepechinellidae: a, Lepechinella bierii J. L. Barnard (1957c). Accessory
flagellum: b, L. bierii. Mouthparts, Lepechinella chrysotheras Stebbing (1908a): c, man-
dible; d, lower lip; ¢,f, maxillae 1, 2; g, maxilliped. Mandible: h, Paralepechinella longt-
palpa Pirlot (1933a). Gnathopods 1-2: 7,7, L. bierit. Uropod 3: k, L. bierit. Telson:
1, L. chrysotheras.

288 U.S. NATIONAL MUSEUM BULLETIN 271

Paralepechinella Pirlot
Paralepechinella Pirlot, 1933a.

Type-species: P. longipalpa Pirlot, 1933a (original designation).
Species: 1, Indonesia, bathyal (1,301 m).

Ficure 113.—Leucothoidae: a, Leucothoe spinicarpa (Abildgaard) (Sars, 1895, pl. 100);
b, Leucothoella bannwarthi Schellenberg (1928); c, Leucothoides pottst Shoemaker (1933a).
Upper lip: d, Leucothoe. Mandible: e, Leucothoe; f, Leucothoides.

MARINE GAMMARIDEAN AMPHIPODA 289

Leucothoidae

Figures 113, 114

Diaenosis.—Accessory flagellum vestigial, 1- or 2-articulate, very
small; mandible lacking molar; outer plates of maxilliped very small,
probably never larger than inner plates; telson entire; gnathopod 1
carpochelate. See Anamixidae, Sebidae, Amphilochidae, Cressidae,
Thaumatelsonidae.

DescripTion.—Body lacking dorsal processes; accessory flagellum
vestigial; epistome probably always produced in front; upper lip
probably always asymmetrically incised; mandibles lacking molar,
palp 1- or 3-articulate; maxillae basic; maxilliped with very small
outer plates, partially fused inner plates and long slender palp;
gnathopod 1 carpochelate, immovable finger long and narrow, gnatho-
pod 2 larger than 1, subchelate, article 5 elongate and guarding
article 6 posteriorly; coxae variable; uropod 2 shortened; uropod 3
with styliform rami, peduncle elongate, rami not longer than peduncle;
telson entire, not longer than peduncle of uropod 3.

ReuatTionsHip.—The Anamixidae lack maxillae and recognizable
mandibles.

The Cressidae, Thaumatelsonidae, and Stenothoidae have unira-
mous third uropods and subchelate first gnathopods.

The Sebidae have uniramous third uropods and chelate second
enathopods.

Key to the Genera of Leucothoidae

1. Coxa 2 covering most of coxa 1 (fig. 113c), mandibular palp 1-articulate
Gils, WISI) ie os . . . Leucothoides

Coxa 1 not concealed, Sibu male Be icuiate ae NBT AOD Masse ties Ais 074

2. Coxa 2 very short, broader than long, acute anteriorly (fig. 1130).
Leucothoella

Coxa 2 at least as long as broad, rounded below and anteriorly (fig. ll3a). .3

3. Outer plate of maxilliped reaching less than halfway along palp article 1
Gig, WMA SG a . . . . . Leucothoe

Outer plate of Tealliped P aehine almnoet 46 ond of palp article 1.
Paraleucothoe

Genera of Leucothoidae

Leucothoe Leach

Leucothoe Leach, 1814a.—Stebbing, 1906.
Cuvieria Leach, 1814a (nomen nudum).
Lycesta Savigny, 1816.
Type-species: Gammarus spinicarpus Abildgaard, 1789 (monotypy
and subsequent synonymy). See Sars, 1895.

290 U.S. NATIONAL MUSEUM BULLETIN 271

Coxa 2 at least as long as broad, rounded ventrally and anteriorly,
coxa 1 not concealed; mandibular palp 3-articulate; outer plate of
maxilliped reaching less than halfway along palp article 1. Species:
25, cosmopolitan, littoral to abyssal.

Ficure 114.—Leucothoidae: Leucothoe spinicarpa (Abildgaard) (Sars, 1895, pl. 100):
a, lower lip; b,c, maxillae 1, 2; d, maxilliped; ¢, uropod 3; f, telson; g,h, gnathopods 1, 2.
Leucothoides pottst Shoemaker (1933a): 7, telson; 7, gnathopod 2.

Leucothoella Schellenberg
Leucothoella Schellenberg, 1928b.

Type-species: ZL. bannwarthi Schellenberg, 1928b (monotypy)-

Coxa 2 very short, broader than long, acute anteriorly, coxa 1 not
concealed; mandibular palp 3-articulate; outer plate of maxilliped
reaching less than halfway along palp article 1. Species: 1, tropics,
Red Sea to Fiji, littoral.

Leucothoides Shoemaker
Leucothoides Shoemaker, 1933a.

Type-species: L. pottsi Shoemaker, 1933a (monotypy).

Coxa 2 at least as long as broad, rounded ventrally and anteriorly,
coxa 2 covering most of coxa 1; mandibular palp l-articulate; outer
plate of maxilliped reaching less than halfway along palp article 1.
Species: 2, tropical Pacific, Caribbean, littoral.

MARINE GAMMARIDEAN AMPHIPODA 9291

Paraleucothoe Stebbing

Paraleucothoe Stebbing, 1899d.—Stebbing, 1906.

Type-species: Leucothoe novaehollandiae Haswell, 1880c (original
designation).

Coxa 2 at least as long as broad, rounded ventrally and anteriorly,
coxa 1 not concealed; mandibular palp 3-articulate; outer plate of
maxilliped reaching almost to end of palp article 1. Species: 1,
Australia, littoral.

Liljeborgiidae
Fiacures 115, 116

Draanosis.—Molar of mandible poorly developed, not triturative,
accessory flagellum 2- or more articulate. See Gammaridae, Eusiridae,
Pleustidae, Haustoriidae, Astyridae, Vitjazianidae.

Descriprion.—Accessory flagellum always present, 2- or more
articulate; rostrum inconspicuous; coxae long or of medium length;
upper lip possibly slightly emarginate, not conspicuously so; mandibles
always bearing 3-articulate palp with article 1 often elongate and as
long as article 3 (except Jdunella forming a link to Gammaridae);
molar of mandible obsolescent, nontriturative; remaining mouthparts
basic; gnathopods powerfully subchelate; uropod 3 with flattened,
lanceolate rami; telson deeply cleft, not greatly elongate, apices of
lobes bifid.

RELATIONSHIP.—In the Gammaridae only the genus Parelasmopus
has an elongate mandibular palp article 1 but its mandibular molar
has a well-developed grinding surface, unlike that of Liljeborgiidae.
Apart from the mandible the Liljeborgiidae are like the Gammaridae.

The Astyridae differ from the Liljeborgiidae by the overall appear-
ance and the presence of inner lobes or a broad medial space on the
lower lip (see figures 32 and 41). Astyrids have feeble gnathopods,
whereas those of liljeborgiids are powerfully developed. The rami of
uropod 3 of Astyridae are much more elongate than those of Lilje-
borgiidae.

Some genera of Pleustidae have poorly developed mandibular
molars as in Liljeborgiidae and the lower lips are similar. Pleustids,
however, have vestigial or no accessory flagella and usually have
uncleft telsons, but one species of Austropleustes confounds the defini-
tion. See Gammaridean Family Key G for specific distinctions.

The Eusiridae always have the accessory flagellum 2-articulate or
less and this conflicts only with the genus Listriella in the Liljeborgiidae.
But in contrast to the Eusiridae, Listriella usually has a 2-articulate

292

U.S. NATIONAL MUSEUM BULLETIN 271

outer ramus of uropod 3 with an elongated article 1 of the mandibular
palp. Lusirella and Eusiropsis are the only eusirids having an obsoles-
cent mandibular molar and they are clearly eusirids for their possession
of calceoli. Eusirids have the outer ramus of uropods 1 and 2 shortened;
in liljeborgiids this shortening is slightly evident only on uropod 2.

One genus of Amphilochidae, Pseudamphilochus, bears resemblance
to Liljeborgiidae for its unreduced coxa 1, cleft telson, weak mandi-
bular molar, and moderately powerful gnathopods. Pseudamphilochus

presumably lacks an accessory flagellum, has a large rostrum, and
nonbifid apices of the telson.

oe
an
as
ee
a

—eN
<5
ol
BE
Se

Ficure 115.—Liljeborgiidae: a, Liljeborgia brevicornis (Bruzelius) (Sars, 1895, pl. 187,
as L. pallida); b, Idunella aequicornis (Sars, 1895, pl. 190).

MARINE GAMMARIDEAN AMPHIPODA 293

Key to the Genera of Liljeborgiidae

1. Gnathopod 1 larger than gnathopod 2 (fig. 115b) . . . . .. . +. =. Idunella

Gnathopod 1 smaller than or equal to gnathopod 2 (fig. 115a) . ES AD,

2. Article 5 of gnathopods 1-2 strongly produced, slender and elongate (fig. 1169) ;

outer ramus of uropod 3 always simple (fig. 1167) .... . . Liljeborgia

Article 5 of gnathopods 1-2 weakly produced, thick and blunt (fig. 116));
outer ramus of uropod 3 usually composed of two articles (fig. 116k)

Listriella

1

Ficure 116.—Liljeborgiidae: Liljeborgia brevicornis (Bruzelius) (Sars, 1895, pl. 187, as
L. pallida): a, upper lip; b, mandible; c, lower lip; d,e, maxillae 1, 2; f, maxilliped; g,
gnathopod 2; h, telson; 1, uropod 3. Gnathopod 2: 7, Listriella goleta J. L. Barnard (1959b).
Uropod 3: k, Listriella diffusa J. L. Barnard (1959b); l, Listriella ertopisa J. L. Barnard
(1959b).

Genera of Liljeborgiidae

Idunella Sars

Idunella Sars, 1895.—Stebbing, 1906.—J. L. Barnard, 1959b.
Sextonia Chevreux, 1920.

Type-species: Liljeboria aequicornis Sars, 1876 (monotypy). See
Sars, 1895.

Species: 4, subarctic, warm-temperate of N. Atlantic, tropical
Indian Ocean, Japan, littoral to bathyal (763 m).

294 U.S. NATIONAL MUSEUM BULLETIN 271

Liljeborgia Bate

Iduna Boeck, 1861 (homonym, Aves).
Liljeborgia Bate, 1862.—Stebbing, 1906.—K. H. Barnard, 1932.
Microplax Liljeborg, 1865 (homonym, Hemiptera).
Lilljeborgiella Schellenberg, 1931.
Type-species: Gammarus pallidus Bate, 1857a (monotypy of Bate,
1862). See Bate and Westwood, 1863.
Species: 28, bipolar and amphiboreal, littoral to abyssal.

Listriella J. L. Barnard

Listriella J. L. Barnard, 1959b.

Type-species: JL. goleta J. L. Barnard, 1959b (original designation).
Species: 8, tropical and warm-temperate Atlantic and E. Pacific,
littoral, often inhabiting tubes of maldanid polychaetes.

Lysianassidae
Figures 117-132

Diacnosis.—Accessory flagellum usually present and multiarticu-
late; article 3 of gnathopod 2 elongate (fig. 3a), gnathopod 2 other-
wise of the typical lysianassid kind and with a fuzz of scales or setules
on articles 5 and 6 (fig. 129); peduncle of antenna 1 short and stout.
See Sebidae, Ochlesidae, Stegocephalidae, Didymocheila (incertae
sedis).

Description.—Accessory flagellum usually present and more than
3-articulate but occasionally vestigial or few in segments; peduncle
of antenna 1 short and stout, articles 2 and 3 usually much shorter
than 1 and occasionally scarcely distinguishable because of tele-
scoping; pereon generally lacking processes, often urosomal segment
1 processiferous; body compact, chitin usually very smooth, porcel-
lanous; mouthparts enormously variable, either chewing or piercing,
often with reduction in mandibular molars, variable in position of
mandibular palps; upper lip and epistome invaluable for identification,
either or both often produced in variable configurations; maxilla 1
generally with 2-articulate palp but occasionally reduced or absent;
maxilla 2 occasionally poorly developed; maxilliped occasionally with
reduced plates and palps; gnathopods usually quite small, 2 mitten-
shaped terminally, with article 3 elongate, article 5 generally with a
“pineapple cushion”’, i.e., posterior margin bulbous and with micro-
scopic texture of a pineapple, structure composed of scales or coarse
setules, article 6 generally covered with coarse setules or tasseled
setules; pereopods generally stout, occasionally prehensile; coxae
usually long, first one or two or three occasionally reduced in size or

MARINE GAMMARIDEAN AMPHIPODA 295

hidden by following coxae; rami of uropod 3 lanceolate, outer often
2-articulate, inner occasionally reduced in size, rarely both absent;
telson cleft, emarginate or entire, long or short; rarely absent.

Gnathopod 2 is an unmistakable mark of this family. Nevertheless,
Sebidae have a chelate gnathopod 2 with elongate article 3 but they
are distinguished from Lysianassidae by the elongate peduncle of
antenna | and the elongate and uniramous uropod 3. Some talitroideans
have a mitten-shaped gnathopod 2 but article 3 is not elongate. Some
Stegocephalidae also have long third articles but all of them have a
characteristic shape of coxae 1-4, the first acuminate, the second
and third narrower than 1, the fourth extremely broad, all four coxae
fitting together tightly, forming a ventral semicircle and not over-
lapping. Stegocephalid mouthparts are highly characteristic although
a few Lysianassidae have mouthparts resembling them. Stegocephalid
mandibles usually have some foliaceous parts, rarely occurring in
Lysianassidae. Stegocephalid mouthparts are always formed into a
conical bundle, rarely occurring in Lysianassidae. Gnathopod 2 is
always simple, whereas few Lysianassidae have a completely non-
subchelate gnathopod 2.

Nomenclatural Changes in Lysianassidae

Alicella scotiae Chilton (1912) is removed to Paralicella.

Aruga Holmes (1908) is synonymized with Lysianassa.

Arugella Pirlot (1936) is synonymized with Lysianassa.

Lakota adversicola K. H. Barnard (1925) is removed to Schisturella.

Lakota rotundatus K. H. Barnard (1925) is removed to Schisturella.

Lysianassa anomala Nicholls (1938) is removed provisionally to
Parambasia.

Lysianopsis Holmes (1903) is synonymized with Lysianassa.

[Pronannonyz] Schellenberg (1953) is also synonymized with Lysian-
assa.

Shoemakerella Pirlot (1936) is synonymized with Lysianassa.

Socarnopsis obesa Chevreux (1927) is removed to Socarnes in view
of Hurley’s (1963) classification, because uropod 3 has the outer
ramus 2-articulate.

Tryphosites cozalis J. Li. Barnard (1962d) is transferred to Tryphosella
Bonnier, and is reflected in Key I, triplicut 7 (p. 304).

Key to the Genera of Lysianassidae

A basic key partitions the genera into several groups further developed
in keys A to Q. This method brings together genera or portions of
genera in different places in order to signal their complicated, net-like

296

U.S. NATIONAL MUSEUM BULLETIN 271

relationships and obviates the use of a long key with more than 100
couplets. Key N is provided as a separate entity for a special group of

genera.
BASIC LYSIANASSID KEY

ie aelsonvabsents (hess W22 ail 24 a) Ne eet sine cnet re ee enh Once eae av

Telson present . . RT AN Hee ses Ae uA SEM ALE tr ert

2. Mandible without distineth incisor iGo 126c) ..... . . .Kerguelenia

Mandible with distinct incisor (figs. 126a,b,d) .... . ii) CaaS

3. From lateral view, mouthparts group strongly and comically ancien below

head, more or less styliform (fig. 123e) ... . Pan aeersenune aan 2" 6}

Mouthparts group usually not strongly or eonielhy projects below head,

parts not styliform [if so, uropod 3 lacking rami ae SVAN iis eects ncn Ae

4. Gnathopod 1 chelate (figs. 129a,e,h,0). . . . : 4 Visuaoaae KCN

Gnathopod 1 subchelate or simple, rarely appearing areen ele Ben ice aD

5. Mandibles: middle of incisor sharply dentate (fig. 1260) ... . .Key D

Mandibles: middle of incisor not dentate .............. 6

Oo Maxilla tilackane: palpy (ies 127b))i ie AGIAN e ms bond (ONY, IS

Maxilla 1 bearing 1- or 2-articulate palp (fig. 126a) . Sioa 1 2 gol Meee TE

7. Both coxae 1 and 2 (and occasionally 3) very small or distinctly smaller

10.

Ii.

12.

13.

than following coxae, largely hidden by following coxae (fig. 122c).
Key F
Neither coxae 2 nor 3 distinctly reduced, both visible (fig. 122d) (occasionally
coxa 1 reduced alone) . . . . . . . Choose either couplet 8 or 21!
Telson entire or emarginate, no syniteall slit (iigss 128pp)) ae ee yaGr
Telson cleft enough to provide slit or facing edges at angle of less than
45 degrees (figs. 128n,0,t) . . . SPUN GAA On ae aE ear
Article 2 of pereopod 3 deeply anGlantaniadl (Ge 1310) ... . Glycerina
Article 2 of pereopod 3 not deeply indentured . . use either couplet 10 or 11
[Branchiae with large plaits (fig. 127s) (use Key H for rapid identification of
select group of genera only, otherwise continue to couplet 11) . . Key H]
[Branchiae lacking plaits (if in doubt use this couplet). ...... 414]
Article 7 of gnathopod 1 minute, shrouded among a large and long tuft of
setae or cirri attached to either or both propodus and dactyl (figs. 129n,
ES O70) a ne ene peer os od Sy IEG

Article 7 of paaoped 1 leary eaiple! not ehroudcd:

use either couplets 12 or 19
Upper lip produced forward lobately and usually conspicuously in front of
epistome (epistome occasionally also projecting forward alongside dorsal

edge of labral lobe) (figs. 125a-d) ... .. . Mena anints ) IK LY
Upper lip not conspicuously produced forward lobe, or epistome and
upper lip coalesced even if produced. . . pseu lay
Epistome produced forward acutely and seoematalhy ioral appee lip (fig. 1250).
14

Epistome, if produced, not acute or conical or epistome and upper lip co-
alescedieven though produced sac) ied iach)

1 Couplet 8 leads to characters of ease in identification whereas couplet 21 leads to a possible natural group-
ing by use of mandibular structures.

2 Key I generally contains taxa with produced upper lip (labrum) even if coalesced with epistome as long
as labral portion recognizable by its position; couplet 13 generally contains taxa with epistome and upper
lip together lobate and separated by a notch even though some of these taxa occur in key I for comparative
purposes.

14.

15.

16.

17.

18.

19.

20.

ile

22.

23.

24.

MARINE GAMMARIDEAN AMPHIPODA 297

Mandibular palp attached level with molar or at its proximal corner. . 15
Mandibular palp attached strongly proximal to molar.

(exceptional) Orchomene
Coxa 1 tapering slightly and partially covered by coxa 2, inner ramus of

WEOPOG) eZ uasimiple rns) ae me . .. . Tryphosella
Coxa 1 fully expanded distally and: freely, “eels, t inner ramus of uropod 2
deeply constricted... . . Tryphosites
Telson cleft less than one fonean ai Ais fener Gre. 128n, OGM) sche ) WEN]
Telson cleft one third or more of its length (figs. 128s,t,v) . . . ... 17

Coxa 2 concealing anteroventral corner of coxa 1; or otherwise shielding
coxa 1; or coxa 1 reduced in size or subtriangular (figs. 119b, 122d, 1236).

Key L
Coxae 1 and 2 both large and visible and not triangular (figs. 117a, 118a@) . 18
Mandible lacking palp...,.......... . . .Stephensenia
Mandible bearing palp. . . . Gai wee NE

(from couplet 11.) Coxa 1 dlicitiy or Perrone pedieca in size or tapering
distally (figs. 119b, 122d, 1236), often partially covered by coxa 2. Key O
Coxa 1 of normal size, anterior and posterior edges parallel or coxa 1 distally

expanded... . nS CERT SLI Baa MeL Meili ir carne CoN P Rng Snare ene AD
Mandible lacking snl CATENIN epUNE OS Retccict Saas ame Sara _ Stephensenia
Mandible bearing palp. . . . Patan Stardaan ee (eye) SOV ANNE
(from couplet 7.) Mandible neta walls . 2... . . . . Stephensenia
Mandible bearing palp .... aber ee
Article 7 of gnathopod 1 minute, simondles among eciae or cirri attached to

AGCICIES a/MOL iO elec: Sete Oe oe erage . Key K
Article 7 small or large, Higiislle, neues, Sona ey OA, oe Ry 8)

Pereopod 1 prehensile and resembling a large patnonod)
Endevoura and Ensayara
Pereopod 1 simple ..... GaGa ur Pao
Mandibular palp attached diecinedly -onodianell 40 mola or ait molar afer
palp distinctly proximal in position. . . . pea le eae
Mandibular palp attached level with or distal io inoler, or if molar absent
palp attached distinctly toward distal end of mandible. . . . . . Key Q

KEY A

These are highly aberrant genera, not certainly placed in the Lysianas-
sidae. Shoemaker (1945a) believed Thoriella to be in the family
Talitridae but the second gnathopod is distinctly that of a lysianassid.
Chevreuriella and Danaella bear resemblance to cyphocarids (key F)
by reason of their coxae.

Il,

to

1.

Maxillipedal palp absent (fig. 1282); pereonites 3-6 hugely inflated, forming a

slobular body, m4 (heal Zila) aan 5 ene . . . . . Danaella

Maxillipedal palp present even if degraded: Wercomires 3-6 not larger than

other body segments (fig. 1176). . . . . EO Oca aa as ye,

Urosome composed of two segments (fig. 1240) . SPO een oE Uoheweuxiells

Urosome composed of three segments (fig. 122a). . . ... . . . Thoriella
KpyY BA

i)

HKyes and gnathopod 1 together powerfully developed (fig. 123a) .
Eyes and gnathopod 1 not together powerfully developed .

(es)

298 U.S. NATIONAL MUSEUM BULLETIN 271

2. Coxa 1 half as long as coxa 2 (fig. 123a), palp of maxilla 1 very short (fig.

LTE) wie ane ee : Trischizostoma
Coxa 1 nearly as Ns & as coxa 2, jello Bf saocaill 1 noneie . . . Normanion
3. Uropod 3biramous . . . SAY UAE CS pha ce VEC Seats as ale eR
Uropod 3 not biramous, rami aia aalsie PUTA ee mime Oa Ud See Ny
4. Maxilla 1: palp well developed, 2-articulate ......2.2.2.2.2.2.. 5
Maxilla 1: palp vestigial (fig. 127g)... ....... . . Acidostoma
5. Maxilliped: article 4 of palp well developed. .......2.2.2.2.2. 6
Maxilliped: article 4 of palp vestigial (fig. 128d) ........2.2.2.7
6. Telson deeply cleft, gnathopod 1 stout. ....... Shackletonia
Telson entire, gnathopod 1 very slender, flagellate. . . . . Azotostoma
7. Gnathopod 1 simple. .. . be betes tea Ee ie EP HOxOSstona
Gnathopod 1 strongly pubenelates POE OE SIN edo as Des NOR MaAnion
8: "Gnathopod df ‘simples. ee an he Oe Ron pa
Gnathopod -1 ‘chelate’; 4.050% 50.0. . bo ee ee Se Deriugiana
o> Dactyl of gnathopod'2) present) (yy, 20 es eee ieee)
Dactyl of gnathopod 2 absent... . . . . Ocosingo

10. Coxae 1-2 strongly reduced in size and nivel iidden Ph large coxa 3.
Lepidepecreella
Coxae 1-2 large and visible. . . . . . Acontiostoma and Stomacontion

1 See also Parawaldeckia kidderi (Smith) (p. 357).

KEY C

The first gnathopods of Valettia, Normanion, Cheirimedon and occa-
sionally of Orchomene and Hirondellea and females of Onesimoides are
weakly chelate. These genera are also included in other keys. Gnatho-
pod 1 of Microlysias is vaguely chelate but the genus is not included
here.

1.” Uropod! dijlackingrami 2.3 us, 618 aa oe ee ee enjugiana
Uropod 3 withrami .. . 2

2. Mandible: cutting edge ohenly touted (though sue minutely) (ig
P26 Oi es ae Pre Rea SAE 4g : 3
Mandible: einen pds net toothed! Giz 126h) . HPs eye ee eS

3. Pereopod 3: article 2 deeply indentured (fig. 131b) . . . . . . Podoprion
Pereopod, ajarticle 2 not indentured: 4 vis) 6) ve ee

4. Inner ramus of uropod 8 vestigial, telson entire . . . . . . Onesimoides
Rami of uropod 3 subequal to each other, telson cleft. . . . . . Valettia

5. | Maxillipedall palp)3-articulate (fig. 1289). a) ts =
Maxillipedalpalp 4-articulate 22.) s gmk tk

6. Melsoniclefts ox mencan Whatua ay atte ronnie hy runs Dien ene OC ODEONICES
Telson entire .. . LiSeh si gS Aiea eal Sn RCIA NSt AR dele a atc aN ae le

7. Maxilla 1 lacking Palpe slated al de Bec gun cle Mt | et hee ene ee eae aC Lite lle
Maxilla 1 bearing palp .... : ae 8

8. Pereopods 3-5: article 2 strongly andienined pal p of Peeciined nen eeaadiing
end of outer plate. .... . . . . . Podoprionella
Pereopods 3-5: article 2 not Sadana: wal of maxilliped shorter than
outer plate (fig. 128g) .... . . . Normanion

9. Coxa 1 reduced in size or tapering distally andl santelie hidden by coxa 2. 10
Coxa 1 of normal size and subquadrate shape, and not hidden by coxa 2. 14

MARINE GAMMARIDEAN AMPHIPODA 299

10. Inner plate of maxilla 1 with large sickle-shaped setae (fig. 127d).

Hirondellea

Inner plate of maxilla 1 with normal setae... . sii ep beh eae a

11. Mandibular molar obsolescent or absent, gnathopod 1 eeronely: chelate or
lleWeuey wae ens ctl elie bY
Mandibular polar Ww ell developed: rieed or asnelloxs, Eatinonod 1 weakly
chelate (fig. 129a) . . . ; Aenea li

12. Gnathopod 1 very slender, anal aries (fig. 1290), anions si aionoaly.
Euonyx

Gnathopod 1 powerful, article 6 very broad, chela formed by deeply excavate
palm (fig. 129e), epistome unproduced . . . . Opisa

Gnathopod 1 powerful, article 6 very broad and disiellly wexonniling palm
protuberant, not excavate (fig. 1291), epistome and labrum each with

shallow rounded lobe ... . . . . Cheirimedon latimanus Sars, 1895

13. Epistome with anterior bulb separated from protrusion of upper lip by long,

flat margin... ... . . Aristiopsis
Epistome rounded anal papresced t0 Proton oF Mane lip (exceptional).

Schisturella

14. Mandibular molar well developed, cuboid or ovate, setulose or ridged . . 15

Mandibular molar conical and unridged, obsolete or absent. . . .. . 19

SMe SOmsaclehts neu eeumalsunesen lewis: taal = aanae sole is .... . Gainella

Telson entire... . . . . some species of Orchomene

16. Gnathopod 2 tapering tor a point (but Rate chelate) (fig. 1312).
(alternate position) Gainella

Gnathopod 2 distally blunt. . . OS RSS ERE oh er Ta ocd LG
17. Maxillipedal palp article 4 cestisial Ge. 128¢) 1 aoe ae ne Pachynus
Maxillipedal palp article 4 normal, claw-shaped. . .. . ee ON car ae LSS
18. Outer plate of maxilliped large, exceeding end of palp article 2 Pg een ase gs Nadas)

Outer plate of maxilliped small, not or scarcely exceeding palp article 1 . 20
19. Eyes very large (losing pigment in alcohol), accessory eae 4+ -articulate,

telson longer than broad. ... . Jon Moroga

Eyes absent, accessory flagellum 1- maaeiee, telson as broad as long.
Figorella
20. Spines on outer plate of maxilla 1 reduced to two or three fused members,
outer ramus of uropod 3 uniarticulate ... . . . . . . Sophrosyne
Spines on outer plate of maxilla 1 of normal humber (9) and articulate,
Guberramus of uropodss biarticulate!. 0... 62) 4.00.0. 4. eiyska

KEY D

Most lysianassids have the primary cutting edge of the mandible
bearing at least one notch subtended by a tooth but these notches
are always lateral and never in the middle of the cutting edge; the
following species have teeth along the middle margin of the primary
mandibular plate.

Valettia is also included in Key B, because of its slightly chelate
first gnathopod. Podoprion and Onesimoides in Key C and Aristiopsis
in Keys C and N also have a toothed incisor. Clepidecrella may have
an extremely minute incisorial dentition. Nicholls’ (1938) figure of
Adeliella has a slight indication of incisorial teeth.

285-135 O - 69 - 20

300 U.S. NATIONAL MUSEUM BULLETIN 271

1) ¢Gnathopod Aysimple a evs OM nnn Chin MG has aes ate Alice lal
Gnathopod 1 subchelate. . . . . ia SLM Daa) ni aan Ni Ato
2. Telson entire, inner ramus of anopod 3 sjestioval ... . . . Onesimoides
Telson cleft, rami of uropod 3 subequal to each other. ........ 3

3. Coxa 2 perfectly rectangular and elongate, partially covering coxa 1, latter
shortened, subtriangular; inner plates of maxillae 1-2 fully lined with
setae medially. .... .. . . . Valettiopsis

Coxa 2 short, broad, cupreumded soareely sonesaline any portion of coxa 1,
latter not shorter than coxa 2 and of similar shape; inner plates of maxillae
1-2 with mostly terminal setae, medial edges scarcely or not setose.

Valettia

KEY E
Maxilla 1 lacks a palp, but the mouthparts are arranged in a quadrate
bundle, gnathopod 1 is not chelate, and the mandibular cutting edge

is not toothed. See Key A, Acidostoma in Key B and Prachynella in
Key C.

1. Gnathopod 1 very large, subchelate (fig. 130a), ‘‘telson entire’ . Pachychelium

Guathopod 4d smallisimpley {telson clebti@. 4 92) se ee

2. Antenna 1: article 2aslongasarticle1] ...... . . . Bathyamaryllis

Antenna I article: 2;abouthalfaswongyas Wei) ea en eee

3. Anteroventral corner of coxa 4 acutely produced ....... . . Vijaya

Anteroventral corner of coxa 4 rounded .......... .. Amaryliis
KEY F

These are the cyphocarids, characterized by the distinctly reduced
first and second coxae (and occasionally the third), often hidden
behind coxa 3 (fig. 122c) but not always, depending on the anterior
prolongation of the third coxae (or fourth). Coxae 3-4 of Metacyclocaris
do not fit this key because of their small size but the genus is included
for other comparisons. See note of Key A.

1. Urosomites 2 and 3 coalesced (fig. 122c). . . . . . . .Crybelocyphocaris
Wrosomites.2 andis separatel, m0 suis hy way ene ne ce nuit le dae eam:
2.5 Mandible lacking: paler tai ce wives sie Ree Miele eeu enn aS.
Mandible bearing palp ... . at SES eae

3. Pereopods 4—5 prehensile, article 6 of Pneihopod 1 enone then article 5.
Mesocyclocaris
Pereopods 4-5 not prehensile, articles 5 and 6 of gnathopod lsubequal . . 4
4. Pelson entired sn, 0 ie to es Sl am ee 2 ee = Cry belocephalus
Telson cleft... . . . . . . . Metacyphocaris
5. Uropod 3: inner ramus speomeley neetneadl or aneent (fig 32h) 2) eee
Uropod 3: rami subequal to each other, well developed. . ..... =. 8
6. Pereopods simple (fig. 117a). .......... . . . Lepidepecreella
Pereopods prehensile a LSEG) ORRIN Oe SS ot 8a ae
(eel SOnnClehtineiie Manin (chu iain ninn Winer aaah ea Me SoCyDROeanls
Telson entire... . . . . Cebocaris
8. Coxa 2 much larger pan 1 eval covering Hf bout coxae 90 4 small, not covering
anterior coxae. . . . . . . . Metacyclocaris

Some of coxae 1-2 or 1- 3 porered ener a 3 or i coxa 4 large) 5) oul

MARINE GAMMARIDEAN AMPHIPODA 301

9. Article 2 of pereopod 3 deeply indentured (fig. 1316) or with very long, simple
posterodistal tooth; coxae 1-3 all small and covered by coxa 4.

Cyphocaris

Article 2 of pereopod 3 not deeply indentured, only coxae 1—2 reduced and
COMELEMUDYACOKAC ES LOGLA Mie We aia eu) styrcliby sr hes il eea nls Meant AI ae ganna KO)
NOsaSomespercopods premensile yi ism seep sega) ccna etnies fol) Baracehocnnie
Pereopods simple ... . foc aaaunrasis ella!

11. Mandible lacking distinct nals or aria wonicclarminets amd unridged anallar.
Cyclocaris

Mandible bearing triturative molar (fig. 126g) . . . . . . . Procyphocaris

KEY G

The genus ‘“‘Onisimus” auct. (=Boeckosimus, new genus, herein)
has always been poorly defined and confusing (see Stebbing, 1906,
key p. 9, couplet 14). Some of its species overlap with ‘““Pseudalibrotus”’
auct. (=Onisimus as senior synonym) to which I believe they should
be transferred. The genus also appears in Keys J and M.

“Onisimus’”’ auct. comprises a series ranging from entire telson (as
in some species of ‘‘Pseudalibrotus” auct.) to slightly split (as in Key
J) to half split (in Paronesimus and Paratryphosites (= Hippomedon)
in Key M). The proportions of articles 5 and 6 of gnathopod 1 vary
slightly, enough to cross couplets in Key M.

1. Pereopod 1 formed into a subchelate grasping organ similar to a gnathopod

(ieanlG0D) sate olen sonia nse) el ey 2 Mndevoura andy Ensayara
Pereopod 1simple ... . Sve eee

2. Upper lip formed into a lanes ibeel: iftke lobe wrens stronely projecting in
front of epistome (fig. 125c) . ... . LNG Neg evan

Upper lip usually small, obscure, andlor scaroaly mroicctine in front of
epistome (figs. 125g,n,0), or epistome and upper lip coalesced, upper lip

if lobate equalled in extent by lobe of epistome. . . . pte gaiecne Ml

3. Uropod 3: inner ramus less than half as ie as outer, invele 5 of male
antenna 2 dilated (fig. 124j)}. .............. Parawaldeckia
Uropod 3: inner ramus subequal to or more than half as long as outer,
[article 5 of male antenna 2 not dilated, article 4 rarely dilated] . . . . 4

4. Gnathopod 1 with broadly transverse palm (fig. 129a) .. . . . Koroga
Gnathopod 1 minutely but distinctly subchelate (fig. 129b) . ..... 5
Gnathopod ysimple (ig 129m) ta ne ee ey cree
HeArticle 4-ofantenmar2 tumiducnen set ae ee iysianella
Article 4 of antenna 2 slender. ..... =.=. +... .. . Paralysianopsis

6. Article 4 of male antenna 2 dilated. ......... .. . Lysianella
Article 4 of male antenna 2 not dilated... . . ERM in phn Noes an ooeed ana

7. Uropod 3: peduncle expanded distally into a narrow, olavelike projecting
siiveliige (fiicasl'322))) ines ee BN Nera AUR ATE I
Uropod 3: peduncle simple, sabeyiindriedl Ge 1329) . . . . Lysianassa !

8. Maxilla 1 with two kinds of spines on inner plate. .......... 9
Maxilla 1 with one kind ofspineoninnerplate. .........-. .- 10

9. Inner ramus of uropod 2 constricted (fig. 117a). .. . . . [Arugella‘]
Inner ramus of uropod 2 unconstricted. . . .. . . . [Pronannonyx']

10. Mazxilla 2: inner plate nearly twice as broad as outer . . [Shoemakerella ']

Maxilla 2: inner plate equal to outer in width . [Lysianopsis '] and [Aruga ']

1 Tam amalgamating all of these genera in Lysianassa, but Pronannonyz is questionable and also retained
as a genus in alphabetical order.

13.

14.

15.

16.

bw
ou

U.S. NATIONAL MUSEUM BULLETIN 271

Uropod 3: inner ramus vestigial orabsent. ... . Bios tania. euiey iy
Uropod 3: inner ramus more than half as long as outers TFAMNUS eee eet
Uropod 3: inner ramus vestigial, article 1 of antenna 1 flagellum 5 times as
long as next article. .... . . . . . Onesimoides 2
Uropod 3: inner ramus absent or nprented Bye a spine, article 1 of antenna 1
flagellum not longer than next article. . . . . . . . . Paronesimoides
Uropod 3: inner ramus vestigial, article 1 of antenna | flagellum not longer
than wnextiarticle:: ia) ha ee see Saar nee Clepidectellags
Gnathopod 1 simple. .. . oy eo et ae
Gnathopod 1 distinctly, fyoueh Biten Prince pubehelate SS OLS Ry Dia Hy 1.7
Mandibular palp attached level with molar (fig. 126m), coxa 4 scarcely
excavate posteriorly (fig. 117a) . . . . . Paralibrotus and Menigrates
Mandibular palp attached well proximal of molar (fig. 126h), coxa 4 excavate
posteriorly (fig. 119b) .... ia Se eiecu mM Rea inet tetany Boi ry 3 DLED)
Palp article 4 of maxilliped tiny Gre, 128k) BG Sd tee oN TTT ONYX
Palp article 4 of maxilliped well developed (fig. 128c). . . ...... 16
Epistome and upper tear together strongly produced and separated by a
Siti ean ke ‘ . . . Aruga (in part) [—Lysianassa]

(and ORs [Pronannonyx] and see Adeliella)
Epistome and upper lip together not strongly produced, and apparently

comlesced yaar Spelotel ah oak Lailvics ast Mules eauley eat ARAM ASI
Maxillipedal palp 3- smowilaie ete ere imma eis 3 a. LS
Maxillipedal palp 4-articulate. . . . SESE Oe 2 ot Nas eae Ne
Maxilliped: inner plates obsolescent (Gis, 128)) . oa. oe Rerrierella
Maxilliped: inner plates as long as palp (fig. 128g). . . . . . Normanion
Gnathopod 1: palm excavate (mouthparts unknown) . . . Pseudambasia
Gnathopod 1: palm not excavate .. . PUREE Ni io, oe) AU)
Mandibular palp attached level with mole Ge. 126m) . Foyle) ag vee eaareammead|
Mandibular palp attached proximal to molar (fig. 126h) or molar absent. 23
Maxilla 2 strongly setose (fig. 127p), coxa 1 visible . ..... 2... 22
Maxilla 2 poorly setose (fig. 127k), coxa 1 nearly concealed by coxa 2.
Adeliella
Mandibular molar small, conical, Cia: setulose, gnathopod 1 article 6
enlarge dime cae : I ee os oroga

Mandibular molar of eesti size, etimatioe’ gnalionod 1 not enlarged.
“Pseudalibrotus’”’ (—Onzsimus) and ‘‘Onisimus”’ [—Boeckosimus (in part) |
Rami of uropod 3 reduced in size, inner ramus about half as long as peduncle

Gare NAVE) gg . . Nannonyx
Rami of uropod 3 as iene as or longer shea me danele (Ge 1324) . ese ue:
Article 6 of gnathopod 1 very stout, enlarged (fig. 1291) ....... 25

Article 6 of gnathopod 1 small, rectangular (fig. 129d).
some species of Orchomene and [Allogaussia]
Mandibular molar of medium size, triturative, article 4 of maxillipedal palp

Shortaeeueiaas .. . . . Pseudokoroga
Mandibular molar sell conical etnonetadl, eeralece article 4 of maxilli-
pedal palpi loma:s ee Sy ieee ik Vik edn pets aiRu lem neSl aes a nen O ee

2 See also Adeliella.
3 See also Parawaldeckia kidderi (Smith) (p. 357), distinguished from Clepidecrella by extreme proximal
location of mandibular palp.

MARINE GAMMARIDEAN AMPHIPODA 303
KEY H

These genera have gills bearing swellings or plaits. Since so many
lysianassids have not been investigated for this characteristic and
because gills of the following genera are not always clearly plaited
the genera have been added to subsequent keys. Hence key H simply
represents a quick way for the taxonomist to determine familiar
genera with obviously plaited gills. If this key is unsatisfactory one
must proceed to couplet 11 of the Basic Key. Many species of Hip-
pomedon have indistinctly plaited gills. Gills of Socarnella are unknown.

1. Coxa 1 reduced and hidden by coxa 2 (fig. 123b) . . . . . . Schisturella
Coxa 1 of normal dimensions and not hidden by coxa 2 . : Bilas aia) 7
2. Gnathopod 1 subchelate 3
Gnathopod 1 simple 7
3. Mandibular palp attached rexel! to mole Gin, 1267) . 4
Mandibular palp attached level with molar (fig. 126e) : 5)

4. Article 5 of male antenna 2 extremely tumid (like article 4 of fie 124h).
Microlysias
Article 5 of male antenna 2 slender or not relatively stouter than article 4.
Orchomene
5. Mandibular molar imperfectly conical or attenuate, setulose, not ridged
Giger 267) Ge .. . Anonyx
Mandibular molar cylindrical or ncaboical: mdeed @e 126g) . Beene 6
6. Dactyl of gnathopod 1 very small, similar to dactyl of gnathopod 2, Alp
article 4of maxilliped stout, short... . . . Pseudoanonyx
Dactyl of gnathopod 1 and palp article 4 of meciliped long and claw-like.
Hippomedon

7. Dactyl of gnathopod 1 setose or spinose on posterior edge (fig. 18072), upper lip
poorly delineated (fig. 125g), mandibular palp attached level with molar.
Ichnopus

Dactyl of gnathopod 1 not setose or spinose on posterior edge, upper lip
strongly delineated (figs. 125a,c), mandibular ae attached distinctly

proximal) tojmolar =.) a eee site

8. Outer ramus of uropod 3 Drareiculates or eicompoccd of one syareicle and large
terminal spine .. . SUES a er Ua emai aoe Ue SOCALNES
Outer ramus of uropod 3 amieeticulates AEM Es ca Sera ie eee)

9. Telson emarginate; mandible long and narrow, malp aricley2 Thoin 5 times as
long as article 1 and more than 2 times as long as article 3. . . Socarnella

Telson cleft more than halfway; mandible short and stubby, palp article 2
only 3 times as long as article 1 and about 1.5 times as long as article 3.
Socarnopsis

KEY I

Genera of Key H are included here owing to possible confusion in
Key G concerning gills. Koroga is repeated in case its telson is con-
sidered as slightly cleft.

1. Coxa 2 concealing anteroventral corner and most of coxa 1 (fig. 1196), coxa 1

slightly or strongly reduced in size or not quadrate. ........ 2

304

10.

11.

12.

13.

14.

U.S. NATIONAL MUSEUM BULLETIN 271

Coxa 2 not concealing anteroventral corner of coxa 1 (fig. 117a), coxa 1
normal and quadrate or distally expanded . 8
Coxa 1 half as long as its article 2 (fig. 130k) . . Late, By aac ne eee
Coxa 1 as long as or longer than its article 2 (fig. 180g). ....... 6
Gnathopod 1 subchelate (often ee OT MR haa 0 4
Gnathopod 1 simple 5
Lobes of maxilla 2 equally Broad or inner Tope: narrower “hema amie, “qacmadtion
lar molar short, blunt, or weakly laminate (fig. 126e), inner plate of maxilla
1 with two setae... . . . . .Metambasia and Schisturella
Inner lobe of maxilla 2 soul Tprond a than outer, mandibular molar conico-
laminate (fig. 126f), inner plate of maxilla 1 densely setose (4 or more

Setaey te rs ... . . Aristias

Lobes of maxilla 2 not or eerecly. eesostin inner sae Orowdes than outer.
Metambasia

Lobes of maxilla 2 gaping (fig. 1277), inner much broader than outer.

Aristias
Maxilliped: outer plate lacking spines . ........ . . . Ambasiopsis
Maxilliped: outer plate spinose .. . Peary les titty enc
Telson deeply cleft (three fourths or aRGr), outer lobe of smassle 1 poorly
spinose (fig. 127c), inner ramus of uropod 2 simple. . . . . Neoambasia
Telson deeply cleft, outer lobe of maxilla 1 normally spinose, inner ramus of
uropod 2simple.... . . . . . . Tryphosella
Telson cleft one third, outer lobe Gr smosalla 1 nonnalls spinose, inner ramus
of uropod 2 constricted .... . . . .Schisturella (—Pseudonesimus)
Guuthopod Teimple) Ue) kh GE) aia a alt
Gnathopod 1 subchelate .. . Be ice iran vin aR amaelsn iG: AL IL
Pereopod 5 much longer than neetonod 4 GC A ae Gia ae Te hnopus
Pereopod 5 not longer than pereopod4......... Boner gh tO)
Inner ramus ofjuropod!2inciseds =) iis ae a I Soenrnideet

Inner ramus of uropod 2 simple.
Waldeckia, Socarnes, and Socarnella (see Key P, couplet 20)
Head lacking distinct lateral lobes, anterolateral cephelic margins straight

(fig. 121b); coxa 5 as long as coxa 4... .. . .. .. .Lepidepecreoides
Head bearing lateral lobes, anterolateral ae margins bulbous or convex;
coxa 5 shorter than coxa 4 .. 2 AGUA CR Nt ee
Article 4 of antenna 2 very tumid Go 124k) . Lise a Ae iLysianella
Article 4 of antenna 2 of normal dimensions. . . . Ste Aes sete) Les}

Mandibular molar large but not strictly Cabordain sien subconical or
attenuated or basally bulbous and covered densely with setules (fig. 1267)

14

Mandibular molar large or of medium size, cuboidal, ridged, setules if rarely
present very sparse (fig. 1269)... . Bae is auccee ILS)
Article 6 of gnathopod 1 enlarged (fig. 1291), felcon apical DGtened Korsga
Article 6 of gnathopod 1 not enlarged, telson deeply cleft. . . . . . Anonyx
Uropod 2 with inner ramus strongly incised, epistome with extremely large,
acute anterior process... . . .. . . . Tryphosites

Uropod 2 with simple inner ramus, eaions produced!
“Tmetonyx”’ nobilis (Stimpson) (see Shoemaker, 1930) and Hippomedon
Uropod 2 with inner ramus strongly incised, epistome and upper lip coalesced
(not separated by notch from lateral view), apparent upper lip portion
strongly produced and acute.
‘“Tryphosa’”’ carinata Schellenberg, 1926a

1 And see “Tryphosa”’ carinata Schellenberg, 1926a (couplet 15) and Orchomene (p. 353).

MARINE GAMMARIDEAN AMPHIPODA 305

KEY J

Both Koroga and Orchomene (=?Allogaussia) are repeated here, in
the event their quantitatively expressed characters were overlooked
in previous keys.

iy 1 @oxail nearly concealed by coxa 2-°%)0-0-0.0.5. 23.0... 2 4. Adeliella
Coxa fully) visible.) 5 <7. Be ne Cer ee RR nO

2. Mandibular palp attached level with molar (fg. tea). Ns ae Wea Resale ra SUR
Mandibular palp attached proximal to molar (fig. 1267). . .......5

3. Gnathopod 1 subchelate, inner ramus of uropod 3 as long as article 1 of outer.
4

Gnathopod 1 simple, imner ramus of uropod 3 vestigial. . . . Clepidecrella

4. Mandibular molar triturative........ “Onisimus”’ [= Boeckosimus]
Mandibular molar simple... . .... . . .Menigrates

5. Gnathopod 1: article 6 not wider fae article 2) SRN Y: See O

Gnathopod 1: article 6 distally twice as wide as article 3 (fig. 1291).
Koroga and Pseudokoroga (see Key G for distinction)
6. Gnathopod 1 subchelate ...... . . . . Orchomene [=? Allogaussia]
Gnathopod 1 simple. ........ .. .. .. .Socarnella and Menigrates

KEY K

Gnathopod 1 is slender, the dactyl hidden in a tuft of setae or cirri
(fig. 130m). Ichnopus bears resemblance to this group as a genus with
a posterior or distal brush of setae on the dactyl. Pseudoanonyz is only
marginally assignable to this group.

1. Gnathopod 1 stout and minutely and transversely subchelate or chelate, palp

article 4 of maxilliped stout and subclavate. . . . . . . Pseudoanonyx
Gnathopod 1 slender and simple, or palm very oblique, palp article 4 of
maxilliped slender and claw-like. . . . 2

2. Mandible lacking molar, article 6 of pereouods) 12 2 slightly longer har
articles 4 and 5 combined (fig. 131c) and slightly prehensile.

Scopelocheiropsis

Mandible bearing molar, article 6 of pereopods 1-2 shorter than or equal to
articles 4 and 5 combined (fig. 117a). . ... : A oho

3. Lower lip with major lobes a excavate aeinlie forming a medial gap
dig: 125s) 5. . . . Bathycallisoma
Lower lip with ede adles of major Nokes Elosely appressed ( ve WAR A) os e:

4. Gnathopod 2 minutely chelate (fig. 131f). ........ Bice He ameepeta)
Gnathopodli2isubchelatenCipadoily)ire st ei ahkie nr Weert (i700

5. Epistome flat in front (fig. 1250) .. . Neon ei each et hee AT OUT
Epistome grossly lobate in front (fig. 12510) .... . . . Scopelocheirus

6. Inner plate of maxilla 1 setose only terminally and subterminally, dactyl of
gnathopod 1 not distinct from cirri (fig. 129m) . . . . Paracallisomopsis

Inner plate of maxilla 1 setose medially, dactyl of gnathopod 1 distinct from

cirri (fig. 130m) (see other characters under Paracallisomopsis diagnosis) . 7

7. Article 2 of gnathopod 1 linear, not glandular (fig. 130m), coxa 5 much larger
than and partially covering coxa6.. . . . . Paracallisoma
Article 2 of gnathopod 1 swollen, glandular fe 1299), coxa 5 not much larger
than and not covering coxa6.... . ..... . . . Eucallisoma

306

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U.S. NATIONAL MUSEUM BULLETIN 271

KEY L
Pereopods 1—5 with long dactyl folded back on an inflated article 6, thus
prehensile (fig. 130d) .... . .. . . . Metacyclocaris
Pereopods 1—5 normal, dactyls lnc anal path ticles unexpanded. . . 2

Maxilla 2: lobes gaping and/or inner much broader than outer (fig. 1277);
mandible with molar, combined with nearly simple gnathopod 1 . Aristias
Maxilla 2: lobes appressed and of similar shape, other characters not in
combination. . . [MENTE ERM AGAS GEE RO. UE ya ne en am eS
Mandible lacking moles (pat hopodl 1 Emplel SNE 4
Mandible bearing molar, gnathopod 1 subchelate Gun in iNfciambascom 5)
Article 2 of maxillipedal palp as broad as or broader than outer plate.
Ambasiella
Article 2 of maxillipedal palp less than half as broad as outer plate. Ambasia
Inner plate of maxilla 1 with one or two immense, falcate setae (fig. 127d).
Hirondellea !

Inner plate of maxilla 1 with small, slender setae. . ......... 6
Maxillayisinner plates with tivetoramore Sevaciis sich) cat icicles enn)
Maxilla 1: inner plate with three or fewer setae. . . . AR Ae seh tS)
Mandibular molar immense (fig. 126d), inner plate of aT 1 setose only

terminally, article 3 of gnathopod 1 not elongate .. . . . Eurythenes

Mandibular molar slender but strongly projecting, inner plate of maxilla 1
setose medially and terminally, article 3 of gnathopod 1 elongate [see 2

species of Cheirimedon] . . . . Se IE ene tal ee aralice lla

Inner ramus of uropod 2 Coneenictca AGRA URM ENS SER cE Mom tM mast ti ph i a <i v8)

Inner ramus of uropod 2 simple. . .. . | Vase Rates: Weal
Gnathopod 1 slightly chelate, epistomal margin daneoyenteallly elongate.

Aristiopsis

Gnathopod 1 subchelate or simple (Metambasia), epistomal margin dorso-

ventrally compressed even though lobate (normal) ........ 10

Mandibular molar poorly developed, weakly triturative or fuzzy, palp
attached proximal to molar, gnathopod 1 subchelate.

“Orchomenella’”’ groenlandica (Hansen)

Mandibular molar strongly triturative, palp attached level with molar,

gnathopod 1 subchelate . .. . .. . . . Schisturella
Mandibular molar weakly tritur ative or 4 fiom bat very large, palp attached

level with molar, gnathopod 1 simple. . ..... . . . Metambasia
Outer ramus of uropod 3 uniarticulate . . .. . . . . . . Tryphosoides
Outer ramus of uropod 3 biarticulate ........ Sok een Lez,
Mandibular molar lacking setules =) 2.5 24 5 2. hepidenecreunace
Mandibular molar bearing setules. . . . Berea ists alot AL)

Gnathopod 1 with article 6 enlarged and dieealle qantas, nearly twice as
broad as article 2, palm protuberant, long, and slightly chelate.
Cheirimedon latimanus Sars, 1895
Gnathopod 1 with article 6 unexpanded, palm not chelate . .... . 14
Head very small,? epistome and upper oe weakly developed and incon-
spicuous. . . .. . . . Uristes [—Uristoides]
Head of normal size, , aotone andl | upper lip strongly developed, epistome
generally shightly lobateici) 2h) os ie fee rei Ona ra Se eee to)

1 See note, Key N, couplet 23 (p. 310).
2 Occasionally head ventrally extended and covering base of antenna 2 as in ‘‘ Uristes’”’ lepidus J. L.
Barnard (1964a).

15.

10.

Il,

12.

13.

MARINE GAMMARIDEAN AMPHIPODA 307

Article 7 of gnathopod 1 with accessory tooth.
some species of Tryphosella [—‘‘Tmetonyzx’’ auct. ]

Article 7 of gnathopod 1 simple . . . . Tryphosella [—‘‘7ryphosa”’ auct.|
KEY M
Palp of mandible attached proximal to molar (fig. 1267) . .. . A

Palp of mandible attached level with molar or molar absent (figs. 126j,k) .
Article 1 of antenna 1 nee produced distally and dorsally La

Gigs 4d) es irks 2 . . . . . Lepidepecreum
Article 1 of antenna 1 yearly or nee menaced discal, not dorsally crested

(tags WOO A So aides San ORMOND MMU ae LUC rss IOUS DE Nadel ts hanciteareD
Gnathopod 1 simple andl oun sai ed ass detec Suse Meee
Gnathopod 1 subchelate (or very large aed coominely sfimoralls) Seale Vaan eee
Outer ramus of uropod 3 uniarticulate ...... =... . .Socarnopsis
Ourerramus,omunropodsoppiarpiculave sii ly nceyle Westie nel euiu ian lesa. O
Inner ramus of uropod 2 constricted. ....... .. . . .Socarnoides
Inner ramus of uropod 2 simple. . . . .6

Telson cleft one third of its length, Soma Doris not onomninend Ge 1254),
articles 2-3 of antenna 1 peduncle half as long as article 1 of primary
flagellum .... . . . . .Menigrates

Telson cleft faleeete or tore mreoucen rpms anouainent (fig. 1250), articles
2-3 of antenna 1 as long as article 1 of primary flagellum.

Waldeckia and Socarnes (see Key P, couplet 21)

Article 3 of gnathopod 1 elongate and slender as in typical lysianassid
gnathopod 2, articles 5-6 also elongate and slender (fig. 1297).

Pseudorchomene
Articles 3,5,6 of gnathopod 1 stout, not relatively elongate (fig. 129d) . .8
Article 4 of antenna 2 very tumid, article 5 slender . . . . . . Lysianella

Article 5 of [male] antenna 2 very tumid, stouter and longer than article 4.
Microlysias !
Neither of articles 4-5 of antenna 2 expanded . . i
Articles 5 and 6 of gnathopod 1 not immensely cnlemaed Ge. 1294) (also
check Lepidepecreum).
Orchomene [=Orchomenella] and juveniles of Pseudokoroga

Articles 5 and 6 of gnathopod 1 immensely enlarged (fig. 1291) . . . .10
Maxillae slender (fig. 1271), maxillipedal palp article 4 short, mandibular
molar triturative ... . . . . . Pseudokoroga
Maxillae stout (fig. 1270), ranselltraadlel sopalfay ontniells 4 elongate, mandibular
molar nontriturative . .. . ee A ECOROZa
Mandibular molar laminate, ponical nee euiotcely Grince simple, lacking
ridges, often weakly setulose, or molar absent . ... . sete GULP

Mandibular molar cuboidal or columnar, surface usually beseine conspicuous
ridges and spines, or distinct punctations, or densely setulose if not fully
cuboidal and slightly attenuate (figs. 126e,9,7,m) . . . arcuate ser 0)

Gnathopod 1: article 6 very large and expanded, more ‘her 3 times as long
as article 5 (fig. 1291). . .. . . . . . .Koroga and Cheirimedon

Gnathopod 1: article 6 small, never more than 1.5 times as long as article 5. 13

Inner lobe of maxilla 1 bearing five or moresetae.......... 14

Inner lobe of maxilla 1 bearing two or fewer setae. ......... 15

1 Possibly article 5 of Microlysias has been misidentified.

308 U.S. NATIONAL MUSEUM BULLETIN 271

14.) Gnathopodsisimple) 22s Meeat itey el) on) Mice eae eee Ce lia
Gnathopods subchelatee ane ey one ee aralicella

152 sGnathopod Lsimipleyy ia eau aac: Saeeayeia bel ant ean ipa Guile eres Meatee ammaa
Gnathopod 1 subchelate. ... . SBE etsy aia. lve 11 7/

16. Rami of uropod 3 subequal to each onhen in Tenet» . . . . . Ichnopus ?
Inner ramus of uropod 8 vestigial, scale-like. ... . . . . Clepidecrella

17. Head globular, maxilla 1 with sickle-shaped setae on inner lobe (fig. 127d).
Hirondellea

Head other than globular, maxilla 1 with normal setae on inner lobe. . 18

18. Maxillipedal palp article 4 reduced in size (fig. 128a). . . . Centromedon
Maxillipedal palp article 4 of normal size. . . . Set) tera ee peeteas LG)

19. Coxa 1 normally quadrate, untapering, ganhelion leasing Tater lobes or
strongly truncate anteriorly (fig. 121b), coxa 5 as long as coxa 4.

Lepidepecreoides

Coxa 1 normally quadrate, untapering, cephalon with lateral lobes, coxa 5
much shorter than coxa4 . . . . . proceed to 24

Coxa 1 tapering distally, slightly “-qnosioncdl ‘aaellon with lateral lobes,
coxa 5 much shorter than coxa 4 . . Lepidepecreopsis and Tryphosella

20. Coxa 1 very slightly triangular, tapering distally (figs. 129c,k) .... 21
Coxa 1 quadrate, or rounded, expanded or unexpanded distally (figs. 129

CH TOS aU Cie Ne nie MATAR DI NMR cad Ye 3 Say

21. Outer ramus of meopad 3 aantieulate SAR UGA NsERE AN eh aaa iunyphocuides
Outer ramusofuropod:s biarticulates ~ 1. 4.40070) fai) oO) een ee

22. Head small, epistome and upper lip inconspicuous.
Uristes ? [= Uristoides] ?some species of Cheirimedon
Head large, epistome and upper lip prominent from lateral view .. . 23
23. Dactyl of gnathopod 1 with inner accessory tooth.
some species of ‘‘Tmetonyx” auct. [= 7 ryphosella]
Dactyl of gnathopod 1 lacking accessory tooth.
Tryphosella 3 [=‘‘Tryphosa’’ auct.]
24. Telson not longer than broad, always cleft less than halfway.
Rifcus and Boeckosimus [=‘“‘Onisimus’’ auct.]

Telson longer than broad (or if short, then cleft halfway or more). . . 25
25. Gna thopod jh Ssimiple nity ay spin) ease mnie Neola UUs OM Rr he DO
Gnathopod 1 subchelate .... : Baer eA AAS)

26. Gnathopod 1 perfectly simple, srecallinaedtal dace samntied. - Menigratopsis
Gnathopod 1 very slightly subchelate, palm small, maxillipedal dactyl

clongate ea wn. Bou Minuet eny ese BONA MES BPA TF

27. Uropod 2 with simple sh innerramus .. . ie iparoncsitnns (in part)
Uropod 2 with incised inner ramus . i onmonyace cicadoides Stebbing, 1888

28. Mandibular molar densely covered with setules . .. . BS ei ia KG)
Mandibular molar ridged or weakly punctate, setules aqanernllly absent or
inconspicuous . . . pes ler oO)

29. Coxa 1 expanded dligiclt. npenion dated of rametinonadl 1 rigireilby lacking
accessory tooth, upper lip slightly or strongly protruding in front of
epistome, inner ramus of uropod 2 often incised, palp article 4 of maxilliped
UIT GUE OLIN pe ie sane eee) Rul ave CU ey ee UIA Cin su aN Meta Fis eee ANT O TTLVAXG

2 Menigrates angustipes Gurjanova (1962) may key to this position inadvertently but note diagnosis of
Ichnopus does not otherwise fit Menizrates.
3 See also ‘“‘Orchomene’’ takoradia J. L. Barnard (1961); ‘‘Uristes’”’ velia J. L. Barnard (1961).

30.

Bl,

32.

33.

o4,

MARINE GAMMARIDEAN AMPHIPODA 309

Coxa 1 unexpanded distally, unguiform dactyl of gnathopod 1 bearing inner
accessory tooth, epistome slightly protruding in front of upper lip, inner
ramus of uropod 2 simple, palp article 4 of maxilliped unguiform.

type-species of Tmetonyx

Coxa 1 slightly expanded distally, dactyl of gnathopod 1 with accessory
tooth but reduced to size of dactyl on gnathopod 2, prebuccal parts
apparently like Anonyz of triplicut 29, inner ramus of uropod 2 simple,

palp article 4 of maxilliped tumid, subclavate. . . . . . Pseudoanonyx
Dactyl of gnathopod 1 very small and similar in size and morphology to
dactyl of gnathopod2 ... . . . . . Pseudoanonyx
Dactyl of gnathopod | long, alanis, thus aan Seng santana leas Wea inaeae
Article 5 of gnathopod 1 longer than article6 .. . a hea
Article 6 of gnathopod 1 longer than or subequal to mols 5 SG ks GO
Head without distinct lateral lobes, anterolateral margin convexly truncate,
coxa Daslongascoxa4..... . . . . . Lepidepecreoides

Head with small but distinct lateral lobes, .¢ coxa 5 shorter than coxa 4.
Hippomedon [= Paratryphosites, ? Paracentromedon, ? Elimedon|
Outer ramus of uropod 3 uniarticulate ...... .. . . Tryphosoides
Outer ramus of uropod 3 biarticulate . . . . te aie aa OA
Articles 5 and 6 of gnathopod 1 subequal in fenet “atiels 6 not saoaacled!,
palm obsolescent ... . . . .[Paracentromedon] and Paronesimus
Article 6 of gnathopod 1 large, distally expanding, more than 3 times as long
as article 5 (fig. 1291), palm strong ... . . . . .Cheirimedon (type)

KEY N

All lysianassid genera with coxa 1 distinctly reduced in size and/-
or often largely hidden by coxa 2 are assembled here for ease of com-
parison. Those species of Key F, in which both coxae 1 and 2 are small
and hidden by following coxae, are not included. See also notes on

Stomacontion.
1. Pereopods 1-5 with long dactyls folded back on inflated article 6, thus pre-
hensile (fig. 130d)... . . . . . . . Metacyclocaris
Pereopods 1-5 normal, dactyls shen’ ail sein articles unexpanded. . . 2

ou

Upper lip strongly lobate and projecting in front of epistome (figs. 125c,d) . 3

_ Upper lip scarcely or not projecting in front of epistome, latter often large

and dominating upper lip (figs. 125h,7,k) or epistome and upper lip not
distinct from one another . US Rae 8)

Coxa 1 half as long as its second paticle Ge. 130) . SOs IG ety Boren
Coxa 1 as long as or longer than its second article (fig. 1309) . Bi Ree Nn Ie)
Gnathopod 1 subchelate even ee weakly... . . . . Schisturella
CGnathopodi la simples. (9 4) .0). . . . . . Metambasia
Outer plates of Se lacking spines, “messi 1 with seven spines on

GQUIEE OND 5 6 4 es .. . . . Ambasiopsis
Outer plates of maillyped spinose, masa 1 nora with 9+ spines on outer

Ko) oveypre i enAe Oe .. . . . Schisturella
Outer plates of anenclliaed spinose, peceilln 1 ant spines on outer lobe

REGUCeGtOptiVe; 4.2. ..... . . . Neoambasia
Mandibular incisor walthicleretiarta (fig. 1269). A rieh islet La ceed ea Le NGO END TE
Mandibular incisor not dentate... . 8

Mandible lacking molar, gnathopod 1 cnetnte: seule 2 of Dercopad 3 deeply
imdentunedi (fies SND) ae Wo re ol oes o Podoprion

310

10.

11.

13.

14.

15.

16.

18.

U.S. NATIONAL MUSEUM BULLETIN 271

Mandible with triturative molar, Chia 1 subchelate, article 2 of pereo-

pod 3 not deeply indentured. ... . ... . . Valettiopsis
Lobes of maxilla 2 broadly separated, inner lobe aoineine medially and of

different shape or much broader than outer lobe (fig. 1277) . . .... 9
Lobes of maxilla 2 appressed and similar in shape (fig. 12717)... .. . 10
Telson cleft... . . . Ambasiella and Aristias (see Key O, couplet 2)
Telson entire... . . . Perrierella
Eyes and panenopods 1 sarventtallly developed Ge. 123) . 2 eit GD
Eyes and gnathopod 1 not together powerfully developed. . .... . 13

Gnathopod 1 with complete torsion distal from article 4 (fig. 130h),
dactyl and palm thus posterior, mandibular molar needle-like (fig. 126b),

telson searcely longer than broad (fig. 128w) . . . . . Trischizostoma
Gnathopod 1 untwisted, normal, mandibular incisor broad, flat, telson
elongate, muchlongerthan broad .. . Berita Wer 74
Gnathopod 1 chelate, palm thus deeply oats (Ge. 1296) . . . Opisa

Gnathopod 1 cheliform only, palm convex and protuberant (fig. 1291).
some species of Cheirimedon

Palp article 4 of maxilliped vestigial (figs. 128d,e,7) ......... 14
Palp article 4 of maxilliped well developed, claw-like. ........ 17
Lobes of maxilla 2 strongly setose, telson deeply cleft... ..... 15
Lobes of maxilla 2 with one and four setae, respectively (fig. 127k), telson

scarcely.) cleft ei adi cult Gre Ri MAAC UV aa 9) REA ep aerate sme Mem@Ar elke lta)
Mt pal ley TL waaeloy VOM Gh ee are oh aka ey oo be eT A eG
Maxilla 1 lacking palp ... . Prot! feoutinite to couplet 21

Epistome and labrum aoelenead, iabral aonvion not projecting (epistome
often projecting), mandible without molar . Ambasia and Ambasiella
Epistome and labrum separated by notch, labral portion slightly projecting,
mandible with molar .. . . . . Centromedon
Mandibular molar distinctly agounedl al riiecal (Ge, 1269) ii se ees
Mandibular molar untoothed and unridged (fig. 1267), or ridges forming
marginal girdle only (fig. 126d), often fuzzy, or molar absent ... . 19

Inner ramus of uropod 2 incised, outer ramus of uropod 38 biarticulate.
Aristiopsis and Schisturella !

Inner ramus of uropod 2 simple, outer ramus of uropod 3 uniarticulate.

Tryphosoides
Gnathopod 1 chelate (chela and gnathopod slender) . Podoprion and Euonyx
Gnathopodt subchelaterorsimyple sit Wai ye ins e o) cele nua an nee, ()
Gnathopod 1 simple. ... . Mapes Nh bee fie 24dl
Gnathopod 1 subchelate (often ofeawoteely <0) or r chelate . Rees ret ey hey) 1)
Anteroventral corner of coxa 4 acutely produced ....... . Vijaya
Anteroventral corner of coxa 4rounded or quadrate. . ....... =. 22
Article 2 of antenna 1 half as long as article 1 ... . . . . Amaryllis
Article 2 of antenna 1 fully as long as article 1 . . . . . Bathyamaryllis

Inner plate of maxilla 1 with one or two immense falcate setae (fig. 127d).
(cf. Cheirimedon pectinipalma K. H. Barnard, 1925) Hirondellea
Inner plate of maxilla 1 with normal setae 24
Inner plate of maxilla 1 with five or more setae Ganlts)
Inner plate of maxilla 1 with three or fewer sctae .

ww we
“JI or

| And see Key L, couplet 10, for ‘“Orchomenella’’ groenlandica (Hansen).

25.

26.

27.

28.

29.

10.

MARINE GAMMARIDEAN AMPHIPODA 311

Setae of inner plate on maxilla 1 mostly terminal, mandibular molar ex-
tremely broad even though laminate and subconical, with girdle of ridges
or setules (pleonite 3 with dorsal notch), mandible when flattened,
with deep S-shaped insinuation between palp and incisor (fig. 126d).

Eurythenes

Setae of inner plate on maxilla 1 medial only or medial and terminal, man-
dibular molar ‘‘absent”’ or long, narrow, conical and lacking distinct
girdle (apparently pleonite 3 dorsally smooth), mandible when flattened

with shallow coneavity between palp and incisor. . . . piteg Nau a7A0)
Coxa 1 longer than broad, ovatorectangular, as long as its prorele 2, lobes of
maxilla 2 equally broad. ... . . . . . . Paralicella

Coxa 1 as broad as long and auipaniadiante, andl oni wo thirds as long as its
article 2, inner lobe of maxilla 2 much broader than outer (lobes often

ORNS OM OVER) Serer) Gaara Pape Cee hens Ure not PATISUAS
Mandibular molar ineaine aetulese ... . . . . . . . Lepidepecreopsis
Mandibular molar strongly setulose . .. . St Nv Geel! mae a Ao}
Head small, epistome and upper lip small antl inconspicuous . . . Uristes
Head of normal size, epistome and upper lip a and conspicuous, epistome

usually slightly iopane ir eelaa ESE NEI ROD VNB EA Gah a Ce ih Pam a ae)

Article 7 of gnathopod 1 with accessory “oot
some species of ‘“‘Tmetonyx” auct. [= 7ryphosella]

Article 7 of gnathopod 1 simple . . . . Tryphosella [=‘“7ryphosa” auct.]
KEY O

Pereopods 1-5 prehensile (fig. 130d)... .... .. . . Metacyclocaris

Pereopods 1—5 not prehensile. . . . . Py eRe aise

Lobes of maxilla 2 gaping, and/or inner ane accel proaden than outer
(fig. 1277), mandible with molar and gnathopod 1 nearly simple. Aristias
Lobes of maxilla 2 appressed and of intersimilar shape, characters not in

SimMilarmcomtoimatiomexts joke eis teAuiery arma Pernciie EDS Poa hRiheeetns. ee
Gnathopod 1 simple .. . gripes, suk A TR Se kd Aree ema sca To ee
Gnathopod 1 subchelate, even einonen WwW DT BANG Oe se a ee Ser
Mandible lacking molar (or if rudiment present, palp aiiecinad Sosa to

TOMOMENE)) gs 6 oe i eV het tier on SAeU NR Tae ae Fad Toe (Wy SA let coca ds bese tad)
Mandible bearing olan ue ed ened a)

Article 2 of maxillipedal palp ec) ‘ahem ell? as locale as outer “ile.
Centromedon and Ambasia
Article 2 of maxillipedal palp much broader than outer plate . Ambasiella
Outer plates of maxilliped with spines. . . ... . . . . Metambasia!
Outer plates of maxilliped lacking spines. . . .. . . . . Ambasiopsis
Inner plate of maxilla 1 with one or two immense faleate setae (fig. 127d).
Hirondellea ?

Inner plate of maxilla 1 with small, slender setae. ..........8
Inner ramus of uropod 2 dorsally notched or constricted. . ...... 9
IMnMereramUusyOL UTOpOG..2) simple wy Meade tee sey mend igeumeets ta) LoL
Gnathopod 1 subchelate. .. . A UIMOLU URC SIE ES Mee APR gh iti ti Seda WC]
Gnathopod 1 weakly chelate Ge. 1292). Po en 2 BAe VAPIstiopsis

Coxa 1 tryphosid in outline (fig. 129k).
“Orchomenella’’ groenlandica (Hansen)

1 See Schisturella diagnosis (p. 361).
2 See note, Key N, couplet 23.

312

11.

12.

13.

14.

15:

16.

17.

18.

19.

U.S. NATIONAL MUSEUM BULLETIN 271

Coxa 1 hemioval or subtriangular (fig. 129c).
Schisturella [= Pseudonesimus]

Outer lobe of maxilla 1 degenerate and Sane distally, bearing only

five or six spines. . . BEE ean) mien cam angel
Outer lobe of maxilla 1 aun) distill bearing seven ormore spines. . .13
Apices of maxilla 2 lobes densely setose, with nine or more setae each.
Neoambasia
Apices of maxilla 2 lobes sparsely setose, inner with one seta, outer with
about four setae (fig. 127k)... . . . . . . . Adeliella
Medial edge of inner lobe of maxilla 1 fully aeiase, enfticlle 3 of poenoned
1 elongate, as long as article 6..... . . . . . Paralicella
Only distal end of inner lobe of maxilla 1 aeines article 3 of gnathopod
1 not elongate... . Pua Sees

Mandibular molar an immense, [mend panier seumlese, girdled lemma
(fig. 126d), article 1 of antenna 2 enlarged and circular in adults (fig. 1247),
inner plate of maxilla 1 with five or more setae. . . . . . Eurythenes

Mandibular molar of medium or small size, cuboid, subspheroid, or conically
laminate, article 1 of antenna 2 not subcircular or eae inner plate of

maxilla 1 with three or fewer setae. . . Heat es UNS alo
Article 4 of maxillipedal palp vestigial (fig. 1284) . . . . .Centromedon
Article 4 of maxillipedal palp not vestigial. . .. . Reet a ICG)

Mandibular molar very small, sublaminate, conical or ‘hava, article 6 of
gnathopod 1 enlarged (fig. 1291).
some species now assigned to Cheirimedon
Mandibular molar of normal size and cuboidal dimensions, article 6 of gnatho-
pod small es Atte artes rear le Gh
Outer ramus of uropod 3 cinfantenlate, Tedibulen molar nideed)
Tryphosoides
Outer ramus of ee 3 biarticulate, mandibular molar unridged but
setulose;. 9.23 Sia Sabin ttc lke
Epistome from tera view oxiondlins anh fareher hen nel projecting
convexly in front of upper lip (fig. 1251).
Tryphosella (‘‘Tryphosa’’? auct. and some species of “‘7'’metonyx’’ auct.)
Epistome small, short, unprojecting, occasionally dominated by upper
lip (figs. DEO) Ne J NSE reek)
Upper lip and epistome sane Mreoncpicnee Ges. l2soa))
some species of Uristes

Upper lip dominating epistome (fig. 125n) (head ?large, coxa 1 an elongate

triangle in type-species, variable in others). . . . . . . Ambasiopsis
KEY P

Gnathopod! I subchelateseven minutely.) 915.2240 ee
Gnathopod 1 simple. . . . Reimann ws cous ld
Hither article 4 or 5 of smieama 2 Fer nnd (Ge 124k). Moe Sis 4) ch i@)
Neither article: 4 or 5) of, antenna: 2 tumids jos ese shee
Broad epistomal lobe dominating labrum. . . .. . . . . . Microlysias
Labrum strongly projecting in front of epistome ... . . . . Lysianella
Epistome and upper lip amalgamated even though their parts produced,

parts not separated by notch (fig. 125k). . .. . Bee even 0)
Epistome and upper lip separated by deep notch (figs. 125¢ if) Bina parity Ce

Maxillipedal palp 3-articulate, peduncle of uropod 3 elongate, nearly as long
as pleonite 4 and peduncles of uropods 1-2 and longer than rami of uro-
DOGS ess Sr Ee SERS Tet as sen Ue re un ove ea NOT STRATE TY

10.

11.

12.

13.

14.

15.

16.

21.

22.

MARINE GAMMARIDEAN AMPHIPODA 313

Maxillipedal palp 4-articulate, peduncle of uropod 3 short, subequal to
pleonite 6 and much shorter than peduncles of uropods 1-2. ... . 6
Epistomal portion of prebuccal complex slightly produced, or flush, inner
ramus of uropod 2 simple, inner ramus of uropod 3 half or less as long as
outerramus... . ... . . Nannonyx
Labral portion of erepuccall someon neately) mrodced! inner ramus of
uropod 2 constricted, inner ramus of uropod 3 exceeding three fourths

length of outer ramus. .. . . . . . . Paralysianopsis
Inner ramus of uropod 2 gemsanciedl Ge 131d) Ly CN ol eee oe Mike em Mir eet Aor )
Ininersramus of Uropodi2 simples sgn fea eye ete es ean
Coxa 1 normally quadrate . . . . . . Pseudokoroga

Coxa 1 tapering distally and Partially elton ie coxa 2.
“Orchomenella”’ groenlandica (Hansen)
Article 1 of antenna 1 dorsodistally crested or toothed (fig. 124d), body
often crested dorsally. . .. . . . . . . Lepidepecreum
Article 1 of antenna 1 simple, body spoon or wiesios only on pleonite 4. 10
Article 3 of gnathopod 1 normally short and stout, articles 5 and 6 normally

stout (ig. 129d)... . . . . Orchomene
Article 3 of gnathopod 1 dloneste andl Bender like osnls 3 of prerhonod Ds

articles 5-6 elongate and slender (fig. 1297) . . . . . Pseudorchomene
Telson entire or emarginate ... . dg Rami etak RS US es Ae a aK A Pst NL
Telson cleft one third or more .. . cihike Bias) Sr ae ne ge LD)
Inner ramus of uropod 3 half or less ironed af Outer Ms Parawaldeckia
Inner ramus of uropod 3 three fourths or more length of outer . . . . 13

Upper lip prominently lobate, occasionally epistome also lobate (fig. 125c).
Lysianassa, ?Socarnella, [Pronannonyx]

Upper lip not prominently lobate ... . Slee Santee We
Telson distinctly longer than wide, inner ramus ai moped 2 constricted.!
Parambasia
Telson not longer than wide, inner ramus of uropod 2 simple.

Menigratopsis
Article 2 of pereopod 3 deeply indentured (fig. 131b), article 3 of gnathopod 1
elongate and slender like article 3 of gnathopod 2... . . Glycerina
Article 2 of pereopod 3 with small teeth or none, article 3 of gnathopod 1
not elongate .. . Aad aces eat ALG)
Coxa 1 tapering aictally or cnonrened andl ear ial frdden by coxa 2
(Aig eI Obie eae aes Ane ba cleat a bh
Coxa 1 large, subquadrate or pearcely tapering, riety Sr eiples icon Van cl Ka)
Palp article 2 of maxilliped half or less as broad as outer plate . . Ambasia

Palp article 2 of maxilliped as broad as or broader than outer plate.
Ambasiella
Inner ramus of uropod 2 constricted... ...... . . . Socarnoides
Innerjranis ofirepod 2rsimple i. fv je) ol ie) Cena tyae a (LO
Outer ramus of urepod Ss) umiarticulate i.) Gena sa ee 20
Outer ramus of uropod 3 biarticulate. . . . . iimedclstneee ly Pall
Palp article 2 of mandible about 3 times as lone as eral 1, article 3 two
thirds length of article 2... .°. . . . . . Socarnopsis
Palp article 2 of mandible about 5 Aime as fone as article 1, article 3 about
halfas lone as article 2,204) 0 ks. wale ect ye.) Socarnella
Gillsistrongly:plaited) (figs 2s). 5 ot ye ee Socaknes
Gills minutely ridged or smooth . 22

Telson about as long as wide, cleft less (han hear pills suave

Menigrates
Telson about twice as long as wide, cleft more than halfway, some gills with
basalzaccessoryslobes 8 24) Win ay ye Waldeckia

! Assumed in type-species.

314

10.

11.

16.

U.S. NATIONAL MUSEUM BULLETIN 271

KEY Q
Mandibular molar distinctly triturative and usually columnar, cylindrical or
cuboidal, bearing ridges or cusps for grinding (figs. 126d,g,m) . . . . 2
Mandibular molar not triturative or absent, often setulose or bearing a few
articulate spines (figs. 126e,h,7,j) . . . Bete Nance a 17
Coxa 1 noticeably reduced and partially nvidldlen onl coxa o or coxa | tapering
Or wrlenoepene 55 4G spe! SLA aie cane eam
Coxa 1 large, visible, nacional ¢ or alistetlse spared. Si ei lela a
Mandibular molar with triturative aac molar immense and not columnar
(ficemlt2 OG) eee . . . . . Eurythenes

Mandibular molar nor anol aoitnnmen or 2 adie raped and medium in size. 4
Telson emarginate or cleft one sixth or less, maxilla 2 with setae reduced to

One, to fours pernloben (igen 27/1) i weenie .. .. . Adeliella
Telson cleft one third or more, maxilla 2 si ‘iipibes strongly setose
CATR AED CSN cS ae eeuG Ue MAUNA Og 3 | &

Inner ramus of uropod 2 simple.
Tryphosella, Tryphosoides, and ‘‘Pseudotryphosa’”’ (?= Uristes)

Inner/ramus) of uropod!2) constrictediy 4) ey sed eee ee
Epistomal region not dorsoventrally elongate. . . . . . . . Schisturella
Epistomal region dorsoventrally elongate . ..... . .. . . Aristiopsis
Gnathopod di simple wh esa ea eS
Gnathopod 1 subchelate. . . . Be dessa Hessen el 1h G)
Telson as broad as long, oremnate BAI a Vonicinmele Pecudaieroeial
Telson longer than broad, deeply cleft . .... . . . . Menigratopsis

Telson as broad as long, cleft halfway.
Rifcus and Paronesimus (sec diagnoses)
Inner ramus of uropod 3 vestigial or absent ane 132k), always less than

half length of outerramus... . Rete Manta NA i oT)
Inner ramus of uropod 3 subequal to outer 7 LATO See ibis eee ihie dk!
Inner ramus of uropod 3 present, article 1 of both flagella of antenna 1

elongate and that of accessory flagellum expanded . . . . Onesimoides
Inner ramus of uropod 3 absent, article 1 of both flagella of antenna 1 of

normal size .. . . 2 a. ss a ss . . . Paronesimoides
Accessory flagellum 2- Perle: eRe eM ta erie gg!» LLP
Accessory flagellum 3+--articulate . . . Maer eee oh lee
Spines on outer plate of maxilla 1 Poomeed 6 eae or six. . . . .Rifcus
Outer plate of maxilla 1 with nine or more spines. . . a oe nae ie)
Hand of gnathopod 1 enlarged and expanding distally . Ponenimedan Gane
Hand of gnathopod 1 of normal unexpanded dimensions. . . Tryphosoides

Telsonic apices with seven or more spines each (fig. 128s).
Hippomedon [= Paratryphosites]
Telsonic apices with three or fewer spines or telsonentire. ..... . .15
Pelson cleft jhaliway ermorel oo pani ais. th See erie dae
Telson cleft one third or less.
Boeckosimus and Onisimus [= ‘“‘Onisimus’’ and ‘‘Pseudalibrotus’’}
Inner ramus of uropod 2 simple, epistome unproduced and slightly dominated
by labrum (from lateral view) (fig. 125n).
Hippomedon ! and [Paracentromedon] !
Inner ramus of uropod 2 deeply constricted, epistome acutely produced
ig 12Gb) i as ina ae a co Ge Oe an a ie Ree Gu nM OSITes

! And see Cheirimedon femoratus (Pfeffer) and C. fougneri Walker distinguished from these genera by the
transverse palm of gnathopod 1 (sce pp. 334-335).

17.

18.

IS).

20.

21.

22.

23.

24.

25.

26.

27.

28.

29.

30.

él.

32.

MARINE GAMMARIDEAN AMPHIPODA 315

Gnathopod 1 simple?. . . By Pt od eae ey cua LTH Ga crt ont Ame ne)
Gnathopod 1 distaois pubchelate ra eas Brea RUC
Coxa 1 reduced or distinctly eerie from nase aul Sian strongly hidden
lOs7 COkG Boo 65 co Sa tes eee! Re)
Coxa 1 large, TaviemeEe: aml not fiddent fener if resent occurring in
onlysdistalithird of ‘coxays 2 2a Pia oe hf ud ae seisunae) vase 26
Rereopods) prehensile) sive aie sy eis se eh ee victacelacaris
Pereopods simple. . . Ha) We pbuivees see «20
Maxillipedal palp with iheeel catiotes telecon eaaie) . .. . . . Perrierella
Maxillipedal palp with four articles, telson cleft. . ......... .2!21
Maxillipedal palp article 4 vestigial (fig. 128a). .......... .22
Mazxillipedal palp article 4 well developed and claw-like. . . .... . .238
Mandibular molar present, upper lip and epistome coalesced, epistomal
region strongly produced ..... ... . .Centromedon
Mandibular molar absent, upper lip and soitome separated by notch,
upper lip slightly protruding .. . . . . .Ambasia
Inner plate of maxilla 2 much broader Aten oniar plate aaa usually gaping
mediad. .... ... . . .Aristias
Plates of maxilla 2 seemed in Raden nal not pironely, CApINGeees eet. 1. 24:
Upper lip and epistome not differentially produced, small and inconspicuous.
Uristes
Upper lip projecting as lobe in front of epistome (from lateral view) . . 25
Outer plates of maxilliped lacking spines, gnathopod 1 weakly subchelate,
coxa 1 nearly as long as article 2 of gnathopod 1... . . Ambasiopsis
Outer plates of maxilliped spinose, gnathopod 1 perfectly simple, coxa 1
about half as long as article 2 of gnathopod1 .... . . Metambasia
Inner plate of maxilla 1 fully lined medially with eight or more setae.
Alicella
Inner plate of maxilla 1 with three or fewer terminal setae . . . . . . 27
Inner ramus of uropod 3 vestigial, less than one third length of outer.
Clepidecrella
Inner ramus of uropod 3 subequal to outer. . ......2.2.2.2.~. 28
shelsonicleftxone| third or more! 292 >. S3ee si. Bt. aie. ee eee 29
Telson entire or emarginate . . ahh. coer 0)
Gnathopod 1 perfectly simple, pieader “oiibgiviitemn, dacel strong and
bearing posterior setal brush near base (fig. 1307) . . . . . . Ichnopus
Gnathopod 1 perfectly simple, slender, dactyl strong and lacking posterior
setal brush. .... . . . . . Menigratopsis

Gnathopod 1 weakly pubele lates ee raediuen sienna, dactyl strong but
lacking posterior setal brush although bearing small accessory tooth.

Paronesimus
Labral region of prebuccal complex with strongly acute anterior lobe [telson
longer than broad]. ...... . . . . . Paralysianopsis

Labral region of prebuccal complex aieh convened rounded keel or unpro-
duced [telson as broad as long in Paralibrotus only].

Menigrates (in part), Paralibrotus

Coxa 1 reduced or distally tapering from base and often strongly hidden by

COXA) 275 a ee SAS OES, Oe
Coxa 1 large, receaanlee inal aah hidden by coxa 2, panes if Berea occurring
only in distal third of coxa... . NSRORILGE., Na mim nada cman ASD
Inner plate of maxilla 1 with three or more Botae UNS ee SB RU SeR en OO

Inner plate of maxilla 1 with two or fewer setae (or inner plate absent). 35

2 Some latitude is expressed in this division as certain genera are found in both pathways.

285-135 O - 69 - 21

316
33.

34.

30.

36.

37.

38.

39.

40.

41.

42.

43.
44.
45.

46.

47.

48.

U.S. NATIONAL MUSEUM BULLETIN 271

Maxillipedal palp with three articles (fig. 1287) .... . . . Perrierella
Maxillipedal palp with four articles . . . ; URE:
Plates of maxilla 2 subequally broad and nae capstan, mtiiclle 1 of antenna 2

very enlarged and subspherical (adults), telson elongate . . Eurythenes

Inner plate of maxilla 2 much broader than outer, inner plate often strongly
gaping mediad (fig. 1277), article 1 of antenna 2 normally small, telson

not elongate... . WR bo) a APIStaSs
Palp article 4 of amped ood rests Ge 128k) . . . . . Centromedon 2
Palp article 4 of maxilliped well developed and claw-like. ..... . 36

Inner plate of maxilla 1 with at least one large falcate seta (fig. 127d).
Hirondellea 4

Inner plate of maxilla 1 with normal setae ... . MeEESe DERE ay Cai aI
Setae of maxilla 2 reduced to one-two and four ee inner ianid outer lobes

CED eyes seis ceils eae eo deliell a!
Setae of maxilla 2 numerous 4 per tobe): BU ae is PVRS LER NEC tee ool Pea teS
Outer plate of maxilla 1 with spines reduced to five or fomee . Neoambasia
Outer plate of maxilla 1 with seven or more spines... . Sots Oe esa)

Gnathopod 1 enlarged, article 5 very short and with small, narrow posterior
lobe, article 6 expanding distally, very broad, palm transverse or nearly

chelate (fig. 1291) . . . .. . . . . Cheirimedon latimanus Sars, 1895
Gnathopod 1 of normal size, article 5 rarely lobate, article 6 not expanding
distally even though palm rarely transverse. . . . ic iente 40
Labrum with distinct lobe projecting in front of epierome (irom lateral view).
Ambasiopsis
Labrum not dominating epistome . . . Be a ole vege tice a
“Head small’ (fig. 1236), epistome ond mpsa el gine, inconspicuous,
neither dominant . . . “ana e) SanUEIStes
“Head large’’ (fig. 122d), Prebacenl complex pone pienGne: epistome usually
dominating upper lip ..... .. . . . Tryphosella
Inner plate of maxilla 1 with eight or more aeuae le medial edge.
Paralicella
Inner plate of maxilla 1 with three or fewer setae... . foe: Auentee tee
Coxa dS as\longias4 (fe 21D) ) >. = eee Tepidepeceenidas
Coxa 5 much shorter than4 .... . BBY es SOG LAM RS ek
Telson entire or cleft less than one sein its lenenh PR MMR i ekey dk oo)
Telson cleft one eighth or moreitslength. ....... Seer Senet aC
Outer ramus of uropod 3 uniarticulate ......... Ubeeudarabacis
Outer ramus of uropod 3 biarticulate . . . . Sei eG

Labrum rounded anteriorly and scarcely produced nerond iepietoue’ inner
ramus of uropod 2 simple, article 6 of gnathopod 1 coe and stout,
palm transverse (fig. 129a) . . . . ... . . Koroga

Labral area of prebuccal complex nanialy ‘prodiiead| tt in front of epistomal
area, inner ramus of uropod 2 constricted, article 6 of gnathopod 1 slender,

palm obsolescent, oblique. . . . . . . Paralysianopsis
Palp article 4 of maxilliped very stout ind supeluate. dactyl of gnathopod 1
as short and stout as dactyl of gnathopod 2. . . . . . Pseudoanonyx
Maxillipedal palp article 4 typically unguiform (fig. 128c). .... . 48
Epistome with acute anterior process dominating upper lip. . Tryphosites
Hpistomedlationroundedvanteriorly 6 45 ane ee

3 Compare Adeliella.
4 See note, Key N, couplet 23.

MARINE) GAMMARIDEAN AMPHIPODA 317

49. Gills plaited (deeply folded), labral lobe slightly dominating epistome.
Anonyx
Gills simple (at most striate), epistome equivalent to or dominating labral
IWoY YS): sisi ced aa te anes ISIC eam MR ANS Sous sg ated Aer Tmetonyx (type only)
Gills simple (at most striate), labral lobe slightly dominating epistome.
Paronesimus

Genera of Lysianassidae

The following characters are not mentioned in the diagnoses unless
they have classificatory value; if not mentioned they conform to the
typical condition as stated in sequel: gnathopod 2 either minutely
chelate or subchelate; mazilla 1 with 2-articulate palp, inner lobe
poorly setose; maailliped with well-developed lobes and 4-articulate
palp, the fourth article large and claw-like; coxae 1 or 2 large, visible;
uropod 8 of medium length, outer ramus 2-articulate, inner slightly
shorter than outer, rami lanceolate; accessory flagellum multiarticulate;
inner ramus of uropod 2 unconstricted or unnotched; mouthparts not
formed into a cone-shaped bundle; pereopod 3, article 2 not deeply
indentured.

Acidostoma Liljeborg

Acidostoma Liljeborg, 1865.—Stebbing, 1906.

Type-species: Anonyx obesus Bate, 1862 (monotypy). See Sars,
1895.

Mouthparts forming a ventral conical bundle, styliform; antenna
1 very stout in male; upper lip and epistome continuous; mandible
usually lacking molar, small when present, palp attached quite
proximally; palp of maxilla 1 vestigial, 1-articulate, outer plate with
hook-like spines; palp of maxilliped scarcely exceeding outer plate,
article 4 vestigial; gnathopod 1 simple, article 6 slightly longer than 5;
gnathopod 2 lacking article 7; uropod 3 short, outer ramus 1-articulate;
telson short, cleft or emarginate. Species: 7, boreal, littoral to
bathyal (1,200 m).

Acontiostoma Stebbing

Acontiostoma Stebbing, 1888; 1906.

Type-species: A. marionis Stebbing, 1888 (original designation).

Mouthparts forming a ventral conical bundle, styliform; mandible
lacking molar, palp attached quite proximally; palp of maxilla 1
uniarticulate; maxillipedal palp scarcely exceeding outer plate, article
4 very small; gnathopod 1 simple, article 6 slightly longer than 5;
coxa 1 projecting over side of head; uropod 3 lacking rami; telson
short, entire or emarginate. See Stomacontion for remarks. Species:
1, subantarctic, littoral.

318 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 117.—Lysianassidae: a, Onisimus (=Pseudalibrotus) litoralis (Kréyer) (Sars, 1895,
pl. 35); b, Stomacontion pepinii (Stebbing, 1888), note coxa 1 overlapping head, and eye
dimly showing through.

MARINE GAMMARIDEAN AMPHIPODA 319

all
Zipp [fh
yf

NN

Figure 118.—Lysianassidae: a, Lepidepecreum umbo (Goés) (Sars, 1895, pl. 39); b, Kergue-
lenia borealis Sars (1895, pl. 40).

320 U.S. NATIONAL MUSEUM BULLETIN 271

Adeliella Nicholls

Adeliella Nicholls, 1938.

Type-species: A. laticornis Nicholls, 1938 (original designation).

Upper lip and epistome not differentially produced; mandible with
“distinct molar”, palp apparently attached level with molar; maxilli-
pedal palp article 4 rather small; maxillae poorly spinose or setose;
gnathopod 1 subchelate, short, articles 5 and 6 equal in length; telson
short, emarginate or notched apically. Species: 1, antarctic, bathyal
(420 m).

Alicella Chevreux

Alicella Chevreux, 1899.—Stebbing, 1906.

Type-species: A. gigantea Chevreux, 1899 (original designation).
See Chevreux, 1935.

Upper lip and epistome not differentially produced; mandibular
cutting edge with small teeth, molar conicolaminate, setulose, palp
attached level with molar; inner plate of maxilla 1 densely setose on
medial edge; maxillipedal palp article 4 very small; gnathopod 1 feeble,
simple, articles 5 and 6 equal in length; article 6 of gnathopod 2 as
long as article 5, both very elongate and linear; rami of uropod 3 long,
equal, lanceolate, apparently outer ramus 1l-articulate; telson of
medium length, deeply cleft. Species: 1, N. Atlantic, abyssal.

[Allogaussia Schellenberg]
Allogaussia Schellenberg, 1926a.—K. H. Barnard, 1932.

Type-species: A. paradoxa Schellenberg, 1926a (selected by
Stasek, 1958).

Type-species distinct from type-species of Orchomene by entire
telson, produced posterior lobe of coxa 5, asymmetrical posterior
lobe of pereopod 3. Species that have been assigned on basis of
telson alone: 6 (also included in Orchomene).

Amaryllis Haswell
Amaryllis Haswell, 1880b.—Stebbing, 1906.

Type-species: A. macrophthalmus Haswell, 1880b (selected by
Pirlot, 1933b; indicated by Stebbing, 1888, p. 702!, through monotypy
by synonymy).

Epistome apparently projecting in front of upper lip; mandibular
molar feeble, setulose, palp attached near proximal end of molar;
maxilla 1 lacking palp; gnathopod 1 simple, article 6 longer than 5;
coxa 1 partially hidden, following coxae increasingly larger; inner
ramus of uropod 2 constricted; telson of medium length, half or

MARINE GAMMARIDEAN AMPHIPODA 321

deeply cleft; article 2 of antenna 1 half as long as article 1 (Bathya-
maryllis) ; anteroventral corner of coxa 4 rounded (Vyaya). Species: 2,
antiboreal, littoral.

Ambasia Boeck

Ambasia Boeck, 1871.—Stebbing, 1906.

Type-species: Gammarus atlanticus Milne Edwards, 1830 (mono-
typy and subsequent synonymy). See Sars, 1895.

Epistome and upper lip coalesced, but epistomal part greatly and
sharply produced in front; mandibular molar absent, palp attached
to middle of mandible; marillipedal palp article 2 half or less as broad
as outer plate (Ambasiella), article 4 very small; gnathopod 1 simple,
article 5 slightly longer than 6; coxa 1 slightly shortened and partially
hidden by coxa 2; inner ramus of uropod 3 half as long as outer;
telson of medium length, deeply cleft. Species: 1, subarctic N.
Atlantic, littoral and bathyal (to 1,400 m).

Ambasiella Schellenberg

Ambasiella Schellenberg, 1935.

Type-species: Ambasia murmanica Briiggen, 1905 (monotypy).
See Gurjanova, 1951.

Epistome and upper lip not differentially produced; mandibular
molar absent, palp attached proximally; mazillipedal palp article 2
much broader than outer plate (Ambasia, Hirondellea), article 4 vestig-
ial; gnathopod 1 simple, article 5 longer than 6; coxa 1 slightly
shortened, coxa 2 concealing most of it; telson of medium length,
deeply cleft. Species: 1, arctic, littoral.

Ambasiopsis K. H. Barnard

Ambasiopsis K. H. Barnard, 1931; 1932.

Type-species: A. georgiensis K. H. Barnard, 1931 (original
designation) ; 1932.

Upper lip lobately produced in front of epistome; mandibular
molar large, truncate, not ridged, setulose, palp attached level with
molar; outer plate of maxilliped lacking spines (Neoambasia) ; gnathopod
1 poorly subchelate; coxa 1 two thirds as long as coxa 2 and partially
hidden by it, but as long as article 2 of gnathopod 1; telson deeply
cleft. Species: 2, subantarctic, littoral (to 342 m).

U.S. NATIONAL MUSEUM BULLETIN 271

vik ;
A/ i ‘

a

SEF
V7 RY)
ae ti

i Ce

fa” hye Mee VS

=

Ficure 119.—Lysianassidae: a, Eucallisoma glandulosa J. L. Barnard (1961); b, Ambasia

atlantica (Milne Edwards) (Sars, 1895, pl. 17, as 4. danielssent); c, Pachychelium davidis
Stephensen (1925a).

MARINE GAMMARIDEAN AMPHIPODA 323

VAN Uh
Tp x ,
YY) Ns) 5} f a

4 [ ) [>
Soe lim
(| cl WS
| Ol Lyf ft ff
y,
Seated «| al
Figure 120.—Lysianassidae: a, Acid b (Bate) (Sars, 1895, pl. 114); b, Tryphos-

esum
Barnard, 1962d).

324 U.S. NATIONAL MUSEUM BULLETIN 271

Anonyx Kr¢yer

Anonyz Kr¢gyer, 1838.—Stebbing, 1906.—Gurjanova, 1962.—Hurley, 1963.
Chironesimus Sars, 1895.
Lakota Holmes, 1908.

Type-species: Cancer nugax Phipps, 1774 (elimination and syn-
onymy to result in monotypy by Boeck, 1871, but firmly selected
by Boeck, 1876, with subsequent synonymy). See Sars, 1895.

Upper lip projecting slightly or strongly in front of epistome;
mandibular molar large, or of medium size, subconical, unridged
(Boeckosimus), setulose, palp attached level with molar; gnathopod
1 subchelate, articles 5 and 6 variable in length; inner ramus of
uropod 2 constricted or not; telson of medium length, deeply cleft;
gills plaited except in Lakota; eyes present; pereopods 1-2 unrth distal
locking spines on article 6 at base of dactyls; article 4 of mazillipedal
palp slender and claw-like (Pseudoanonyz). Species: 30+, arctic-
boreal, littoral to abyssal.

Aristias Boeck
Aristias Boeck, 1871.—Stebbing, 1906.

Type-species: Anonyx tumidus Krgyer 1846 (monotypy). See
Sars, 1895.

Upper lip either projecting or not projecting beyond epistome,
occasionally lobate, angular or poorly produced; mandibular molar
a long unridged conical lamina, palp attached level with molar;
inner lobe of maxilla 1 strongly setose (5+ setae) and spinose; lobes
of maxilla 2 often gaping, inner much broader than outer; gnathopod
1 nearly simple or with minute palm, article 7 overlapping palm,
articles 5 and 6 equal, article 3 not elongate (Paralicella); coxa 1
short, half as long as its article 2, partially hidden by coxa 2; telson
of medium length or short, deeply cleft. Differs from Eurythenes
by: shorter telson, broader inner plate of maxilla 2, narrow mandibular
molar, longer article 2 of pereopod 8. Species: 16, primarily bipolar,
amphiboreal (one tropical), littoral to bathyal.

Aristiopsis J. L. Barnard
Aristiopsis J. L. Barnard, 1961.

Type-species: A. tumidus J. L. Barnard, 1961 (original desig-
nation).

Both epistome and upper lip with short anterior humps separated
by a long, flat, unnotched margin; mandible with well-developed,
ridged molar, palp attached over posteroproximal corner of molar;

MARINE GAMMARIDEAN AMPHIPODA 325

gnathopod 1 stout, slightly chelate; coxa 1 short, half as long as its
article 2, partially concealed by coxa 2; uropod 2 with inner (not
outer = error of J. L. Barnard, 1961) ramus slightly constricted;
telson short, cleft one third its length. Species: 1, Pacific, bathyal-
abyssal.

Ficure 121.—Lysianassidae: a, Danaella mimonectes Stephensen (1925b); b, Lepidepecreoides
xenopus K. H. Barnard (1932).

326 U.S. NATIONAL MUSEUM BULLETIN 271

VES
POOR i
Ȥ Cee oe

Ficure 122.—Lysianassidae: a, Thoriella islandica Stephensen (1915), offset showing dorsal
urosome; b, Onesimoides chelatus Pirlot (J. L. Barnard, 1961); c, Crybelocyphocarts tattersalli
Shoemaker (1945a); d, Tryphosella sarsi Bonnier (Sars, 1895, pl. 27, as Tryphosa nana).

MARINE GAMMARIDEAN AMPHIPODA 327

Ficure 123.—Lysianassidae: a, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as
T. raschi); b, Uristes umbonatus (Sars, 1895, pl. 29); c, Hippomedon denticulatus (Bate)
(Sars, 1895, pl. 20); d, Centromedon pumilus (Liljeborg) (Sars, 1895, pl. 34); e, Acidostoma
obesum (Bate) (Sars, 1895, pl. 14).

328 U.S. NATIONAL MUSHUM BULLETIN 271

Aroui Chevreux
Arouti Chevreux, 191 1a.

Type-species: A. setosus Chevreux, 191la (monotypy).

Upper lip and epistome not differentially produced, epistome flat in
front (Scopelocheirus); mandibular molar small, cylindrical, ridged,
palp attached slightly proximal to molar; inner plate of maxilla 1
extremely setose; lobes of maxilla 2 subcircular, outer shorter than
inner; article 6 of gnathopod 1 terminated by short shroud of setae;
article 7 vestigial; gnathopod 2 minutely chelate (Paracallisoma); telson
of medium length, deeply cleft. Species: 1, Algeria, littoral.

Azotostoma J. L. Barnard
Azotostoma J. L. Barnard, 1965.

Type-species: <A. fusta J. L. Barnard, 1965 (original designation).

Mouthparts forming a ventral conical bundle, styliform; mandible
lacking molar, palp attached very proximally; palp of maxilla 1
large, exceeding outer plate, bent, 2-articulate; inner plate of maxilla
2 expanded proximally, distal tapered portion with three stout spines;
inner plates of maxilliped styliform, outer very large and flagellate,
palp 4-articulate, scarcely exceeding outer plate, article 2 produced
distally; gnathopod 1 simple, flagelliform, article 6 much longer than
5, article 7 elongate but not as long as article 6; uropod 3 biramous,
rami equal, 1-articulate; telson short, entire. Species: 1, Micronesia,
littoral.

Bathyamaryllis Pirlot

Bathyamaryllis Pirlot, 1933a.

Type-species: JB. perezii Pirlot, 1933a (original designation).

Epistome projecting slightly in front of upper lip; mandible with
long, subconical, setulose molar, palp attached level with molar; palp
of maxilla 1 absent; gnathopod 1 simple, article 6 longer than 5; coxa
1 small, partially hidden, subsequent coxae increasing in size; uropod
3 with l-articulate outer ramus; telson of medium length, deeply
cleft; article 2 of antenna 1 equal to article 1 (Amaryllis). Species: 4,
probably cosmopolitan, bathyal to abyssal.

Bathycallisoma Dahl

Bathycallisoma Dahl, 1959. Synonym of Scopelocheirus fide Birstein and
Vinogradov, 1964. See also Paracallisoma.

Type-species: B. pacifica Dahl, 1959 (monotypy).
Upper lip and epistome not differentially produced; mandibular

MARINE) GAMMARIDEAN AMPHIPODA 329

molar small, conical, unridged, palp attached level with molar but
both rather proximal; lower lip with major lobes excavate medially,
forming medial gap (combining character); inner plate of maxilla 1
densely setose; gnathopod 1 simple, article 7 small, article 6 tipped
with short setal shroud; gnathopod 2 subchelate but not distinctly
chelate (Scopelocheirus); telson of medium length, deeply cleft.
Species: 1, central Pacific, hadal.

Ficure 124.—Lysianassidae: a, Chevreuxiella metopoides Stephensen (1915), offset showing
dorsal urosome (3, 4, 5-6; U=uropod). Antenna 1: b, Bathyamaryllis perezit Pirlot
(1933a); c, Lysianassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); d, Lepidepecreum
longicorne (Bate and Westwood) (Sars, 1895, pl. 38, as L. carinatum); e, Ambasia atlantica
(Milne Edwards) (Sars, 1895, pl. 17, as 4. danielsseni); f, Socarnotdes kergueleni Stebbing
(1888); g, Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); h, Normanion sarsi Stebbing
(Sars, 1895, pl. 13, as N. quadrimana). Antenna 2: 1, Eurythenes gryllus (Lichtenstein)
(Sars, 1895, pl. 30); 7, Parawaldeckia kidderi (Smith) (Tattersall, 1922); k, Lystanella
petalocera Sars (1895, pl. 18).

330 U.S. NATIONAL MUSEUM BULLETIN 271

Boeckosimus, new genus

Type-species: Anonyx edwardsi Krgyer, 1846 (present selection).
This genus embraces those species formerly included in the genus
Onisimus Boeck, 1871, since the time of Sars (1895) and Stebbing
(1906). The genus Onisimus must include those species having gone
in the genus Pseudalibrotus Della Valle, 1893, since the time of Steb-
bing (1906). Boeck (1876) designated Anonyzx litoralis Krgyer, 1845,
as type of Onisimus Boeck, 1871, and Pseudalibrotus Della Valle
is a junior synonym of Onisimus because of monotypy, A. litorahs
also being the type-species of Pseudalibrotus. Sars (1895) designated

Uy

Ficure 125.—Lysianassidae: Upper lip-epistome complex, lateral view, left sides: a,
Socarnoides kerguelent Stebbing (1888); b, Tryphosites longipes (Bate and Westwood)
(Sars, 1895, pl. 28); c, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); d,
Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); e, Eurythenes gryllus (Lichtenstein)
(Sars, 1895, pl. 30); f, Orchomene batei Sars (1895, pl. 22); g, Ichnopus spinicornis Boeck
(Sars, 1895, pl. 15); 4, Ambasia atlantica (Milne Edwards) (Sars, 1895, pl. 17, as 4. dani-
elsseni); 1, Lepidepecreum longicorne (Bate and Westwood) (Sars, 1895, pl. 38, as L.
carinatum); 7, Orchomene amblyops Sars (1895, pl. 25); k, Nannonyx goest (Boeck) (Sars,
1895, pl. 24); 1, Orchomene minuta (Kr¢yer) (Sars, 1895, pl. 24); m, Scopelochetrus crenatus
Bate (Sars, 1895, pl. 19); », Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); 0, Orcho-
mene obtusa (Sars, 1895, pl. 26). Lower lips: p, Hippomedon; q, Nannonyx; r, Normanton;
5, Eurythenes; t, Ichnopus; u, Opisa eschrichti (Krgyer) (Sars, 1895, pl. 14); v, Trischizo-
stoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi). Upper lip, anterior view: w,
Acidostoma obesum (Bate) (Sars, 1895, pl. 14); x, Trischizostoma.

v

MARINE GAMMARIDEAN AMPHIPODA 331

A. edwardsti as type of Onisimus, erroneously, and I follow his prece-
dent in using A. edwardsii as type of Boeckosimus. Gurjanova (1951)
listed Anonyx plautus Kroyer, 1845, as type of Onisemus but this
is also erroneous. The genus Alibrotus Milne Edwards, 1840, may
be a senior synonym either of Boeckosimus or Onisimus but its status
has remained unclear. Stebbing (1906) lists it as a dubious genus.
Its type-species, Lysianassa chauseica Milne Edwards, 1830 (mono-
typy), probably cannot be a member of either Onisimus (= Pseudali-
brotus) or Boeckosimus (=‘‘Onisimus”’ auct.) because no species of
either genus has been recorded as far south as Isles Chauseay in the
English Channel.

Upper lip and epistome not prominent; mandibular molar ridged
(Anonyzx), palp attached level with molar; gnathopod 1 subchelate or
poorly subchelate, articles 5 and 6 subequal or article 6 longer than
5; inner ramus of uropod 2 constricted or not; telson short (Hippome-
don), variable, entire or cleft one third; outer plate of maxilla 1 with
nine or more spines; palp of mazilliped elongate and slender, outer
plate of maxilliped reaching to end of palp article 2 (Rifcus). Species:
16, arctic-boreal, mostly N. Atlantic, littoral to abyssal.

Note: This genus (‘Onisimus” auct.) and Onisimus (= Pseudali-
brotus auct.) are very close. Sars (1895) distinguished them on the
more powerful gnathopod 1 and uropod 3 and longer antennae of
“Pseudalibrotus”’ and Gurjanova (1962) distinguished them on the
absence of bent spines on the apices of the maxillipedal outer plates
in ‘“Pseudalibrotus.’’ Most species of ‘“Onisimus’’ have slightly cleft
telsons and various species of ‘‘Pseudalibrotus’”’ have the epistomal
portion of the prebuccal complex dominating the labral portion.
All of these seem to be very weak generic characters. If the genera
were synonymized the unfortunate nomenclatural problems would be
solved.

Cebocaris J. L. Barnard
Cebocaris J. LL. Barnard, 1964a.

Type-species: OC. grutesca J. L. Barnard, 1964a (original designa-
tion).

Epistome large, broad, convex, but not projecting beyond upper
lip from lateral view; head deformed; mandibular molar absent;
gnathopod 1 simple, articles 5 and 6 subequal; article 5 of gnathopod 2
only slightly longer than article 6, appendage simple; coxae 1 and 2
small, but coxa 3 only slightly enlarged and covering coxa 2 only
partially; article 2 of pereopod 3 not indentured; pereopods 1-5
strongly prehensile; telson short, entire. Species: 1, Caribbean,
abyssopelagic.

285-135 O - 69 - 22

Ficure 126.—Lysianassidae: Mandible: a, Centromedon pumilus (Liljeborg) (Sars, 1895,
pl. 34); b, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); c, Kergu-
elenia borealis Sars (1895, pl. 40); d, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl.
30); e, Tryphosella sarst Bonnier (Sars, 1895, pl. 27, as Tryphosa nana); f, Aristias neglectus
Hansen (Sars, 1895, pl. 17, as A. audouinianus); g, Hippomedon denticulatus (Bate)
(Sars, 1895, pl. 20); h, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); 1,
Orchomene batei Sars (1895, pl. 22); 7, Anonyx nugax (Phipps) (Sars, 1895, pl. 31); &,
Acidostoma obesum (Bate) (Sars, 1895, pl. 14); 7, Menigrates obtusifrons (Boeck) (Sars,
1895, pl. 38); m, Boeckosimus edwardsi (Krgyer) (Sars, 1895, pl. 36); n, Normanion sarst
Stebbing (Sars, 1895, pl. 13, as MN. quadrimana); 0, Alicella gigantea Chevreux (1899);
p, Ichnopus spinicornis Boeck (Sars, 1895, pl. 15); g, Valettiopsis multidentata J.L.
Barnard (1961), distal portion to show teeth of incisor.

MARINE GAMMARIDEAN AMPHIPODA 333

Centromedon Sars

Centromedon Sars, 1895.—Stebbing, 1906.

Type-species: <Anonyx pumilus Liljeborg, 1865 (monotypy).

Upper lip and epistome small, but separated by a notch, upper lip
slightly projecting; mandibular molar conicolaminate, unridged, palp
attached level with molar; maxillipedal palp article 4 very small;
enathopod 1 poorly subchelate, articles 5 and 6 equal; coxa 1 slightly
tapering distally; telson of medium length, deeply cleft; head small,
lateral lobes acute (Hirondellea), mouthparts hidden by coxae. Species:
1, subarctic and boreal N. Atlantic, littoral.

Cheirimedon Stebbing
Cheirimedon Stebbing, 1888; 1906.

Type-species: C. crenatipalmatus Stebbing, 1888 (monotypy).

Upper lip and epistome not differentially produced; mandibular
molar very small, sharp or blunt, ridged (type) or poorly ridged and
sublaminate, palp attached level with molar; gnathopod 1 subchelate,
article 6 broadly expanded (type) or poorly expanded, occasionally
slightly chelate in appearance; coxae normal (type) or coxa 2 con-
cealing part of slightly shortened (and occasionally strongly narrowed)
coxa 1; telson of medium length, deeply cleft (type) or cleft as
little as one third. Some species resembling Anonyz, Boeckosimus
(=“Onisimus” auct.), Schisturella (= Pseudonesimus), and Tryphosella
(inel. “‘7ryphosa”’ auct.). Species: 5, amphiboreal, littoral to bathyal.

The type-species, Cheirimedon crenatipalmatus Stebbing (1888) is
characterized by: lateral outline of mouthpart group quadrate;
epistome and upper lip not differentially produced; head normal;
mandibular palp attached level with molar, molar process columnar,
triturative; other mouthparts normal; coxa 1 quadrate, not narrowed
distally or covered by coxa 2; gnathopod 1 subchelate, with article
6 longer than 5 and slightly expanded distally; telson cleft more than
three fourths of its length; uropod 3 normal, outer ramus 2-articulate.

Cheirimedon crenatipalmatus thus differs from Anonyz in (1) the
nonsetulose mandibular molar which in Anonyx is bulging or sub-
laminate but lacking ridges and teeth and (2) in the expanded article
6 of gnathopod 1. It differs from Tryphosella (=“Tryphosa’”’ auct.)
in the nontapering coxa 1, and the distally expanding article 6 of
enathopod 1. The type-species is joined by C. femoratus (Pfeffer) (=C.
dentumanus Chevreux, 1906b) which is distinguished generically only
by the telson being cleft halfway and by the poor expansion of gnath-
opod 1.

The other species that have been assigned to this genus depart
radically from the above brief diagnosis. Cheirimedon latimanus

334 U.S. NATIONAL MUSEUM BULLETIN 271

Sars, 1895, has the expanded article 6 of gnathopod 1 but coxa 1
tapers strongly to a subacute point and is partially covered by coxa 2;
the mandibular molar is like that of some species of Anonyzx in its
sublaminar condition lacking teeth and ridges. The telson is deeply
cleft. Except for gnathopod 1 this species could be assigned to Uristes,
although Uristes has a mandibular molar more like that of Cheirimedon
crenatipalmatus. Cheirimedon latimanus also bears comparison to
Centromedon pumilus on technical grounds, and is very close to
“Tryphosa” auct. except for the first gnathopodal expansion.

Figure 127.—Lysianassidae: Maxilla 1: a, Lysianassa heterodonta (Pirlot, 1936); »,
Bathyamaryllis perezii Pirlot (1933a); c, Trischizostoma nicaeense (Costa) (Sars, 1895,
pl. 12, as T. rascht) d, Hirondellea trioculata Chevreux (1900); e, Scopelochetrus crenatus
Bate (Sars, 1895, pl. 19); f, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae);
g, Acidostoma obesum (Bate) (Sars, 1895, pl. 14); h, Aristias neglectus (Bate) (Sars, 1895,
pl. 17); 1, Orchomene batei Sars (1895, pl. 22). Maxilla 2: 7, Perrierella; k, Adeliella
laticornis Nicholls (1938); 1, Orchomene; m, Trischizostoma; n, Acidostoma; 0, Scopelocheirus;
p, Lysianassa plumosa. Coxa1: q, Anonyx nugax (Phipps) (Sars, 1895, pl. 31). Coxa 4:
r, Paralibrotus setosus Stephensen (1923). Gills: 5, Ichnopus spinicornis Boeck (Sars,
1895, pl. 15). Pleonites 3-6 (5-6 fused), lateral: ¢, Crybelocyphocaris tattersalli Shoemaker
(1945a).

MARINE GAMMARIDEAN AMPHIPODA 335

Cheirimedon fougnerr Walker (1903) differs from the type-species
of Cheirimedon by the short cleft of the telson and thus resembles some
species of “‘Onisimus’’ auct. and Paronesimus. Cheirimedon pectini-
palma K. H. Barnard (1925) resembles C. latimanus in coxa 1 and
mandible but has a poorly cleft telson, and is distinguished from
Uristes primarily by the telson. Because these species assigned to
Cheirimedon confound its crisp definition and bridge so many other
genera it would seem prudent to isolate at least C. latimanus in a genus
of its own and assign C. pectinipalma to it provisionally. The latter
also has affinities with Hirondellea except in its head. The three species
then remaining in Cheirimedon, C. crenatipalmatus, C. fougneri, and
C. femoratus should be examined from time to time in light of a viewpoint
that they simply represent minor departures in the evolutionary
scheme of certain genera such as Anonyz (C. crenatipalmatus and C.
femoratus) and ‘‘Onisimus’”’ (C. fougnerr) while those species of the new
genus, with C. latimanus as type, should be considered as possible
subgeneric members of Uristes and ‘‘Tryphosa.’’ A parallel may be
seen in Pseudokoroga rima J. L. Barnard which is simply an Orchomene
with enlarged first gnathopod. Perhaps many of the lysianassids with
enlarged first gnathopods are the rubble left behind in the differentia-
tion of genera, now so remote from their ancestors, that their origins
are unrecognizable.

Cheirimedon is mentioned often in the foregoing keys to Lysianas-
sidae because the several species confound the couplets at vital points.

Chevreuxiella Stephensen

Chevreuxiella Stephensen, 1915.

Type-species: C. metopoides Stephensen, 1915 (monotypy).

Accessory flagellum absent; upper lip and epistome not differen-
tially produced but both large and helmet-shaped; mandible lacking
molar and palp; maxilliped of diverging interpretation: possibly
composed of small inner and outer plates and an immense opercular
palp composed of one article supporting a second minute proximal
article; possibly the opercular palp article is the outer plate and
the second article is the palp; gnathopod 1 simple, short, article 5
longer than 6, article 7 with apical spine; gnathopod 2 simple; coxae
1 and 2 much smaller than coxae 3 and 4, coxa 2 partially hidden by
3 and coxa 1 partially hidden by 2; coxa 4 large, not posteriorly ex-
cavate; uropods 1 and 2 flattened, inner ramus less than half of outer;
telson and uropod 3 absent; urosome composed of two segments.
Species: 2, Atlantic, antarctic, bathy-abyssopelagic.

Frcure 128.—Lysianassidae: Maxilliped: a, Centromedon pumilus (Liljeborg) (Sars,
1895, pl. 34); b, Kerguelenia borealis Sars (1895, pl. 40); c, Lystanassa plumosa Boeck
(Sars, 1895, pl. 16, as L. costae); d, Acidostoma obesum (Bate) (Sars, 1895, pl. 14); e,
Pachynus chelatum Bulycheva (Gurjanova, 1962); f, Lystanassa heterodonta (Pirlot,
1936); g, Normanion sarsi Stebbing (Sars, 1895, pl. 13, as N. quadrimana); h, Trischizos-
toma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); 1, Ambasia atlantica (Milne Ed-
wards) (Sars, 1895, pl. 17, as 4. danielsseni); 7, Perrierella audouiniana (Bate) (Chevreux
and Fage, 1925); k, Nannonyx goest (Boeck) (Sars, 1895, pl. 24), /, Danaella mimonectes
Stephensen (1925b), palp absent. Telson: m, Acidostoma; n, Onisimus edwardsi (Kr¢yer)
(Sars, 1895, pl. 36); 0, Orchomene batei Sars (1895, pl. 22); p, Lysianassa; q, Ichnopus
spinicornis Boeck (Sars, 1895, pl. 15); 7, Opisa eschrichti (Krdyer) (Sars, 1895, pl. 14);
5, Hippomedon abysst (Goés) (Gurjanova, 1962); t, Eurythenes gryllus (Lichtenstein)
(Sars, 1895, pl. 30); u, Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); 2, Boeckosimus
normani (Sars, 1895, pl. 36); w, Trischizostoma.

Ficure 129.—Lysianassidae: Gnathopod 1: a, Koroga megalops Holmes (Stephensen, 1923);
b, Paracentromedon crenulatus (Chevreux) (Chevreux and Fage, 1925); c, Schisturella
(=Pseudonesimus) abyssi (Chevreux, 1926); d, Orchomene bate: Sars (1895, pl. 22);
e, Opisa eschrichti (Krdyer) (Sars, 1895, pl. 14); f, Nannonyx goesi (Boeck) (Sars, 1895,
pl. 24); g, Eucallisoma glandulosa J. L. Barnard (1961); h, Gainella chelata Chevreux
(1912b); 7, Pseudorchomene coatst (Chilton, 1912); 7, Tryphosella sarsi Bonnier (Sars, 1895,
pl. 27, as Tryphosa nana); k, Uristes umbonatus (Sars, 1895, pl. 29); 7, Chetrimedon latimanus
Sars (1895, pl. 13); m, Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); », Paracal-
lisomopsis beljaevi Gurjanova (1962); 0, Euonyx chelatus Norman (Sars, 1895, pl. 40).

338 U.S. NATIONAL MUSEUM BULLETIN 271

Clepidecrella J. L. Barnard
Clepidecrella J. L. Barnard, 1962d.

Type-species: C. cabinda J. L. Barnard, 1962d (original designa-
tion).

Head small, partially hidden by coxa 1; article 1 of antenna 1
carinate; upper lip and epistome not differentially produced; mandib-
ular molar vestigial, palp very long, attached level with molar;
outer plates of maxilliped rather small, palp very long; gnathopod 1
simple, articles 5 and 6 equal; uropod 3 very short, inner ramus short,
scale-like; telson short, apically emarginate. Species: 1, S. Atlantic,
abyssal.

Crybelocephalus Tattersall
Crybelocephalus Tattersall, 1906.

Type-species: C. megalurus Tattersall, 1906 (original designa-
tion). See Shoemaker, 1945a.

Accessory flagellum absent; head somewhat deformed; mandible
lacking both molar and palp; gnathopod 1 simple, articles 5 and 6
subequal; gnathopod 2 simple; coxae 1 and 2 small and fully hidden
by coxa 3; pereopods 1 and 2 subprehensile; telson large, entire. Spe-
cies: 3, cosmopolitan, bathy-abyssopelagic.

Crybelocyphocaris Shoemaker
Crybelocyphocaris Shoemaker, 1945a.

Type-species: C. tattersalli Shoemaker, 1945a (original designa-
tion).

Accessory flagellum absent; head deformed; mandible lacking molar,
palp attached rather proximally; gnathopod 1 simple, articles 5 and
6 equal; gnathopod 2 simple; coxae 1 and 2 small and partially hidden
by coxa 3; pereopods 1 and 2 prehensile, pereopod 3 subprehensile;
uropod 3 small, outer ramus equal to peduncle, 2-articulate, inner
ramus one third as long as outer; urosomites 2 and 3 coalesced. Spe-
cies: 1, Atlantic, bathypelagic.

Cyclocaris Stebbing

Cyclocaris Stebbing, 1888; 1906.

Type-species: C. tahitensis Stebbing, 1888 (monotypy).

Apparently upper lip lobately produced in front of epistome; head
somewhat deformed and globular; mandible flattened, possibly with
elongate, lanceolate, setulose molar or none, palp attached medially

MARINE GAMMARIDEAN AMPHIPODA 339

or level with molar; gnathopod 1 slender, simple; coxae 1 and 2 small
and partially hidden by coxa 3; telson long, deeply cleft. Species: 2,
arctic to Pacific tropics, epi- to ?abyssopelagic.

Cyphocaris Boeck

Cyphocaris Boeck, 1871.—Stebbing, 1906.

Type-species: C. anonyx Boeck, 1871 (monotypy). See Stebbing,
1888 (as C. micronyz); Schellenberg, 1926b, 1927.

Epistome and upper lip not differentially produced; head deformed;
mandibular molar of medium size, ridged, palp attached level with
molar; gnathopod 1 simple, articles 5 and 6 equal; gnathopod 2
nearly simple, article 5 only slightly longer than article 6; coxae 1,
2, and 3 small, partially hidden by large coxa 4; article 2 of pereopod
3 deeply indentured or not, or with long posterior cusp; pereopods
1-4 decreasingly prehensile or not prehensile; telson long, deeply
cleft. Species: 7, cosmopolitan, bathy-abyssopelagic.

Danaella Stephensen
Danaella Stephensen, 1925b.

Type-species: D. mimonectes Stephensen, 1925b (monotypy, origi-
nal designation in title).

Body globular, with pereonites 3-6 especially enlarged; accessory
flagellum absent; epistome and upper lip not prominent; mandible
simple, lacking molar and palp; maxilliped composed of large oper-
cular outer plates covering smaller curved inner lobes; gnathopod 1
simple, dactyl very short; gnathopod 2 simple; coxae 1 and 2 small
and hidden by following coxae; inner rami of uropods 1 and 2 short
and spiniform; uropod 3 a vestigial microscopic bud; telson probably
absent; urosome possibly consisting of two segments with a vestigial
third. Species: 1, Atlantic, bathypelagic.

Derjugiana Gurjanova
Derjugiana Gurjanova, 1962.

Type-species: JD. insolita Gurjanova, 1962 (monotypy).

Mouthpart group conically produced, mouthparts styliform; man-
dible lacking molar, palp attached proximally; maxillipedal palp not
exceeding outer plate, article 4 short and not claw-shaped; gnathopod
1 chelate, article 6 longer than 5; inner ramus of uropod 2 slightly
constricted, uropod 3 lacking rami; telson short, entire. Species: 1,
Okhotsk Sea, littoral.

340 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 130.—Lysianassidae: Gnathopod 1: a, Pachychelium davidis Stephensen (1925a);
c, Valettiopsis dentatus Holmes (1908); ¢, Lysianassa plumosa Boeck (Sars, 1895, pl. 16,
as L. costae); f, Valettia coheres Stebbing (1888); g, Neoambasia tumicornis (Nicholls, 1938) ;
h, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); 1, Ichnopus spinicornts
Boeck (Sars, 1895, pl. 15); 7, Kerguelenia borealis Sars (1895, pl. 40); k, Aristias neglectus
Hansen (Sars, 1895, pl. 17, as 4. audouinianus); 1, Sophrosyne robertsoni Stebbing and
Robertson (1891); m, Scopelocheirus crenatus Bate (Sars, 1895, pl. 19). Pereopod 1:
b, Endeavoura mirabilis Chilton (1921); d, Paracyphocaris praedator Chevreux (Shoemaker,
1945a).

MARINE GAMMARIDEAN AMPHIPODA 341

Ficure 131.—Lysianassidae: Pereopod 3: a, Paracyphocaris praedator Chevreux (Shoe-
maker, 1945a); b, Glycerina tenuicornis (Haswell) (Pirlot, 1936). Pereopod 2: c, Scopelo-
chetropsis abyssalis Schellenberg (1926a). Uropod 2: d, Tryphosites longipes (Bate and
Westwood) (Sars, 1895, pl. 29); e, Orchomene groenlandica (Hansen) (Sars, 1895, pl. 26).
Gnathopod 2: f, Scopelocheirus crenatus Bate (Sars, 1895, pl. 19); g, Hirondellea brevicau-
data Chevreux (1910); 4, Lysianassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); 1,
Gainella chelata Chevreux (1912b); 7, Tryphosites; k, Trischizostoma nicaeense (Costa)
(Sars, 1895, pl. 12, as T. raschi); 1, Lepidepecreum longicorne (Bate and Westwood) (Sars,
1895, pl. 38, as L. carinatum); m, Anonyx debruynii (Hoek) (Sars, 1895, pl. 37); n, Hippo-
medon denticulatus (Bate) (Sars, 1895, pl. 20). Urosome: 0, Danaella mimonectes Stephen-
sen (1925b), showing long uropod 1, shorter uropod 2 and arrow pointing to vestigial
uropod 3 tubercles.

342 U.S. NATIONAL MUSEUM BULLETIN 271

Endevoura Chilton
Endevoura Chilton, 1921.

Type-species: EH. mirabilis Chilton, 1921 (original designation).

Antennal flagella very short, 2- to 5-articulate; [upper lip and
epistome not described]; mandibular molar well developed, ridged,
palp attached proximal to molar; maxillipedal palp article 4 bulbous;
enathopod 1 simple, dactyl very short; pereopod 1 prehensile, subche-
late, large; telson entire, short. Species: 1, Bass Strait, littoral.

Ensayara J. L. Barnard
Ensayara J. L. Barnard, 1964d.

Type-species: EE. ramonella J. L. Barnard, 1964d (original desig-
nation).

Antennal flagella very short, about 5-articulate; upper lip and
epistome inconspicuous, latter projecting shghtly; mandibular molar
smooth, palp attached about level with molar; maxillipedal palp
article 4 claviform; gnathopod 1 simple, dactyl short; pereopod 1
prehensile, subchelate, large; telson entire, short. Species: 1, Baja
California, littoral.

Eucallisoma J. L. Barnard
Eucallisoma J. L. Barnard, 1961.

Type-species: Ff. glandulosa J. L. Barnard, 1961 (original desig-
nation).

Head very small but not deformed; antenna 1 immense, stout,
conical, article 1 of flagellum elongate, flagellum bearing only three
articles, accessory flagellum appressed to body of antenna (Paracal-
lisomopsts) ; epistome and upper lip each convex in front; mandibular
molar small, conical, unridged, palp attached level with molar;
enathopod 1 simple, not elongate, article 7 vestigial, shrouded by
short cirri, article 2 immensely swollen; telson of medium length,
deeply cleft. Species: 1, tropical Atlantic, abyssal.

Euonyx Norman

Euonyx Norman, 1867b.—Stebbing, 1906.
Leptochela Boeck, 1876 (homonym, Decapoda).

Type-species: EE. chelatus Norman, 1867b (monotypy). See Sars,
1895.

Epistome acutely produced in front of upper lip; mandibular
molar obsolete or absent, palp attached distally; gnathopod 1 slender,

MARINE GAMMARIDEAN AMPHIPODA 343

chelate, thumb narrow; article 7 of gnathopod 2 rather large and
curved; coxa 1 short, nearly hidden by coxa 2; telson of medium
length, deeply cleft. Species: 7, cosmopolitan, epi- to bathypelagic.

Eurythenes Smith

Eurytenes Liljeborg, 1865 (homonym, Hymenoptera).
Eurythenes Smith, 1884.—Stebbing, 1906.

Euryporeia Sars, 1895.

Katius Chevreux, 1905c.

Type-species: Gammarus gryllus Lichtenstein, 1822 (monotypy).
See Sars, 1895.

Article 1 of antenna 2 tumid in adults; epistome broadly lobate
in front of upper lip; mandibular molar immense, extremely broad,
laminate, apically ridged, palp attached level with molar; gnathopod
1 subchelate, article 7 overlapping palm, article 6 longer than 5,
article 3 not elongate (Paralicella); coxa 1 shortened, partially con-
cealed by coxa 2; telson long, deeply cleft. See Arzstias. Species:
2, cosmopolitan, bathyal to abyssal.

Figorella J. L. Barnard
Figorella J. L. Barnard, 1962d.

Type-species: F. tanidea J. L. Barnard, 1962d (original desig-
nation).

Accessory flagellum 1-articulate; [upper lip and epistome not
described]; mandibular molar absent, palp attached rather distally;
palp of maxilla 1 uniarticulate; gnathopod 1 slightly chelate, article
6 somewhat enlarged, much longer than article 5; uropod 3 very
short, inner ramus shortened; telson short, apparently entire.
Species: 1, S. Atlantic, abyssal.

Gainella Chevreux
Gainella Chevreux, 1911c; 1912a; 1912b.

Type-species: G. chelata Chevreux, 1912a, 1912b (designated by
Chevreux, 1912a).

Epistome and upper lip not differentially produced; mandibular
molar small but ridged, palp attached proximal to molar; inner
plates of maxilliped styliform, palp very long, slender, article 4
densely setulose apically; gnathopod 1 chelate, thumb narrow and
short, article 6 much longer than 5; gnathopod 2 styliform but mi-
nutely chelate; uropod 3 short, inner ramus a short scale, one third
as long as outer ramus; telson short, deeply cleft. Species: 1, antarc-
tic, littoral (297 m).

344. U.S. NATIONAL MUSEUM BULLETIN 271

i ee A\N

Ficure 132.—Lysianassidae: a, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl. 30); b
Metacyphocaris helgae Tattersall (1906). Uropod 3: c, Danaella mimonectes Stephensen
(1925b); d, Stomacontion pepinii (Stebbing, 1888); ¢, Aristias tumidus (Kr@yer) (Sars,
1895, pl. 18); f, Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); g, Lysianassa plumosa
Boeck (Sars, 1895, pl. 16, as L. costae); h, Lepidepecreella ctenophora Schellenberg (1926a);
1, Lysianassa alba (Holmes) (Shoemaker, 1933b); 7, Acontiostoma marionis Stebbing
(1888); k, Nannonyx goesi (Boeck) (Sars, 1895, pl. 24).

MARINE GAMMARIDEAN AMPHIPODA 345

Glycerina Haswell

Glycera Haswell, 1880b (homonym, Polychaeta).
Glycerina Haswell, 1882.—Stebbing, 1906.

Type-species: Glycera tenuicornis Haswell, 1880b (monotypy). See
Pirlot, 1936.

Upper lip acutely produced in front of epistome; mandibular molar
prominent, setulose, palp attached proximal to molar; gnathopod 1
simple, slender, article 5 slightly longer than 6; article 2 of pereopod
3 deeply indentured ; inner ramus of uropod 2 constricted; telson deeply
cleft. Species: 1, Coral Sea to E. Australia, littoral.

Hippomedon Boeck

Hippomedon Boeck, 1871.—Stebbing, 1906.—Gurjanova, 1962.
Platamon Stebbing, 1886.

Paratryphosites Stebbing, 1899d.

?Paracentromedon Chevreux and Fage, 1925.

?Hlimedon J. L. Barnard, 1962d.

Type-species: Anonyx holboll1 Krgyer, 1846 (selected by Boeck,
1876; note, the A. holbolli of Boeck, 1871 and 1876=Hippomedon
denticulatus [Bate, 1857a] but the type-species should stand as A.
holbolla Krgyer). See Sars, 1895.

Head small, largely hidden by cora 1 (‘‘Tryphosa’’ auct.), with
distinct lateral lobes (Lepidepecreoides); upper lip either distinctly
or not lobate in front of epistome but both projections small, incon-
spicuous; mandibular molar large, ridged, palp attached level with
molar; gnathopod 1 subchelate or nearly simple, article 5 longer
than or equal to 6; telson short or medium, half or deeply cleft.
See also Paronesimus. Species: 41, cosmopolitan, cold water, littoral
to abyssal.

Hirondellea Chevreux

Mirondellea Chevreux, 1889.—Stebbing, 1906.—K. H. Barnard, 1930.
Tetronychia Stephensen, 1923.

Type-species: Hirondellea trioculata Chevreux, 1889 (original
designation). See Chevreux, 1900.

When visible, eyes triple; head globular or with large lateral lobes
rounded, palp article 4 of mazilliped claw-shaped (Centromedon) ;
mandibular molar conical, setulose, palp attached at proximal end of
molar; inner plate of mazilla 1 with at least one enlarged sickle-seta
or seta with strong attenuation or constriction, palp article 2 often
notched medially; mazillipedal palp article 2 half or less as broad as
outer plate (Ambasiella); gnathopod 1 subchelate or slightly chelate,
articles 5 and 6 equal; coxa 1 rather small, often partially hidden by

346 U.S. NATIONAL MUSEUM BULLETIN 271

coxa 2; inner ramus of uropod 2 constricted or not; telson of medium
length or short, deeply or shortly cleft. Species: 6, cosmopolitan,
primarily bathyal to abyssal, pelagic.

Ichnopus Costa

Ichnopus Costa, 1853c; 1857.—Stebbing, 1906.

Type-species: J. taurus Costa, 1853c (monotypy). See Chevreux
and Fage, 1925.

Upper lip slightly produced in front of epistome; mandibular molar
laminate, scarcely ridged, palp attached level with molar; gnathopod
1 simple, article 7 with a posterior or distal bundle of setae; telson of
medium length, cleft; gills plaited on both sides, pereopod 5 much longer
than pereopod 4 (Socarnes). Species: 5, probably cosmopolitan,
littoral to abyssal, pelagic.

Kerguelenia Stebbing
Kerguelenia Stebbing, 1888; 1906.

Type-species: KK. compacta Stebbing, 1888 (monotypy).

Upper lip and epistome small, inconspicuous; mandible lacking
distinct cutting edge and molar, palp attached distally; maxillae and
maxilliped poorly developed, stunted; gnathopod 1 simple, styliform;
uropod 3 tiny, rami usually shorter than peduncle, inner often vestigial
or absent, outer 1- or 2-articulate; telson short, entire. Species: 6,
bipolar, amphiboreal, littoral to bathyal.

Koroga Holmes
Koroga Holmes, 1908.

Type-species: K. megalops Holmes, 1908 (original designation).
See Gurjanova, 1962.

Upper lip projecting lobately in front of epistome; mandibular
molar short, conical or laminate, setulose, palp attached slightly
proximal to molar; mazillae stout, mavzillipedal palp article 4 long
(Pseudokoroga); gnathopod 1 subchelate, palm broad, transverse,
article 6 slightly broadened, much longer than article 5; telson apically
notched. Species: 1, cosmopolitan, epipelagic.

Kyska Shoemaker
Kyska Shoemaker, 1964.

Type-species: K. dalli Shoemaker, 1964 (original designation).
Upper lip not projecting in front of epistome; mandibular molar of

MARINE GAMMARIDEAN AMPHIPODA 347

medium size, conical, setulose, lacking ridges, palp attached distal to
molar; gnathopod 1 chelate, article 6 and chela slender, article 6 much
longer than 5; inner ramus of uropod 2 unconstricted; telson of medium
length, deeply cleft; gills plaited on both sides. Like Anonyz but
enathopod 1 chelate. Species: 1, Alaska, littoral.

Lepidepecreella Schellenberg

Lepidepecreella Schellenberg, 1926a.
Paracyclocaris K. H. Barnard, 1930.
Type-species: JL. ctenophora Schellenberg, 1926a (monotypy).
Epistome forming immense anterior keel on head, projecting in
front of upper lip but together coalesced; mouthpart field thus giving
conical appearance, mandible slender, molar bulge present, palp
attached proximal to molar; gnathopod 1 simple, styliform, article 7
apically (and marginally) setose; coxae 1 and 2 small, partially hidden
by coxa 3; outer ramus of uropod 3 shorter than peduncle, 1- or
2-articulate, inner ramus one third (rarely one half) as long as outer
or absent; telson short, entire. Species: 5, bipolar, littoral to bathyal.

Lepidepecreoides K. H. Barnard

Lepidepecreoides K. H. Barnard, 1931; 1932.

Type-species: L. xenopus K. H. Barnard, 1931 (original desig-
nation); 19382.

Head deeply concavotruncate from lateral view (Hippomedon); upper
lip lobately projecting in front of epistome; mandibular molar small,
palp attached level with molar; gnathopod 1 subchelate, article 5
longer than 6; article 7 of gnathopod 2 rather large, palm slightly
chelate; coxa 5 as deep as coxa 4 (Hippomedon); telson of medium
length, deeply cleft. Species: 1, subantarctic, littoral.

Lepidepecreopsis Stephensen
Lepidepecreopsis Stephensen, 1925a.

Type-species: L. biloba Stephensen, 1925a (monotypy). See
Tmetonyz Stebbing and Tryphosella Bonnier.

Epistome broadly rounded and dominating upper lip, latter slightly
produced, both separated by deep incision; mandibular molar sub-
columnar, not triturative (Tryphosella), palp attached level with
molar or at proximal corner; coxa 1 slightly tapering distally and
partially hidden by coxa 2; gnathopod 1 subchelate, articles 5 and 6
subequal in length or article 5 shorter than 6; telson of medium length,
deeply cleft. Species: 1, subarctic Atlantic, abyssal.

285-135 O - 69 - 23

348 U.S. NATIONAL MUSEUM BULLETIN 271

Lepidepecreum Bate and Westwood

Lepidepecreum Bate and Westwood, 1868.—Stebbing, 1906.

Type-species: Anonyx longicornis Bate and Westwood, 1863
(monotypy and subsequent synonymy). See Sars, 1895.

Article 1 of antenna 1 usually carinate (combining characteristic)
but occasionally not; accessory flagellum absent (type) or present;
epistome broadly produced in front of upper lip; body often dorsally
carinate or toothed; mandibular molar a setulose, quadrate lamina,
palp attached proximal to molar; gnathopod 1 subchelate, articles 5
and 6 subequal; telson long, deeply cleft. Some species approach
Orchomene complex. Species: 19, bipolar, amphiboreal, littoral to
bathyal.

Lysianassa Milne Edwards, provisional synonymy

Lysianassa Milne Edwards, 1830.—Stebbing, 1906.

Lystanassina Costa, 1867.

Lysianax Stebbing, 1888.

Lysianopsis Holmes, 1903; 1905. Type-species: L. alba Holmes, 1903 (mono-
typy). See Shoemaker, 1933b.

Aruga Holmes, 1908. Type-species: A. oculata Holmes, 1908 (monotypy).

Arugella Pirlot, 1936. Type-species: A. heterodonta Pirlot, 1936 (original designa-
tion).

Shoemakerella Pirlot, 1936. Type-species: Lysianassa nasuta Dana, 1853
(original designation). Hurley (1963) notes that Pirlot’s diagnosis of Shoe-
makerella fits Lysianax cubensis Stebbing, 1897, and not -L. nasuta. However,
the latter should remain as type-species because it was so designated.

?Pronannonyx Schellenberg, 1953. Type-species: P. minimus Schellenberg, 1953
(monotypy). [See also in alphabetical order.]

Type-species: JL. costae Milne Edwards, 1830 (denoted by elimina-
tion of Milne Edwards, 1840, and selected by Boeck, 1876). Species
obscure. Genus based here on L. plumosa Boeck (L. costae of Sars,
1895, pl. 16, fig. 1).

Upper lip strongly lobate in front of epistome or epistome pro-
duced forward along with upper lip; mandibular molar large, coni-
colaminate, unridged or nearly obsolete, palp attached proximal to
molar; gnathopod 1 simple, articles 5 and 6 subequal; inner ramus of
uropod 2 unconstricted (L. plumosa), or constricted (L. longicornas) ;
outer ramus of uropod 3 uniarticulate; telson entire, emarginate, or
minutely notched. Species, including synonymous genera: 24,
cosmopolitan, littoral (rarely in bathyal). Species with aberrant
maxilla 2: L. hypocrita Ruffo.

MARINE GAMMARIDEAN AMPHIPODA 349

Lysianella Sars
Lysianella Sars, 1882.—Stebbing, 1906.

Type-species: JL. petalocera Sars, 1882 (original designation). See
Sars, 1895.

Article 4 of antenna 2 tumid, article 5 slender; upper lip lobately
produced in front of epistome; mandibular molar vestigial, palp at-
tached proximal to molar; gnathopod 1 subchelate, often weakly,
articles 5 and 6 equal; inner ramus of uropod 2 constricted or not;
telson of medium length, entire or cleft 25 percent. Species: 3,
boreal and subtropical Atlantic and Mediterranean, littoral to abyssal.

Menigrates Boeck
Menigrates Boeck, 1871.—Stebbing, 1906.

Type-species: Anonyz obtusifrons Boeck, 1861 (monotypy). See
Sars, 1895.

Upper lip and epistome not differentially produced, inconspicuous;
mandibular molar obsolescent, an undifferentiated lamina, palp at-
tached slightly proximal to molar; gnathopod 1 simple, article 6
longer than 5; telson short, cleft one third (type) or a quarter or less;
peduncular articles 2 and 3 of antenna 1 short, half as long as article 1
of flagellum (Waldeckia and Socarnes). See Paralibrotus. Species: 3,
subarctic, littoral.

Menigratopsis Dahl
Menigratopsis Dahl, 1945.

Type-species: MM. svenillsoni Dahl, 1945 (monotypy).

Epistome and upper lip indistinct from each other; mandibular
molar well developed, but possibly not triturative, palp attached
level with molar; gnathopod 1 simple, article 5 slightly longer than
article 6; telson of medium length, deeply cleft. Species: 1, the
Sound between Sweden and Denmark, littoral.

Mesocyclocaris Birstein and Vinogradov
Mesocyclocaris Birstein and Vinogradov, 1964.

Type-species: MM. gracilis Birstein and Vinogradov, 1964 (mono-
typy).

Accessory flagellum absent; mandible lacking molar and palp;
gnathopod 1 simple, article 5 longer than 6; coxae 1 and 2 much
smaller than coxae 3 and 4 and probably partially hidden by coxa 3;
pereopods 1-5 prehensile; uropod 3 elongate, rami interequal, outer
1-articulate; [telson unclear]. Species: 1, Indian Ocean, bathypelagic.

350 U.S. NATIONAL MUSEUM BULLETIN 271

Mesocyphocaris Birstein and Vinogradov
Mesocyphocaris Rirstein and Vinogradov, 1960.

Type-species: M/. longicaudatus Birstein and Vinogradov, 1960
(original designation).

Accessory flagellum 1-articulate; head somewhat deformed; man-
dible lacking molar, palp attached medially on mandibular body;
enathopod 1 simple, articles 5 and 6 subequal; coxae 1 and 2 much
smaller than coxae 3 and 4 and probably partially hidden by coxa 3;
pereopods 1-4 prehensile; uropod 3 elongate, outer ramus 2-articulate,
inner very short and scale-like; telson short, cleft one third its length.
Species: 1, Pacific, abyssopelagic.

Metacyclocaris Birstein and Vinogradov
Metacyclocaris Birstein and Vinogradov, 1955.

Type-species: MM. polycheles Birstein and Vinogradov, 1955 (orig-
inal designation).

Mandible lacking molar, palp attached rather distally; gnathopod
1 simple, article 5 longer than 6, gnathopod 2 slender, nearly simple;
coxa 2 much larger than coxa 1 and covering it, coxae 3 and 4 not
enlarged as in other cyphocarid genera; pereopods all prehensile;
telson long, deeply cleft. Species: 1, N.W. Pacific, abyssopelagic

Metacyphocaris Tattersall
Metacyphocaris Tattersall, 1906.

Type-species: MM. helgae Tattersall, 1906 (original designation).

Head deformed; accessory flagellum 1-articulate; mandible with
very small, smooth molar, palp absent; gnathopod 1 simple, articles
5 and 6 equal; gnathopod 2 simple; coxae 1 and 2 small, partially
hidden by coxa 3; pereopods 1-3 prehensile; outer ramus of uropod 3
elongate, 2-articulate, inner ramus very short, scale-like; telson of
medium length, cleft one fourth. Species: 1, cosmopolitan,
bathypelagic.

Metambasia Stephensen
Metambasia Stephensen, 1923.

Type-species: MM. faeroensis Stephensen, 1923 (monotypy).

Article 3 of male antenna 2 tumid; upper lip projecting slightly
in front of epistome; mandibular molar a long conical lamina, palp
attached level with molar; gnathopod 1 simple, article 5 longer
than 6; coxa 1 short, half as long as its second article, partially
hidden by coxa 2; inner ramus of uropod 2 constricted; telson cleft
halfway or more. See Schisturella. Species: 1, N. Atlantic, bathyal.

MARINE GAMMARIDEAN AMPHIPODA 351

Microlysias Stebbing
Microlysias Stebbing, 1918.

Type-species: MM. xenokeras Stebbing, 1918 (monotypy). See
K. H. Barnard, 1937.

Article 4 of male antenna 2 very tumid; epistome slightly protruding
in front of upper lip; mandibular molar feeble, palp attached quite
proximally; maxillipedal palp 3-articulate, possibly with a minute
fourth article; gnathopod 1 subchelate, article 6 slightly longer
than 5; telson of medium length, deeply cleft; gills plaited. Species:
2, S. Africa, S. Arabian coast, littoral.

Nannonyx Sars
Nannonyz Sars, 1895.

Type-species: Orchomene goesit Boeck, 1871. (monotypy). See
Sars, 1895.

Mouthparts substyliform; epistome and upper lip coalesced,
epistomal portion projecting slightly in front of upper lip; mandibular
molar vestigial, palp attached proximally; maxillipedal palp scarcely
exceeding outer plate, article 4 very small (except N. spinimanus
Walker); gnathopod 1 appearing simple but with microscopic palm,
articles 5 and 6 short and stout; rami of uropod 3 equal to or shorter
than peduncle; telson entire or slightly emarginate. Species: 4,
boreal-subtropical N.E. Atlantic and Mediterranean, Kerguelen
Island, littoral.

Neoambasia Dahl
Neoambasia Dahl, 1959.

Type-species: Ambasiopsis tumicornis Nicholls, 1938 (monetypy).

Upper lip projecting slightly in front of epistome; mandibular
molar apparently well developed, setose, palp attached level with
molar; outer plate of maxilla 1 poorly spinose (Pseudambasia) ; outer
plate of maxilliped spinose (Ambasiopsis); gnathopod 1 subchelate,
article 6 slightly longer than 5; coxa 1 not greatly shortened, but
almost concealed by coxa 2, about as long as article 2 of gnathopod
1; rami of uropod 3 not longer than peduncle; telson short, deeply
cleft. Species: 1, antarctic, bathyal.

Normanion Bonnier

Normania Boeck, 1871 (homonym, Ostracoda).
Normanion Bonnier, 1893 (new name).—Stebbing, 1906.

Type-species: Opis quadrimana Bate and Westwood, 1868 (mono-
typy). See Sars, 1895. Even though Boeck’s identification of O.

352 U.S. NATIONAL MUSEUM BULLETIN 271

quadrimana was in error, the type should remain that species for
purposes of stability. Boeck’s material was named Normanion sarsi
by Stebbing (1906).

Mouthparts forming pseudoconical bundle or not; epistome appla-
nated, not distinct from upper lip; mandibular molar conical or
nearly obsolete, not ridged (?except N. abyssi Chevreux), palp
attached proximal to molar; outer plates of maxilliped almost or
ereatly exceeding palp, palp essentially 3-articulate, scarcely or
slightly exceeding inner plates, article 4 obsolete; gnathopod 1 mod-
erately powerful, subchelate, palm transverse; peduncle of uropod 3
elongate; telson short, quadrate, entire. Species: 3, N.E. Atlantic,
Mediterranean, littoral to abyssal (some fish commensalism).

Ocosingo J. L. Barnard
Ocosingo J. L. Barnard, 1964b.

Type-species: 0. borlus J. L. Barnard, 1964b (original designation).

Mouthparts substyliform, forming ventral subconical bundle; man-
dible bearing vestigial molar ridge, palp attached quite proximally;
palp of maxilla 1 uniarticulate; maxillipedal palp scarcely exceeding
outer plate, 3-articulate, ultimate article claviform; gnathopod 1
simple, article 6 longer than 5; coxa 1 projecting over side of head;
uropod 3 lacking rami; telson small, entire. Species: 1, California,
littoral.

Onesimoides Stebbing
Onesimoides Stebbing, 1888; 1906.

Type-species: O. carvnatus Stebbing, 1888 (monotypy).

Upper lip and epistome not differentially produced, mandibular
molar ridged, palp attached level with molar; gnathopod 1 subchelate
or slightly chelate, article 5 much longer than 6, gnathopod 1 rather
enlarged in males; outer ramus of uropod 3 biarticulate, inner ramus
half or less as long as outer; telson entire; article 1 of first antennal
flagellum 5 times as long as neat article (Paronesimoides). Species:
3, Indo-Pacific tropics, bathyal to hadal.

Onisimus Boeck, new synonymy

Onisimus Boeck, 1871.
Pseudalibrotus Della Valle, 1893.—Stebbing, 1906.

Type-species: Anonyzx litoralis Krgyer, 1845 (selected by Boeck,
1876). Stebbing, Sars, and Della Valle apparently overlooked the
fact that Boeck (1876) selected Anonyz litoralis as type of Onisumus.
Della Valle erected Pseudalibrotus and cited only A. litoralis, thus
making it type of the genus by monotypy. All species of Pseudali-

MARINE GAMMARIDEAN AMPHIPODA 353

brotus, having gone under that name for over 60 years, must now be
transferred to Onisumus and the 15 or more species of Onisimus,
having gone under that name for nearly 100 years, must now be
transferred to a new appellation, which I designate as Boeckosimus
in honor of Axel Boeck, one of the first students of Amphipoda to
understand the importance of type designations.

The genus Alibrotus Milne Edwards (1840) with its monotype, A.
chausercus Milne Edwards (1840) may be a senior synonym of either
Onisimus (auct. =Pseudalibrotus) or Boeckosimus, new name, but
the problem has never been clarified and may be insoluble. The
problem should be reviewed at least once more by a student of arctic-
boreal faunas and perhaps submitted to ICZN in order to cancel the
genus Alibrotus.

Epistome broadly rounded and projecting slightly in front of upper
lip; mandibular molar ridged, palp attached level with molar; gnath-
opod 1 subchelate, articles 5 and 6 equal; inner ramus of uropod 2
constricted or not; telson short, entire. See Boeckosimus. Species: 7,
arctic littoral, Caspian Sea, glacial relicts.

Opisa Boeck

Opis Krgyer, 1842 (homonym, Mollusca).
Opisa Boeck,1876.—Stebbing, 1906.

Type-species: Opis eschrichtia Kroyer, 1842 (monotypy and sub-
sequent synonymy; Boeck (1876) cited O. typica Krgyer as type-
species, probably as a technical error for he recognized it was a junior
synonym of O. eschrichtir).

Upper lip rounded, slightly projecting in front of epistome; mandib-
ular molar obsolete, palp attached proximally; gnathopod 1 powerful,
chelate, chela broad, palm excavate to form thumb; coxa 1 slightly
shortened and partially hidden by coxa 2; telson long, deeply cleft.
Species: 2, boreal, littoral (to 432 m).

Orchomene Boeck, new synonymy

Orchomene Boeck, 1871.—Stebbing, 1906.

Tryphosa Boeck, 1871.—Stebbing, 1906.

Orchomenella Sars, 1895.*—Stebbing, 1906.

Orchomenopsis Sars, 1895.—Stebbing, 1906.

?Allogaussia Schellenberg, 1926a. [See also in alphabetical order.]

Type-species: Anonyx serratus Boeck, 1861 (selected by Boeck,
1876). See Sars, 1895* (as Orchomenella ciliata). Regrettably Sars and
Stebbing overlooked the fact that Boeck (1876) had selected the type-
species of Tryphosa as Anonyx nanus Krgyer (1846) a species un-

*This portion of Sars’ monograph was published in 1891.

354 U.S. NATIONAL MUSEUM BULLETIN 271

doubtedly congeneric with the type-species of Orchomenella, Anonyx
minutus Kroyer, 1846 (original designation). Sars had incorrectly
identified Anonyz nanus and his material later was named Tryphosella
sarsi by Bonnier (1893); indeed Sars had in hand Anonyz nanus but
he erected for the material the name Orchomenella ciliata, later shown
by Stebbing (1906) to be a synonym of A. nanus. Unfortunately
Stebbing did not therefore synonymize Orchomenella with ‘‘Tryphosa,”
since the two type-species were congeneric. Thus, Orchomenella must
fall to Tryphosa and all species of Tryphosa (auct.), so well known
since the time of Sars (1895) and Stebbing (1906) must be given a
different generic name, which is 7ryphosella Bonnier.

Epistome generally broadly lobate in front, occasionally acute, or
flat and unproduced, lobe of upper lip generally prominent but rarely
projecting far in front of epistome; mandibular small, poorly ridged or
weakly setulose, palp attached proximal to molar; gnathopod 1
subchelate, article 6 longer than 5; telson variable, entire (Adllogaussia)
or minutely and deeply cleft. See Lepidepecreum for intergrading
species. Species (including Allogaussia): 61-+-, cosmopolitan, cold-
water, littoral to abyssal. ‘“Orchomenella’”’ groenlandica (Hansen) is
treated in Keys L, N, O, and P as a distinct generic taxon.

Pachychelium Stephensen
Pachychelium Stephensen, 1925a.

Type-species: P. davidis Stephensen, 1925a (original designation).

Antenna 1 very stout, body slender; mandibular molar absent, palp
attached rather distally; maxilla 1 lacking palp; maxilla 2 with
vestigial inner plate; maxilliped lacking inner plate or inner plate
very small; enathopod 1 enlarged, subchelate, article 6 large, ovate;
telson short, entire. Species: 3, bipolar, littoral to bathyal.

Pachynus Bulycheva
Pachynus Bulycheva, 1955.

Type-species: P. chelatum Bulycheva, 1955 (original designation).

Upper lip and epistome apparently not differentially produced;
mandibular molar absent, palp attached rather distally; maxillipedal
palp 3-articulate, or with a small fourth article, article 3 elongate
or not; gnathopod 1 chelate; telson short, entire. Species: 2, boreal
Pacific, littoral to bathyal.

Paracallisoma Chevreux
Paracallisoma Chevreux, 1903.

Type-species: P. alberti Chevreux, 1903 (original designation).

MARINB GAMMARIDEAN AMPHIPODA 355

[Upper lip and epistome not studied]; mandibular molar small,
simple, palp attached level with molar, both molar and palp attached
proximally; inner plate of maxilla 1 densely setose; gnathopod 1
simple, with apical setal shroud, article 7 vestigial; gnathopod 2
subchelate (Aroui, Scopelocheirus); telson long, deeply cleft. Species:
2, Atlantic-Pacific, bathypelagic.

Paracallisomopsis Gurjanova
Paracallisomopsis Gurjanova, 1962.

Type-species: P. beljaevi Gurjanova, 1962 (monotypy).

Head lacking lateral lobes; upper lip and epistome not differenti-
ally produced; mandibular molar conical, unridged, palp attached
level with molar; gnathopod 1 simple, article 7 vestigial, shrouded
with cirri; telson of medium length, deeply cleft.

Differs from Eucallisoma J. L. Barnard (1961) by: the larger head,
smaller first antenna with its less elongate flagellar base, the acces-
sory flagellum conspicuous (in Hucallisoma it is appressed to inner
face of the immense antenna and is brush-like); the less styliform
first gnathopod, the cirri more dense, dactyl not distinct from cirri;
more spines on the outer plate of the maxilliped; the shorter second
article on the outer ramus of uropod 3. Species: 1, Bering Sea,
?epipelagic.

[Paracentromedon Chevreux and Fage]

Paracentromedon Chevreux and Fage, 1925. Presumed synonym of Hippomedon.
Elimedon J. L. Barnard, 1962d.

Type-species: Centromedon crenulatum Chevreux, 1900 (original
designation).

Head small, partially hidden by coxa 1 (Tryphosella); upper lip and
epistome not differentially produced or inconspicuous; mandibular
molar ridged, palp attached level with molar; gnathopod 1 nearly
simple, articles 5 and 6 equal, article 7 greatly overlapping minute
palm; telson of medium length, deeply cleft. See Paronesimus. Spe-
cies: 3, Atlantic, bathyal-abyssal.

Paracyphocaris Chevreux
Paracyphocaris Chevreux, 1905a.

Type-species: P. praedator Chevreux, 1905a (original designation).

Head deformed; mandible lacking molar, palp attached medially;
enathopod 1 simple or very minutely subchelate, article 5 slightly
longer than 6; gnathopod 2 nearly simple; coxae 1 and 2 small, par-
tially hidden by coxa 3; pereopods 1-4 (or 5) prehensile; telson long,
deeply cleft. Species: 2, Atlantic-Pacific, bathy-abyssopelagic.

356 U.S. NATIONAL MUSEUM BULLETIN 271

Paralibrotus Stephensen
Paralibrotus Stephensen, 1923.

Type-species: P. setosus Stephensen, 1923 (monotypy).

Epistome and upper lip not differentially produced; mandibular
molar small but distinct, possibly triturative, palp attached over
proximal end of molar; gnathopod 1 simple, rather stout, article 6
slightly longer than 5; article 2 of outer ramus on uropod 3 minute;
telson short, entire. Merges with some species of Menigrates. Species:
1, subarctic, littoral.

Paralicella Chevreux
Paralicella Chevreux, 1908a.

Type-species: P. tenuipes Chevreux, 1908a (original designation).

Upper lip and epistome not described; mandible lacking molar in
type-species or bearing conical setulose lamina in other species;
palp attached rather distally; inner plate of mavilla 1 densely setose,
with 10 or more setae (combining character); gnathopod 1 subchelate,
article 6 slightly longer than 5, article 3 elongate, nearly as long as
article 6 (Aristias, Eurythenes); telson of medium length, deeply
cleft. Species: 4, Atlantic-Pacific, bathyal-abyssal.

Paralysianopsis Schellenberg

Paralysianopsis Schellenberg, 1931.
Austronisimus K. H. Barnard, 1931.

Type-species: P. odhneri Schellenberg, 1931 (monotypy).

Upper lip produced acutely in front of epistome, but almost fully
coalesced with epistome; mandibular molar obsolescent, palp attached
slightly proximal to molar; gnathopod 1 nearly simple, with minute
oblique palm; inner ramus of uropod 2 constricted; telson entire,
with minutely notched apex. Species: 1, subantarctic, littoral.

Parambasia Walker and Scott
Parambasia Walker and Scott, 1903.

Type-species: P. forbesi Walker and Scott, 1903 (monotypy).

Diagnosis based partially on P. rossi Stephensen (1927) assuming
it to be congeneric. [Epistome and upper lip poorly known], probably
fused together; mandibular molar small, setulose, palp attached
proximal to molar; gnathopod 1 simple, article 5 slightly longer than
article 6; inner ramus of uropod 2 constricted (possibly not on type) ;
outer ramus of uropod 3 uniarticulate; telson entire. Species: 2,
Arabian Sea, Auckland Islands, littoral. Possibly congeneric: Lyst-
anassa anomala Nicholls, 1938, subantarctic.

MARINE GAMMARIDEAN AMPHIPODA 357

Parawaldeckia Stebbing

Parawaldeckia Stebbing, 1910.

Type-species: Nannonyz thomson Stebbing, 1906 (monotypy).

Article 5 of male antenna 2 slightly dilated and longer than article
4; [prebuccal area and mandible unknown but presumably similar to
Waldeckia]; gnathopod 1 simple, articles 5 and 6 equal; uropod 3
short, outer ramus not longer than peduncle, peduncle sublamelliform,
inner ramus scarcely half as long as outer; telson entire or slightly
emarginate. Species: 2,* antiboreal, littoral.

*Parawaldeckia kiddert (Smith).—Generic diagnosis: to some extent mouth-
parts bundled conically, upper lip and epistome confluent, slightly convex ante-
riorly, lacking sinus; mandibular molar conical, unridged, distally setose, palp
attached very proximally; gnathopod 1 simple; uropod 3 with vestigial inner
ramus; telson short, entire. Diagnosis based on Monod (1926) and cotypes
examined in U.S. National Museum; general appearance of cotypes, without
dissection, similar to Monod’s figures and not to concepts of Chilton (1909,
1921), Stephensen (1927), and Tattersall (1922) in the following characters:
general appearance including coxae and pleon, (antenna 2 slightly shorter than
shown by Monod), prebuccal region, telson, and uropod 3.

Paronesimoides Pirlot

Paronesimoides Pirlot, 19338a.

Type-species: P. lignivorus Pirlot, 1933a (original designation).

Apparently upper lip projecting and dominating epistome; man-
dibular molar not strongly produced, setulose, poorly ridged, palp
attached at proximal end of molar; gnathopod 1 subchelate, palm
transverse, article 6 slightly enlarged and much larger than article
5; uropod 3 very small, outer ramus 2-articulate, equal to peduncle,
inner ramus reduced to a minute spiniform process; telson short,
entire; articles 1 and 2 of flagellum of antenna 1 equal in length
(Onesimoides). Species: 1, Indonesia, abyssal.

Paronesimus Stebbing

Paronesimus Stebbing, 1894; 1906.

Type-species: P. barentsi Stebbing, 1894 (monotypy). See
Gurjanova, 1962.

[Upper lip and epistome not described, presumably like Onisimus];
mandibular molar small, weakly ridged, palp attached level with
molar; gnathopod 1 with very small palm, appearing simple, articles
5 and 6 subequal; telson of medium length, cleft about halfway.
Very close to Hippomedon. See Paracentromedon. Species: 2, arctic,
littoral.

358 U.S. NATIONAL MUSEUM BULLETIN 271

Perrierella Chevreux and Bouvier

Perrierella Chevreux and Bouvier, 1892.
Pararistias Robertson, 1892.

_ Type-species: Lysianassa audowimana Bate, 1857a (original des-
ignation and subsequent synonymy). See Sars, 1895.

Upper lip and epistome not differentially produced; mand bular
molar a vestigial hump, palp attached level with molar; lobes of
maxilla 2 slightly gaping, inner broader than outer; maxillipedal palp
not exceeding outer plate, 3-articulate or with vestigial fourth article,
inner plates vestigial; gnathopod 1 nearly simple, with poorly defined
palm; coxa 1 very small and largely hidden by coxa 2; telson of me-
dium length, entire, apically truncate. Species: 1, boreal and warm-
temperate E. Atlantic and Mediterranean, littoral.

Phoxostoma K. H. Barnard
Phozostoma K. H. Barnard, 1925.

Type-species: P. algoense K. H. Barnard, 1925 (monotypy).

Mouthparts forming conical bundle; upper lip and epistome con-
tinuous; mandible slender, molar obsolete, palp attached quite prox-
imally; maxillipedal palp slightly exceeding outer plate, article 4
very small; gnathopod 1 simple, article 6 longer than 5; telson short,
deeply insinuate. Species: 1, 8. Africa, littoral.

Podoprion Chevreux
Podoprion Chevreux, 1891.—Stebbing, 1906.

Type-species: P. bolwari Chevreux, 1891 (monotypy). See
Chevreux and Fage, 1925.

Epistome and upper lip not differentially produced; mandible lack-
ing molar, palp attached rather distally, ?cutting edge dentate; gnath-
opod 1 chelate, thumb narrow; article 2 of pereopod 3 deeply indentured;
coxa 1 shorter than coxa 2 and partially hidden by it; telson of medium
length, deeply cleft. See Huwonyx. Species: 1, warm-temperate E.
Atlantic, littoral.

Podoprionella Sars

Podoprionella Sars, 1895.—Stebbing, 1906.

Type-species: P. norvegica Sars, 1895 (monotypy).

Epistome and upper lip not differentially produced; mandible lack-
ing molar, palp attached rather proximally; maxillipedal palp shorter
than outer plate, 3-articulate; gnathopod 1 chelate; article 2 of pere-

MARINE GAMMARIDEAN AMPHIPODA 359

opods 38-5 deeply indentured; outer ramus of uropod 3 uniarticulate;
telson entire. Species: 1, boreal N. E. Atlantic, littoral.

Podoprionides Walker
Podoprionides Walker, 1906a.

Type-species: P. incerta Walker, 1906a (monotypy). See Walker,
1907.

Epistome and upper lip not differentially produced; mandible lack-
ing molar, palp attached rather proximally; maxillipedal palp not ex-
ceeding outer plate, 3-articulate; gnathopod 1 chelate; article 2 of
pereopods 3-5 deeply indentured; telson of medium length, cleft more
than halfway. Species: 1, antarctic, littoral.

Prachynella J. L. Barnard
Prachynella J. L. Barnard, 1964b.

Type-species: P. lodo J. L. Barnard, 1964b (original designation).

Upper lip and epistome small, not differentially produced; man-
dibular molar absent, palp attached about in middle of mandibular
body; maxilla 1 lacking palp; maxillipedal palp with three articles,
article 3 long and claw-like; gnathopod 1 chelate; telson short, entire.
Species: 1, California, littoral.

Procyphocaris J. L. Barnard
Procyphocaris J. L. Barnard, 1961.

Type-species: P. primata J. L. Barnard, 1961 (original designa-
tion).

Head not deformed; epistome and upper lip not differentially
produced; mandibular molar ridged, palp attached level with molar;
enathopod 1 scarcely subchelate, palm very oblique; coxae 1 and 2
small, largely hidden by coxa 3; telson long, deeply cleft. Species:
1, Australia, bathypelagic.

[Pronannonyx Schellenberg]
Pronannonyx Schellenberg, 1953. Presumed synonym of Lysianassa.

Type-species: P. minimus Schellenberg, 1953 (monotypy).

Upper lip lobately produced forward (possibly including the epi-
stome); mandibular molar vestigial, palp attached quite proximally;
maxilla 1 unth two kinds of spines on the outer plate (combining charac-
ter); gnathopod 1 simple, article 6 longer than 5; uropod 3 with
l-articulate outer ramus equal to peduncle, peduncle with plate-like
expansion; telson short, entire; inner ramus of uropod 2- simple.
Species: 1, S. W. Africa, littoral.

360 U.S. NATIONAL MUSEUM BULLETIN 271

Pseudambasia Stephensen
Pseudambasia Stephensen, 1927.

Type-species: P. bipartita Stephensen, 1927 (monotypy).

Article 2 of antenna 1 elongate, two thirds as long as article 1;
epistome obtusely produced in front of upper lip; [mandible and
maxilla 1 undescribed]; gnathopod 1 subchelate, article 6 slightly
longer than 5, palm excavate (combining character); outer ramus of
uropod 3 uniarticulate, rami equal to each other and peduncle; telson
of medium length, entire; urosomal segments apparently coalesced.
Species: 1, Auckland Islands, littoral.

Pseudoanonyx Kudrjaschov

Pseudoanonyx Kudrjaschov, 1965.

Type-species: P. caecus Kudrjaschov, 1965 (original designation):

Apparently upper lip projecting slightly in front of epistome [from
analogy to Anonyx]; mandibular molar large, unridged (Onisimus),
setulose, palp apparently attached level with molar; gnathopod 1
subchelate, articles 5 and 6 stout, subequal in length, dactyl very
small and like dactyl of gnathopod 2; inner ramus of uropod 2 simple;
telson of medium length, deeply cleft; gills plaited; eyes absent;
pereopods 1-2 without distal locking spines on article 6 at base of dactyl;
article 4 of mazillipedal palp very short, tumid (Anonyz). Species:
1, Okhotsk Sea, littoral.

Pseudokoroga Schellenberg

Pseudokoroga Schellenberg, 1931.

Type-species: P. barnardi Schellenberg, 1931 (monotypy).

Upper lip and epistome produced forward together; mandibular
molar small, ridged, palp attached proximal to molar; mazilae
slender, article 4 of mazillipedal palp short (Koroga); gnathopod 1
subchelate, article 6 enlarged, much longer than article 5; inner
ramus of uropod 2 constricted; telson of medium length, entire.
Species: 2, subantarctic, Baja California, littoral.

Pseudorchomene Schellenberg

Pseudorchomene Schellenberg, 1926a.

Type-species: Orchomenopsis coats Chilton, 1912 (monotypy).

Epistome broadly lobate in front of upper lip; mandibular molar
small, poorly ridged, palp attached proximal to molar; gnathopod 1
poorly subchelate, articles 5 and 6 equal, article 3 as long as 5 or 6,

MARINH GAMMARIDEAN AMPHIPODA 361

all these articles slender and elongate (combining character); telson
deeply cleft. Species: 1, antarctic, littoral.

Rifcus Kudrjaschoy
Rifcus Kudrjaschov, 1965.

Type-species: FR. auspicatus Kudrjaschov, 1965 (original designa-
tion).

Upper lip and epistome apparently not prominent; accessory flagel-
lum 2-articulate; mandibular molar ridged (Anonyz), palp attached
level with molar; gnathopod 1 weakly subchelate, articles 5 and 6
subequal in length; inner ramus of uropod 2 not constricted; telson
short (Hippomedon), cleft less than halfway; outer plate of maxilla 1
with only five spines (Tryphosoides); palp of mazilliped very stout and
short, outer plate of mazilliped reaching nearly to end of palp article 3
(Boeckosumus). Species: 1, Okhotsk Sea, littoral.

Schisturella Norman, new synonymy

Schisturella Norman, 1900a.
Pseudonesimus Chevreux, 1926.

Type-species: Zryphosa pulchra Hansen, 1887 (monotypy). See
Shoemaker, 1930.

Upper lip lobately produced in front of epistome or not (Pseudo-
nesimus) but both separated by a notch; mandibular molar well
developed or small, blunt, ridged, palp attached level with or at
proximal end of molar; gnathopod 1 weakly or strongly subchelate,
article 5 longer than or equal to 6; coxa 1 slightly or strongly shortened,
often half as long as its article 2, largely hidden by coxa 2; inner
ramus of uropod 2 constricted; telson of medium length, cleft one
third to three quarters of its length; (article 3 of antenna 2 slightly
tumid). See Metambasia. Species: 3, Atlantic-Pacific, coldwater,
littoral to hadal.

Scopelocheiropsis Schellenberg
Scopelocheiropsis Schellenberg, 1926a.

Type-species: S. abyssalis Schellenberg, 1926a (monotypy).

Epistome lobately produced slightly in front of upper lip; mandible
lacking molar, palp attached rather distally; inner plate of maxilla
1 densely setose; maxillipedal palp article 4 vestigial; gnathopod 1
linear, simple, articles 5 and 6 equal, dactyl vestigial and shrouded
with a short tuft of setae; pereopods 1-2 subprehensile, article 6 longer
than articles 4-5 combined (combining character); telson deeply cleft.
Species: 1, tropical Atlantic, abyssal (?pelagic).

362 U.S. NATIONAL MUSEUM BULLETIN 271

Scopelocheirus Bate

Callisoma Costa, 185la, b (homonym, Coleoptera) [type species: Callisoma hopei
Costa, 185la, b, selected by Boeck, 1876].

Scopelocheirus Bate, 1857a.—Stebbing, 1906.—Birstein and Vinogradov, 1964.

Bathycallisoma Dahl, 1959 [see also this genus in alphabetical order].

Type-species: SS. crenatus Bate, 1857a(monotypy). SeeSars, 1895.

Epistome slightly and lobately produced in front of upper lip
(Arouz); mandibular molar conical, simple, palp attached level with
molar; inner plate of maxilla 1 densely setose on inner edge; gnathopod
1 simple, linear, article 7 shrouded in setae or cirri, article 6 longer
than 5; gnathopod 2 minutely chelate (Paracallisoma, Bathycallisoma) ;
telson medium to long, deeply cleft. Species: 3 or 4, probably
cosmopolitan, bathy-abyssopelagic.

Shackletonia K. H. Barnard
Shackletonia K. H. Barnard, 1931; 1932.

Type-species: S. robusta K. H. Barnard, 1931 (original designa-
tion) ; 1932.

Mouthparts presumed to be produced in a ventral conical bundle;
epistome and upper lip not differentially produced, apparently
coalesced; mandibular molar slightly conical, palp attached slightly
proximal to molar; outer plate of maxilla 1 with hook-like spines; gnath-
opod 1 simple, stout; rami of uropod 3 ovate, outer with a minute,
spine-like article 2; telson of medium length, deeply cleft. Species:
1, antarctic, deep littoral.

Socarnella Walker
Socarnella Walker, 1904.

Type-species: S. bonniera Walker, 1904 (monotypy).

[Upper lip and epistome not described]; mandibular molar absent,
palp attached proximally; gnathopod 1 simple, article 6 longer than 5;
outer ramus of uropod 3 uaiarticulate; telson short, emarginate or
slightly cleft; mandibular palp article 2 about 5 tumes as long as article 1
(Socarnopsis). Related to Lysianassa [=Lysianopsis, Arugella, etc.]
Species: 1, Ceylon, littoral.

Socarnes Boeck

Socarnes Boeck, 1871.—Stebbing, 1906 (in part, not Ephippiphora White, see
Waldeckia herein).

Type-species: Lysianassa vahlii Krdyer, 1838 (monotypy). See
Sars, 1895.

MARINE GAMMARIDEAN AMPHIPODA 363

Diagnosis based only on type-species. Upper lip lobately produced
in front of epistome or both prominently produced; mandibular molar
conicolaminate, unridged, palp attached proximal to molar; gnathopod
1 simple, articles 5 and 6 equal; outer ramus of uropod 3 biarticulate
(Socarnopsis) ; telson of medium length, cleft slightly more than half-
way; gills plaited on one side only (Ichnopus and Socarnopsis), [or two
sides in S. obesa (Chevreux)]; pereopod 5 not longer than pereopod 4
(Ichnopus). See Waldeckia. Species: 5 or 6, amphiboreal, subtropical,
littoral to bathyal.

Socarnoides Stebbing

Socarnoides Stebbing, 1888; 1906.—Schellenberg, 1931.
Acidostomella Schellenberg, 1926b.

Type-species: S. kerguelent Stebbing, 1888 (monotypy).

Epistome forming broad, flabellate lobe in front of somewhat
lobate upper lip (type) or upper lip forming lobe exceeding unproduced
epistome; mandibular molar low, moderately long, apparently ridged
(type) or setulose, palp attached proximal to molar; maxillipedal palp
scarcely exceeding outer plate (type only) ; gnathopod 1 simple, articles
5 and 6 subequal; inner ramus of uropod 3 constricted; telson of
medium length, cleft one third (type) or deeply. Species: 4 or 5,
possibly cosmopolitan (except polar), littoral.

Socarnopsis Chevreux
Socarnopsis Chevreux, 191la.

Type-species: 8S. crenulata Chevreux, 1911a (monotypy).

Upper lip and epistome together produced forward lobately; man-
dibular molar blunt, ridged, palp attached slightly proximal to molar;
enathopod 1 simple; outer ramus of uropod 3 uniarticulate; telson
of medium length, deeply cleft; gills plaited on both sides (Socarnes) ;
mandibular palp article 2 about 3 times as long as article 1 (Socarnella).
Species: 1, eastern Atlantic, littoral.

Sophrosyne Stebbing

Sophrosyne Stebbing, 1888; 1906.
Paropisa Stebbing, 1899d.

Type-species: S. murrayi Stebbing, 1888 (monotypy).

Upper lip and epistome not differentially produced; mandibular
molar absent, palp attached rather distally; outer plate of maxilla
1 poorly spinose; gnathopod 1 slightly chelate, article 6 longer than
5; telson short, deeply cleft. Species: 3, boreal-antiboreal, littoral
to bathyal, coldwater.

285-135 O - 69 - 24

364 U.S. NATIONAL MUSEUM BULLETIN 271

Stephensenia Schellenberg
Stephensenia Schellenberg, 1928a.

Type-species: S. haematopus Schellenberg, 1928a (monotypy).

Upper lip and epistome not differentially produced; mandibular
molar large, blunt, palp absent; gnathopod 1 nearly simple, article
5 longer than 6; outer ramus of uropod 3 uniarticulate; telson of
medium length, deeply cleft. Species: 1, antiboreal, littoral.

Stomacontion Stebbing
Stomacontion Stebbing, 1899d; 1906.

Type-species: <Acontiostoma pepini Stebbing, 1888 (original
designation).

Mouthparts forming conical bundle, styliform; mandibular molar
absent, palp attached very proximally; palp of maxilla 1 biarticulate
but very small; maxillipedal palp article 4 vestigial; gnathopod 1
simple, article 6 longer than 5; coxa 1 projecting over side of the head;
uropod 3 uniramous, ramus vestigial; telson short, apically notched.
[Based only on type-species. |

Note: Species assigned to this genus intergrade with Acontiostoma
and two of the species have coxa 1 ventrally subconical. Stomacontion
capense K. H. Barnard (1916) lacks any first maxillary palp and S.
insigne K. H. Barnard (1932) has the first maxillary palp 1-articulate
but combined with a vestigial fourth maxillipedal palp article, and
these characters obfuscate the generic limits of Acontiostoma and
Stomacontion. Both species, however, have a subconical first coxa,
perhaps in itself a useful generic or subgeneric character. Species:
5, subantarctic, antiboreal, littoral.

Thoriella Stephensen
Thoriella Stephensen, 1915.

Type-species: 7. islandica Stephensen, 1915 (original designation).

Accessory flagellum absent; flagellum of antenna 2 with especially
inflated articles; apparently epistome broadly rounded and produced
in front of upper lip; mandibular molar obsolete, palp absent; maxil-
liped opercular, inner plates apparently large and subrectangular,
outer plates narrower and triangular, palp apparently composed of
single large opercular article with deep lateral incision [open to dif-
ferent interpretation]; gnathopod 1 simple, short; gnathopod 2 simple;
coxae small, short, not touching serially, coxa 4 not excavate pos-
teriorly ; uropod 3 lacking rami; telson absent. Species: 1, N. Atlantic,
Indian, abyssopelagic.

MARINE GAMMARIDEAN AMPHIPODA 365

Tmetonyx Stebbing

Hoplonyzx Sars, 1895 (homonym, Coleoptera).
Tmetonyx Stebbing, 1906 (new name).

Type-species: Oniscus cicada O. Fabricius, 1780 (original desig-
nation). See Sars, 1895.

This genus was synonymized with Tryphosa by J. L. Barnard,
1962d, but Tryphosa must be synonymized with Orchomenella, of
which Orchomene is a senior synonym, because Boeck designated as
type-species of 7ryphosa a taxon later proved to be congeneric with
the designated type-species of Orchomenella. The Tryphosa concept is
now given the next available name, 7ryphosella Bonnier, but 7'met-
onyx is reinstated on the condition of coxa 1 to include only its
type-species.

Epistome broadly rounded, upper lip broadly rounded, both pro-
jecting together slightly (Anonyzx), but separated by distinct notch;
mandibular molar subcolumnar, slightly attenuated, apparently
mostly setulose and poorly or not ridged, palp attached level with
molar; cora 1 perfectly quadrate, neither expanded nor tapered distally
(Anonyz, Tryphosella); gnathopod 1 subchelate, articles 5 and 6
subequal in length; telson of medium length, deeply cleft; head of
normal size (Uristes); dactyl of gnathopod 1 with wnner tooth (Anonyz).
Species: 1 or 2, N.K. Atlantic Ocean, littoral to abyssal.

Trischizostoma Boeck

Guerinia Costa, 1853b, 1857 (homonym, Diptera).
Trischizostoma Boeck, 1861.—Stebbing, 1906.
Guerina Della Valle, 1893 (new name for Guerinia).

Type-species: Guerinia nicaeensis Costa, 1853b (monotypy and
subsequent synonymy). See Sars, 1895.

Mouthparts forming a ventral conical bundle, styliform; epistome
and upper lip not differentially produced; mandibular molar absent,
palp attached about medially; lobes of lower lip sublanceolate, lack-
ing mandibular processes; palp of maxilla 1 very short; gnathopod 1
very powerful, articles 6 and 7 generally inverted in adult so that
article 7 closes on article 6 from below; coxa 1 short, dominated by
coxa 2; telson short, entire or cleft to middle; branchiae apparently
plaited; eyes enormous. Species: 6, cosmopolitan, bathy-abyssopelagic.

Tryphosella Bonnier (=‘“‘Tryphosa’’ auct.)

Tryphosella Bonnier, 1893 [not Stebbing, 1906].
Tryphosa.—Sars, 1895.*—Stebbing, 1906 (not Boeck, 1871).

*Sars’ book is cited herein under the date 1895 but it was printed in parts from
1891 to 1895 and the section on Tryphosa was printed in 1891, two years before
Bonnier’s work.

366 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: Tryphosella sarsi Bonnier, 1893 (present selection),
(=Tryphosa nana of Sars, 1895, not of Boeck, 1861). Bonnier in-
tended the name Tryphosella to replace Sars’ usage of the name
Tryphosa for a group of species distinct from Orchomenella Sars
(=Orchomene Boeck). Sars did not realize that those species he called
Orchomenella were congeneric with Jryphosa. Stebbing (1906) also
failed to recognize Bonnier’s perception and restricted Tryphosella to
a single species, Tryphosa barbatipes Stebbing, 1888, which later has
been considered to be congeneric with Uristes Dana. Thus J. L.
Barnard’s (1962d) move of Tryphosella to Uristes was illegal because
the type-species of Tryphosella had never been selected. Legaliza-
tion of Bonnier’s intent by selection of the type-species that he
obviously would choose through his comments and his listing of it
in primary position now permits reinstatement of a “tryphosa”’
group to be called a “‘tryphosella”’ group of species. They are close
to Tmetonyx Stebbing, 1906, but have coxa 1 distinctly tapering
distally. Lepidepecreopsis Stephensen may be asynonym of Tryphosella
as Gurjanova (1951) removed the former to Tryphosa. See
discussion with T'metonyz.

Epistome broadly rounded and dominating upper lip, both sepa-
rated by a notch; mandibular molar subcolumnar or cuboid, weakly
setulose and poorly ridged if at all (Lepidepecreopsis), palp attached
level with molar; cora 1 slightly shortened, distinctly tapering
(Anonyx, Tmetonyx) and partially hidden by coxa 2; gnathopod 1
subchelate, articles 5 and 6 subequal in length; telson of medium
length, deeply cleft; head of normal size (Uristes) ; (dactyl of gnathopod
1 with or without inner tooth). Species: ca. 63, cosmopolitan cold-
water, littoral to hadal, in low latitudes primarily bathyal-abyssal.
Composition: all species formerly known under ‘‘Tryphosa”’ and
‘““Tmetonyz’”’ since 1895 except their type-species.

Tryphosites Sars
Tryphosites Sars, 1895.—Stebbing, 1906.

Type-species: Anonyzx longipes Bate and Westwood, 1863 (orig-
inal designation).

Epistome acutely produced in front of upper lip, upper lip also
slightly lobate; mandibular molar ridged, palp attached level with
molar; gnathopod 1 subchelate, article 5 longer than 6; inner ramus
of uropod 2 constricted; telson of medium length, deeply cleft. Species:
5, bipolar, coldwater submergent, littoral to abyssal.

MARINE GAMMARIDEAN AMPHIPODA 367

Tryphosoides Schellenberg

Tryphosoides Schellenberg, 1931.

Type-species: T. falcata Schellenberg, 1931 (monotypy).

Epistome not produced (apparently not as in “7ryphosa’’) ; mandib-
ular molar ridged, palp attached level with molar; gnathopod 1
subchelate, article 6 longer than 5; coxa 1 scarcely narrowed, slightly
beveled anteroventrally; outer ramus of uropod 3 uniarticulate (‘‘Try-
phosa’”’); accessory flagellum vestigial, 2-articulate; telson of medium
length, deeply cleft. Species: 1, antiboreal S. America, littoral.
Possibly synonymous with Uristes. See Rifcus.

Uristes Dana

Uristes Dana, 1849.—Stebbing, 1906.
?Pseudotryphosa Sars, 1895.
Uristoides Schellenberg, 1931.

Type-species: U. gigas Dana, 1852a; 1853 (monotypy of Dana,
1852a). See Stebbing, 1888 (as Tryphosa antennipotens).

Head small, mouthparts largely covered by corae 1-2 (Tmetonyx and
Tryphosella); epistome and upper lip generally small and incon-
spicuous, often slightly produced differentially (Tmetonyzx and Try-
phosella); mandibular molar setulose, palp attached level with molar;
enathopod 1 subchelate or nearly simple, article 6 longer than 5
(Tmetonyx usually); coza 1 narrowing slightly distally (Hippomedon) ;
telson of medium length, deeply cleft. Species: 23, cosmopolitan,
coldwater submergent, littoral to abyssal.

Valettia Stebbing

Valettia Stebbing, 1888; 1906.

Type-species: V. coheres Stebbing, 1888 (monotypy).

Apparently epistome and upper lip not prominent; mandibular
molar ridged, primary cutting edge of mandible toothed, palp attached
level with molar; setae of inner plate of mazilla 1 mostly terminal
(Valettiopsis); outer plates of maxilliped acutely produced apically;
cora 2 short, broad, subrounded, scarcely concealing coxa 1, latter not
Shorter than coxa 2 and of similar shape (Valettiopsis); gnathopod
1 weakly chelate, palm transverse; gnathopod 2 rather stout, article 5
shorter than 6; inner ramus of uropod 2 slightly constricted; telson
short, deeply cleft. Species: 1, antarctic, abyssal.

368 U.S. NATIONAL MUSEUM BULLETIN 271

Valettiopsis Holmes
Valettiopsis Holmes, 1908.

Type-species: V. dentatus Holmes, 1908 (original designation).

Apparently upper lip and epistome not prominent; mandibular
molar ridged, primary cutting edge of mandible toothed, palp attached
level with molar; inner edge of inner plate of maxilla 1 densely setose;
coxa 2 rectangular and elongate, almost covering coxa 1 completely,
latter short, subtriangular (Valettia); gnathopod 1 subchelate; telson
of medium length, deeply cleft. Species: 3, Atlantic-Pacific,
bathyal-abyssal.

Vijaya Walker
Vijaya Walker, 1904.—Gurjanova, 1962.

Type-species: V. tenuipes Walker, 1904 (monotypy).

[Epistome and upper lip unknown]; mandibular molar presumably
feeble, palp attached ?level with molar; maxilla 1 presumably lacking
palp; gnathopod 1 simple, article 6 longer than 5, article 3 slightly
elongate; coxa 1 presumably partly hidden, following coxae increas-
ingly larger; inner ramus of uropod 2 possibly constricted; telson of
medium length, cleft one third; article 2 of antenna 1 half as long as
article 1 (Bathyamaryllis); anteroventral corner of coxa 4 acutely pro-
duced (Amaryllis). Species: 1, Ceylon, littoral.

Waldeckia Chevreux

Ephippiphora White, 1847b (homonym, Lepidoptera).
Charcotia Chevreux, 1905b (homonym, Mollusca).
Waldeckia Chevreux, 1906b (new name).

Type-species: Charcotia obesa Chevreux, 1905b (monotypy).

Epistome broadly rounded and slightly produced in front of upper
lip, both together prominent; mandibular molar subconical, smooth,
palp attached proximal to molar; gnathopod 1 simple, article 6 longer
than 5; telson long, deeply cleft; gills wnplaited but with accessory
lobes (Menigrates). Species: 2, antiboreal, antarctic, littoral. See
Socarnes.

Melphidippidae
FI@uRE 133

Draenosts.—Accessory flagellum vestigial or multiarticulate; pe-
duncle of uropod 3 greatly elongate, as long as or longer than rami of
uropods 1-2, and more than twice as long as telson and peduncle of

MARINH GAMMARIDEAN AMPHIPODA 369

uropod 2, uropod 8 also greatly exceeding apices of uropods 1-2,
rami of uropod 3 elongate, subequal and nearly as long as rami of
uropods 1-2; gnathopods feeble; body posterodorsally carinate and
toothed. See Gammaridae, Oedicerotidae.

Description.—Accessory flagellum vestigial or multiarticulate;
body slender, posterodorsally carinate and toothed; rostrum poorly
developed; head with lateral ocular bulges; coxae short and scarcely
touching serially; mouthparts basic; gnathopods feeble, intermediate
between subchelate and simple; pereopods greatly elongate; uropod
3 immensely elongate as in diagnosis above and Gammaridea Key A,
couplet 7 (p. 107); telson of medium length, deeply cleft or emarginate.

ReLationsHip.—Uropod 3 usually falls off of preserved individuals
of this family as it often does on individuals of Oedicerotidae and
Megaluropus (Gammaridae). Some oedicerotids have uropod 3 elon-
gate as in melphidippids but they lack posterodorsal pleonal teeth.
Melphidippids may also be recognized by the lateral ocular bulges of
the cephalon and the very short coxae. Gammaridae with elongate
uropod 3 have one or more of the following exceptions to the melphi-
dippid pattern: inner ramus of uropod 8 shortened and scale-like;
uropod 3 not exceeding apices of uropods 1-2; peduncle of uropod 3
shorter than rami of uropods 1-2 and not twice as long as telson or
urosomite 3; rami of uropod 3 much shorter than peduncle. Melphi-
dippids lack sexual dimorphism in the gnathopods common to most
Gammaridae.

The Vitjazianidae have a short peduncle of antenna 1.

My interpretation of Walker’s description of Hornellia is that it
lacks the melphidippid condition of uropod 3, for he mentions the
medium length of the uropods and that the inner ramus of uropod 3
is twice as long as the peduncle; if the peduncle were of the normal
melphidippid length the total uropod would be two-thirds the length
of the body, a condition so remarkable that Walker would have
mentioned it. Indeed, Hornellia Walker was removed to the Gam-
maridae by Pillai (1957).

The calliopiid Metaleptamphopus may belong in the Melphidippidae.

Key to the Genera of Melphidippidae

1. Accessory flagellum long, 2- or more articulate (fig. 133b) . . Melphidippa
Accessory flagellum very short, l-articulate (fig. 138c)......... 2
Zomeehelson. sclekt: i(figs USI): annual Meath aloo a oh be Ge Melphidippella
Melson’ entire! ‘emarginate (ig 33) ek oe a cere Melphisana

370 U.S. NATIONAL MUSEUM BULLETIN 271

“tly
Pod Iit
ZC ip oa)

“il

Ficure 133.—Melphidippidae: a, Melphidippa macrura Sars (1895, pl. 170). Accessory
flagella: b, Melphidippa goesi Stebbing (Sars, 1895, pl. 169, as M. spinosa); c, Melphidip-
pella macra (Norman) (Sars, 1895, pl. 171). Mouthparts, M. goest: d, upper lip; e,
mandible; f, lower lip; g,h, maxillae 1, 2; 7, maxilliped. M. goesi: j,k, gnathopods 1, 2;
1, uropod 3; m, telson. Telson: n, Melphisana bola J. L. Barnard (1962b).

MARINE GAMMARIDEAN AMPHIPODA Si
Genera of Melphidippidae

Melphidippa Boeck
Melphidippa Boeck, 1871.—Stebbing, 1906.

Type-species: Melphidippa goest Stebbing, 1899a (=new name
for homonym, Gammarus spinosus Goés, 1866) (type indicated by
Boeck, 1876, considered as type by Sars, 1895, but selected by Gurja-
nova, 1951). See Sars, 1895.

Accessory flagellum long, 2- or more articulate; mandibular palp
article 3 twice as long as article 1; maxillipedal palp article 4 claw-
shaped; gnathopod 2 with linear fifth and sixth articles; telson cleft.
Species: 7, bipolar, littoral to bathyal (750 m).

Melphidippella Sars

Melphidippella Sars, 1895.—Stebbing, 1906.

Type-species: Atylus macer Norman, 1869a (original designation).

Accessory flagellum short, 1-articulate; mandibular palp article
3 equal to article 1; maxillipedal palp article 4 claw-shaped; gnathopod
2 with linear fifth and sixth articles; telson cleft. Species: 2, boreal
N. E. Atlantic, Japan, littoral.

Melphisana J. L. Barnard

Melphisana J. L. Barnard, 1962b.

Type-species: MM. bola J. L. Barnard, 1962b (original designation).

Accessory flagellum short, 1l-articulate; mandibular palp article 3
equal to article 1; maxillipedal palp article 4 short, stout, not claw-
shaped; gnathopod 2 with articles 5 and 6 intermediate between linear
and oval; [uropod 3 unknown]; telson entire, emarginate. Species: 2,
California, Japan, littoral.

Ochlesidae

FIGURE 134

Driacenosis.—Accessory flagellum absent; palp of marilliped absent;
mouthpart group projecting subconically; mandible lacking molar,
coxae 1-4 subacuminate; telson entire. Monogeneric. See Lysi-
anassidae, Acanthonotozomatidae.

Description.—Body massive, processiferous; accessory flagellum
absent; flagella of antennae reduced to a few articles; rostrum large;
coxae 1-4 subacuminate; mouthparts projecting in a subconical

3/2 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 134.—Ochlesidae: a, Ochlesis levetzowi Schellenberg (1953). Antennae 1, 2: 5, O.
innocens Stebbing (1910). Lower lip: c, O. lenticulosus K. H. Barnard (1940). 0. in-
nocens (after Pirlot, 1936): d, mandible; ¢, maxilla 1; f, maxilla 2; g, maxilliped; h,2,
gnathopods 1, 2. Uropod 3: 7, O. innocens Stebbing (1910). Telson: k, O. innocens
(after Pirlot, 1936); J, O. lenticulosus.

MARINE GAMMARIDEAN AMPHIPODA 373

bundle, elongate; upper lip elongate; mandible lacking molar, palp
long, 3-articulate, article 1 elongate; lower lip lacking inner lobes,
outer lobes apically acuminate; maxilla 1 with minute, vestigial
palp or none; maxilla 2 elongate; maxillipeds lacking palps, inner
plates acuminate, outer lobes operculiform; gnathopods simple or
mero- and carpochelate; uropod 3 biramous; telson of medium length,
entire.

ReELATIONSHIP.—Of the suborder Gammaridea, only the Ochlesidae
and the genus Danaella (see also Thoriella and Chevreuziella) in
the Lysianassidae lack a maxillipedal palp. This lack is characteristic
of all members of the suborder Hyperiidea but Ochlesidae appear
in other respects to be so closely related to gammarideans that they
are retained therein. The large coxae are especially characteristic
of gammarideans but the general body shape resembles that of the
Acanthonotozomatidae and Stilipedidae.

Genera of Ochlesidae

Ochlesis Stebbing

Ochlesis Stebbing, 1910.

Type-species: 0. innocens Stebbing, 1910 (monotypy).
Species: 3, S. Africa, Indonesia, Australia, littoral.

Oedicerotidae

FIGURES 135-138

Diacnosis.—Accessory flagellum absent or vestigial; telson entire
or emarginate, very short; eyes when present usually coalesced
dorsally; head often massive and galeate (see glossary); anoculate
members assigned to the family on basis of aspects similar to shallow
water genera having coalesced eyes; pereopod 5 immensely longer
than pereopod 4; rami of uropod 3 scarcely longer than the elongate
peduncle. See Calliopiidae, Pleustidae, Isaeidae, Ampithoidae, Syn-
oplidae, Laphystiopsidae.

Description.—Accessory flagellum absent; head often galeate
and massive, conspicuous rostrum usually present but occasionally
completely absent; mouthparts basic except for mandibular molar
which varies from strongly triturative to nontriturative and nearly
obsolescent; one genus lacking mandibular palp; gnathopods powerful
to weak, highly variable; coxae of medium size; rami of uropod 3
narrowly lanceolate, scarcely longer than the elongate peduncle;

374 U.S. NATIONAL MUSEUM BULLETIN 271

telson entire or emarginate; pereopods strongly setose (fossorial),
pereopod 5 immensely longer than pereopod 4.

ReELATIONSHIP.—In gross morphology, this family is scarcely dis-
tinct from the Calliopiidae and Pleustidae in which the telson is

ay ry. ae

//

Sc
\

a lh
7 samred

foaointhic
Ni x SW) | 2. a
ae ee ‘ eee
>

Ficure 135.—QOedicerotidae: a, Oediceros saginatus Kroyer (Sars, 1895, pl. 102); b, Synchel-
idium halpocheles (Grube) (Sars, 1895, pl. 112); c, Monoculodes carinatus (Bate) (Sars,

1895, pl. 105).

MARINE GAMMARIDEAN AMPHIPODA 375

also entire and the accessory flagellum vestigial or absent. The rel-
atively longer peduncle of uropod 3 in oedicerotids is a reliable
character as is the aspect of those oedicerotids having massive heads
and projecting rostra; often they have pereopods adapted for bur-
rowing by enlarged and spinose articles and pereopod 5 is much longer
than pereopod 4. Since the eyes are lost in many species the taxono-
mist requires some experience in differentiating these families.

Ficure 136.—Oedicerotidae: a, Oediceroides apicalis K. H. Barnard (J. L. Barnard, 1961);
b, Aceroides latipes Sars (1895, pl. 120). Telson: c, Paroediceroides sinuata Schellenberg
(1931); d, Monoculodes borealis Boeck (Sars, 1895, pl. 106); ¢, Monoculodes tesselatus
Schneider (Sars, 1895, pl. 106); f, Oediceros saginatus Kr@yer (Sars, 1895, pl. 102). Uro-
pod 3: g, Oediceros. Coxa 4: h, Perioculodes longimanus (Bate and Westwood) (Sars,
1895, pl. 110).

376 U.S. NATIONAL MUSEUM BULLETIN 271

The Synopiidae and Pardaliscidae usually have a well-developed
accessory flagellum in contrast to its absence in oedicerotids.

The Isaeidae (=Photidae) differ from Oedicerotidae in their
glandular pereopods, fleshy telson, and tube-dwelling habits but some
Oedicerotidae, lacking rostra, are morphologically similar to photids
and differ from them by disparate shortening of pereopods 3-4 in
relation to pereopod 5.

Oedicerotids need extensive generic revision. Ocular shapes are
useful for those genera having oculate type-species but deep-sea
species lacking eyes often are difficult to classify. If eyes were to be
ignored various genera might be combined, such as Oediceropsis and
Paroediceroides (but note that the type-species of Oediceropsis has
a swollen article 1 of antenna 1); Paroediceroides might be included
with Oediceroides except for its acutely produced coxa 4 (a feature
which is present or absent in species of other genera); Paraperioculodes
might be congeneric with Oediceroides and Paroediceroides as coxa 4
is not specifically described, or with Oediceropsis as antenna 2 article
1 is not described; Oediceroides brevirostris may belong with Paraperi-
oculodes (fide Ruffo, 1949); Oediceropsis proxima may belong with
Paroediceroides. Numerous other examples could be cited.

Metoediceros Schellenberg (1931) lacks a mandibular palp, uropod
3 has a simple peduncle without rami, appears to have lateral, unco-
alesced eyes and probably should form the type of a new family with
affinities near the Dogielinotidae.

Key to the Genera of Oedicerotidae '

This key attempts to minimize the characteristics of eyes but several
genera cannot remain separate without some reference to eyes, as seen
in the later portion of the key.

1 (Gnathopod:2 chelate (igs l38k) ho) a. ee ee ee
Gnathopod 2 subchelate or simple . . . 3

2. Mandibular molar triturative (fig. 137k), Panthened 2 ef lone tebe on
article 5 guarding article 6 ..... .. . . . Pontocrates
Mandibular molar degraded, not triturative, ehen beanies articulate spine(s),
article 5 of gnathopod 2 not lobate (fig. 1371) . . . . . . Synchelidium

3. Primary cutting edge of mandible short, poorly projecting, untoothed
Gigs lS 7Tk)iea eae 4

Primary cutting edge of sapritialle paonely oneieaans, footed Ge 1371) . 8
4. Gnathopod 2 with posterior lobe of article 5 partially or fully guarding

article 6 (figs. 1387,7,l). . . . 6
Gnathopod 2 with posterior lobe af article) 5 jornectiins at saeth sipettag ner
guarding article 6 (figs) 1S8dih mm) vey na eae

1 Oedicerina Stephensen (1931) is not included; many of its features are not described because the type is
a broken specimen.

11.

12.

13.

14.

15.

16.

17.

18.

WG

20.

21.

22.

23.

24.

25.

26.

MARINE GAMMARIDEAN AMPHIPODA 377

Gnathopod 1 palm oblique . . . . . . . Westwoodilla and Bathymedon
Gnathopod 1 palm transverse .. . ...... . Carolobatea
Mandibular molar unridged, with a fen nO. .... . . . Perioculopsis
Mandibular molar large, ridged. . . 7
“Mandibular palp article 2 strongly sree” (Ge. 137h), arial 2 of prrcane:
1 as long as or longer than article]. . . . .. . . Arrhis
“Mandibular palp article 2 straight’ (fig. 1371) (type species), article 2 of
antenna 1 shorter than article] .... . . . Aceroides
Uropod 2 reaching only to end of peduncle at Tropa 3. ER en St O
Uropod 2 reaching halfway or more ane rami of uropod 3 (or rami of
uropod 3 absent) ... . RANG ata MON YS oly re uiereae else MN ANZ,
Gnathopods 1-2: Metiele sie ronely lobate: Loupe ar ciitibnnn Glee canmiecug. IE
Gnathopods 1-2: article 5 not lobate (figs. ee BN ee Pe eas wy Seen UO)
Gnathopod 2 simple (fig. 1387). . ..... Uae ina Nes ibaihwnoreiapis
Gnathopod 2 subchelate ... . . . . . . Parhalimedon
Eyes contiguous, appearing as one Body, BE rcapods 1 and 2 with article 7.
Halicreion
Eyes not contiguous, pereopods 1 and 2 lacking article 7 . . . Exoediceros

Mandible lacking palp, uropod 3 lacking rami.
Metoediceros (Incertae Sedis)

Mandible bearing palp, uropod 3 bearing rami. . . . Bie tie eat)
Mandibular molar lacking triturating surface, often ah preiculate spine(s)

(Giles Wa) Son ip BS eA aeons cdnaeanig Leg
Mandibular molar hearts teeth ema dee: (Ge. 137k) . aS Nay ap iase's Coe agiae ho)
Wowe»r lip: imner lobes coalesced (fig. 137t). .0 4. su.) Sa TS
lower lips innerclobes separate; (fig-13/w)> ci. ee i a 16
Melsonventire (fie 136f). 50202 2 8. ee ws) Perioculodes
Telson emarginate (fig. 186a) . ............ . . Perioculopsis
Gnathopods—2 structurally salike: ys (uaa, ie gees ee a Ng,
Gnathopods 1-2 structurally dissimilar. . . ... . . . . Paroediceros
Gnathopods lacking posterior lobe on article5 . .. . . . Exodiceropsis
Gnathopods bearing posterior lobe on article 5 . . Bb oicties ei OME Weo)

Gnathopods 1-2: lobe of article 5 long, fully guarding article! 6. Arrhinopsis
Gnathopods 1-2: lobe of article 5 short, scarcely guarding article 6.

Oediceros
Gnathopod 1, palm transverse ............ +. . Carolobatea
Gnathopod 1, palm oblique. . . . Bue iad Ra Ma colette eer 7D)
Gnathopods 1-2: article 5 not deanery lobate 2 EE al eta Matas MOEN eT
Gnathopods, 1—2: article 5 grossly lobate |. . 3.20: 0-5) 2... 2 28
Gnathopod’2isubchelate: 70.26) ait. thei. 2 ee ene Hie ee aes V4
Gnathopod 2 simple (fig. 1887) ... . SR aa fpuihyporeianus
Eyes unpaired, coalesced; article 2 of nereonod 5 broadly lobed postero-
distally ue: . . . Methalimedon
Eyes paired; article 2 oF Mereonod 5 cincuaty narrowed posterodistally.
Parhalimedon
Back multicarinate ... . ... . . Acanthostepheia
Back not multicarinate, te aaliy, mouth Bec mionally tuberculate . . . 24
Coxa 4 produced subacutely posterodistally ........2.2... 25
Coxai 4 not produced posterodistally jn) 3 3 es 28
Eyes coalesced dorsally or absent . ....... Oe ae 26
Myvesspairedvand lateral’. i) ec od l@ediceconsic (typical)
Article 1 of antenna 2 swollen .... Cregeen OM

Article 1 of antenna 2 not swollen. . Oediccreides ana Paroedicertides

378 U.S. NATIONAL MUSEUM BULLETIN 271

27. Posterior lobe on article 5 of gnathopod 2 guarding article 6 (fig. 1381).

Monoculodes

Posterior lobe on article 5 of gnathopod 2 projecting at right angles.
Oediceropsis
28. Pereopods 1-2: article 7 absent. ......... ... .. .. Exoediceros
Pereopods) 1-2 article: 7-presemtic: five 1 coien etek har Ed Caleta eee)
29. Hyes Pomplctels coalescediaeg an) a. SETURL ath Rea} ()
Kyes separated by a median line or epee or = foram PeCmmemme rs teeter adn co) lb
30. Eyes forming asemicircular ring. ......... ... . Gulbarentsia
Eyes forming a circle or oval . . . . . Paraperioculodes
31. Gnathopod 2: lobe of article 5, if lene! anefectine erectly at nearly right
angles, not guarding article6.. . 55 a
Gnathopod 2: lobe of article 5 DrOVeCtne distally at aravlle of 45° or less,
guarding article6... . SAL ae Se A ERG AAS
32. Eyes paired and lateral (@heorctical Moretti) . . . . . . Oediceropsis
Byes:contiguous:at midline or absentiaiy <a 4) ai fs ye cele cee
33. Maxilla 2, outer plate lacking stout spine . . PPR aA SG Es coves
Maxilla 2, outer plate bearing stout spine (often pifid) . . Anoediceros

34. Basal portion of flagellum on antenna 2 swollen.
Subgenus Lopiceros (see Oediceroides)
Basal portion of flagellum on antenna 2 slender. ... . OEE ao
35. Coxae 3 or 4 excavate ventrally (fig. 136b), antennae pube qual in length.
Aceroides (incl. subgenus Patozdes)
Coxae 3 or 4 not excavate ventrally, antenna 1 very disproportionately

shorter than antenna2 ... . . . . Oediceroides (= Oediceropsoides)

36. Coxa 4 excavate ventrally, rostrum very small or obsolete (fig. 137d), eyes
Absent mn mee ... . . Aceroides

Coxa 4 not excavate “capetnalli either rostaai lerees extending more than
halfway along article 1 of antenna 1, oreyes present. ....... .37

37. Article 3 of antenna 1 less than half as long as article1. . . . Monoculodes
Article 3 of antenna 1 as long as article 1. .... . . . .Monoculopsis

Genera of Oedicerotidae

Acanthostepheia Boeck
Acanthostepheia Boeck, 1871.—Stebbing, 1906.

Type-species: Amphithonotus malmgreni Goés, 1866 (monotypy).
See Stephensen, 1938.

Cutting edge of mandible projecting and toothed, molar apparently
ridged; inner lobes of lower lip separate; gnathopods similar to each
other, subchelate, stout, article 5 with large posterior lobe projecting
at right angles; uropod 2 reaching end of uropod 3; body dorsally
multicarinate (combining character). Species: 3, arctic, littoral to
bathyal (552 m).

Aceroides Sars
Aceroides Sars, 1895.—Stebbing, 1906.

Aceropsis Sars, 1895.
(Patoides) J. L. Barnard, 1964a (subgenus).

MARINE GAMMARIDEAN AMPHIPODA 379

Type-species: Halicreion latipes Sars, 1882 (monotypy). See Sars,
1895.

Cutting edge of mandible scarcely projecting, teeth obsolescent,
molar large, ridged; inner lobes of lower lip separate; gnathopods
similar, subchelate, moderately stout, article 5 with sharp posterior
lobe projecting distalwards; uropod 2 reaching end of uropod 3;
mandibular palp article 2 straight (Arrhis). Probably Aceroides limicola
K. H. Barnard, 1925, is incorrectly assigned here; its mandible
apparently has cutting teeth. Species: 4, arctic-boreal, littoral to
bathyal (1,110 m) and S. Atlantic (1,280 m).

Ficure 137.—Oedicerotidae: Head: a, Oediceropsis brevicornis Liljeborg (Sars, 1895, pl.
114); 5, Monoculodes longirostris (Goés) (Sars, 1895, pl. 108); c, Oediceros saginatus Krdyer
(Sars, 1895, pl. 102); d, Aceroides latipes (Sars, 1895, pl. 120); e, dorsal, Oediceros; f,
Monoculodes carinatus (Bate) (Sars, 1895, pl. 105); g, Arrhis phyllonyx (M. Sars) (Sars,
1895, pl. 119); h, Westwoodilla caecula (Bate) (Sars, 1895, pl. 115, as Halimedon mulleri);
1, Westwoodilla acutifrons (Sars, 1895, pl. 116); 7, Monoculodes packardi Boeck (Sars, 1895,
pl. 109). Mandible: k, Westwoodilla caecula; 1, Oediceros; m, Synchelidium haplocheles
(Grube) (Sars, 1895, pl. 112, as S. brevicarpum). Upper lip: n, Oediceros. Maxilliped:
0, Oediceros. Maxillae 1, 2: p,q, Oediceros. Gnathopod 1: r, Westwoodilla caecula; s,
Synchelidium. Lower lip: t, Perioculodes longimanus (Bate and Westwood) (Sars, 1895,
pl. 110); u, Oediceros.

285-135 O - 69 - 25

380 U.S. NATIONAL MUSEUM BULLETIN 271

Anoediceros Pirlot

Anoediceros Pirlot, 1932.

Type-species: A. hanseni Pirlot, 1932 (original designation).

Cutting edge of mandible projecting, toothed, molar medium,
ridged; inner lobes of lower lip separate; outer plate of maxilla 2 bearing
a stout spine (combining character); gnathopods similar, subchelate,
stout, article 5 with posterior lobe projecting at right angles; uropod 2
reaching end of uropod 3. Species: 1, Indo-Pacific, bathyal-abyssal.

l m
Ficure 138.—Oedicerotidae: Gnathopod 1: a, Aceroides latipes (Sars, 1895, pl. 120); 3,
Paroediceros lynceus (M. Sars) (Sars, 1895, pl. 103); c, Exoediceropsis chiltoni Schellen-
berg (1931); d, Oediceros saginatus Kréyer (Sars, 1895, pl. 102); e, Monoculodes tenutro-
stratus Boeck (Sars, 1895, pl. 109); f, Bathyporeiapus magellanicus Schellenberg (1931).
Gnathopod 2: g, Monoculodes carinatus (Bate) (Sars, 1895, pl. 105); h, Oediceros; 1,
Bathyporeiapus; j, Aceroides; k, Synchelidium haplocheles (Grube) (Sars, 1895, pl. 112);

1, Monoculodes tenuirostratus; m, Westwoodilla caecula (Bate) (Sars, 1895, pl. 115, as
Halimedon mulleri).

MARINE GAMMARIDEAN AMPHIPODA 381

Arrhinopsis Stappers
Arrhinopsis Stappers, 1911.

Type-species: <A. longicornis Stappers, 1911 (monotypy). See
Gurjanova, 1951.

Cutting edge of mandible projecting, toothed, molar bulging,
setulose, but unridged; inner lobes of lower lip separate; gnathopods
similar to each other, subchelate, moderately stout, lobe of article 5
long and guarding article 6; uropod 2 reaching end of uropod 3.
Species: 1, arctic, littoral.

Arrhis Stebbing

Aceros Boeck, 1861 (homonym, Aves).
Arrhis Stebbing, 1906 (new name).

Type-species: Leucothoe phyllonyx M. Sars, 1858 (original designa-
tion and subsequent synonymy). See G. O. Sars, 1895.

Cutting edge of mandible not projecting, untoothed, molar large,
ridged; inner lobes of lower lip separate; gnathopods similar, subchelate
moderately stout, article 5 with sharp posterior lobe pointing distally;
uropod 2 reaching end of uropod 3; mandibular palp article 2 strongly
curved (Aceroides). Species: 3, arctic, littoral to abyssal.

Bathymedon Sars
Bathymedon Sars, 1895.—Stebbing, 1906

Type-species: Halimedon longimanus Boeck, 1871 (original desig-
nation). See Sars, 1895.

Cutting edge of mandible not projecting, untoothed, molar large,
ridged; inner lobes of lower lip separate; gnathopods somewhat
similar to each other, usually gnathopod 2 more slender, posterior
lobe of article 5 moderately developed in gnathopod 1, becoming
obsolescent occasionally in gnathopod 2; uropod 2 reaching end of
uropod 3; “article 2 of mandibular palp not as curved as in Westwood-
illa.”” Species: 16, cosmopolitan coldwater, littoral to abyssal.

Bathyporeiapus Schellenberg
Bathyporeiapus Schellenberg, 1931.

Type-species: B. magellanicus Schellenberg, 1931 (monotypy).

Cutting edge of mandible projecting, toothed, molar large, ridged;
inner lobes of lower lip separate; gnathopods somewhat dissimilar,
slender, gnathopod 1 subchelate, gnathopod 2 simple; uropod 2
apparently only slightly exceeding peduncle of uropod 3. Species:
1, antiboreal, littoral.

382 U.S. NATIONAL MUSEUM BULLETIN 271

Carolobatea Stebbing
Carolobatea Stebbing, 1899d.—Stebbing, 1906.

Type-species: Halimedon schneideri Stebbing, 1888 (original desig-
nation).

Cutting edge of mandible projecting slightly, teeth obsolescent,
molar medium, cup-shaped, dentate; inner lobes of lower lip separated;
enathopods similar to each other, subchelate, small, article 5 with
small posterior lobe, palm of gnathopod 1 transverse (combining charac-
ter); uropod 2 reaching end of uropod 3. Species: 1, New Zealand,
littoral.

Exoediceropsis Schellenberg
Exoediceropsis Schellenberg, 19381.

Type-species: . chaltoni Schellenberg, 1931 (monotypy).

Cutting edge of mandible projecting, toothed, molar small, leaf-
like, produced into a spine; inner lobes of lower lip separate; gnatho-
pods similar to each other, subchelate, slender, article 5 not lobed;
uropod 2 reaching end of uropod 3; coxa 4 produced acutely posteriorly
(combining character). Species: 1, antiboreal, S.W. Atlantic, littoral.

Exoediceros Stebbing
Exoediceros Stebbing, 1899d.—Stebbing, 1906.

Type-species: Oedicerus fossor Stimpson, 1855 (original designa-
tion). See Haswell, 1880c (as Oedicerus arenicola).

Cutting edge of mandible projecting, toothed, molar large, ridged;
{lower lip unknown]; gnathopods similar to each other, subchelate,
article 5 lobed; uropod 2 apparently not reaching far along uropod 3;
eyes not contiguous (Halicreion); article 7 of pereopods 1 and 2 absent.
Species: 2, S.E. Australia, littoral.

Gulbarentsia Stebbing

Barentsia Stebbing, 1894 (homonym, Bryozoa).
Gulbarentsia Stebbing, 1894 (new name, footnote, p. 2).

Type-species: Barentsia hoeki Stebbing, 1894 (monotypy). See
Gurjanova, 1951.

Cutting edge of mandible projecting slightly, toothed; molar
medium, cup-shaped, dentate; inner lobes of lower lip separate;
enathopods similar to each other, subchelate, stout, lobe of article 5
projecting strongly but not especially guarding article 6; uropod 2
reaching end of uropod 3; eyes completely fused, forming a semicircular
ring (Paraperioculodes). Species: 2, arctic, littoral.

MARINE GAMMARIDEAN AMPHIPODA 383

Halicreion Boeck
Halicreion Boeck, 1871.—Stebbing, 1906.

Type-species: Oediceros aequicorms Norman, 1869a (monotypy
and subsequent synonymy). See Sars, 1895 (as H. longicaudatus).

Cutting edge of mandible projecting slightly, poorly but distinctly
toothed, molar large, ridged; inner lobes of lower lip separate; gnatho-
pods similar to each other, subchelate, stout, article 5 strongly lobate,
lobe on gnathopod 2 sufficiently long to appear to be guarding article
6; uropod 3 very long, uropod 2 not exceeding peduncle of uropod 3;
eyes contiguous (Hxoediceros). Species: 3, bipolar, coldwater littoral
to bathyal (730 m, S. Atlantic).

Methalimedon Schellenberg
Methalimedon Schellenberg, 1931.

Type-species: MM. nordenskjoldi Schellenberg, 1931 (monotypy).

Cutting edge of mandible projecting, toothed, molar large, ridged;
inner lobes of lower lip separate; gnathopods similar to each other,
feeble, subchelate, article 5 not lobate; uropod 2 apparently reaching
end of uropod 3; eyes unpaired, coalesced, base of pereopod 5 broadly
lobed distally (Parhalimedon). Species: 1, subantarctic, littoral (to
310 m).

Monoculodes Stimpson

Monoculodes Stimpson, 1853.—Stebbing, 1906.
Kroyera Bate, 1857a, 1858a.

Type-species: MM. demissus Stimpson, 1853 (monotypy). Species
obscure.

Antenna 1 not longer than antenna 2, article 3 less than half as long
as article 1 (Monoculopsis); antenna 2 neither enlarged nor elongate
(Oediceroides); cutting edge of mandible projecting, toothed, molar
large, ridged; gnathopods dissimilar, subchelate, gnathopod 1 usually
stout, lobe of article 5 long or short, but usually not appearing to
guard article 6; gnathopod 2 usually slender, lobe of article 5 usually
very long and guarding article 6; both gnathopods merging to those of
Oediceroides; uropod 2 reaching end of uropod 3. Species: 46, cosmo-
politan in N. Hemisphere, littoral to bathyal (only 2 species in S. Hem-
isphere, Tasman Sea, 610 m, and S. Georgia area).

Monoculopsis Sars
Monoculopsis Sars, 1895.—Stebbing, 1906.

Type-species: Monoculodes longicornis Boeck, 1871 (monotypy).

384 U.S. NATIONAL MUSEUM BULLETIN 271

Antenna 1 longer than antenna 2, article 3 as long as article 1 (Mono-
culodes); cutting edge of mandible projecting and sharp but poorly
toothed, molar large, ridged; inner lobes of lower lip separate;
gnathopods dissimilar or similar, subchelate, article 5 with lobe
guarding article 6 on gnathopod 2, lobe not guarding as closely on
gnathopod 1; uropod 2 reaching end of uropod 3. Species: 2,
bisubpolar, littoral.

Oedicerina Stephensen
Oedicerina Stephensen, 1931.

Type-species: 0. ingolfi Stephensen, 1931 (monotypy). See Gur-
janova, 1951.

Cutting edge of mandible slightly projecting, toothed, molar large,
ridged; inner lobes of lower lip separate; [gnathopods unknown]; coxa
4 with a large blunt posterior lobe. Type specimen badly damaged.
Species: 1, N. Atlantic (1,802 m).

Oediceroides Stebbing

Oediceroides Stebbing, 1888; 1906.
Oediceropsoides Shoemaker, 1925.
(Lopiceros) J. L. Barnard, 1961 [a valid subgenus].

Type-species: Oecdiceropsis rostrata Stebbing, 1883 (selected by
Pirlot, 1932).

Cutting edge of mandible projecting, toothed, molar large, ridged;
inner lobes of lower lip separate; gnathopods similar to each other,
stout, subchelate, posterior lobes of article 5 well developed, pro-
jecting at right angles or nearly so, or not guarding article 6 (Monocu-
lodes), lobes becoming obsolescent; uropod 2 reaching end of uropod
3; inner plate of maxilla 1 with three or more setae (Paroedicerordes) ;
antenna 2 usually enlarged and elongate (Monoculodes). Species: 24,
cosmopolitan cold-water, littoral to abyssal. Oediceroides cystrfera
and O. brevirostris Schellenberg (1931) probably incorrectly assigned
here.

Oediceropsis Liljeborg

Oediceropsis Liljeborg, 1865.—Stebbing, 1906.

Type-species: 0. brevicornis Liljeborg, 1865 (monotypy). See Sars,
1895.

Eyes lateral; cutting edge of mandible projecting, toothed, molar
large, ridged; inner lobes of lower lip separate; gnathopods similar
to each other, subchelate, stout, lobe of article 5 projecting at right
angles; uropod 2 reaching end of uropod 3. Article 1 of antenna 2
swollen in type-species. Bridged through 0. prozima Chevreux (1908c)
to Parodiceroides. Species: 2, boreal N. Atlantic, bathyal.

MARINE GAMMARIDEAN AMPHIPODA 385

Oediceros Kr¢éyer
Oediceros Krgyer, 1842.—Stebbing, 1906.

Type-species: O. saginatus Krgyer, 1842 (monotypy). See Sars,
1895.

Cutting edge of mandible projecting and well toothed, molar lack-
ing ridges; inner lobes of lower lip separate; gnathopods similar to
each other, subchelate, lobes of article 5 well developed and project-
ing at right angles; uropod 2 reaching end of uropod 3. Species:
3, arctic-boreal N. Atlantic, littoral.

Paraperioculodes K. H. Barnard
Paraperioculodes kK. H. Barnard, 1931; 1932.

Type-species. P. brevimanus K. H. Barnard, 1931 (original des-
ignation) ; 1932.

Eyes completely fused, forming a circle or oval (Gulbarentsia, Oedi-
ceroides); cutting edge of mandible apparently projecting, toothed,
molar ridged; inner lobes of lower lip separate; gnathopods similar
to each other, moderately stout, subchelate, posterior lobes of article
5 projecting somewhat distally; uropod 2 reaching end of uropod 3;
cora 4 apparently only bluntly and poorly or not produced posteriorly
(Paroediceroides). Species: 3, antarctic, littoral to bathyal (569 m).

Parhalimedon Chevreux
Parhalimedon Chevreux 1906a; 1906b.

Type-species: P. turqueti Chevreux, 1906a (original designation) ;
1906b.

Eyes paired; base of pereopod 5 sharply narrowed distally (Methali-
medon); cutting edge of mandible projecting, toothed, molar large,
ridged; inner lobes of lower lip separate; gnathopods similar to each
other, feeble, subchelate, elongate, fifth articles lacking posterior
lobes; uropod 2 reaching only to end of peduncle of uropod 3. Spe-
cies: 2, antarctic littoral and tropical bathyal (230 m).

Paroediceroides Schellenberg
Paroediceroides Schellenberg, 1931.

Type-species: P. sinuata Schellenberg, 1931 (monotypy).

Cutting edge of mandible projecting, toothed, molar large, ridged;
inner lobes of lower lip separate; gnathopods similar to each other,
stout, subchelate, posterior lobe of article 5 well developed, par-
tially guarding article 6; uropod 2 reaching end of uropod 3; inner
plate of maxilla 1 with one seta (Oediceroides) ; telson deeply emarginate
in type-species. Species: 2, subantarctic littoral and E. Pacific
bathyal.

386 U.S. NATIONAL MUSEUM BULLETIN 271

Paroediceros Sars
Paroediceros Sars, 1895.—Stebbing, 1906.

Type-species: Oediceros lynceus M. Sars, 1858 (original designa-
tion).

Cutting edge of mandible projecting, toothed, molar unridged;
inner lobes of lower lip separate; gnathopods dissimilar, subchelate,
article 5 of gnathopod 1 scarcely lobed, of gnathopod 2 with a long
posterior lobe guarding article 6; uropod 2 reaching end of uropod 3.
Species: 5, arctic-subarctic, littoral to deep bathyal.

Perioculodes Sars
Perioculodes Sars, 1895.—Stebbing, 1906.

Type-species: Monoculodes longimanus Bate and Westwood, 1868
(monotypy).

Cutting edge of mandible projecting, toothed, molar very small,
conical, unridged; inner lobes of lower lip coalesced; gnathopods
similar to each other, subchelate, article 5 produced into a lobe
guarding article 6; uropod 2 reaching end of uropod 3; telson entire
(Perioculopsis). Species: 3, Ceylon, Red Sea, boreal E. Atlantic,
littoral.

Perioculopsis Schellenberg
Perioculopsis Schellenberg, 1925b.

Type-species: P. lophopus Schellenberg, 1925b (monotypy).

Cutting edge of mandible projecting, not distinctly toothed, molar
small, unridged; inner lobes of lower lip apparently coalesced; gnatho-
pods similar to each other, subchelate, article 5 produced into a lobe
guarding article 6; uropod 2 reaching end of uropod 3; telson emarginate
(Perioculodes). Species: 1, W. Africa, littoral.

Pontocrates Boeck
Pontocrates Boeck, 1871.—Stebbing, 1906.

Type-species: Kroyera arenaria Bate, 1858a (selected as Oedicerus
norvegicus Boeck, 1861, by Boeck, 1876, a species now synonymous
with K. arenaria). See Sars, 1895 (as P. norvegica).

Cutting edge of mandible projecting, toothed, molar medium,
ridged; inner lobes of lower lip poorly developed but separated by
an incision; gnathopods dissimilar, gnathopod 1 stout, subchelate,
article 5 produced into a long posterior lobe partially guarding article
6, gnathopod 2 slender, chelate, long lobe of article 5 guarding article
6; uropod 2 reaching end of uropod 38. Species: 3, subantarctic,
boreal, littoral.

MARINE GAMMARIDEAN AMPHIPODA 387

Synchelidium Sars
Synchelidium Sars, 1895.—Stebbing, 1906.

Type-species: Kroyeria haplocheles Grube, 1864b (selected by
Chevreux and Fage, 1925). See Sars, 1895 (as S. brevcarpum).

Cutting edge of mandible projecting, toothed, molar small, conical,
with apical spine, unridged; inner lobes of lower lip flat but separate;
enathopods dissimilar, gnathopod 1 subchelate, stout, article 5 form-
ing a lobe guarding article 6, gnathopod 2 slender, chelate, article 5
unproduced ;* uropod 2 reaching end of uropod 3. Species: 7, boreal,
littoral to bathyal.

*Possibly a long lobe of article 5 has been amalgamated with the posterior
margin of article 6.

Westwoodilla Bate

Westwoodia Bate, 1857a (homonym, Hymenoptera).
Westwoodilla Bate, 1862 (new name).—Stebbing, 1906.
Halimedon Boeck, 1871.

Type-species: Westwoodia caecula Bate, 1857a (monotypy). See
Sars, 1895 (as Halimedon mullert).

Cutting edge of mandible poorly produced, poorly toothed, molar
large, ridged; ‘‘article 2 of palp more strongly curved than in Bathy-
medon”; inner lobes of lower lip separate; gnathopods similar to
each other, feeble, subchelate, article 5 scarcely lobed; uropod 2
reaching end of uropod 3. Species: 9, arctic, warm-temperate, littoral
to abyssal.

Pagetinidae
FicurE 139

Diacnosis.—Accessory flagellum absent; mandibular molar eva-
nescent; coxae short, all visible; outer lobes of maxillipeds poorly
developed, inner obsolete; uropod 3 uniramous; telson short, entire;
urosomites 2-3 coalesced. See Stenothoidae, Thaumatelsonidae,
Sebidae, Phliantidae, Cressidae, Kuriidae.

Description.—Accessory flagellum absent; urosomites 2-3 coa-
lesced; coxae short, all visible; upper lip probably incised; mandible
with 3-articulate palp, molar evanescent; lower lip as in Pleustidae,
with tilted, oval outer lobes astride coalesced inner lobes; maxilla
1 weak, with 1-articulate palp, inner lobe possibly absent, outer
weakly setose! or spinose; maxillipeds slender, outer lobes small,
inner lobes vestigial; gnathopods small, subchelate; uropod 3 uni-
ramous, telson short, entire.

388 U.S. NATIONAL MUSEUM BULLETIN 271

RELATIONSHIP.—The Stenothoidae, Cressidae, and Thaumatel-
sonidae have enlarged coxae 2-4, more elongate telsons than do
Pagetinidae and coxa 1 is small and hidden by the following coxae.

The Pardaliscidae have biramous third uropods.

The Eophliantidae and Phliantidae lack mandibular palps, the
former having cylindrical bodies, the latter having depressed bodies.

The Pagetinidae bear some resemblance to Sebidae in mouthparts,
uropods, telson, and coxae but Sebidae have chelate gnathcopods,
elongate third artcles of gnathopod 2 (with one exception) and
well-formed plates of the maxillipeds.

Key to the Genera of Pagetinidae

These genera, possibly synonymous, are based on characters which
may be conditions due to different methods of preservation, mounting
technique, and observation.

0

Ficure 139.—Pagetinidae: a, head and gnathopod 1 of Pagetina genarum K. H. Barnard
(1932). Upper lip: b, Heterocressa monodi Nicholls (1938). Mandible: c, Heterocressa.
Maxilla 1: d, Pagetina, e, Heterocressa. Maxilliped: f, Pagetina; g, Heterocressa. Pereo-
pods 1,3,5: h,j,k, Pagetina. Gnathopod 2: 1, Pagetina. 1, Pagetina, dorsal view of tel-
son and uropod 3. Uropod 3: m, Heterocressa. Telson: n, Heterocressa. Lower lip:
o, Pagetina.

MARINE GAMMARIDEAN AMPHIPODA 389

PU ToOpod ao withi2-arviculave ramus ely). 15) ee yes aes Heterocressa
Wropodisawithyl-articulate ramus 4) ote he es Pagetina

Genera of Pagetinidae

Heterocressa Nicholls
Heterocressa Nicholls, 1938.

Type-species: HH. monodi Nicholls, 1938 (original designation).
Species: 1, Macquarie Island, littoral.

Pagetina K. H. Barnard
Pagetina K. H. Barnard, 1931; 1932.

Type-species: P. genarum K. H. Barnard, 1931 (original designa-
tion) ; 1932.
Species: 1, S. Georgia Island, littoral.

Paramphithoidae
{including Amathillopsidae Pirlot, 1934]

Figures 140-142, rroNTISPIECE

Dracenosis.—One or more of coxae 1-4 (usually all) acutely pointed
midventrally, occasionally bifid, accessory flagellum vestigial or
absent, at most composed of two articles. See Acanthonotozomatidae,
Lepechinellidae, Pleustidae, Calliopiidae (especially Oradarea), Eu-
siridae, Haustoriidae, Astyridae.

DescriptTion.—Rostrum usually well developed; body often pro-
cessiferous; accessory flagellum vestigial or absent; coxae large, 1-4
usually acuminate, or coxae slightly excavate ventrally to form weak
anterior-posterior points (weak bifidation); mouthparts variable,
mandible always with molar (except Bathypanoploea) and 3-articulate
palp, molar varying in extent and degree of trituration surface; lower
lip with or without inner lobes; plates of maxillipeds well developed
but palp varying from three to four articles; gnathopods feeble, either
subchelate or simple; uropod 3 with elongate lanceolate rami; telson
entire or partially cleft, slightly elongate or not.

Rewationsuip.—Most Acanthonotozomatidae, also with pointed
coxae, differ from Paramphithoidae by the lack of a distinct mandi-
bular molar and the conically grouped mouthparts. The genus Bathy-
panoploea (Acanthonotozoma australis Chilton, 1912) in the Acan-
thonotozomatidae does not have conically grouped mouthparts and so

390

U.S. NATIONAL MUSEUM BULLETIN 271

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ro.
re

=o, aad
Soper

Ficure 140.—Paramphithoidae: a, Actinacanthus tricarinatus (Stebbing, 1888);
b, Amathillopsis spinigera Heller (Sars, 1885).

MARINE GAMMARIDEAN AMPHIPODA 391

Ficure 141.—Paramphithoidae: a, Epimeria cornigera (Fabricius) (Sars, 1895, pl. 128);
b, Paramphithoe hystrix (Ross) (Sars 1895 pl. 130. as Acanthozone cuspidata).

392 U.S. NATIONAL MUSBUM BULLETIN 271

confounds the distinctions between the two families. I include that
genus also in the key of Paramphithoidae although retaining Schellen-
berg’s (1931, 1939) assignment of it to the Acanthonotozomatidae.

A few paramphithoids like Amathillopsis grevei J. L. Barnard (1961)
have very weak acuminations on coxae 1—4 and might be keyed into
the Calliopiidae. Clerppides of the latter family may also have sub-
acuminate coxae.

The Lepechinellidae differ from Paramphithoidae mainly by the
slender second articles of pereopods 3-5, the possession of coalesced
inner lobes on the lower lip, and apparently the fusion of pleonites
5 and 6.

The genus Amathillopleustes Pirlot (1934) described in the Amathil-
lopsidae was preoccupied by Cleonardopsis K. H. Barnard (1916).
Since this genus bears rounded coxae it cannot be retained in the
Paramphithoidae (=Amathillopsidae); it bears an elongate 1-artic-
late accessory flagellum and cleft telson, hence should be transferred
to the Eusiridae. Note that the Eusiridae and Pontogeneiidae have
been amalgamated.

The Pleustidae are very closely related to Paramphithoidae but have
characteristic lower lips. Mesopleustes is an intermediate genus because
of its weakly pleustid lower lip and tendency to coxal acumination.

Haustoriids have a well-developed accessory flagellum and fossorial
appendages.

Eusiridae lack fully acuminate coxae. Rhachotropis, however,
intergrades Eusiridae and Paramphithoidae.

Oradarea Walker, originally described in the Calliopiidae but
transferred to Paramphithoidae by Pirlot (1934) usually lacks the
pointed coxae of paramphithoids although male O. shoemaker Pirlot
has slightly acuminate coxae, Oradarea should be returned to the
Calliopiidae. Parepimeriella Schellenberg (1931) also has rounded,
untapering coxae, bears the lower lip characteristic of Pleustidae, and
should be transferred to that family.

Eelysis K. H. Barnard (1932) and Epimeriella Walker (1907)
probably should be assigned to the Astyridae and the Paramphithoidae
amended to include only genera with triturative molars. Helysis is
particularly close to Astyra, differing from the latter in the absence
of an accessory flagellum.

One species of Gammaridae, Weyprechtia heuglini (Buchholz), has
coxae 4-5 forming a ventral crescentic curve like some paramphithoids
but that genus may be distinguished in its multiarticulate accessory
flagellum.

MARINE GAMMARIDEAN AMPHIPODA 393
Nomenclatural Changes in Paramphithoidae

Amathillopsis Heller (1875) is removed from Gammaridae and
Amathillopsidae and assigned to Paramphithoidae, thus resulting
in amalgamation of Amathillopsidae with Paramphithoidae.

Amathillopleustes Pirlot (1934) is a synonym of Cleonardopsis K. H.
Barnard (1916) and they are removed to the Eusiridae.

Oradarea Walker (1903) is removed to the Calliopiidae.

Parepimeriella Schellenberg (1931) is removed to the Pleustidae.

Key to the Genera of Paramphithoidae

1. Body densely covered with articulate spines (see ee . Uschakoviella

Body lacking articulate’spines . °°) 2 2455. 2.4 5. 52

2. Molar of mandible absent .... St ane iathypanaploes i
Molar of mandible only a setose evnerirn or cone Ge. LADD) es cre nea eS
Molar of mandible well developed, large, ridged (fig. 1427)... .....4

3. Pereopod 5 as long as or longer than pereopod4...... ... . Eclysis
Pereopod 5 shorter than pereopod4 ... . . . . Epimeriella

4. Maxillipedal palp bearing only three articles 2 Gre 1429) eee aeRO di WRES 5
Maxillipedal palp bearing four articles (fig. 142m). ...........6

5. Lower lip with inner lobes (fig. 1420)... .... .. . . Parepimeria ?
Lower lip lacking inner lobes (fig. 142k). . . . A . . . Metepimeria

6. Coxae 4-5 together forming a crescentic curve Below, Ge ATC) ea Malis
Coxae 4—5 together not forming a crescentic curve below (fig. 1406) . . .8

7. Gnathopods subchelate, dactyl slender, simple (fig. 142b) . . . Epimeria ‘4
Gnathopods simple, dactyl stout, spinose as De a ERA is ree s

8. Accessory flagellum present .. . aes ts SUSAN NE Naa oi Attn er oe USE)
Accessory flagellum absent ... . i oer sir’ 10

9. Accessory flagellum long and spiniform, aathopods Bronely subche-
MACE len . . . . Amathillopsis
Accessory dpe Teta hor, aeolteliee or earl like, gnathopods simple
even though portions occasionally stout. . . . .. . . . Parepimeria

10. Article 5 of gnathopods shorter than article6 .. . . . . Paramphithoe
Article 5 of gnathopods much longer than article6 ... . Actinacanthus

1 Assigned to Acanthonotozomatidae provisionally.

2 Chevreux (1912b).

3K. H. Barnard (1932) states maxillipedal palp article 4 is present.

4 Note resemblance in coxae 4-5 of Weyprechtia (Gammaridae).

5 See also Halirages stebbingi Schellenberg (1931) (=H. huzleyanus Stebbing, 1888, not Bate), in the Callio-
piidae, a species with acuminate anterior coxae; it differs from paramphithoids and other members of Hali-
rages in its stout gnathopods with ovate articles.

Genera of Paramphithoidae

Actinacanthus Stebbing

Acanthechinus Stebbing, 1888 (homonym, Echinodermata).
Actinacanthus Stebbing, 1906 (new name).

394 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: Acanthozone tricarinata Stebbing, 1883 (monotypy).

Accessory flagellum absent; mandibular molar large, ridged; lower
lip lacking inner lobes; maxillipedal palp 4-articulate; gnathopods
poorly subchelate, articles 5 and 6 slender and very elongated,
article 5 longer than 6 (Paramphithoe); telson entire. Species: 1,
S. Indian Ocean (282 m).

Amathillopsis Heller

Amathillopsis Heller, 1875.—Stebbing, 1906.—Pirlot, 1934—J. L. Barnard,
1964c.
Acanthopleustes Holmes, 1908.

Type-species: A. spinigera Heller, 1875 (original designation).
See Sars, 1885; Gurjanova, 1951.

Accessory flagellum spine-like, 2-articulate, article 2 tiny; mandib-
ular molar large, ridged; lower lip apparently with coalesced inner
lobes; maxillipedal palp 4-articulate; gnathopods subchelate, articles
5 and 6 ovoid, article 5 with posterior lobe; telson slightly emarginate.
Species: 7, cosmopolitan, (cold littoral) bathyal and abyssal.

Eclysis K. H. Barnard
Eclysis K. H. Barnard, 1932.

Type-species: FH. sumilis K. H. Barnard, 1932 (monotypy).

Accessory flagellum absent; mandibular molar a setulose conical
lamina; lower lip lacking inner lobes; maxillipedal palp 4-articulate;
genathopods scarcely subchelate, article 5 with small posterior lobe;
pereopod & equal to or longer than pereopod 4 (Epimeriella); telson
cleft halfway. Species: 1, S. Georgia Islands, littoral.

Epimeria Costa

?Vertumnus White, 1847a (nomen nudum).
Epimeria Costa, 1851b.—Stebbing, 1906.

Type-species: Gammarus corniger J. C. Fabricius, 1779 (mono-
typy and subsequent synonymy). See Sars, 1895.

Accessory flagellum absent; mandibular molar large, ridged; lower
lip lacking inner lobes; maxillipedal palp 4-articulate; gnathopods
subchelate, dactyl slender, simple (Pseudepimeria); telson emarginate
or notched; cozae 4-5 together forming a crescentic curve below (com-
bining character). Species: 19, cosmopolitan cold-water, littoral to
abyssal.

MARINE GAMMARIDEAN AMPHIPODA 395

Epimeriella Walker
Epimeriella Walker, 1906b; 1907.

Type-species: EH. macronyx Walker, 1906b; (monotypy) 1907.

Accessory flagellum absent; mandibular molar a setose conical
lamina; [lower lip unknown]; maxillipedal palp 4-articulate; gnatho-
pods subchelate, short article 5 with slight posterior lobe; pereopod
5 shorter than pereopod 4 (Eelysis); telson incised. Species: 3, ant-
arctic, littoral to bathyal.

Ficure 142.—Paramphithoidae: Gnathopod 1: a, Parpimeria crenulata Chevreux (1912b);
b, Paramphithoe hystrix (Ross) (Sars, 1895, pl. 130, as Acanthozone cuspidata); c, Eclysis
similis K, H. Barnard (1932); d, Actinacanthus tricarinatus (Stebbing, 1888). Gnathopod
2: e, Pseudepimeria grandirostris Chevreux (1912b); f, Paramphithoe; g, Actinacanthus.
Uropod 3: h, Paramphithoe. Mouthparts, Paramphithoe: i, upper lip; j, mandible;
k, lower lip; /jm, maxillae 1, 2; 7, maxilliped. Lower lip: 0, Parepimeria. Mandible:
p, Eclysis. Maxilliped: g, Parepimeria. Telson: r, Paramphithoe; s, Epimeria loricata
Sars (1895, pl. 129); ¢, Eelysis.

285-135 O - 69 - 26

396 U.S. NATIONAL MUSEUM .BULLETIN 271

Metepimeria Schellenberg
Metepimeria Schellenberg, 1931.

Type-species: M. acanthurus Schellenberg, 1931 (monotypy).

Accessory flagellum absent; mandibular molar large, ridged;
lower lip lacking inner lobes; maxillipedal palp 3-articulate; gnatho-
pods simple; telson deeply notched. Species: 1, antiboreal, littoral.

Paramphithoe Bruzelius

Acanthosoma Ross, 1835 (homonym, Hemiptera).
Paramphithoe Bruzelius, 1859.—Stebbing, 1906.
Acanthozone Boeck, 1871.

Type-species: Acanthosoma hystrix Ross, 1835 (selected by Bate,
1862, p. 377!; possible synonym of Oniscus cuspidatus Lepechin,
1780, fide Gurjanova, 1951; et al).

Accessory flagellum absent; mandibular molar large, ridged;
lower lip lacking inner lobes; maxillipedal palp 4-articulate; gnatho-
pods subchelate, article 5 shorter than article 6 (Actinacanthus) ; telson
truncate. Species: 4, arctic, littoral to bathyal.

Parepimeria Chevreux
Parepimeria Chevreux 1911c; 1912a; 1912b.

Type-species: P. crenulata Chevreux, 1912a; 1912b (designated
by Chevreux, 1912a).

Accessory flagellum 1-articulate; mandibular molar large, ridged;
lower lip with inner lobes; maxillipedal palp 3-articulate (or 4-articu-
late: K. H. Barnard, 1932); gnathopods simple, articles 5 and 6
slightly elongate; telson entire. Species: 3, antarctic, littoral to
bathyal.

Pseudepimeria Chevreux
Pseudepimeria Chevreux, 1912a; 1912b.

Type-species: P. grandirostris Chevreux, 1912a (original designa-
tion) ; 1912b.

Accessory flagellum 1-articulate; mandibular molar large, ridged;
lower lip lacking inner lobes; maxillipedal palp 4-articulate; gnatho-
pods simple, dactyl stout, spinose (Epimeria); telson cleft; corae 4-5
together forming a crescentic curve below (combining character). Species:
1, antarctic, bathyal.

MARINE GAMMARIDEAN AMPHIPODA 397

Uschakoviella Gurjanova
Uschakoviella Gurjanova, 1955.

Type-species: U. echinophora Gurjanova, 1955 (original designa-
tion). See Shoemaker, 1964.

Body covered with articulate spines; accessory flagellum absent;
lower lip lacking inner lobes; mandibular molar large, ridged; maxilli-
pedal palp 4-articulate; gnathopods scarcely subchelate, but palms
transverse; telson cleft slightly. Species: 1, boreal N. Pacific,
littoral to abyssal.

Pardaliscidae
Fiaures 1438, 144

Draenosis.—Mandible lacking molar; inner plates of maxillipeds
small or vestigial; coxae all short; accessory flagellum present, well
developed, often sexually dimorphic. See Stilipedidae, Acantho-
notozomatidae.

Derscription.—Accessory flagellum multiarticulate, often sexually
dimorphic (“lacking’”’ in one genus), base of primary flagellum often
conjoint in male; rostrum usually prominent, head occasionally like
that of Pseudotiron in Synopiidae; upper lip incised or rounded; man-
dibles flat, almost elytriform; molar of mandible absent, palp 3-
articulate or absent, article 3 often shortened; inner plates of max-
illipeds evanescent, outer plates occasionally small; coxae very short;
enathopods powerful or feeble; rami of uropod 3 elongate, lanceolate
or subfoliaceous; telson rarely elongate but often of medium length,
deeply cleft or entire; mouthparts grouped quadrately (except coni-
cally in Halicella).

ReELationsHip.—The Stilipedidae differ from the Pardaliscidae in
the complete absence of an accessory flagellum. Halicoides, assigned to
Pardaliscidae, also lacks an accessory flagellum (but see Birstein and
Vinogradov, 1960), bears a peculiar process on article 2 of antenna 1
but has other aspects of the Pardaliscidae. Both pairs of maxillae in
stilipedids are highly foliaceous, unlike those of pardaliscids.

Halicella of the Pardaliscidae has its mouthparts formed into a
conical bundle like the Acanthonotozomatidae but its coxae are short,
and the inner plates of its maxilliped are absent. Acanthonotozomatids
have long coxae and well-developed inner plates of the maxillipeds,
as do Astyridae.

Laphystiopsidae have well-developed inner plates of the maxilliped,
a distinct mandibular molar and lack an accessory flagellum.

398 U.S. NATIONAL MUSEUM BULLETIN 271

Ficure 143.—Pardaliscidae: a, Pardalisca cuspidata Krgyer (Sars, 1895, pl. 141); b, Nicippe
tumida Bruzelius (Sars, 1895, pl. 144). Upper lip: c, Pardalisca; d, Nicippe. Accessory
flagella, Pardalisca: e, female; f, male. Mandibles: g,h, right and left, Pardalisca; 1, Halice
abyssit Boeck (Sars, 1895, pl. 145). Maxilla 2:7, Necochea pardella J. L. Barnard (1962d).
Antenna 1, peduncle: k, Pardaliscoides tenellus Stebbing (1888). Telson: /, Parpano
cebus J. L. Barnard (1964a).

MARINE GAMMARIDEAN AMPHIPODA 399

Revisionary Notes on Pardaliscidae

Halicoides Walker poses a problem because pleonites 4—5 of the
type-specimen of the type-species were injured and never clarified. It
was described as lacking an accessory flagellum but bearing a unique
scale on article 2 of antenna 1. Presumably these are also injuries. If

Ficure 144.—Pardaliscidae: a, Pardisynopia tambiella J. L. Barnard (1961). Lower lip:
b, Pardaliscella boecki (Malmgren) (Sars, 1895, pl. 143); c, Pardalisca cuspidata Krgyer
(Sars, 1895, pl. 141). Pardalisca: d,e, maxillae 1-2; f, maxilliped. Maxilliped: g, Halicella
parasitica Schellenberg (1926a). Gnathopod 1: h, Pardalisca; 1, Pardaliscella; 7, Nicippe
tumida Bruzelius (Sars, 1895, pl. 144). Gnathopod 2: k, Pardalisca; l, Nicippe; m,
Pardaliscella. Telson: n, Pardalisca.

400 U.S. NATIONAL MUSEUM BULLETIN 271

an accessory flagellum does exist and if pleonites 4—5 do bear large
teeth then the genus would be referable to Halice. Meanwhile it is
treated as distinct in the following key. Birstein and Vinogradov
(1960, 1962, 1964) have referred their species, Halice nana and H.
indica, to Halicoides; and Synopioides tertia Stephensen (1931) to
Halice (see Birstein and Vinogradov, 1962) but suggest the latter also
might be assigned to Halicoides. A different point of view seems more
reasonable until some of these species, including Halice anacantha
K. H. Barnard (1925) have been more thoroughly described. Birstein’s
and Vinogradov’s (1962) diagnosis of Halice is reasonable, except that
they include Halice tertia and H. anacantha which spoil the definition,
because H. tertia does not have a conjoint article 1 of the primary
flagellum of antenna 1, and both HZ. tertia and H. anacantha have
obsolescent teeth on pleonites 4—5. The flagellum of H. anacantha is
unknown except for K. H. Barnard’s statement that the complete
antenna resembles that of Halice abyssi. If these two aberrant species
were removed from Halice the diagnosis could be strongly tightened
and should be so done as a provisional measure until further light can
be shed by redescriptions and discovery of expected new abyssal
pardaliscids. Undoubtedly H. tertia is congeneric with Pardisynoma
tambiella J. L. Barnard (1961) and P. synopiae J. L. Barnard (1962b)
but scarcely with Halicoides anomalus (type) if conjointing of the
primary first antennal flagellum and obsolescence of pleonal teeth are
diagnostic. Thus, Pardisynopia is retained as a genus distinct from
Halice and Halicoides which have conjoint flagella. Halice anacantha
K. H. Barnard is temporarily removed to Pardisynopia until the
condition of antenna 1 is verified. Halicoides indica and H. nana also
are provisionally assigned to Pardisynopia, even though uropod 3 of
H. nana appears to lack article 2 on the outer ramus.

Key to the Genera of Pardaliscidae

Tes Melson centinen(oaVASh ee en Sale Mera A ute 2c eso rey Seiiee pease Parpano
Telsondeeply cleft: (fig.;144n)). 6). a 2
2. Articles 4-5 of pereopods 1-2 inflated (fig. 144a) .........2.. 3
Articles 4-5 of pereopods 1-2 not inflated (fig. 143a) ......... 5

3. Antenna 1 lacking accessory flagellum (or bearing scale on article 2).
Halicoides
Antenna lbearing- accessory flagellum <) 20.) 2) 59k) eee 4

4, Antenna 1: basal part of flagellum inflated or narrow, but segmented (fig.
148¢), flagellar article 1 scarcely longer than article 3 of peduncle, urosomal

teeth vestiovall si. ayy 28h org a in Sia aaa aie a ae eect Pardisynopia
Antenna 1: basal part of flagellum inflated, conjoint ! (fig. 143f), thus flagellar
article 1 nearly as long as peduncle, urosomal teeth strong . . . Halice

1 Incompletely segmented.

MARINE GAMMARIDEAN AMPHIPODA A401

5. Mouthparts forming a cone-like bundle below head ... . . . Halicella
Mouthparts not forming a cone-like bundle below head . Baines 6

6. Article 6 of both gnathopods 1 and 2 much longer hen ant fale 5, Erticle 5
with long narrow posterior lobe (fig. 1447) . . . . . . . Nicippe
Article 5 of either gnathopods 1 or 2 subequal to or loner aha 6, but posterior
lobe of article 5 broad and shallow if present . . year iNareesil 5c Sa

7. Antenna 1 with elongate peduncular article 2 (fig. 143h) . . Pardaliscoides
Antenna 1 with short peduncular article 2 (fig. 148a) ......... 8

8. Mandible lacking palp, pereopods prehensile ..... ... . Parahalice
Mandible with palp, pereopods not prehensile ............49

Ome Maxalla; 2 obsolescent Gig. 1439), 0. be. ee ~Necochea
Maxilla 2 well developed (fig. 144e) . ... esata ater] 6)

10. Article 5 of gnathopods much longer than 6 Ga 144h iy states amrgetuaee dL
Articles 5 and 6 of gnathopods subequal, or article 6 slightly longer than 5
(figs. 1442,m) . . . Aa Iga cun U7

11. Gnathopodal dactyls aorainallly leer shaped: nor eamece Gea 1443,)), maxilli-
pedal palp more than 3 times as long as medial edge of outer plate.

Princaxelia

Gnathopodal dactyls short, spinose (figs. 144h,k), medial edge of outer plate on
maxilliped nearly as long as palp (fig. 144f) . . .. . . . . Pardalisca

12. Upper lip grossly incised asymmetrically (fig. 143c) . . . . Pardaliscopsis
Upper lip nearly symmetrically incised (fig. 143d). ......... 18

13. Antenna 1: basal part of flagellum narrow, segmented (fig. 143e), flagellar
article 1 scarcely longer than article 3 of peduncle . . . . Pardaliscella
Antenna 1: basal part of flagellum inflated, unsegmented (fig. 143f), flagellar
articlemlmearlydasilongiasspeduncleynane tary ss et a er yaks:

A enGnathoOpod 2ssimple cry soi es wc a Wen ae eee ou en ene alice
Gnathopod:2sulbchelate 7 os ioe 8 wiser ous oe eee Se Areulfia

Genera of Pardaliscidae

The upper lip and lower lip may have important generic characters
but they have not been adequately described for most species. They
are included when possible in the generic diagnoses to follow.

Arculfia J. L. Barnard

Arculfia J. L. Barnard, 1961.

Type-species: A. trago J. L. Barnard, 1961 (original designation).

Article 2 of antenna 1 shorter than article 1; upper lip rounded
below; inner lobes of lower lip coalesced; gnathopod 1 simple, articles
5 and 6 subequal, article 5 not lobed, gnathopod 2 subchelate, dactyls
simple; telson deeply cleft. Species: 1, Tasman Sea (610 m).

Halice Boeck

Halice Boeck, 1871.—Stebbing, 1906.—Birstein and Vinogradov, 1962.
Synopiordes Stebbing, 1888; 1906.

402 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: H. abysst Boeck, 1871 (selected by Boeck, 1876).
See Sars, 1895.

Article 2 of antenna 1 shorter than article 1, article 1 of primary
flagellum as long as or longer than peduncular article 1 (Pardisynopia) ;
[upper and lower lips not described]; gnathopods simple, article 6
slender, longer than 5, article 5 not lobed, dactyls simple or rarely
half spinose (Hl. macronyz); urosomal teeth strong (Pardisynopia);
telson cleft 40 percent or more; head usually normal in appearance,
with projecting lateral lobes and narrow rostrum but occasionally formed
into blunt anterior ‘‘shark nose” condition like Pseudotiron (Synopiidae).
Species: 11, cosmopolitan, bathy-abyssopelagic.

Halicella Schellenberg
Halicella Schellenberg, 1926a.

Type-species: H. parasitica Schellenberg, 1926a (monotypy).

Mouthparts forming a cone-like bundle below head; article 2 of antenna
1 shorter than article 1; [upper lip and lower lip not described];
enathopods simple, article 6 longer than 5, 5 not lobed, dactyls
simple; telson deeply cleft. Species: 1, antarctic, littoral.

Halicoides Walker
Halicoides Walker, 1896.—Stebbing, 1906.

Type-species: H.anomala Walker, 1896 (monotypy). See Chevreux
and Fage, 1925.

Antenna 1 lacking accessory flagellum (but see Birstein and Vindo-
gradov, 1960), article 2 shorter than article 1; [upper lip and lower
lip not described]; gnathopods simple, articles 5 and 6 subequal,
dactyls simple; telson deeply cleft. Species: ?1+, Bay of Biscay,
littoral (possibly also tropical-bathypelagic). See ‘‘Revisionary Notes
on Pardaliscidae.”’

Necochea J. L. Barnard
Necochea J. L. Barnard, 1962d.

Type-species: WN. pardella J. L. Barnard, 1962d (original designation).

Article 2 of antenna 1 shorter than article 1; [upper and lower lip
not described]; mazilla 2 obsolescent; gnathopods simple, article 6
shorter than 5, 5 not lobed, dactyls with a few setules; telson deeply
cleft; coxa 5 with nearly twice as much surface area as other coxae.
Species 1, subantarctic, abyssal.

MARINE GAMMARIDEAN AMPHIPODA 403

Nicippe Bruzelius
Nicippe Bruzelius, 1859.—Stebbing, 1906.

Type-species: N. tumida Bruzelius, 1859 (monotypy). See Sars,
1895.

Article 2 of antenna 1 shorter than article 1; upper lip slightly
bilobed; lower lip with coalesced inner lobes; gnathopods slightly
subchelate, articles 5 and 6 stout, article 6 longer than 5, 5 with a
large posterior lobe, dactyls simple; telson deeply cleft. Species:
2, bipolar, bathyal.

Parahalice Birstein and Vinogradov
Parahalice Birstein and Vinogradov, 1962.

Type-species: P. mirabilis Birstein and Vinogradov, 1962 (orig-
inal designation).

Article 2 of antenna 1 shorter than article 1; mandible lacking palp;
[upper lip and lower lip not described]; gnathopods simple, article 6
slightly longer than 5, 5 not lobed, dactyls simple; all pereopods pre-
hensile; telson cleft one third. Species: 1, central Pacific, bathyal
or abyssopelagic.

Pardalisca Kr¢éyer
Pardalisca Krgyer, 1842.—Stebbing, 1906.

Type-species: P. cuspidata Kroyer, 1842 (monotypy). See Sars,
1895.

Article 2 of antenna 1 shorter than article 1; upper lip very asym-
metrically lobed; lower lip with inner lobes separate; gnathopods
simple, article 6 much shorter than article 5, dactyls spinose; telson
deeply cleft; medial edge of inner plate on mazilliped nearly as long as
palp (Princazelia). Species: 7, bipolar, littoral to bathyal.

Pardaliscella Sars
Pardaliscella Sars, 1895.—Stebbing, 1906.

Type-species: Pardalisca boeckii Malm, 1871 (monotypy).

Article 2 of antenna 1 shorter than article 1, base of flagellum seg-
mented (combining character); upper lip weakly incised and slightly
asymmetrical; lower lip with inner lobes coalesced; gnathopods
simple, article 6 slightly shorter than 5, dactyl with one spine; telson
cleft only halfway; [rostrum obsolete]. Species: 5, boreal, littoral to
abyssal.

404 U.S. NATIONAL MUSEUM BULLETIN 271

Pardaliscoides Stebbing
Pardaliscoides Stebbing, 1888; 1906.

Type-species: P. tenellus Stebbing, 1888 (monotypy). See Stebbing,
1897.

Article 2 of antenna 1 longer than article 1; [upper and lower lips
not described]; gnathopods simple, article 6 shorter than 5, 5 not
lobed, dactyls with tiny setules; telson deeply cleft. Species: 2, N.
Atlantic, S. Pacific, bathyal to hadal.

Pardaliscopsis Chevreux
Paradaliscopsis Chevreux, 1911b.

Type-species: P. tenuipalpa Chevreux, 1911b (original designa-
tion).

Article 2 of antenna 1 shorter than article 1; upper lip very asym-
metrically lobed; lower lip with coalesced inner lobes; gnathopods
simple, articles 5 and 6 subequal, 5 not lobed, dactyls lacking spines;
telson deeply cleft. Species: 1, N. Atlantic, abyssal.

Pardisynopia J. L. Barnard
Pardisynopia J. L. Barnard, 1961.

Type-species: P. tambiella J. L. Barnard, 1961 (original designa-
tion).

Article 2 of antenna 1 shorter than article 1; article 1 of primary
flagellum half or less as long as peduncular article 1 (Halice and Hali-
coides); upper lip scarcely incised below; lower lip with coalesced
inner lobes; gnathopods simple, article 6 longer than 5 or subequal,
5 searcely lobed posteriorly on gnathopod 1, dactyls simple; pleonites
4-5 with teeth obsolescent (Halice); telson deeply cleft. Species: 6 (3
provisional), probably cosmopolitan, bathyal.

Parpano J. L. Barnard
Parpano J. l. Barnard, 1964a.

Type-species: P. cebus J. L. Barnard, 1964a (original designation).

Article 2 of antenna 1 equal to or shorter than article 1; upper lip
symmetrically lobed; lower lip apparently with coalesced inner lobes;
enathopods simple, article 6 much longer than 5, latter obsolescent,
dactyls simple; telson entire; uropod 3 much reduced in size. Species:
2, Caribbean, abyssal.

MARINE GAMMARIDEAN AMPHIPODA 405

Princaxelia Dahl
Princaxelia Dahl, 1959.

Type-species: P. stephensent Dahl, 1959 (original designation).

Article 2 of antenna 1 shorter than article 1; upper lip scarcely
incised, lobes slightly asymmetrical; [lower lip unknown]; mazillipedal
palp more than three times as long as medial edge of outer plate (Par-
dalisca); gnathopods simple, article 5 longer than 6, dactyls simple;
telson deeply cleft. Species: 2, N. Atlantic, S. W. Pacific, bathyal
to hadal.

Phliantidae, revised

[see J. L. Barnard, 1964c]

Ficures 145-147

Draqnosis.—Accessory flagellum absent; body dorsally depressed,
at least anterior coxae splayed; mandible lacking palp, molar degraded;
palp of maxilla 1 degraded or absent; gnathopods feeble; usually
uropod 3 lacking inner ramus, often lacking both rami. See Prophli-
antidae, Eophliantidae, Kuriidae, Dogielinotidae, Talitroidea, Dex-
aminidae, Pagetinidae, Acanthonotozomatidae, Colomastigidae,
Cressidae.

Description.—Phliantidae are depressed dorsally, except Phlias
which is “‘compressed”’ (Stebbing, 1906), with pleon flexed below re-
mainder of body, segments armored and coxae projecting laterally;
accessory flagellum absent; mandible lacking palp, molar absent or
represented by a smooth protuberance or spine; lower lip with or
without inner lobes; maxilla 1 with degraded palp, 1-articulate or
represented by a spine when present; maxilla 2 small, often with
coalesced lobes; plates of maxilliped well developed, palp 2- to 4-
articulate; gnathopods feeble, usually simple, but subchelate in genera
Plioplateia and Ceina, article 3 of gnathopod 2 elongate; uropod 1
biramous (except Temnophlias), uropod 2 usually biramous (except
Temnophlias and Pereionotus), uropod 3 uniramous in most genera
or rami absent; telson usually entire (or cleft in Cena).

Some students, particularly Chevreux (1911a) have considered that
uropod 2 is missing in Pereionotus and that the last biramous uropod
in that genus is uropod 3. However, Pereionotus as stated by Stebbing
(1906) probably has a small uropod 3 lacking rami, perhaps over-
looked by Chevreux. Uropod 3 has not been clearly described in some
genera (especially Ceina).

Rexatronsuip.—The Prophliantidae have biramous third uropods,
a cleft telson, and often have a degraded flagellum on antenna 2.

406 U.S. NATIONAL MUSEUM BULLETIN 271

Ceina bridges the Phliantidae and Prophliantidae in its subchelate
gnathopods and cleft telson but is retained in Phliantidae because
of its nonbiramous third uropods and presence of a flagellum on
antenna 2,

Figure 145.—Phliantidae: Heterophlias seclusus Shoemaker (1933a): a, dorsal view;
b, lateral view.

The Podoceridae have a mandibular palp and molar. The Tali-
troidea have well-developed triturating mandibular molars (except
for Najna which otherwise has the talitroid facies).

The Kuriidae have a well-developed triturating molar and poorly
developed outer plates of the maxilliped. The Dexaminidae, less
Anatylinae, have well-developed triturating molars. The palp of max-
illa 1 is normal in all dexaminids and segments 2-3 of the urosome
are coalesced.

Ficure 146.—Phliantidae and Calliopiidae: a, Ceina carinata (Pirlot, 1936, as
Periphlias carinatus). Upper lip: b, Pereionotus testudo (Montagu) (Chevreux and Fage,
1925). Lower lip: c, Heterophlias seclusus Shocmaker (1933a). Maxilla 2: d, Pereionotus;
e, Heterophlias. Maxilla 1: f, Heterophlias; g, Plioplateia triquetra K. H. Barnard (1916).
Mandible: h, Heterophlias. Body, dorsal: 1, Sancho platynotus Stebbing (1897) [Callio-
piidae]; 7, Temnophlias capensis K. H. Barnard (1916); », T. hystrix K. H. Barnard (1954).
Maxilliped: 7, Sancho; m, T. capensis. Telson and uropod 3: k, Sancho.

408 U.S. NATIONAL MUSEUM BULLETIN 271

The Acanthonotozomatidae have conically bundled mouthparts,
normal uropod 3, and a mandibular palp.

Gnathopod 2 of some phliantids faintly resembles that of Lysi-
anassidae because of the elongate article 2.

The Temnophlias-Sancho Problem

Gurjanova (1962) discussed at length the interrelationships of
Sancho Stebbing, Chosroes Stebbing, and Temnophlias K. H. Barnard
and their relationships to the Phliantidae. Sancho and Chosroes have
been placed in or near the Calliopiidae despite the clear development of
what Gurjanova calls a dorsal shield which is another way of denoting
their extreme dorsoventral depression. They are even more depressed
than typical phliantids. Gurjanova develops the Phliantidae from

Ficure 147.—Phliantidae: Gnathopod 1: a, Plioplateia triquetra K. H. Barnard (1916);
b, Ceina carinata (Pirlot, 1936 as Periphlias carinatus); c, Heterophlias seclusus
Shoemaker (1933a). Gnathopod 2:d, Ceina. Maxilliped: ¢, Heterophlias; f, Temnophlias
capensis K. H. Barnard (1916); g, Pereionotus testudo (Montagu) (Chevreux and Fage,
1925). Pleopod 3: h, Iphinotus typicus (Thomson) (Stebbing, 1899a, as I. chiltonz);
1, Heterophlias; 7, Quasimodia capricornis Sheard (1936a); k, Iphiplateia whitelegger
Stebbing (1899a); 1, Plioplateia. Pleopod 1: m, Heterophlias. Uropod 3: n, Heterophlias;
o, Iphinotus; p, Temnophlias. Uropod 2: g, Temnophlias. ‘Telson: r, Iphinotus.

MARINE GAMMARIDEAN AMPHIPODA 409

organisms like Sancho and Chosroes, which unlike typical phliantids
still bear triturative mandibular molars, mandibular palps, well-
developed biramous third uropods, free urosomal segments, and, ex-
cept for female Sancho, typical uropods 1 and 2. Sancho and Chosroes,
however, apparently flex the abdomen under the thorax like phliantids
and have somewhat splayed coxae. Temnophlias is linked to Sancho
and Chosroes and Gurjanova suggests that all three should be joined
together as an entity separate from the Phliantidae and Calliopiidae.
Temnophlias has the flexed abdomen, the degradation of the urosome,
its appendages and the simplification of mouthparts unlike Sancho
and Chosroes, but it differs from the Phliantidae in the (presumably
secondary) reduction of the coxae and the extension of the lateral
pereonal margins as pleurae, presumably as a substitute for the reduced
coxae. In this way Temnophlias resembles various isopods like Munna
and Pleurogonium and amphipods like Podocerus. This modification
is far more extreme in the unornamented type-species Temnophlias
capensis K. H. Barnard (1916) than its so-called congener, 7. hystrix
K. H. Barnard (1954) which has a thick, triquetral body with strong
dorsal ornamentation and bifidly acuminate coxae.

I suggest that Sancho and Chosroes together should be allocated to
a new family and Temnophlias to another new family in recognition of
these strong differences from Calliopiidae, Phliantidae, and each other.
Temnophlias can be segregated from Phliantidae by the munnid-like
body which seems to be a secondary depression and lateral pleuroniza-
tion of a cylindrioid body. In this respect Temnophlias may be
more closely related to the cylindrioid Eophliantidae than to the
Phhiantidae.

Key to the Genera of Phliantidae

1. Palp of maxilliped 2-articulate (fig. 147f) ...... . . . Temnophlias
Palpiotmaxilliped: 3-articulate (ig N45 ecg ee mss ey 2
Palpvofmaxilliped 4-articulate:(fig. 147e) 7 2). Sse

Zee WO LOMOdss * DIFAMOUS A yee yk ee nei a el an ae eta Malanth nner ta cree Phlias
Uropodesmot DiramMOuUsi is se os ae ee A Re baht ehhh wl Seema ise Oh 3

3. Lobes of maxilla 2 coalesced (fig. 146d), maxilla 1 lacking palp (fig. 146f).

Pereionotus
Lobes of maxilla 2 separate (fig. 146e), maxilla 1 with spine representing palp.
Palinnotus

4. Gnathopods subchelate or male gnathopod 2 chelate. ....... EGO

Gnathopodsisimiple aie space ey oem cia meee en eke alan) RU eae Oa Det Be 6

on

Maxilla 1 lacking palp; pleopod 3 apparently with rami subequal to each
other and with about seven setose articles each; uropod 3 with one ramus.
Ceina

Maxilla 1 with palp; pleopod 3 with vestigial, short, oval rami lacking setae;
MULOPOCtolaACKIn GATAMNIe Ss Ve i ee Mine Aliig oc thane dois Weal Na lia e Plioplateia

410 U.S. NATIONAL MUSEUM BULLETIN 271

6. Uropod 3: peduncle distinct from ramus (fig. 1470) . ......... 7
Uropod 3: peduncle not distinct from ramus (fig. 147n) ....... =. 8
7. Pleopod 3: inner ramus well developed (fig.147h) ....... Iphinotus
Pleopod 3: inner ramus vestigial (fig. 1477) . . ....... Quasimodia

8. Pleopod 3: inner ramus setose, half as long as outer.
Pariphinotus and Heterophlias
Pleopod 3: inner ramus a tiny nonsetose scale (fig. 147k) . . . Iphiplateia

Genera of Phliantidae

Ceina Della Valle

Ceina Della Valle, 1893.—Stebbing, 1906.—Nicholls, 1939.
Periphlias Pirlot, 1936.

Type-species: Nicea egregia Chilton, 1883 (monotypy).

Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods
subchelate; [pleopod 3 unknown, apparently biramous, rami subequal
to each other]; uropod 3 with one ramus. Species: 2, Sulu Sea, New
Zealand, Juan Fernandez Island, littoral.

Heterophlias Shoemaker
Heterophlias Shoemaker, 1933a.

Type-species: H. seclusus Shoemaker, 1933a (monotypy).

Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods
simple; inner ramus of pleopod 3 half as long as outer; uropod 3 lack-
ing rami. Possibly not distinct from Pariphinotus. Species: 1, pan-
America, tropical, littoral.

[phinotus Stebbing

Iphigenia Thomson, 1882 (homonym, Mollusca).
Iphinotus Stebbing, 1899a; 1906.

Type-species: Iphigenia typica Thomson, 1882 (monotypy and
subsequent synonymy).

Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods
simple; rami of pleopod 3 well developed, equal; uropod 3 uniramous.
Species: 1, New Zealand, littoral.

[phiplateia Stebbing
Iphiplateia Stebbing, 1899a; 1906.

Type-species: I. whitelegger Stebbing, 1899a (monotypy).

Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods
simple; inner ramus of pleopod 3 a tiny, nonsetose scale; uropod 3
lacking rami. Species: 1, E. Australia, littoral.

MARINE GAMMARIDEAN AMPHIPODA 411

Palinnotus Stebbing

Palinnotus Stebbing, 1900; 1906.

Type-species: Pereconotus thomsoni Stebbing, 1899a (original
designation).

Maxilla 1 bearing a spine-like palp; lobes of maxilla 2 separate
(Pereionotus) ; maxillipedal palp 3-articulate; gnathopods simple; rami
of pleopod 3 equal; uropod 3 lacking rami. Species: 3, Japan Sea,
S. Africa, E. Australia, littoral.

Pariphinotus Kunkel
Pariphinotus Kunkel, 1910.

Type-species: P. tuckeri Kunkel, 1910 (monotypy).

Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods
simple; inner ramus of pleopod 3 half as long as outer; “uropod 3
absent.”” Possibly synonymous with Heterophlias. Species: 1, Ber-
muda, littoral.

Pereionotus Bate and Westwood

Pereionotus Bate and Westwood, 1863.—Stebbing, 1906.
Icridium Grube, 1864a.

Type-species: Oniscus testudo Montagu, 1808 (monotypy). See
Chevreux and Fage, 1925.

Maxilla 1 lacking palp; lobes of maxilla 2 coalesced (Palinnotus) ;
maxillipedal palp 3-articulate; gnathopods simple; rami of pleopod 3
equal in length; uropod 3 apparently absent or not detected; uropod 2
with two rami in male, one ramus in female. Species: 1, tropical
EK. Atlantic, Mediterranean, Red Sea, littoral.

Phlias Guerin
Phlias Guerin, 1836.—Stebbing, 1906.

Type-species: P. serratus Guerin, 1836 (monotypy).

Obscure genus resembling Pereionotus. Maxilla 1 lacking palp;
maxillipedal palp 3-articulate; gnathopods simple; pleopod 3 with rami
equal in length; uropod 3 “‘biramous,’”’ but probably confused with
uropod 2. Guerin’s dorsal figure of urosome shows a setose pair, of
“uropods” probably representing a pair of pleopods. Species: 1,
between Iles Malovines* and Port Jackson, Australia, ?littoral.

*Falkland Islands; thus, type-locality of this species obscure.

285-135 O - 69 - 27

412 U.S. NATIONAL MUSEUM BULLETIN 271

Plioplateia K. H. Barnard

Plioplateia K. H. Barnard, 1916.

Type-species: P. triquetra K. H. Barnard, 1916 (monotypy).

Maxilla 1 with palp; maxillipedal palp 4-articulate; gnathopods
subchelate; pleopodal peduncles not expanded; both rami of pleopod
3 vestigial, short, oval, lacking setae; uropod 3 lacking rami. Species:
1, S. Africa, littoral.

Quasimodia Sheard
Quasimodia Sheard, 1936a.

Type-species: @. womersleyi Sheard, 1936a (present selection).

Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods
simple; inner ramus of pleopod 3 vestigial; uropod 3 with a single
ramus, occasionally with a vestigial second ramus. Species: 3, S.
Australia, littoral.

Temnophlias K. H. Barnard

Temnophlias K. H. Barnard, 1916.

Type-species: 7. capensis K. H. Barnard, 1916 (monotypy).

Maxilla 1 lacking palp; maxillipedal palp 2-articulate; gnathopods
simple or parachelate with chela subdistal; rami of pleopod 3 equal
in length; ramus of uropod 3 completely fused to peduncle; body
extraordinarily depressed and unornamented or triquetral and bearing
dorsal segmental teeth, pereonal segments laterally discontiguous as
in munnid isopods, coxae either small and simple or of medium size
and acuminately bifid, pereopods parachelate or simple, chela sub-
distal; antennal flagella of three or four or fewer articles; uropods
1-2 uniramous. Probably should be removed to a new family, see
remarks in ‘‘Relationship.”’ Species: 2, S. Africa, littoral.

Phoxocephalidae

Figures 148-151

Diacenosis.—Accessory flagellum well developed and multiarticu-
late; pereopod 5 shorter than pereopod 4 and structurally dissimilar
to it; head elongate, with overhanging rostrum, usually large, flat,
and hoodlike, occasionally reduced in size or dorsoventrally thickened,
but always distinct; pereopods strongly spinose for burrowing; uropod
3 biramous; telson cleft. See Haustoriidae, Gammaridae.

Description.—Accessory flagellum well developed; peduncles of
antennae very short, flagella very short except occasionally in males,

MARINE GAMMARIDEAN AMPHIPODA 413

antenna 2 peduncle usually heavily spinose; rostrum often hood-
shaped and enveloping base of antenna 1 or narrowed and projecting
between first antennae, usually flattened dorsoventrally ; body without
carinae except on urosome; mouthparts basic, mandibular molar
triturative or not, often spinose; palp of maxilla 1 uni- or biarticulate;
lower lip with inner lobes; gnathopods subchelate, small or of medium

CHC TTT
4 hla:

Ah
\:
‘
1 J?
| TF tors ;
R / y a &
y,\ sy TAT

SS

a

Figure 148.—Phoxocephalidae: a, Harpinia plumosa (Kréyer) (Sars, 1895, pl. 52);
b, Leptophoxus falcatus Sars (1895, pl. 50); c, Mandibulophoxus uncirostratus (Giles)
(Pillai, 1957).

414 U.S. NATIONAL MUSEUM BULLETIN 271

size (in one genus, Joubinella, gnathopod 1 larger than gnathopod 2);
uropod 3 biramous, outer ramus often reduced in size; telson short,
or of medium length, cleft.

ReEiatronsuip.— Most Haustortidae lack a conspicuous rostrum but
intergradations occurring in genera such as Platyischnopus prevent
absolute distinction; many haustoriids have pereopod 5 as long as or
longer than pereopod 4 and of structure similar to each other, but
genera such as Pontoporeia have pereopods 4 and 5 like those of the
Phoxocephalidae. Those haustoriids having semblance of a rostrum
never have pereopod 5 smaller and of different structure than 4 and
are restricted arbitrarily from the Phoxocephalidae. Probably the
families should be reorganized on the basis of pereopod 5 and by
disregarding the condition of the rostrum.

Nomenclatural Changes in Phoxocephalidae

Phoxocephalus capuciatus Oliveira (1955) is synonymized with Platy-
ischnopus gracilipes Schellenberg (1931).

Seven species of Harpinia and Harpinipsis are removed to Pseud-
harpinia; they have a basal ensiform process on antenna 2 and
distal spines or setae on the rami of uropod 2.

Several other species of Harpinia lacking a basal ensiform process
on antenna 2 and presumably bearing elongate male antenna 2 are
removed to Harpimopsis.

Key to the Genera of Phoxocephalidae

1. Article 2 of pereopod 3 more than twice as wide as article 3 (fig. 151c) . . .2
Article 2 of pereopod 3 about as wide as article 3 (fig. 151d). . . . . .10

2.) Palp/of maxilla 1 biarticulate (ig) 150))) 2.275 ss; PSE CN ie tani o>
Palp of maxilla 1 uniarticulate (fig. 1507). . . . 7

3. Gnathopod 1 larger than gnathopod 2 and Bhi cpeecliiee Ge NO), 5 man-
dibular molar triturative (fig. 150e) ... . ... . . .Joubinella
Gnathopods 1 and 2 similar in size, or gnathopod 2 eee than 1, mandibular
molar not triturative (fig. 150f) .... PBN pe)

4. Palp article 4 of maxilliped bearing two fia @istell aBe, nor dlr ire.
(fig. 150m), rostrum minute (fig. 149c). . .. . : Pee fo ta)

Palp article 4 of maxilliped not bearing large distal seine or 7 bannine one nail,
claw-like (fig. 150n), rostrum well developed (figs. 149a,d) ..... .6

5. Mandibular molar vestigial, a small spinose protuberance . Microphoxus
Mandibular molar very large, minutely setulose (fig. 97d).

Urothoides (Haustoriidae)

6. Mandibular palp sickle-shaped, borne on a large process of the mandibular

body, Gigs) 150g) ae: . . . . .Mandibulophoxus

Mandibular palp not Bick leehaned: not Bosna on a large process of the

mandibular body, (fig: 150e).. Basi) Seoes uy a dae araphoxus

MARINE GAMMARIDEAN AMPHIPODA 415

7. Article 3 of maxillipedal palp produced (fig. 1501) . . .....2.2.2. 2.8
Article 3 of maxillipedal palp unproduced (fig. 150n) ......2.2. 2.9

8. Molar of mandible triturative, large (fig. 150e) . . . . . .Leptophoxoides
Molar of mandible nontriturative, small (fig. 150f) . . . . .Leptophoxus

9. Molar of mandible triturative (fig. 150e). .... .. . . Phoxocephalus
Molar of mandible nontriturative, smooth (or ponte eo 150f) . Metaphoxus
ee i ae Hotel ecaibe tcstire nareiu be
Kyes present .. : Sade Meena ica unless sti

11. Antenna 2 with bagel sracitionen y process Ge 1500) Stn Geet pe Bel 2
Antenna 2 lacking basal ensiform process (fig. 1506) . . . . . Harpiniopsis

12. Rami of uropod 2 with apical spines (fig. 151f) . . . . . . Pseudharpinia
Rami of uropod 2 lacking apical spines (fig. 15le) ... . . . .Harpinia

18. Antenna 2 with basal ensiform process (fig. 150a) .. . . . Heterophoxus
Antenna 2 lacking basal ensiform process (fig. 150b). . ...... =.14!

14. Dactyl of maxillipedal palp composed of long curved claw bearing short
nail-like spine .. . . . . Mandibulophoxus
Dactyl of maxillipedal elt ieompoced oF phone quned or straight body
bearing elongate, curved, claw-like spine .... . . . .. .Proharpinia

1 And see Phozocephalus coralis K. H. Barnard (1932) to be relegated to a new genus.

Genera of Phoxocephalidae

Harpinia Boeck

Harpina Boeck, 1871 (homonym, Coleoptera).
Harpinia Boeck, 1876 (new name).—Stebbing, 1906.

Type-species: Phoxus plumosus Kroyer, 1842 (selected by Boeck,
1876). See Sars, 1895.

Antenna 2 with basal ensiform process, male antenna 2 short (Harpin-
topsis); mandibular molar a small, occasionally weakly spinose,
nontriturative protuberance; palp of maxilla 1 biarticulate; maxilli-
pedal palp article 3 not produced, article 4 short, with one or a few
apical spines or setae; article 2 of pereopod 3 slender, scarcely wider
than article 3; eyes absent; rami of uropod 2 lacking distal spines
or setae (Pseudharpinia). Species: about 21, N. Atlantic, ? N.
Pacific, cold-water, primarily deep sublittoral and bathyal.

Harpiniopsis Stephensen
Harpiniopsis Stephensen, 1925a.

Type-species: H. similis Stephensen, 1925a (monotypy).

Antenna 2 lacking basal ensiform process, male antenna 2 as long
as body (Harpinia, Pseudharpinia); mandibular molar a small,
occasionally weakly spinose, nontriturative protuberance; palp of
maxilla 1 biarticulate (often obscurely); maxillipedal palp article 3
not produced, article 4 short, with one or a few apical spines or setae;

416 U.S. NATIONAL MUSEUM BULLETIN 271

article 2 of pereopod 3 slender, scarcely wider than article 3; eyes
absent. Species: about 19, cosmopolitan, bathyal and abyssal.

Ficure 149.—Phoxocephalidae: a, Paraphoxus tridentatus J. L. Barnard (1960); d, Joubinella
traditor Pirlot (1932). Head, dorsal: c, Microphoxus minimus J. L. Barnard (1960);
d, Paraphoxus oculatus Sars (1895, pl. 51).

MARINE GAMMARIDEAN AMPHIPODA 417

Heterophoxus Shoemaker

Heterophozus Shoemaker, 1925.

Type-species: Harpinia oculata Holmes, 1908 (monotypy and
subsequent synonymy).

Mandibular molar a spinulose nontriturative protuberance; palp
of maxilla 1 biarticulate; maxillipedal palp article 3 not produced,
article 4 short, with a stout apical spine and setae; article 2 of pereo-
pod 3 slender, scarcely wider than article 3; eyes present or absent;
antenna 2 with basal ensiform process (Proharpima). Species: 4, E.
Pacific, S. Africa, antarctic, littoral to deep bathyal.

Figure 150.—Phoxocephalidae: Antenna 2: a, Heterophoxus oculatus (Holmes) (J. L
Barnard, 1960); b, Phoxocephalus holbolli (Krdyer) (Sars, 1895, pl. 49). Antenna 1:
c, Phoxocephalus. Upper lip: d, Phoxocephalus. Mandibles and palps: e, Phoxocephalus;
f, Leptophoxus falcatus Sars (1895, pl. 50); g, Mandibulophoxus uncirostratus (Giles)
(J. L. Barnard, 1957a). Lower lip: h, Phoxocephalus. Maxilla 1: 1, Phoxocephalus;
j, Harpinia plumosa (Kréyer) (Sars, 1895, pl. 52). Maxilla 2: k, Phoxocephalus. Maxil-
lipeds: 1, Leptophoxus; m, Microphoxus minimus J. L. Barnard (1960); », Phoxocephalus.
Gnathopod 1: 0, Joubinella strelkovi Gurjanova (1952); p, Phoxocephalus.

A18 U.S. NATIONAL MUSEUM BULLETIN 271

Joubinella Chevreux
Joubinella Chevreux, 1908a.

Type-species: J. ciliata Chevreux, 1908a (original designation).

Mandibular molar well developed, ridged; palp of maxilla 1 bi-
articulate; maxillipedal palp article 3 not produced, article 4 long,
subobtuse or claw-like, setulose or spinulose; article 2 of pereopod 3
more than twice as wide as article 3; gnathopod 1 larger than gnatho-
pod 2; gnathopods resembling those of Eusirus; flagellum of antenna 2
reduced to two or three articles. Species: 5, Atlantic-Pacific, epi- to
bathypelagic.

Ficure 151.—Phoxocephalidae: Gnathopod 2: a, Phoxocephalus holbolli (Krgyer) (Sars,
1895, pl. 49); b, Joubinella strelkovi Gurjanova (1952). Pereopod 3: c, Phoxocephalus;
d, Harpinia plumosa (Krgyer) (Sars, 1895, pl. 52). Uropod 2: e, Phoxocephalus ;
f, Pseudharpinia dentata Schellenberg (J. L. Barnard, 1960). Uropod 3, Phoxocephalus: g,
female; h, male. Telson: 1, Phoxocephalus; 7, Harpinia.

Leptophoxoides J. L. Barnard

Leptophoxoides J. L. Barnard, 1962d.

Type-species: L. molaris J. L. Barnard, 1962d (original designation).

Mandibular molar wel] developed, ridged; palp of maxilla 1 uniartic-
ulate; maxillipedal palp article 3 apically produced, article 4 of medium
length, subclaviform, with two apical] setae; article 2 of pereopod
3 broad, more than twice as wide as article 3; eyes absent. Species:
1, S. Atlantic, abyssal.

MARINE GAMMARIDEAN AMPHIPODA 419

Leptophoxus Sars
Leptophoxus Sars, 1895.—Stebbing, 1906.

Type-species: Phozxus falcatus Sars, 1882 (monotypy).

Mandibular molar a small nontriturative protuberance; palp of
maxilla 1 biarticulate; maxillipedal palp article 3 apically produced,
article 4 long, claw-like; article 2 of pereopod 3 more than twice as
wide as article 3; eyes absent. Species: 1, subarctic-boreal, littoral to
abyssal.

Mandibulophoxus J. L. Barnard
Mandibulophozus J. L. Barnard, 1957a; 1960.

Type-species: Phoxus uncirostratus Giles, 1890 (original designation
and subsequent synonymy).

Mandibular molar a setulose, nontriturative protuberance; palp of
maxilla 1 biarticulate; maxillipedal palp article 3 not produced,
article 4 long, claw-like, tipped with short nail; article 2 of pereopod 3
more than twice as wide as article 3 (type) or not; eyes present or
absent; mandibular palp set on extension of mandibular body (combining
character). Species: 2, Indo-E. Pacific tropical, S. Africa, littoral.

Metaphoxus Bonnier
Metaphoxus Bonnier, 1896.—Stebbing, 1906.

Type-species: MM. typicus Bonnier, 1896 (monotypy).

Mandibular molar a setulose, nontriturative protuberance; palp of
maxilla 1 uniarticulate; maxillipedal palp article 3 not produced, article
4 long, claw-like; article 2 of pereopod 3 more than twice as wide as
article 3; eyes present. Species: 4, boreal-warm temperate, littoral
to deep bathyal.

Microphoxus J. L. Barnard
Microphozus J. L. Barnard, 1960.

Type-species: MM. minimus J. L. Barnard, 1960 (original desig-
nation).

Mandibular molar a setulose nontriturative protuberance; palp of
maxilla 1 biarticulate; maxillipedal palp article 3 not produced,
article 4 long, but with two large apical setae; article 2 of pereopod 3
more than twice as wide as article 3; eyes present. Species: 1, Pacific
Costa Rica, littoral.

420 U.S. NATIONAL MUSEUM BULLETIN 271

Paraphoxus Sars

Paraphozus Sars, 1895.—Stebbing, 1906.—J. L. Barnard, 1960.
Pontharpinia Stebbing, 1897.

Parharpinia Stebbing, 1899d.

Protophoxus K. H. Barnard, 1930.

Trichophozus K. H. Barnard, 1930.

Metharpinia Schellenberg, 1931.

Type-species: Phozus oculatus Sars, 1879 (monotypy).

Mandibular molar a setulose nontriturative protuberance; palp of
maxilla 1 biarticulate; maxillipedal palp article 3 not produced,
article 4 long, claw-like; article 2 of pereopod 3 more than twice
as wide as article 3; eyes present. Species: 46, cosmopolitan, mainly
littoral, occasionally bathyal in high latitudes.

Phoxocephalus Stebbing

Phorus Kr¢gyer, 1842 (homonym, Coleoptera).
Spinifer Krgyer, 1842 (homonym, Mollusca).
Phoxocephalus Stebbing, 1888 (new name); 1906.

Type-species: Phoxus holbolli Kréyer, 1842 (selected by Boeck,
1876). See Sars, 1895.

Mandibular molar well developed, ridged; palp of maxilla 1 uni-
articulate; maxillipedal palp article 3 not produced, article 4 long,
claw-like; article 2 of pereopod 3 more than twice as wide as article
3; eyes present or vestigial. Species: 7 (one dubious), cosmopolitan
cold-water, littoral to bathyal.

Proharpinia Schellenberg
Proharpinia Schellenberg, 1931.

Type-species: P. antipoda Schellenberg, 1931 (monotypy).

Mandibular molar a setulose nontriturative protuberance; palp of
maxilla 1 biarticulate; maxillipedal palp article 3 not produced,
article 4 of medium length or short, with a long apical spine and
accessory setae, but appearing long and claw-like because of the
long spine; article 2 of pereopod 3 slender, scarcely wider than
article 3; eyes present; antenna 2 lacking basal ensiform process
(Heterophorus). Species: 4, Galapagos, antiboreal, littoral.

Pseudharpinia Schellenberg
Pseudharpinia Schellenberg, 1931.

Type-species: P. dentata Schellenberg, 1931 (monotypy).
Antenna 2 urth basal ensiform process, male antenna 2 unknown in
type and elongate in some species (Harpiniopsis); mandibular molar a

MARINE GAMMARIDEAN AMPHIPODA 421

small, occasionally weakly spinose, nontriturative protuberance;
palp of maxilla 1 biarticulate (often obscurely); maxillipedal palp
article 3 not produced, article 4 short, with one or a few apical
spines or setae; article 2 of pereopod 3 slender, scarcely wider than
article 3; eyes absent; rami of uropod 2 bearing distal spines or setae
(Harpinia and Harpiniopsis). Species: about 8, antarctic littoral
and probably cosmopolitan in bathyal-abyssal.

Pleustidae
Fieures 152, 153

Driaenosis.—Accessory flagellum composed of a small scale or ab-
sent; lower lip formed of two oval outer lobes obliquely astride nearly
coalesced inner lobes; telson entire; upper lip incised. See Calliopiidae,
Paramphithoidae, Liljeborgiidae, Isaeidae, Eusiridae, Ampithoidae,
Amphilochidae, Ischyroceridae, Laphystiopsidae, Oedicerotidae.

Description.—Body with or without dorsal processes; rostrum
long or inconspicuous; accessory flagellum vestigial; upper lip incised ;
mandibles bearing a 3-articulate palp, molar triturative or nontritura-
tive; lower lips characteristic (see diagnosis) ; remaining mouthparts
basic, article 3 of palp of maxilliped occasionally produced; gnathopods
subchelate, feeble or powerful; coxae of medium length or long;
uropod 3 with rami much longer than peduncle, lanceolate, outer
ramus usually shorter than inner; telson entire or apically sculptured,
cleft nearly one fifth in one species of Austropleustes.

RELATIONSHIP.—One genus of Amphilochidae, Amphilochoides, pos-
sesses a lower lip like those of pleustids but most amphilochids can
be characterized by the small, partially hidden coxa 1, and some
amphilochids have both coxae 1 and 2 concealed. Pseudamphilochus
however has a normal coxa 1 and cleft telson and thus resembles one
species of Austropleustes. See family key G for distinctions.

Only the shape of the lower lip separates this family from the
Calliopiidae. Two genera, Harpinioides and Mesopleustes, have lower
lips intermediate between the two families.

The Laphystiopsidae are very similar to the Pleustidae and further
study is needed to clarify their systematic position.

Key to the Genera of Pleustidae

1. Mandibular molar with a triturating surface composed of ridges and teeth
(Giese ISTE Pee a ee bls al Cl aia a ona ial ennui ranula ND muita ty eMeSetrn tac 2
Mandibular molar a simple, small, smooth hump (fig. 1538c) ...... 5
2. Maxillipedal palp article 3 bearing a distal process (fig. 15372).

Stenopleustes
Maxillipedal palp article 3 lacking a distal process (fig. 153h). . . .. . 3

422 U.S. NATIONAL MUSEUM BULLETIN 271

3. Gnathopods simple, slender (fig. 1530) . . . .. =. =... . Parepimeriella
Gnathopods subchelate, stout (figs. 153l,m). . . . Pee eae ye

4, Rostrum less than one fourth as long as article 1 of amnienne L.
Pleusymtes, new genus

Rostrum nearly as long as article 1 of antenna 1... . . . Mesopleustes
5. Maxillipedal palp article 3 bearing a distal process (fig. 1532) . . . . . . 6
Maxillipedal palp article 3 lacking a distal process (fig. 153h) . . . . 7

6. Uropod 3 with a large peduncular process supporting the rami (fig. 153g).
Austropleustes
Uropod 3 lacking accessory peduncular process .. . . . . Neopleustes
7. Gnathopods with distally bent, slender, tapering fin articles, essentially
simples (fig4i20) ane . . . . . Harpinioides !

Gnathopods subchelate, with arate c or solneotanrallar: unbent sixth articles. 8

8. Antennae short, gnathopods bearing lobate article 5 (fig. 153m) . Pleustes ?
Antennae long, gnathopods usually lacking lobate article 5 (fig. 153p).

Parapleustes 2

1 See this genus in the Calliopiidae.
2 The differences between Pleustes and Parapleustes are subjective. See Barnard and Given (1960, p. 39)
for discussion.

Genera of Pleustidae

Austropleustes K. H. Barnard
Austropleustes K. H. Barnard, 1931; 1932.

Type-species: A. cuspidatus K. H. Barnard, 1931 (original desig-
nation) ; 1932.

Mandibular molar a small smooth protuberance; article 3 of maxil-
lipedal palp distally produced; gnathopods slender but subchelate;
uropod 3 with a large peduncular process supporting the rami (Neo-
pleustes). Species: 2, antarctic, deep littoral.

Mesopleustes Stebbing
Mesopleustes Stebbing, 1899d; 1906.

Type-species: Pleustes abyssorum Stebbing, 1888 (original desig-
nation).

Mandibular molar well developed, ridged; article 3 of maxillipedal
palp not distally produced; gnathopods subchelate; rostrum nearly
as long as article 1 of antenna 1 (Pleusymtes). Because lower lip is
not typical of other pleustids this genus probably should be assigned
to the Calliopiidae, noting that it also has affinities with Param-
phithoidae because of the tendency of the coxae toward acumination.
Species: 1, probably cosmopolitan, bathyal-abyssal.

MARINE GAMMARIDEAN AMPHIPODA 423

f}
f}
4,
t}
t}
t}
t}
i}
|
1
i)
!
\
\
\
‘i
i
i

Figure 152.—Pleustidae: a, Stenopleustes malmgreni (Boeck) (Sars, 1895, pl. 125); b, Pleustes
panopla (Kr¢yer) (Sars, 1895, pl. 121).

424 U.S. NATIONAL MUSEUM BULLETIN 271

Neopleustes Stebbing

Neopleustes Stebbing, 1906.

Type-species: Amphitoe pulchella Krgyer, 1846 (original designa-
tion). See Sars, 1895 (as Paramphathoe).

Mandibular molar a small smooth protuberance; article 3 of maxil-
lipedal palp produced distally; gnathopods subchelate; uropod 3
lacking a large peduncular process (Austropleustes). Species: 6, arctic-
boreal, littoral to bathyal (1416 m).

Figure 153.—Pleustidae: Upper lip: a, Pleustes panopla (Kr@yer) (Sars, 1895, pl. 121).
Mandible: b, Sympleustes glaber (Boeck) (Sars, 1895, pl. 126); c, Pleustes. Lower lip:
d, Pleustes; e, Mesopleustes abyssorum (Stebbing, 1888). Maxillae 1, 2: fig, Pleustes.
Maxillipeds: h, Pleustes; 1, Stenopleustes latipes (M. Sars) (Sars, 1895, pl. 127); 7, Meso-
pleustes, palp. Gnathopod 1: k, Austropleustes cuspidatus K. H. Barnard (1932); J,
Neopleustes pulchellus (Kr@yer) (Sars, 1895, pl. 122); m, Pleustes. Gnathopod 2: n,
Pleustes; 0, Parepimeriella irregularis Schellenberg (1931); p, Neopleustes. Uropod 3: gq,
Austropleustes, showing peduncular process [shaded]; 7, Pleustes. ‘Telson: 5, Neopleustes;
t, Pleustes.

MARINE GAMMARIDEAN AMPHIPODA 425

Parapleustes Buchholz

Parapleustes Buchholz, 1874.—Sexton, 1909.
Incisocalliope J. L. Barnard, 1959a.

Type-species: [P. glacilis Buchholz, 1874, =—lapsus] P. gracilis
Buchholz, 1874 (monotypy). See Sars, 1895 (as Paramphithoe brevi-
corns) .

Mandibular molar a small smooth protuberance; article 3 of maxil-
lipedal palp not distally produced; gnathopods subchelate; antennae
“Jong,” gnathopods usually lacking a lobe on article 5, rostrum reaching
only one third along article 1 (Pleustes). Species: 14, arctic-boreal,
littoral.

Parepimeriella Schellenberg
Parepimeriella Schellenberg, 1931.

Type-species: P. irregularis Schellenberg, 1931 (monotypy).
Mandibular molar large, ridged; article 3 of maxillipedal palp not
distally produced; gnathopods simple, slender. Species: 1, Falkland
Islands, littoral.
Pleustes Bate

Pleustes Bate, 1858b.—Stebbing, 1906.

Type-species: Amphithoe panopla Krgyer, 1838 (monotypy and
subsequent synonymy). See Sars, 1895.

Mandibular molar a small smooth protuberance; article 3 of
maxillipedal palp not distally produced; gnathopods subchelate, stout ;
antennae “‘short,”’ gnathopods bearing a strong posterior lobe on article
5, rostrum usually reaching apex of peduncle of antenna 1 in typical
species [except Pleustes behningi (Gurjanova, 1938) which should be
transferred to Parapleustes]. Species: 7, arctic-boreal, littoral.

Pleusymtes, new genus

Type-species: Amphithopsis glaber Boeck, 1861 (present selec-
tion). See Sars, 1895.

Species included: All species listed by Barnard and Given (1960,
p. 40) under the name “‘Sympleustes.”’

Mandibular molar large, ridged; article 3 of maxillipedal palp not
produced; gnathopods subchelate; rostrum extending less than one
quarter along article 1 of antenna 1 (Mesopleustes). Species: 17,
arctic-boreal, littoral to abyssal.

Stenopleustes Sars, new synonymy

Stenopleustes Sars, 1895.—Stebbing, 1906.
Sympleustes Stebbing, 1899d (type: Amphithoe [sic] latipes M. Sars, 1858).

426 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: Amphithopsis malmgreni Boeck, 1871 (selected by
Chevreux and Fage, 1925).

Mandibular molar large, ridged; article 3 of maxillipedal palp
apically produced; gnathopods subchelate. Species: 7, arctic-boreal,
littoral to bathyal.

Podoceridae

Figures 154-156

Diaanosis.—Accessory flagellum variable, often absent; urosome
markedly depressed, segment 1 more than twice as long as segment 2,
often considerably longer; telson entire; pereopods not glandular.
See Corophiidae, Cheluridae.

Description.—Accessory flagellum variable, multiarticulate, ves-
tigial or absent; body variable, broadly depressed or slender and
subcylindrical, often carinate and toothed, urosome strongly de-
pressed, segment 1 more than twice as long as any following segment,
occasionally segment 6 apparently fused with segment 5, segments
5 and 6 reduced in length; coxae usually small and serially discon-
tiguous, occasionally splayed; mouthparts basic, except upper lip
slightly bilobed; gnathopods of medium size or large, subchelate;
uropod 1 normal; uropods 2 and 3: one or the other often missing,
vestigial or abnormal; uropod 3 biramous only in Jcilius, generally
with one or no rami, telson circular or oval, short, entire; pereonites
6-7 coalesced in Dulichia and Paradulichia.

Revationsuip.—The elongation of pleonite 4 distinguishes this
family from the Corophiidae.

Podoceridae often have the subcylindrical bodies of Eophliantidae
and Colomastigidae but the latter two families may be distinguished
by their degraded mandibles lacking palps.

Runanga J. L.. Barnard (1961) is removed to the Corophiidae.

Key to the Genera of Podoceridae

ft Antennal lackime accessory, flagellum) salem eine il sein

Antenna 1 bearing accessory flagellum . . . ao

Pleon with only five distinct segments pr eredine ‘ihe telecon Ge 156a).
Laetmatophilus !

to

Pleon with six distinct segments preceding the telson (fig. 156b) . . . .3
3. Pleonite 5 carrying well-formed uropods ...... . .. . .Cyrtophium
Pleonite > lacking wropeds. 2) 3.25.4 es a Leipsuropus?

1 See Dryopoides (Aoridae or Ischyroceridae) bearing distinct uropod 3.

2 Analysis of Leipswropus may be defective; the loss of uropod 2 but the retention of uropod 3 seems il-
logical; the so-called third uropod may indeed be the second and this genus would then be referable to
Laetmatophilus.

MARINE GAMMARIDEAN AMPHIPODA 427

4. Three pairs of uropods present (third often minute) (fig. 156b), pereonites

G-(itreer ae oe: Bala ese)

Two pairs of sexeinosls miccent Ge 156), sonuontes 6- 7 conieccede arenes)

eNO ropod:3 biramous* (igs. lo6c9)) 2.) 2k so me urge Vole CRS
Uropod 3 uniramous or lacking rami (figs. 156a 1) Hive Sts acme Sakae te iat)

6. Antenna 1 shorter than antenna 2 (fig. 154c) . . .. . . . . . Podocerus
Antenna 1 as long as or longer than antenna 2 (figs. 154a,b) . .....7

“I

Coxae 1-4 tiny, separated from Gren other by a distance morethan their
width (fig. 1546), pereopods 1—2 shorter than the gnathopods (fig. 1546).
Neoxenodice‘
Coxae small, separated from each other by a distance of one half their width
(fig. 155a), pereopods 1-2 longer than the gnathopods (fig. 155a).

Xenodice‘
Se Wropode2sbiramous lance (igalloGe)inye ten a) oe eee ee Danlichia
Uropod 2 uniramous, very small (fig. 156c). . . .. . . . . Paradulichia

3 Inner ramus of uropod 3 fragile and often missing from specimens but its presence is indicated by a
supporting process of the peduncle.
4 Uncinotarsus (Aoridae) keys to this position but gnathopod 1 is larger than 2.

Genera of Podoceridae

Cyrtophium Dana

Cyrtophium Dana, 1852b.—Stebbing, 1906.

Type-species: ©. orientale Dana, 1853 (selected by monotypy of
Dana, 1853). See Bate, 1862.

Accessory flagellum absent; antennae subequal in length; pleon
with six segments and three pairs of uropods; uropod 2 with two
rami, uropod 3 lacking rami. Species: 2, Singapore, E. Australia.
littoral.

Dulichia Kroyer
Dulichia Kr¢éyer, 1845.—Stebbing, 1906.
Dyopedos Bate, 1857a,b.

Type-species: D. spinosissima Krgyer, 1845 (monotypy). See Sars,
1895.

Accessory flagellum present; antenna 1 slightly longer than 2;
pleon with only five segments and only two pairs of uropods; uropod
2 biramous; uropod 3 absent; pereonites 6-7 coalesced. Species: 17,
arctic-boreal, littoral to abyssal.

Icilius Dana

Icilius Dana, 1849; 1852a.—Stebbing, 1906.

Type-species: J. ovalis Dana, 1852a (selected by monotypy of
Dana, 1852a). See Stebbing, 1888 (as I. danae, I. ellipticus, and I.
australis).

285-135 O - 69 - 28

428 U.S. NATIONAL MUSEUM BULLETIN 271

V
ee)
Nh
; ;

Figure 154.—Podoceridae: a, Dulichia spinosissima Kréyer (Sars, 1895, pl. 228); b, Neo-
xenodice caprellinoides Schellenberg (J. L. Barnard, 1962d); c, Podocerus variegatus Leach
(Chevreux and Fage, 1925).

MARINE GAMMARIDEAN AMPHIPODA 429

Accessory flagellum present; antenna 1 much shorter than 2; pleon
with six segments and three pairs of uropods; uropod 2 biramous;
uropod 3 “‘biramous,” inner ramus fragile, often missing but indicated
by a supporting process on peduncle. Species: 4, Indonesia, Australia,
littoral.

el
i)

XK Ce ff eh

al Ree
Hy It

Ficure 155.—Podoceridae: a, Xenodice frauenfeldti Boeck (Sars, 1895, pl. 227); b, Laetmato-
philus tuberculatus Bruzelius (Sars, 1895, pl. 226). Upper lip: c, Laetmatophilus. Mandi-
ble: d, Paradulichia typica Boeck (Sars, 1895, pl. 232); ¢, Laetmatophilus. Laetmatophilus:
f, lower lip; g,h, maxillae 1, 2.

430 U.S. NATIONAL MUSEUM BULLETIN 271

Laetmatophilus Bruzelius

Laetmatophilus Bruzelius, 1859.—Stebbing, 1906.

Type-species: L. tuberculatus Bruzelius, 1859 (monotypy). See
Sars, 1895.

Accessory flagellum absent; antennae subequal; pleon with only
five segments and only two pairs of uropods; uropod 2 lacking rami,
uropod 3 absent. Species: 7, cosmopolitan, littoral to bathyal.

Ficure 156.—Podoceridae: Urosome, uropods and telson, dorsal view: a, Laetmatophilus
tuberculatus Bruzelius (Sars, 1895, pl. 226); b, Podocerus chelonophilus (Chevreux and de
Guerne) (Chevreux and Fage, 1925); c, Paradulichia typica Boeck (Sars, 1895, pl. 232);
d, Dulichia spinosissima Kr@yer (Sars, 1895, pl. 228); ¢, Icilius ovatus Dana (Stebbing,
1906). Maxilliped: f, Laetmatophilus. Gnathopods 1, 2: g,h, Laetmatophilus. Uropod
3: 1, Icilius danae Stebbing (1910); 7, Icilius punctatus Haswell (Stebbing, 1910); ,
Laetmatophilus.

Leipsuropus Stebbing
Leipsuropus Stebbing, 1899d; 1906.

Type-species: Cyrtophium parasiticum Haswell, 1880b, 1885
(monotypy).

Accessory flagellum absent; antenna 1 shorter than 2; pleon with
(6?) segments: “uropod 2 absent, uropod 3 short, foliaceous, ovate,
lacking rami’’: these facts probably erroneous; probably uropod 3 1s
absent, uropod 2 lacks rami, hence referring the genus to Laetmatophilus.
Species: 1, E. Australia, littoral.

MARINE GAMMARIDEAN AMPHIPODA 431

Neoxenodice Schellenberg

Neoxenodice Schellenberg, 1926c.

Type-species: NN. caprellinoides Schellenberg, 1926c (monotypy).

Accessory flagellum present: antennae equal in length; pleon with
sIx segments and three pairs of uropods; uropod 2 with two rami,
uropod 3 lacking rami; corae tiny, separated by a distance more than
their width, pereopods 1-2 shorter than the gnathopods (Xenodice).
Species: 1, antiboreal, abyssal.

Paradulichia Boeck

Paradulichia Boeck, 1871.—Stebbing, 1906.

Type-species: P. typica Boeck, 1871 (monotypy). See Sars, 1895.

Accessory flagellum present; antenna 1 longer than 2; pleon with
only five segments and two pairs of uropods; uropod 2 uniramous,
very small, uropod 3 absent; pereonites 6-7 coalesced. Species: 3,
arctic-boreal, N. Atlantic, littoral to bathyal.

Podocerus Leach

Podocerus Leach, 1814b.—Stebbing, 1906.
Platophium Dana, 1852b.
Dexiocerella Haswell, 1885.

Type-species: P. variegatus Leach, 1814b (monotypy). See Chevy-
reux and Fage, 1925.

Accessory flagellum present; antenna 1 shorter than 2; pleon with
six segments and three pairs of uropods; uropod 2 with two rami;
uropod 3 lacking rami. Species: 21, cosmopolitan, littoral (rarely
bathyal, 569 m in antarctic).

Xenodice Boeck
A enodice Boeck, 1871.—Stebbing, 1906.

Type-species: X. frauenfeldti Boeck, 1871 (monotypy). See Sars,
1895.

Accessory flagellum present; antenna 1 slightly longer than 2;
pleon with six segments and three pairs of uropods; uropod 2 with
two rami; uropod 3 lacking rami; corae small, separated by a distance
half their width, pereopods 1-2 longer than the gnathopods (Neoxrenodice).
Species: 1, subarctic, N. Atlantic, littoral to bathyal.

432 U.S. NATIONAL MUSEUM BULLETIN 271

Prophliantidae, revised
Fiegur#s 157, 158

Diaenosis.—Accessory flagellum absent; body laterally compressea;
mandible lacking palp, molar smooth or evanescent; gnathopods
feeble but clearly subchelate; telson deeply cleft; uropod 3 biramous;
all of urosomal segments coalesced; antennae very short, not longer
than head. See Phliantidae, Eophliantidae, Dexaminidae, Atylidae,
Colomastigidae, Kuriidae.

Description.—Body laterally compressed, urosomites 1-3 coa-
lesced; mandible lacking palp, molar smooth or evanescent; lower
lip with inner lobes; palp of maxilla 1 uniarticulate; maxilla 2 normal;
maxillipedal inner plates small, outer greatly elongate; gnathopods
feeble, subchelate; coxae of medium size or large (coxa 5); antennae
very short, not longer than head, antenna 2 in type-genus lacking
flagellum, in second genus bearing flagellum; uropod 3 biramous;
telson deeply cleft.

RELATIONSHIP.—The Eophliantidae have cylindrical bodies and
reduced coxae and the gnathopods are simple or parachelate, not
subchelate.

The Talitroidea have distinct urosomal segments and a well-
developed mandibular molar.

The Phliantidae have uniramous or aramous third uropods. The
genus Ceima in that family forms a bridge to the Prophliantidae.

The Kuriidae have a flagellum on antenna 2 and a uniramous
third uropod.

The Atylidae have a mandibular palp.

The Dexaminidae (including Anatylidae) have only urosomites 2
and 3 fused.

The Prophliantidae are characterized especially by coalesced uro-
somal segments, biramous third uropods, lack of mandibular palps,
and cleft telsons.

Haustoriopsis Schellenberg, 1938, is removed from the Haustoriidae
to this family. It resembles the type-genus in all respects except that
it bears a 3-articulate antennal flagellum whereas Prophlias lacks a
flagellum. The coalesced urosomal segments, large coxa 5, similar
maxuliped with small inner plates and elongate outer plates, 1-
articulate first maxillary palp, absence of mandibular palp, smooth
and bulging mandibular molar, all point to the close relationship of
these two genera. Ruffo (1959) suggested that Haustoriopsis be re-
moved to the Dexaminidae but I believe that the Prophliantidae
should be segregated from that family on the distinctive urosome.
Presumably Prophliantidae have been derived from dexaminid-like
ancestors.

MARINE GAMMARIDEAN AMPHIPODA 433
Nomenclatural Changes in Prophliantidae

The following genera are removed to the Eophliantidae: Biancolina
Della Valle (1893), Bircenna Chilton (1884), Oeinina Stephensen
(1933), Cylindryllioides Nicholls (1938), Hophliantis Sheard (1936a),
and Wandelia Chevreux (1906a).

Cerna Della Valle (1893) is removed to the Phliantidae.

Kuria Walker and Scott (1903) is removed to the Kuriidae.

Figure 157.—Prophliantidae: a, Prophlias anomalus Nicholls (1939); b, Haustoriopsis
reticulatus Schellenberg (1938); c, Prophlias, antenna 2.

434 U.S. NATIONAL MUSEUM BULLETIN 271

Key to the Genera of Prophliantidae

1. Antenna 2 with flagellum (fig. 157b) ..... =. =... .. . Haustoriopsis
Antenna 2 lacking flagellum (fig. 157a) . . ... =.=... . .. . Prophlias

€

Figure 158.—Prophliantidae: Prophlias anomalus Nicholls (1939): a,b, gnathopods 1, 2;
c, head; d, mandible; e, lower lip; f,g, maxillae 1, 2; A, maxilliped; 7, uropod 3; 7, telson.
Haustoriopsis reticulatus Schellenberg (1938): k, mandible; /, maxilla 1; m, uropod 3.

Genera of Prophliantidae

Haustoriopsis Schellenberg

Haustoriopsis Schellenberg, 1938.

Type-species. H. reticulatus Schellenberg, 1938 (monotypy).
Species: 1, Bismarck Archipelago, littoral.

Prophlias Nicholls

Prophlias Nicholls, 1939.

Type-species: P. anomalus Nicholls, 1939 (monotypy).
Species: 1, W. Australia, littoral.

MARINE GAMMARIDEAN AMPHIPODA 435

Sebidae

Figure 159

Draenosis.—Accessory flagellum 2-articulate, elongate; molar of
mandible obsolescent; plates of maxillipeds of medium size; gnatho-
pods 1 and 2 chelate, gnathopod 1 larger than 2, article 3 of gnatho-
pod 2 elongate; antenna 1 peduncle elongate; uropod 3 uniramous;
telson entire; urosomites 2-3 coalesced. Monogeneric. See Lysi-
anassidae, Leucothoidae, Anamixidae, Pagetinidae.

Description.—Accessory flagellum 2-articulate, elongate; body
smooth or with poorly developed carinae, urosomites 2—3 coalesced;
mandibular palp article 3 short, molar evanescent; remaining mouth-
parts basic although plates of maxillipeds somewhat small; coxae of
medium length, rounded-quadrate below; gnathopods chelate, rather

ty)

Ficure 159.—Sebidae: Seba armata Chevreux (1900): a, lateral view; b, upper lip; c,
mandible; d, maxilla 1; ¢, maxilliped; f,g, gnathopods 1, 2; h, telson; 7, antenna 1; 7, uropod
3. Seba saundersi Stebbing (1888): k, lower lip.

436 U.S. NATIONAL MUSEUM BULLETIN 271

slender, first larger than second, and occasionally weakly chelate,
article 3 of gnathopod 2 elongate; uropod 3 with its single ramus
longer than the short peduncle; telson entire.

RELATIONSHIP.—The Leucothoidae have biramous third uropods.
The Lysianassidae have a short peduncle of antenna 1 with article 2
much shorter than 1 whereas the Sebidae not only have an elongate
peduncle but article 2 is much longer than 1. Few Lysianassidae have
a uniramous uropod 3 and when so it is never elongate.

The Stenothoidae have subchelate gnathopods and coxa 1 is con-
cealed by coxa 2, but mouthparts and uropod 3 indicate a close rela-
tionship of the two groups.

Platyischnopus in the Haustoriidae resembles the Sebidae in its
subchelate gnathopods but differs from sebids in its biramous third
uropods, much larger outer plates of the maxilliped and cleft telson

Genera of Sebidae

Seba Bate

Seba Bate, 1862.—Stebbing, 1906.—K. H. Barnard, 1957.
Teraticum Chilton, 1884.

Grimaldia Chevreux, 1889.

Paravalettia K. H. Barnard, 1916.

Type-species: S. innominata Bate, 1862 (monotypy).
Species: 7, probably cosmopolitan cold-water, littoral to bathyal.

Stegocephalidae

Ficures 160, 161

DraGgnosis.—Mouthparts projecting in a conical bundle below
head; mandible lacking palp and molar; accessory flagellum 1-
or 2-articulate; gnathopods feeble, simple or weakly subchelate. See
Amphilochidae, Lysianassidae.

DescripTion.—Peduncle of antenna 1 short; mouthparts variable;
upper lip bilobed, mandible Jacking palp and molar, lower lip without
inner lobes; palp of maxilla 1 large or small, with one or two articles,
maxilla 2 usually with two plates, outer plate occasionally absent,
often set on extended base so as to appear geniculate; maxillipedal
palp with three or four articles; gnathopods feeble, simple or scarcely
subchelate; coxae quadrate or rounded below, or first three often
acuminate below, first four usually forming a continuous shield,
coxa 1 never hidden by coxa 2; uropod 3 variable, peduncle long or
short, rami long or short; telson short, or medium, entire or cleft.

MARINE GAMMARIDEAN AMPHIPODA 437

Ficure 160.—Stegocephalidae: a, Stegocephalus inflatus Krdyer (Sars, 1895, pl. 69). Head:
b, Stegocephalus. Upper lip: c, Stegocephalus; d, Stegocephalina ingolf' Stephensen
(1925a). Mandible: ¢, Stegocephalus; f, Andaniella pectinata Sars (1895, pl. 72); g, pair,
Andaniexis abyssi (Boeck) (Sars, 1895, pl. 71). Lower lip: h, Stegocephalus. Maxilla
1: 1, Andaniexis; 7, Stegocephalus;, k, Phippsia gibbosa (Sars, 1895, pl. 71); J,
Stegocephalina.

438 U.S. NATIONAL MUSEUM BULLETIN 271

RELATIONSHIP.—The loss of both mandibular palp and molar is
approached only in the Phliantidae, Eophliantidae, and Pro-
phliantidae but in those families the accessory flagellum is absent.
The characteristic coxae and globular shape of stegocephalids plus
the conical grouping of the mouthparts are unmistakable characters.

Ficure 161.—Stegocephalidae: Maxilla 2: a, Phippsia gibbosa (Sars, 1895, pl. 71); 8,
Stegocephalina ingolfi Stephensen (1925a); c, Stegocephalus inflatus Krdyer (Sars, 1895,
pl. 69); d, Andaniexis abyssit (Boeck) (Sars, 1895, pl. 71). Maxilliped: e, Phippsia; f,
Andaniotes simplex K. H. Barnard (1930); g, Andaniella pectinata Sars (1895, pl. 72);
h, Stegocephalus. Telson: 1, Stegocephalus; 7, Andaniopsis nordlandica (Boeck) (Sars,
1895, pl. 72). Pereopod 4:k, Andaniopsis; l, Stegocephalus. Pereopod 5: m, Andaniopsis;
n, Stegocephalus; 0, Tetradeion crassum (Chilton, 1924). Pereopod 5: p, Stegocephalus.
Uropod 3: q, Stegocephalus; r, Andaniella. Pereopod 2: 5, Parandaniexis mirabilis Schel-
lenberg (1929b). Gnathopod 1: t, Stegocephalus. Gnathopod 2: u, Stegocephalus.

MARINE GAMMARIDEAN AMPHIPODA 439

Some Stenothoidae lack a mandibular palp and all have a shield-
like coxa 4 but the first coxa is always very small and hidden by the
second coxa and the accessory flagellum is a vestigial scale or absent.

The Amphilochidae have a small coxa 1 partially hidden by follow-
ing coxae and the mouthparts project in a quadrate bundle.

The general aspect, head, coxae, body shape, mouthpart bundle,
telson, uropods, pereopods, and gnathopods of Stegocephalidae suggest
affinities with the Acanthonotozomatidae but the absence of mandib-
ular palps and molars and the elytriform mandibular body distin-
guish the Stegocephalidae.

Stegocephalidae have considerable resemblance to Lysianassidae.
Some stegocephalid species have an elongate article 3 on gnathopod
2 which resembles that of those Lysianassidae having gnathopod 2
simple (not subchelate). However, most lysianassids have a mitten-
shaped or minutely chelate article 6. All Stegocephalidae lack mandib-
ular palps and molars, have some foliaceous portions on the maxillae,
have a 1- or 2-articulate accessory flagellum and a highly characteristic
configuration of coxae 1-4. This combination of characters never
occurs in Lysianassidae.

Andaniotes ingens Chevreux (1906c) probably should form the type
of a new genus because of its reduced first maxillary palp and narrow
article 2 of pereopod 4 in combination but in other special characters
it fits Huandania more than Andaniotes and so is transferred to the
former. Andaniotes simplex KK. H. Barnard (1930) also should be
distinguished as a new genus by its 3-, not 4-articulate maxillipedal
palp, having articles 1 and 2 coalesced.

Key to the Genera of Stegocephalidae

1. Mandible bearing multitoothed (often minutely) incisor (fig. 160f) .. . 2
Mandible bearing smooth incisor (fig. mie Fe NUE aes Ad UNIO aR TOMI IH inaete td Eve
2eeehelson cleht, (fe slGlle)ac Ausra es ye RAE SpE NECROSS Nene eae hey cers Fine
Melsonventuiren(fies Gl). nnn oe ete ek cere eee nae Swen cao aay
Be Waxilla 2 lacking outer plater. =.) 8) eR. h i) see fie ipuchivstecorenhalie
Maxilla 2 bearing outer plate... . Sita Wy oy ee isla ing
4. Palp article 2 of maxilliped produced sNsonnastiiliy (i 161e) . . Phippsia
Palp article 2 of maxilliped not produced . He SR Ae chet 3)
5. Palp of maxilla 1 biarticulate (figs. 1607,h) 6
Palp of maxilla 1 uniarticulate (fig. 1607) 8
6. Article 2 of pereopod 4 broad (fig. 1611) ies i
Article 2 of percopod 4 slender (fig. 1614) ..... . Gre rocentalopia
7. Outer plate of maxilla 2 gaping and geniculate (figs. 161a,b,c) . . Phippsiella
Outer plate of maxilla 2 normal (fig. 161d)... . . . Pseudandaniexis !
8. Article 2 of pereopod 4 slender (fig. 161k) PPM e Oy die He ROU aE one
Auuicle Ziolpercopod4ubroad (ie Ol) sys 2. ee ene eee lO

1 Telson unknown.

440

es

10.

11.

12.

13.

14.

15.

16.

17.

U.S. NATIONAL MUSEUM BULLETIN 271

Outer plate of maxilla 2 gaping and geniculate (figs. 161a,b,c).

Stegocephaloides
Outer plate of maxilla 2 normal (fig. 161d) ...... .. .. . Steleuthera
Upper and lower lips and maxilla 1 not elongate (figs. 160c,h), spines of outer
plate on maxilla 2 with hooks (fig. 16lc) . .... . .. . Stegocephalus
Upper and lower lips and maxilla 1 elongate (fig. 160d), spines of outer plate
on maxilla 2 lacking hooks (fig. 16ld) ..... .. . . Stegocephalina
Pereopod! 5 wath)seven articles Gigs: U6limin)i)2) 2 ee le
Pereopod 5 with three articles (fig. 16lo) .. . : . . . Tetradeion
Inner plate of maxilliped reaching end of palp ariacle 1 Ges, 161h).
Andaniopsis
Inner plate of maxilliped scarcely reaching base of palp article 1 (figs.
NGG) ae er Moegie, us}
Palp of maxilla 1 cmniaratioilate, annals 2 of mercopod 4 dence Ge, 161k).
Andaniella

Palp of maxilla 1 biarticulate, article 2 of pereopod 4 broad (fig. 1611).
Pseudandaniexis !
Te ee
Telson entire or apically slit ... . 5 co eaten
Flagellar article 1 of antenna 1 eabecmal to or ahora? than sacle: pedun-
cular article 5 of antenna 2 subequal to or shorter than article 4; pleonite 6
longer than peduncle of uropod 3. . . . . Andaniotes 2 Mreientanea)
Flagellar article 1 of antenna 1 much longer than peduncle; peduncular
article 5 of antenna 2 longer than article 4; pleonite 6 shorter than peduncle

of uropod3 . . Ba bait ane area
Pereopod 2 deumetly pabenclare Ge 161s) ... .. . . Parandaniexis
Pereopod 2simple .... MA AU tenn near MEAGRE rs rcs LG.
Palp of maxilla 1 Particulate’: SNE TMG aint Ce AN ane XS
Palp of maxilla 1 uniarticulate ........... =... Parandania

Genera of Stegocephalidae

Andaniella Sars

Andaniella Sars, 1895.—Stebbing, 1906.

Type-species: Andania pectinata Sars, 1882 (monotypy).

Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer
plate of maxilla 2 not geniculate nor gaping; inner plate of maxiliped
scarcely reaching base of palp article 1 (Andaniopsis), article 2 of palp
not produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad;
telson entire. Species: 1, boreal N. Atlantic, littoral to bathyal.

Andaniexis Stebbing

Andania Boeck, 1871 (homonym, Lepidoptera).
Andaniexis Stebbing, 1906 (new name).

Type-species: Amdania abyssi Boeck, 1871 (selected by Boeck,
1876). See Sars, 1895.

MARINE GAMMARIDEAN AMPHIPODA 44]

Mandibular incisor not toothed; palp of maxilla 1 biarticulate;
outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of
maxilliped not produced; article 2 of pereopod 3 slender, of pereopods
4-5 broad; telson entire. Species: 6, cosmopolitan, bathy-abysso-
pelagic.

Andaniopsis Sars
Andantopsis Sars, 1895.—Stebbing, 1906.

Type-species: Andania nordlandica Boeck, 1871 (monotypy). See
Sars, 1895.

Mandibular incisor toothed; palp of maxilla 1 uniarticluate; outer
plate of maxilla 2 not geniculate nor gaping; inner plate of mazxilliped
reaching distal end of palp article 1 (Andaniella), palp article 2 not
produced; article 2 of pereopods 3-4 slender, of article 5 broad; telson
entire. Species: 1, boreal N. Atlantic, littoral.

Andaniotes Stebbing. new synonymy

Andaniotes Stebbing, 1897; 1906.
Metandania Stephensen, 1925a.

Type-species: Anonyx corpulentus Thomson, 1882 (monotypy).
See Stebbing, 1888 (as Andania abyssorum).

Mandibular incisor not toothed; palp of maxilla 1 uniarticulate;
outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of
maxilliped not produced; article 2 of pereopod 3 slender, of pereopods
4—5 broad; telson cleft one third; flagellar article 1 of antenna 1 equal to
or shorter than peduncle; peduncular article 5 of antenna 2 equal to or
shorter than article 4; pleonite 6 longer than peduncle of uropod 3 (Hu-
andania). Species: 4, antarctic, N. Atlantic, littoral to bathyal.

Bathystegocephalus Schellenberg

Bathystegocephalus Schellenberg, 1926b.

Type-species: Stegocephalus globosus Walker, 1909 (monotypy).

Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer
plate of maxilla 2 absent; palp article 2 of maxilliped not produced;
article 2 of pereopods 3-4 slender, of pereopod 5 broad; telson cleft
one third. Species: 1, Indo-Pacific, S. Atlantic, bathypelagic.

Euandania Stebbing
Euandania Stebbing, 1899d; 1906.

Type-species: Andania gigantea Stebbing, 1888 (original designa-
tion).

442 U.S. NATIONAL MUSEUM BULLETIN 271

Mandibular incisor not toothed; palp of maxilla 1 uniarticulate;
outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of
maxilliped not produced; article 2 of pereopod 3 slender, of pereopod
4 either broad [or slender H. ingens (Chevreux)], of pereopod 5 broad;
telson cleft one third; flagellar article 1 of antenna 1 much longer than
peduncle; peduncular article 5 of antenna 2 longer than article 4; pleo-
mite 6 shorter than peduncle of uropod 8 (Andaniotes). Species: 2,
cosmopolitan, bathypelagic.

Parandania Stebbing
Parandania Stebbing, 1899d; 1906.

Type-species: Andania boecki Stebbing, 1888 (original designa-
tion).

Mandibular incisor not toothed; palp of maxilla 1 uniarticulate;
outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of
maxilliped not produced; article 2 of pereopod 3 slender, of pereopods
4—5 broad; telson entire. Species: 1, cosmopolitan, bathypelagic.

Parandaniexis Schellenberg
Parandaniexis Schellenberg, 1929b. —

Type-species: P. mirabilis Schellenberg, 1929b (monotypy).
Mandibular incisor not toothed; palp of maxilla 1 biarticulate;
outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of
maxiliped not produced; article 2 of pereopods 3-4 slender, of pereopod
5 broad; telson entire; pereopod 2 subchelate. Species: 1, tropical E.
Pacific, abyssal.
Phippsia Stebbing

Aspidopleurus Sars, 1895 (homonym, Pisces).
Phippsia Stebbing, 1906 (new name).

Type-species: Stegocephalus gibbosus Sars, 1882 (monotypy).
Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer
plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped
produced distomedially; article 2 of pereopods 3-4 slender, of pereopod
5 broad; telson cleft one third. Species: 2, arctic-boreal, bathy-
pelagic.
Phippsiella Schellenberg

Phippsiella Schellenberg, 1925a.

Type-species: Stegocephalus similis Sars, 1895 (monotypy).
Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer
plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped not

MARINE GAMMARIDEAN AMPHIPODA 443

produced; article 2 of pereopod 3 slender, of pereopods 4-5 broad;
telson cleft more than halfway. Species: 7, probably cosmopolitan
coldwater, littoral to abyssal.

Pseudandaniexis Nicholls

Parandaniexis Nicholls, 1938 (homonym, Amphipoda).
Pseudandaniexis Nicholls, 1938, Corrigenda (new name).

Type-species: Parandaniexis mixtus Nicholls, 1938 (original desig-
nation).

Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer
plate of maxilla 1 not geniculate nor gaping; inner plate of maxilliped
reaching proximal end of palp article 1 (Andaniopsis), palp article 2
of maxilliped unproduced; article 2 of pereopod 3 slender, of pereopod
5 broad; [telson broken and not clearly analyzed]. Species: 1,
antarctic, bathyal.

Stegocephalina Stephensen

Stegocephalina Stephensen, 1925a.

Type-species: SS. ingolfi Stephensen, 1925a (monotypy).

Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer
plate of maxilla 2 geniculate and gaping; palp article 2 of maxilliped
not produced; article 2 of pereopod 3 slender, of pereopods 4-5
broad; telson cleft halfway; upper lip, lower lip, and mazilla 1 elongate;
outer plate of marilla 2 lacking hooks (Stegocephalus). Species: 1,
boreal N. Atlantic, bathyal.

Stegocephaloides Sars
Stegocephaloides Sars, 1895.—Stebbing, 1906.

Type-species: Stegocephalus christianiensis Boeck, 1871 (original
designation).

Mandibular incisor toothed; palp of maxilla 1 uniarticulate; owter
plate of maxilla 2 geniculate, gaping (Steleuthera); palp article 2 of
maxilliped not produced; article 2 of pereopods 3-4 slender, of pereo-
pod 5 broad; telson cleft. Species: 7, cold-water, N. and S. Atlantic,
primarily bathyal.

Stegocephalopsis Schellenberg
Stegocephalopsis Schellenberg, 1925a.

Type-species: Cancer ampulla Phipps, 1774 (monotypy). See
Gurjanova, 1962.

285-135 O - 69 - 29

444 U.S. NATIONAL MUSEUM BULLETIN 271

Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer
plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped not
produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad;
telson cleft halfway or more. Species: 3, arctic-boreal, littoral to
bathyal.

Stegocephalus Kr¢éyer

Stegocephalus Kr@éyer, 1842.—Stebbing, 1906.

Type-species: S. inflatus Krgyer, 1842 (monotypy). See Sars, 1895.

Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer
plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped not
produced; article 2 of pereopod 3 slender, of pereopods 4-5 broad;
telson cleft halfway; upper lip, lower lip, and maxilla 1 not elongate;
spines of outer plate of mavilla 2 with hooks (Stegocephalina). Species:
2, subaretic-boreal, littoral to bathyal.

Steleuthera J. L. Barnard
Steleuthera J. L. Barnard, 1964a.

Type-species: S. maremboca J. L. Barnard, 1964a (original
designation).

Mandibular incisor toothed; palp of maxilla 1 untarticulate; outer
plate of maxilla 2 not geniculate (Stegocephaloides); palp article 2 of
maxilliped not produced; article 2 of pereopods 3-4 slender, of pereo-
pod 5 broad; telson poorly cleft. Species: 1, Peru, hadal.

Tetradeion Stebbing
Tetradeion Stebbing, 1899d; 1906.

Type-species: Cyproidia crassa Chilton, 1883 (original designa-
tion). See Hurley, 1955.

Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer
plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped
distomedially produced; article 2 of pereopods 3-4 slender, of pereopod
5 broad; pereopod 5 with only three articles; telson entire. Species: 1,
New Zealand, littoral.

Stenothoidae

FIGURE 162

Diaanosis.—Accessory flagellum absent or composed of one or
two vestigial articles; mandibular molar evanescent; coxa 1 very
small, always partially covered by following coxae; coxa 4 enlarged,

MARINE GAMMARIDEAN AMPHIPODA 445

shield-like, not posterodorsally excavate; uropod 3 uniramous;
outer lobes of maxilliped vestigial; telson entire; pereopod 3 with
article 2 slender. See Amphilochidae, Thaumatelsonidae, Cressidae,
Leucothoidae, Anamixidae, Phliantidae, Pagetinidae.

DescripTion.—Rostrum inconspicuous; accessory flagellum absent
or 1- or 2-articulate, vestigial; body smooth or carinate; coxa 1
small, hidden by following coxae; coxa 4 enlarged, shield-like, not
posterodorsally excavate; upper lip incised; mandible with weak,
sparsely spinose molar, not triturative, palp absent and when present
1-, 2-, or 3-articulate; lower lip usually with inner lobes amalgamated,
outer lobes with blunt extremities; maxilla 1 with 2- or 1-articulate
palp; maxilla 2 small, stout; maxillipeds slender, with outer lobes
vestigial; gnathopods usually powerful, subchelate, occasionally
feeble, gnathopod 1 often simple; uropod 3 uniarticulate, ramus
2-articulate; telson of medium length, entire; pereopod 3 with slender
article 2.

ReLarionsHip.—The Amphilochidae resemble stenothoids but
have a biramous uropod 3 and well-developed outer lobes on the
maxilliped.

The Thaumatelsonidae have some urosomal segments coalesced
and the telson usually strongly thickened. Perhaps the Thaumatel-
sonidae deserve only subfamily status as members of the Stenothoidae.
The Cressidae differ from the Stenothoidae by the coalescence of
the telson with pleonite 6 and the expanded article 2 of pereopod 3.

Classification of Genera of Stenothoidae

The genera of Stenothoidae seem to be segregated from each
other by very unsatisfactory means, such as conditions of mandibular
palp, maxilla 1, accessory flagellum, and to some extent breadth of
article 2 on pereopods 3-5. In other familes these characters (see
following key) often distinguish species-groups into natural genera
but in the Stenothoidae such “pigeon-holing” throws together
species with great diversity in gnathopods. Several genera thus
represent grades of structure rather than clades. In these small
organisms mandibular palps are very difficult to work with taxo-
nomically as palps are hard to observe and may break off during
dissection. There is a strong possibility that some species may indeed
have palps in some populations and lack them in others. It is prudent
to mount the head of the animal on a slide and look for the mandib-
ular palp before dissection. Often the articulation line separating
palp articles 1 and 2 of the first maxillae is difficult to resolve.

Shoemaker (1955) in his key to Stenothoidae shows the logic in
reestablishing the genera Prometopa and Microstenothoe which were

MARINE GAMMARIDEAN AMPHIPODA 447

formerly synonymous with Metopa and Stenothoe, for his key points
out the fact that Proboloides and Metopoides are characterized only
by the presence or absence of an accessory flagellum. Because Pro-
metopa and Microstenothoe bear accessory flagella, it is inconsistent
to reduce them without also eliminating the genera Proboloides and
Metopoides.

Nomenclatural Changes in Stenothoidae

Mesostenothoides Gurjanova (1938) is removed to Stenothordes.

Metopoides Della Valle (1893) (Stebbing, 1906) is reestablished.

Metopella nasutigenes (Stebbing, 1888) is removed to Probolisca.

Microstenothoe Pirlot (1933b) is reestablished.

Parametopella stelleri Gurjanova (1948) omitted from index by J. L.
Barnard (1958a).

Parametopella minuta (Holmes), erroneously assigned to this genus, is
removed to Stenothoe.

Prometopa Schellenberg (1931) is reestablished.

Stenothoe minuta Holmes, (Stebbing, 1906; Kunkel, 1918), erroneously
assigned to Parametopella previously.

Stenula J. L. Barnard (1962c) was erected for species previously
assigned to Mesostenothoides.

Key to the Genera of Stenothoidae
(emended after Shoemaker, 1955)

1. Pereopods 4 and 5: article 2 linear (figs. 162¢,) . .. ren Key vA
Pereopod 4: article 2 linear (fig. 162v); pereopod 5, mite D ended (figs.
IGAS)) 6 5 i HUG 2 cheb aan IONS
Pereopods 4 and 5: “onal 2 oanendad Gres 162a, a Sia De ne Maal iat] Bente
KEY A

lemelalprotamaxidla taunianrticmlaten (hg. G2q)) a) 4) aes mien Gea eign) Sec eee
Palp of maxilla 1 biarticulate (fig. 162h) ...... .. .. . .Probolisca
2. Mandibular palp absent (fig. 162c)......... ... . Parametopella
Mandibular palp l-articulate .. . ..... . . Metopelloides
Mandibular palp 2- or 3-articulate (fiz. 162d) Tae oti an ee ne Ee COpella:

ass

Figure 162.—Stenothoidae: a, Stenothoe marina (Bate) (Sars, 1895, pl. 80). Upper lip:
b, Stenothoe. Mandible: c, Stenothoe; d, Proboloides gregarius (Sars, 1895, pl. 84). Lower
lip: e, Stenothoe; f, Proboloides. Maxilla 1: g, Metopa alderi (Bate) (Sars, 1895, pl. 86);
h, Stenothoe. Maxilla 2:1, Stenothoe. Maxilliped: 7, Stenothoe; k, Metopa; 1, Probolotdes.
Gnathopod 1: m, Metopa pusilla (Sars, 1895, pl. 90); n, Metopa alderi; 0, Metopa robusta
(Sars, 1895, pl. 96). Gnathopod 2: p, Metopella longimana (Boeck) (Sars, 1895, pl. 97); ¢,
Metopa alderi. "Telson: r, Stenothoe. Pereopod 5:5, Mesometopa neglecta (Hansen) (Sars,
1895, pl. 97); t, Metopella; u, Stenothoe. Pereopod4:v, Metopella. Uropod 3: w, Stenothoe.

448

KEY B

1. Palp of maxilla 1 uniarticulate .
Palp of maxilla 1 biarticulate. . Bilan
2. Mandibular palp l-articulate or absent .
Mandibular palp 2- or 3-articulate .

KEY C

1. Palp of maxilla 1 uniarticulate .
Palp of maxilla 1 biarticulate

2. Mandibular palp absent.
Mandibular palp l-articulate
Mandibular palp 2- or 3-articulate .

3. Accessory flagellum absent .
Accessory flagellum l1-articulate

4. Mandibular palp absent
Mandibular palp present

5. Accessory flagellum 1l-articulate
Accessory flagellum absent

6. Mandibular palp 1-articulate
Mandibular palp 2- or 3-articulate .

7. Accessory flagellum absent
Accessory flagellum 1- or 2-articulate

Genera of Stenothoidae

Mesometopa Gurjanova
Mesometopa Gurjanova, 1938.

Type-species:

U.S. NATIONAL MUSEUM BULLETIN

27:1

.2

. Mesoproboloides

. Stenothoides
. Mesometopa

5B

Bes LA
. Parametopa
. Stenula

12 epee

. Metopa

. Prometopa
ao
Microstenothoe
. Stenothoe

. Prostenothoe
En rte of
Proboloides
Metopoides

Metopa esmarki Boeck, 1872 (original designation).

Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate;
article 2 of pereopod 4 linear, of article 5 expanded. Species: 4,

arctic-boreal, littoral.

Mesoproboloides Gurjanova

Mesproboloides Gurjanova, 19388.

Type-species:
designation).

Metopella cornuta Schellenberg,

1926a

(original

Mandibular palp 3-articulate; palp of maxilla 1 biarticulate; arti-
cle 2 of pereopod 4 linear, of pereopod 5 expanded. Species: 2,

subantarctic, littoral.
Metopa Boeck
Metopa Boeck, 1871.—Stebbing, 1906.

Metopina Norman, 1900b (homonym, Diptera).
Sthenometopa Norman, 1902.

Type-species:

1876). See Gurjanova, 1948, 1951; Shoemaker, 1955.

Leucothoe clypeata Kroyer, 1842 (selected by Boeck,

MARINE GAMMARIDEAN AMPHIPODA A49

Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate;
article 2 of pereopods 4—5 expanded; accessory flagellum absent (Pro-
metopa). Species: 49, subarctic-boreal, littoral to abyssal, (one species
Prometopa tuberculata is subantarctic).

Metopella Sars

Metopella Sars, 1985.—Stebbing, 1906.

Type-species: Metopa longimana Boeck, 1871 (selected by Gur-
janova, 1938). See Sars, 1895.

Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate;
article 2 of pereopods 4-5 linear. Species: 8, arctic-boreal, littoral.

Metopelloides Gurjanova
Metopelloides Gurjanova, 1938, 1951.

Type-species: Metopella micropalpa Shoemaker, 1930 (original
designation).

Mandibular palp 1l-articulate; palp of maxilla 1 untarticulate;
article 2 of pereopods 4-5 linear. Species: 9, arctic-boreal, littoral,

Metopoides Della Valle

Metopoides Della Valle, 1893.—Stebbing, 1906.

Type-species: Metopa magellanica Stebbing, 1888 (selected by
Gurjanova, 1938).

Mandibular palp 2- or 3-articulate; palp of maxilla 1 biarticulate;
article 2 of pereopods 4-5 expanded; accessory flagellum 1-articulate
(Proboloides). Species are combined with Proboloides; most Meto-
poides are subantarctic.

Microstenothoe Pirlot
Microstenothoe Pirlot, 1933b.
Type-species: M. ascidiae Pirlot, 1933b (original designation).
Mandibular palp absent; palp of maxilla 1 biarticulate; article

2 of pereopods 4—5 expanded; accessory flagellum 1-articulate (Steno-
thoe). Species combined with Stenothoe.

Parametopa Chevreux
Parametopa Chevreux, 1901b.
Type-species: P. kervillei Chevreux, 1901b (original designation).
See Chevreux and Fage, 1925.
Mandibular palp absent; palp of maxilla 1 uniarticulate; article 2
of pereopods 4—5 expanded. Species: 3, subarctic and warm-temper-
ate in Atlantic, littoral.

450 U.S. NATIONAL MUSEUM BULLETIN 271

Parametopella Gurjanova
Parametopella Gurjanova, 1938, 1951.

Type-species: Stenothoe cypris Holmes, 1905 (original designation).
See Kunkel, 1918.

Mandibular palp absent; palp of maxilla 1 uniarticulate; article
2 of pereopods 4-5 linear. Species: 3, subarctic-boreal, littoral.

Probolisca Gurjanova
Probolisca Gurjanova, 1938.

Type-species: Metopa ovata Stebbing, 1888 (original designation).
Mandibular palp 2- or 3-articulate; palp of maxilla 1 biarticulate;
article 2 of pereopods 4-5 linear. Species: 3, antiboreal, littoral.

Proboloides Della Valle

Proboloides Della Valle, 1893.—Stebbing, 1906.—K. H. Barnard, 1932.
Proboltella Walker, 1906b, 1907.

Type-species: Metopa gregaria Sars, 1882 (selected by Gurjanova,
1938). See Sars, 1895.

Mandibular palp 2- or 3-articulate; palp of maxilla 1 biarticulate;
article 2 of pereopods 4-5 expanded; accessory flagellum absent
(Metopoides). Species (incl. Metopoides): 34, bipolar, littoral to
abyssal.

Prometopa Schellenberg

Prometopa Schellenberg, 1926a.

Type-species: P. tuberculata Schellenberg, 1926a (monotypy).

Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate;
article 2 of pereopods 4—5 expanded; accessory flagellum 1-articulate
(Metopa). Species: 1; included with Metopa.

Prostenothoe Gurjanova
Prostenothoe Gurjanova, 1938.

Type-species: P. sextonae Gurjanova, 1938 (original designation) ;
1951.
Mandibular palp 1-articulate; palp of maxilla 1 biarticulate; article
2 of pereopods 4—5 expanded. Species: 1, Japan, Okhotsk Sea,
littoral.
Stenothoe Dana

Stenothoe Dana, 1852b.—Stebbing, 1906.
Probolium Costa, 1853c, 1857.

Montagua Bate, 1857a (homonym, Decapoda).
Montaguana Chilton, 1883.

MARINE GAMMARIDEAN AMPHIPODA 451

Type-species: S. validus Dana, 1853 (selected by monotypy of
Dana, 1853). See Chevreux and Fage, 1925.

Mandibular palp absent; palp of maxilla 1 biarticulate; article 2
of pereopods 4-5 expanded; accessory flagellum absent (Microstenothoe).
Species: 32, cosmopolitan, littoral to bathyal.

Stenothoides Chevreux

Stenothoides Chevreux, 1900!—J. L. Barnard, 1962c.
Mesostenothoides Gurjanova, 1938.
Type-species: S. perrieri Chevreux, 1900 (monotypy).
Mandibular palp 1-articulate or absent; palp of maxilla 1 uni-
articulate; article 2 of pereopod 4 linear, of article 5 expanded. Spe-
cies: 6, subarctic-boreal, littoral.

Stenula J. L. Barnard
Stenula J. L. Barnard, 1962c.

Type-species: Stenothoides latipes Chevreux and Fage, 1925 (ori-
ginal designation).

Mandibular palp 1-articulate; palp of maxilla 1 uniarticulate;
article 2 of pereopods 4-5 expanded. Species: 10, arctic-boreal,
littoral to bathyal.

Stilipedidae

FicgurEs 163, 164, FRONTISPIECE

Dracnosis.—Accessory flagellum absent or vestigial, less than
3-articulate; mandible broad and lacking molar; maxillae foliaceous
(figs. 163¢,d); gnathopods simple but not especially slender. See
Pardaliscidae Hyperiopsidae, Laphystiopsidae, Astyridae.

DescripTion.—Peduncles of antennae very short, accessory flagel-
lum vestigial or absent; upper lip asymmetrically bilobed; mandible
very broad, pardaliscid in appearance, palp 3-articulate, thin, lacking
molar; lower lips variable; maxilla 1 variable, some parts always
foliaceous; maxilla 2 foliaceous; plates of maxilliped large, palp
variable in length; gnathopods simple but not especially slender;

,coxae long; uropod 3 biramous, peduncle short, rami elongate,
lanceolate; telson apically emarginate, short.

ReELATIONSHIP.— Differing from the Acanthonotozomatidae by the
foliaceous maxillae (except Mazilliphimedia in that family) and the
mouthparts not being arranged in a conical bundle.

The Astyridae differ from the Stilipedidae by the weaker foli-
aceousness of the first maxilla but intergradation occurs in the genus

452 U.S. NATIONAL MUSEUM BULLETIN 2711

Alexandrella assigned to the Stilipedidae, for its first maxillae are not
as foliaceous as those of the type-genus Stilipes. The palp is especially
strongly geniculate, resembling species of Hyperiopsidae. Astyridae
bear a distinct but nontriturative mandibular molar. Astyrovdes is
removed from the Astyridae and synonymized with Alexandrella.

The Laphystiopsidae have normal maxillae, a mandibular molar,
and small maxillipedal plates.

Ficure 163.—Stilipedidae: a, lexandrella dentata Chevreux (1912b). Maxilla 1: 3,
Alexandrella; c, Stilipes sanguineus (Hurley, 1954e). Maxilla 2: d, Stilipes. Maxilliped:
e, Alexandrella; f, Stilipes.

MARINE: GAMMARIDEAN AMPHIPODA 453

The absence of a mandibular molar distinguishes Stilipedidae from
the Synopiidae and Hyperiopsidae. Stilipedids resemble the Par-
daliscidae in the mandible but differ by the long coxae and the
foliaceous second maxillae. The first maxillary palp of some pardali-
scids is foliaceous but not the inner plate. Pardaliscids have a multi-
articulate accessory flagellum (except Halicoides).

Ficure 164.—Stilipedidae: Mandible: a,b, Stilipes sanguineus (Hurley, 1954e). Upper
lip: c, Alexandrella dentata Chevreux (1912b); d, Stilipes sanguineus. Lower lip: e, Stilipes
sanguineus; f, Alexandrella. Gnathopod 2: g, Alexandrella. Gnathopods 1, 2: h,1,
Stilipes sanguineus. Telson: 7, Stilipes distincta Holmes (1908). Uropod 3: k, Stzlipes
distincta.

Key to the Genera of Stilipedidae

1. Palp of maxilliped greatly exceeding outer plate (fig. le upper lip very

weakly emarginate (fig. 164d) ... . . . . . Stilipes
Palp of maxilliped not aes outer plat (i 163¢),. upper lip deeply
inciseds (tig G40) es 3 : ... . . . Alexandrella

Genera of Stilipedidae

Alexandrella Chevreux, new synonymy

Alexandrella Chevreux, 1911c; 1912a; 1912b.
Astyroides Birstein and Vinogradova, 1960.

454 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: A. dentata Chevreux, 1912a; 1912b (designated by
Chevreux, 1912a).
Species: 2, antarctic to N.W. Pacific, littoral to hadal.

Stilipes Holmes

Stilipes Holmes, 1908.
Cacao K. H. Barnard, 1931; 1932.
Type-species: S. disltincta Holmes, 1908 (original designation). See

Shoemaker, 1964.
Species: 3, N.E. Pacific, antiboreal, littoral to bathyal.

Synopliidae, revised
{including Tironidae]

Figures 165-167

Dracnosis.—Head massive,* and often galeate, produced into a
downturned or deflexed rostrum; gnathopods feeble, scarcely sub-
chelate, often simple; telson elongate (rarely short), but always as
long as peduncle of uropod 1; eyes when present coalesced. See
Astyridae, Vitjazianidae, Argissidae, Hyperiopsidae, Oedicerotidae.

Descrietion.—The characters of this family are so subtle that
they are practically indefinable, yet taxonomists have little trouble
in recognizing a synopiid. An inclusive diagnosis has to be inter-
spersed with the words ‘‘most species” since there are so many excep-
tions to any pair of character alternatives, yet the majority of species
shares anyone of the following characters and all species share the
majority of them.

Antenna 1 with multiarticulate accessory flagellum (except Jeddo);
base of primary flagellum conjoint in male; eyes when present coal-
esced dorsally; mandible with poorly developed, slender palp (except
Synopia), molar well developed, often extraordinarily large, smooth
and dominating mandible, article 3 usually shortened considerably
(except in Bruzeliopsis); outer lobes of lower lip frequently notched;
mouthparts otherwise like basic gammaridean; coxa 4 never much
larger than coxa 3; coxae long (as contrasted with Pardaliscidae) ; uro-
pods 1-2 with outer ramus shortened; telson cleft (except in Bruzelia,
Bruzeliopsis, and some Synopia), elongate (except Synopia), as long
as peduncle of uropod 1; head massive, produced into a deflexed or

*As tall as or taller than long (length from posterior margin of head to anterior
margin of lateral lobe), head (except Tiron) as long as pereonites 1-3 combined.

MARINE GAMMARIDEAN AMPHIPODA 455

f ee |

Figure 165.—Synopiidae: a, Tiron spiniferum (Stimpson) (Sars, 1895, pl. 140, as T.
acanthurus); b, Syrrhottes serratus Sars (1895, pl. 137). Accessory flagella: c, Tiron; d,
Syrrhoe crenulata Goés (Sars, 1895, pl. 136). Mandible: e, Synopia variablis Spandl
(J. L. Barnard, 1965).

456 © U.S. NATIONAL MUSEUM BULLETIN 271

ey
yecredg )

Ni \
\ \ | \\

ppygyhy. |

“ie

Figure 166.—Synopiidae: a, Syrrhoe crenulata Goés (Sars, 1895, pl. 136); b, Bruzelia typica
Boeck (Sars, 1895, pl. 138). Upper lip: ¢, Tiron spiniferum (Stimpson) (Sars, 1895, pl.
140, as T. acanthurus). Mandible: d, Bruzelia; e, Tiron. Lower lip: f, Syrrhottes serratus
Sars (1895, pl. 137); g, Tiron. Telson: h, Bruzelia tuberculata Sars (1895, pl. 139).

MARINE GAMMARIDEAN AMPHIPODA 457

downturned rostrum; pereonites 1-4 often very short; gnathopods
feeble, slender, scarcely subchelate; eyes when present dorsally
confluent.

RELATIONSHIP.—Synoplids differ from the basic gammaridean
mainly by the massive head with large rostrum, poorly developed
mandibular palp in most genera, feeble gnathopods, and elongated
telson. From the Eusiridae they differ by the combination of the
same characters, except for the elongate telson. They are similar to
the Oedicerotidae in head, eyes, and general appearance, but all
Oedicerotidae have a vestigial or no accessory flagellum, usually
powerful gnathopods and short, uncleft telson.

Synopiidae differ from Pardaliscidae by the possession of a man-
dibular molar but often appear similar in other ways, except that
the coxae of Pardaliscidae are always very short.

The Vitjazianidae differ from Synopiidae by the small, poorly
rostrate head.

The Argissidae have a normal gammaridean head, practically no
rostrum, and a unique pattern in the shapes of coxae 1-4.

Astyridae resemble Synoptidae closely and differ from them in
the occurrence of just one or two articles in the accessory flagellum,
the setose, laminate, and not triturative mandibular molar. All
synoplids, except Bruzeliopsis, have a reduced palp article 3 on the
mandible, whereas astyrids have an elongated article 3. All Astyridae
have short telsons and widely spaced outer lobes on the lower lip.

A revision of Synopiidae is presented herein and where generic
composition has been changed, the species have been listed.

Key to the Genera of Synopiidae

1. Mandibular molar very large and smooth (minutely setulose or fuzzy oc-
casionally), completely dominating mandible, body of mandible extremely
bulkyacsubclobularydwartinejoalpn (igs GGG) 18 je ln ee 2

Mandible as in first triplet but palp absent . .. . PARKA edd
Mandibular molar of medium size, occasionally emeoth os minutely fuzzy
but usually triturative, body of mandible usually stout but not extra-
ordinarily bulky or subglobular, palp relatively strong (fig. 166e). . . 5

2. Coxa 3 strongly expanded anteroposteriorly and much larger than coxa 4
(fig. 166a) (more than twice as much surface area), coxa 3 with strong
posterodorsal excavation dorsal to expansion, article 1 of antenna 1 very
elongate and bearing long subconical distal process, articles 2 and 3 short
and subequal in length, coxa 4 very small and shaped like a comma (fig.
OOO) eae Ree aii . . . . . Bruzeliopsis

Coxae 3 and 4 atineqinell | in size ‘oad Teneal coxa 3 never with more than 1.5—
times as much surface area as 4, eoxa 3 weakly expanded or not expanded
distally, lacking distinct posterodorsal excavation, coxa 4 adze-shaped or
quadrate, its posteroventral lobe or midposterior cusp distinetly directed

458

10.

U.S. NATIONAL MUSEUM BULLETIN 271

posteriorwards and at right angles to anterior margin, posterodorsal
excavation almost right-angular, articles 2 and 3 of antenna 1 even if
shortened always together as long as article 1, distal tooth of latter if
presenti shortzandihOokc likey men ee any oer amee n n ea
MelSONVeN tires pedicels kena ea dieg ee Aalst 5 tice) Cac Oe SEEN Ta ee es a 4
Telson cleft .... . SORA ah, ae Gyrrhoites
Palms of gnathopods recta: one or ro corrate spines, coxa 4 adze-shaped,
tooth distinctly posteroventral . ... . .. . . . Bruzelia
Palms of gnathopods with 4 nonserrate hint: spines, coxa 4 rectangular and
bearing midposterior cusp . . . . . .Stephobruzelia, new genus
Both telson and peduncle of aoa 3 very short and subequal in length,
mandibular palp extremely stout (fig. 165e). . .... . . . .Synopia
Telson elongate, twice as long as peduncle of uropod 3 even when peduncle
elongate, mandibular palp slender (figs. 166d,e) .... . Pa RAY ck | cro AG)
Coxa 3 strongly expanded distally, some portion of distal half at least twice
as broad as proximal end, expansion directed posteriorly, posterodorsal
margin strongly excavate, coxa 4 much smaller than 3, comma-shaped and
usually with less than half as much surface area as coxa3d...... 7
Coxa 3 rectangular or quadrate, scarcely expanded distally, occasionally
with anteroventral corner attenuated only, posterior margin nearly parallel
with anterior margin, coxa 4 almost as long as 3, its surface area scarcely
smaller than that of coxa 3, coxa 4 adze-shaped or chisel-shaped . . . 9
Coxa 3 rectangular or subquadrate, scarcely expanded distally, coxa 4
noticeably shorter than 3 and weakly comma-shaped .
some species a Tiron
Article 6 of gnathopods linear, slender, ae simple, lacking strong
serrate spines... . .. . . . Pseudotiron
Article 6 of gnathopods, if ample ror feat or 2 Hechenmulan either ovate and
tumid or bearing distinct transverse palms, palms bearing one or two
serrate spines .. . Ae OU eames
Telson cleft more than ellinnee “elias of apeunapades teens . Syrrhoe
Telson cleft less than 20%, palms of gnathopods oblique or indistinct.
Ileraustroe, new genus
Article 6 of both gnathopods linear, slender, een simple, lacking
strong serrate spines .. . . . Tiron
Article 6 of gnathopod 1 tumid but ail 6 of Dthoped 2 2 pertcetly linear
and rectangular . . Syrrhoe fimbriatus! Stebbing and Robertson, 1891
Article 6 of both gnathopods, if simple not linear or rectangular, either ovate
and tumid or bearing transverse palms, palms bearing one or two serrate

spines... Air errcanuises, wll)
Palms of Ereihonodl trmavemel engin Hose 6 one very taree spine giving
chelate appearance to palms. . . . . . . . . Garosyrrhoe
Palms of gnathopods oblique or Gpsolescent Heating one or two serrate
spines but not appearing chelate ........ .. .. . Austrosyrrhoe

' Currently assigned to Austrosyrrhoe.

Genera of Synopiidae

Austrosyrrhoe K. H. Barnard

Austrosyrrhoe K. H. Barnard, 1925.

MARINE GAMMARIDEAN AMPHIPODA 459

Type-species: A. crassipes K. H. Barnard, 1925 (monotypy).
Poorly known species.

Composition: A. septentrionalis Stephensen, 1931.

Mandible of medium size, molar of medium size and not dominating
mandible, apparently weakly triturative or fuzzy; coxa 3 rectangular
or subquadrate, scarcely expanded distally, posterior margin nearly
parallel with anterior margin and not strongly excavate; coxa 4 adze-
shaped, nearly as long as coxa 8, its surface area scarcely smaller than
coxa 3; article 6 of gnathopods not a perfectly linear rectangle, palms
oblique or obsolescent and bearing one or two large serrate spines;
telson elongate and deeply cleft. Species: 2, Atlantic, bathyal-
abyssal.

Bruzelia Boeck

Bruzelia Boeck, 1871.—Stebbing, 1906.

Ficure 167.—Synopiidae: Tiron spiniferum (Stimpson) (Sars, 1895, pl. 140, as 7. acan-
thurus): a,b, maxillae 1, 2; c, maxilliped; d,e, gnathopods 1, 2; f, uropod 3; g, telson.
Syrrhoe crenulata Goés (Sars, 1895, pl. 136): h,i, gnathopods 1, 2. Synopta scheeleana
Bruzelius (Stebbing, 1888): 7,k, gnathopods 1, 2; /, pereopod 2.

285-135 O - 69 - 30

460 U.S. NATIONAL MUSEUM BULLETIN 271

Type-species: 8B. typica Boeck, 1871 (monotypy). See Sars, 1895.

Composition: B. australis Stebbing, 1910; B. diodon K. H. Barnard,
1916; B. tuberculata Sars, 1882, 1895.

Mandible enlarged, molar enlarged, dominating mandible, smooth
or fuzzy; coxa 3 rectangular, not distally expanded except for antero-
ventral cusp, posterior margin parallel with anterior margin and not
excavate; coxa 4 nearly as long as coxa 3, surface area subequal to
coxa 3, adze-shaped; article 6 of gnathopods trapezoidal or subovate,
palms oblique, bearing one or two large serrate spines; telson elongate
and entire. Species: 4, probably cosmopolitan, mainly cold-water,
bathyal-abyssal (B. australis is littoral),

Bruzeliopsis Chevreux

Bruzeliopsis Chevreux, 1911b.

Type-species: B. albertt Chevreux, 1911b (original designation).

Composition: B. turba J. L. Barnard, 1964a; Bruzelia cuspidata
J. L. Barnard, 1962d.

Mandible enlarged, molar enlarged, dominating mandible, smooth
or fuzzy; coxa 3 distally expanded, with broad quadrate posterior
lobe, posterodorsal margin excavate; coxa 4 much smaller than 3,
comma-shaped; article 6 of gnathopods trapezoidal or subovate, palms
oblique, bearing one or two nonserrate or distally flagellate locking
spines; telson elongate and entire or minutely cleft apically; article 1
of antenna 1 elongate and bearing conical distal cusp. Species: 3,
Atlantic, bathyal-abyssal.

Garosyrrhoe J. L. Barnard
Garosyrrhoe J. L. Barnard, 1964a.

Type-species: Syrrhoites bigarra J. L. Barnard, 1962a (original
designation).

Mandible of medium size, molar of medium size and not dominating
mandible, apparently weakly triturative or fuzzy; coxa 3 softly rec-
tangular, posterior margin nearly parallel with anterior margin and
not excavate; coxa 4 expanded midposteriorly, posterodorsal margin
sloping, not concave but giving appearance of posterodorsal excavation,
coxa 4 with larger surface area than coxa 3; article 6 of gnathopods
trapezoidal or subovate, palms transverse and each defined by one large
serrate spine lending chelate appearance; telson elongate and deeply
cleft. Species: 1, California, littoral.

Ileraustroe, new genus

Type-species: Austrosyrrhoe ilergetes J. L. Barnard, 1964a (present
selection).

MARINE GAMMARIDEAN AMPHIPODA 461

Composition: Austrosyrrhoe ?torpens J. L. Barnard, 1964a (not
J. L. Barnard, 1962d) (=new species).

Mandible of medium size, molar of medium size or slightly enlarged
and not dominating mandible [palp large], fuzzy; coxa 3 strongly
expanded distally, with midposterior or posteroventral lobe, postero-
dorsal margin excavate; coxa 4 much smaller than 3, comma-shaped;
article 6 of gnathopods trapezoidal or subovate, palms oblique, bearing
at least one large serrate spine; telson elongate but cleft only 20 percent
or less. Species: 2, Mediterranean, Caribbean, abyssal.

Jeddo J. L. Barnard
Jeddo J. L. Barnard, 1962d.

Type-species: J. sumplisyrrhis J. L. Barnard, 1962d (original
designation).

Mandible enlarged, molar enlarged, dominating mandible, smooth
or fuzzy; mandibular palp absent; coxa 3 distally expanded, with large
posteroventral lobe, posterodorsal margin excavate; coxa 4 much
smaller than 3, hemi-ovate; article 6 of gnathopods essentially linear
and rectangular but not strongly elongate, simple, lacking distinct
locking spines; telson elongate and cleft about halfway. Species: 1,
S. Atlantic, deep bathyal.

Pseudotiron Chevreux

Pseudotiron Chevreux, 1895.—Stebbing, 1906.

Type-species: P. bouvieri Chevreux, 1895 (original designation).

Mandible of medium size, molar of medium size, columnar and
triturative; coxa 3 strongly expanded distally, with quadrate postero-
ventral lobe, posterodorsal margin forming quadrate excavation, coxa 4
much smaller than 3, comma-shaped; article 6 of gnathopods a nearly
perfectly linear rectangle, simple and lacking spine(s) ; telson elongate
and deeply cleft. Species: 3, Indo-Atlantic, Mediterranean, bathy-
abyssopelagic.

Stephobruzelia, new genus

Type-species: Bruzelia dentata Stephensen, 1931 (present selection).

Mandible enlarged, molar enlarged, dominating mandible, smooth
or fuzzy; coxa 3 rectangular, not distally expanded except for antero-
ventral cusp, posterior margin parallel with anterior and not excavate;
coxa 4 as long as and as large as coxa 3, rectangular, with midposterior
cusp; article 6 of gnathopods trapezoidal or subovate, palms oblique,
bearing four large, nonserrate, distally blunt spines; telson elongate
and entire. Named for K. Stephensen. Species: 1, subarctic, bathyal.

462 U.S. NATIONAL MUSEUM BULLETIN 271

Synopia Dana
Synopia Dana, 1852b.—Stebbing, 1906.

Type-species: S. ultramarina Dana, 1853 (present selection).
See Stebbing, 1888 (as S. scheeleana).

Mandible of medium size, molar of medium size, triturative or
fuzzy, palp extremely tumid (fig. 165e); coxa 3 strongly expanded dis-
tally, with quadrate posteroventral lobe, posterodorsal margin
excavate; coxa 4 much smaller than 3, very short, weakly comma-
shaped; article 6 of gnathopods slender or tumid, subovate, not
perfectly linear, simple, posterior margins setose but lacking distinct
locking spines, dactyl of gnathopod 2 vestigial (fig. 167k); telson and
peduncle of uropod 8 both short and subequal in length, telson cleft or
entire, distally castellate in forms with entire telson; mazillipeds
foliaceous. Species: 3, tropical, epipelagic.

Syrrhoe Goés
Syrrhoe Goés, 1866.—Stebbing, 1906.

Type-species: S. crenulata Goés, 1866 (selected by Boeck, 1876).
See Sars, 1895.

Mandible of medium size, molar of medium size and not dominating
mandible, apparently weakly triturative or fuzzy; coxa 3 strongly
expanded distally, with posteroventral lobe, posterodorsal margin
excavate; coxa 4 much smaller than 3, comma- or weakly adze-shaped ;
article 6 of gnathopods trapezoidal, subovate and elongate, palm
nearly transverse and armed with serrate spine not giving chelate
appearance; telson elongate and deeply cleft. Species: 8, probably
cosmopolitan, littoral to bathyal.

Syrrhoites Sars
Syrrhoites Sars, 1895.—Stebbing, 1906.

Type-species: Bruzelia serrata Sars, 1879 (original designation).
See Sars, 1895.

Composition: S. anaticauda K. H. Barnard, 1930; Kinda lorida
J. L. Barnard, 1962d; S. pacificus Nagata, 1965; S. pusilus Enequist,
1950; K. sorpresa J. L. Barnard, 1962d; S. tenellus K. H. Barnard,
1925; S. terceris J. L. Barnard, 1964a; S. walkeri Bonnier, 1896.

Mandible enlarged, molar enlarged, dominating mandible, smooth
or fuzzy; coxa 3 rectangular, not distally expanded, posterior margin
parallel with anterior margin and not excavate; coxa 4 nearly as long
as coxa 3, surface area subequal to coxa 3, adze-shaped; article 6 of
gnathopods trapezoidal or subovate, palms oblique, bearing one or
two large, nonserrate or distally flagellate locking spines; telson

MARINE GAMMARIDEAN AMPHIPODA 463

elongate and deeply cleft. Species: 9, cosmopolitan, cold-water,
deep littoral to abyssal.

Tiron Liljeborg

Tiron Liljeborg, 1865.—Stebbing, 1906.—Shoemaker, 1955.
Tessarops Norman, 1868 (homonym, Arachnida).

Type-species: Lysianassa spinifera Stimpson, 1853 (monotypy
and subsequent synonomy). See Sars, 1895 (as T. acanthurus).

Mandible of medium size, molar of medium size, strongly projecting,
columnar and triturative; coxa 3 softly rectangular, posterior margin
almost parallel with anterior margin and not strongly excavate; coxa
4 variable, typically adze-shaped and almost as long as coxa 3, surface
area of coxa 4 nearly equal to coxa 3; rarely coxa 4 distinctly shorter
and smaller than 3 and weakly comma-shaped; article 6 of gnathopods
a nearly perfectly linear rectangle, simple and lacking distinct locking
spine(s); telson elongate and deeply cleft. Species: 8, cosmopolitan,
littoral (rarely to bathyal).

Superfamily Talitroidea
[includes Talitridae, Hyalidae, and Hyalellidae]
FicurREs 168, 169

Diaenosis.—Accessory flagellum absent; mandible lacking palp;
uropod 3 essentially uniramous; tiny scale-like inner ramus rarely
present. See Phliantidae, Dogielinotidae, Eophliantidae.

Description.—Accessory flagellum absent; body rarely with teeth
or carinations; coxae of medium size; mandible lacking palp, molar
large and strongly triturative (except Najna) ; lower lip without inner
lobes; palp of maxilla 1 often reduced or absent; maxilliped occasion-
ally with fewer than four palp articles; gnathopods usually powerful,
occasionally simple or feeble; uropod 3 small, usually with one ramus,
inner ramus reduced to a scale when present; telson very short, entire,
apically emarginate, cleft, or appearing completely bilobate.

RELAtTIonsuip.—The following families, lacking mandibular palps,
differ from the Talitroidea in their coalesced fifth and sixth pleonites:
Dexaminidae, Prophliantidae, and Kuriidae.

The Dogielinotidae, which probably should be assigned to the
Talitroidea, differ by their haustoriid-phoxocephalid aspect, with
multispinose antennae, short spinose pereopods and shape of head,
bearing a conspicuous but small rostrum. Their epistome has a nasi-
form anterior lobe.

The Eophliantidae differ from talitroids by their cylindrical bodies,

a
cae

Lo rep
FE!

i

rE

Ne

.
8

MARINE GAMMARIDEAN AMPHIPODA 465

short coxae which generally do not touch each other, and degraded
mandibular molars.

The Phliantidae differ from Talitroidea by their degraded man-
dibular molars and the general aspect of their depressed, massive,
often processiferous bodies with laterally splayed coxae. Phliantids*
never have enlarged male second gnathopods as do most talitroids,
especially marine ones.

*See supplement.

Classification of Talitroidea

Bulycheva (1957) partitioned the Talitridae, emended and restricted
the concept of the Talitridae and removed several genera to two new
families, the Hyalidae and Hyalellidae. Morphologically there are
some small conflicts in her arrangement but they detract little from
what appears to be a sensible and logical arrangement, although the
classificatory criteria used are to a large extent not qualitative and
indeed to some extent are concerned with the ecology of the organisms.

As so outlined, in her extensive paper, the Talitridae are confined
to entirely terrestrial genera, many species of which dwell on the
strand but nevertheless are not aquatic. The Hyalidae are exclusively
marine, being characterized by cleft telsons. The Hyalellidae inhabit
both marine and fresh waters, with more genera but few species in the
oceans than in freshwater, where one genus, Hyalella, has 27 species.
Morphologically, it is impossible strictly and qualitatively to separate
the terrestrial Talitridae from the concepts of the other two families.
Bulycheva (1957) in her figure 22 has given an excellent synopsis of
the telsons of the three families, showing Talitridae with generally
uncleft, heavily spinose telsons; Hyalidae with cleft, poorly spinose
telsons; and Hyalellidae with uncleft, poorly spinose telsons. She
shows the fluvial Chiltonia mihiwaka in the Hyalellidae with uncleft
telson but Hurley (1954a) shows it with partially cleft telson, al-
though other chiltonias may have the telson uncleft. Chaltonia is not
assignable to Talitridae because of the poorly spinose telson. Najna
shows both uncleft and minutely cleft telsons in the same species;
Bulycheva assigned the genus to the Hyalidae, but it should be
removed to a new family because of its mandibular molar.

<<

Ficure 168.—Talitroidea: a, Talitrus saltator (Montagu) (Sars, 1895, pl. 9, as T. locusta);
b, Hyale chevreuxi K. H. Barnard (Chevreux 1901la, as H. macrodactylus). Gnathopod 2:
c, Insula antennulella Kunkel (1910); d, Chiltonta mthiwaka (Chilton) (Hurley, 1954a);
e, Hyale bassargint Derjavin (Gurjanova, 1951); f, Talitrus; g, Allorchestes plumicornis
(Heller) (Gurjanova, 1951, as A. ptilocerus). Gnathopod 1: h, Talitrus; i, Insula.

466 U.S. NATIONAL MUSEUM BULLETIN 271

When present the first maxillary palp is supposed to be 2-articulate
in Talitridae and 1-articulate in the other two families, but the marine
Parallorchestes, obviously related to Hyalidae, has a 2-articulate
first maxillary palp, hence bridges the gap to the terrestrial

Talitridae.

ch
[)
LMT a
Hi
t | AQ
ays
i
A
(\ iN
\ N
d
l
n

Ficure 169.—Talitroidea: a, Najna sp. (J. L. Barnard, 1962c). Upper lip: b, Talitrus
saltator (Montagu) (Sars, 1895, pl. 9, as T. locusta). Mandible: c, Talitrus; d, Najna.
Lower lip: e, Talitrus. Maxilla 1: f, Parhyalella batesoni Kunkel (1910); g, Hyale nilssoni
(Rathke) (Sars, 1895, pl. 11); 4, Talitrus. Maxilla 2: 1, Talitrus. Maxilliped: 7, Insula
antennulella Kunkel (1910); k, Parhyale hawaiensis (Dana) (Chevreux, 190la, as Hyale
brevipes); 1, Najna; m, Chiltonia mthiwaka (Chilton) (Hurley, 1954a); n, Talitrus. Uropod
3: 0, Parhyale (Shoemaker, 1956b); p, Chiltonia; q, Talitrus. Telson: r, Talitrus; s,

Hyale.

MARINE GAMMIARIDEAN AMPHIPODA 467

There are exceptions to the neat morphological partitioning of
talitrid-like animals but there are definitely three or more facies and
perhaps three lines of evolution as Bulycheva’s arrangement points
out. She did not assign all existing genera to families, which I do in
the following scheme, considering for the uses of this handbook that
they are subfamilies.

Subfamily Talitrinae: Taltrus, Talorchestia, Orchestia, Orchestoidea.
Subfamily MHyalellinae: Hyalella, Parhyalella, Insula, Chiltonia,
Afrochiltonia, Neobule, Austrochiltonia, [Najna?] but see below.
Subfamily Hyalinae: Hyale, Parhyale, Allorchestes, Parallorchestes.

?(Subfamily Najninae): Nayjna.
?(Subfamily Dogielienotinae): See as a family, p. 207.

Probably Neobule Haswell was improperly described and classified.
It has not been recovered since its original description in 1880. It
has the aspect of a Parhyalella but may belong to another family,
such as the Kuriidae, Prophliantidae, or Lysianassidae. Its ‘“biramous
posterior pleopoda” (presumably uropoda) may be an erroneous
observation and the condition of the mandibular molar was not
stated. It is the only talitrid presumed to have clearly biramous third
uropods and so confuses what would otherwise be a clear diagnosis
of the family.

Key to the Genera of Talitroidea

Neobule Haswell, a dubious genus, is not included.

1. Terminal maxillipedal palp article unguiform (claw-shaped) (figs. 1697,k) . 2

Terminal maxillipedal palp article not unguiform (figs. 1691,m,n).. . .8
ZeUropod:s: with minutesanner ramus) (fig. 1690)". 2 25 9 2. so
Uropod 3 uniramous (fig. 169q) . . . . aarali( bac un pis Gee a ten

3. Palp of maxilla 1 uniarticulate (fig. 169h) . ulate . . . . Parhyale
Palp of maxilla 1 biarticulate (often faintly so) (fig. 1694) . . Parallorchestes

A PEaMclSONeMtiney CIEL G OM iis jen i Milieu se itn ee ere Mauna ance vines Amen MCA a
Telson cleft (fig. 169s)... . Seiarer ran Naren een Naley UR Diner tae Mee ae ARL RITA

5. Maxilla 1 lacking palp (fig. 169f) . WM yeas Cente ac a ee arnyalella
Maxilla 1 bearing palp (figs. 1699,h). ... . peaaaieen tle

6. Male gnathopods 1 and 2 similar to each Other in size sand morphology
(fig. 169a).... . . . .Najna and Insula

Male gnathopod 2 sail Nanece ana of difterent aneratbllores than gnathopod

Te Gan) GIO) 3 eo . . . Hyalella (freshwater)

7. Article 5 of male pantheped 2 aneceaal by ticles 4 and 6 (fig. 168e) . Hyale
Article 5 of male gnathopod 2 produced between articles 4 and 6 (fig.

GSO) tie tes Sheila, ae Bo ea Ne A orchestés

8. Male gnathopod 1 Fimpic ‘Ge. ‘168h) . Sa Waele een) Clam Meccan Mea ea Cut iaen pAWiNG i
Male gnathopod 1 subchelate (fig. 1681) ..... Geel)

9. Male gnathopod 2 feebly chelate (fig. 168f). . . . | Talitrus Ganexiie))

Male gnathopod 2 large and strongly subchelate (fig. 168e).
Orchestoidea (terrestrial)

468 U.S. NATIONAL MUSBHUM BULLETIN 271

10. Female gnathopod 1 simple (fig. 168h) . . . . . Talorchestia (terrestrial)
Female gnathopod 1 subchelate ... . He eegeMecniny nly ay

11. Article 5 of male gnathopod 2 prodiced ipeinasen cmsniies 4 and 6 Ge
NGLA) Go geo) . . . Najna

Article 5 of male paaenenadl 2 iesked peugeenl perieles 4 inl 6 Ge 168e) . 12

12. Antenna 1 shorter than peduncle of antenna 2 (fig. 168a), female gnathopod 2
mitten-shaped (fig. 168f), maxilla 1 usually with palp (even though
minute), uropod 3 with ramus... .. . . .Orchestia (terrestrial)
Antennae 1 and 2 subequal in length, female gnathopod 2 normally sub-
chelate (fig. 168d), maxilla 1 ee palp, uropod 3 usually lacking a

TAMAS (igs OOD) eee hn nene Bair) esti IES}

13. Gnathopods 1 and 2 alike in both ; sexes, ener ole caathonod 2 not en-
larcedinian .. . . .Afrochiltonia (freshwater)
Gnathopod 2 of male onleweradl, mnie Ghatrom femelle iss ree

14. Malepleopodinormal. ... . . . . . . Austrochiltonia Gaestnrratien)
Male pleopod 1 with inner ramus attenuated to backturned, whip-like
Lash) oo eae Si aE oie um hil toniay (iEestwjau els)

Genera of Talitroidea

Afrochiltonia K. H. Barnard
Afrochiltonia K. H. Barnard, 1955.

Type-species: Chiltonia capensis K. H. Barnard, 1916 (original
designation).

Maxilla 1 lacking palp; article 4 of maxillipedal palp short, conical;
gnathopods of both sexes subchelate, male gnathopod 2 not larger
than 1, female gnathopod 2 like gnathopod 1; male pleopod 1 normal
(Chiltonia); uropod 3 lacking inner ramus; telson entire. Species:
1, S. Africa, freshwater.

Allorchestes Dana

Allorchestes Dana, 1849.—Stebbing, 1906.
Aspidophoreia Haswell, 1880d.

Type-species: A. compressa Dana, 1852a (selected by Chevreux
and Fage, 1925). See Stebbing, 1899a.

Maxilla 1 bearing palp; article 4 of maxillipedal palp unguiform;
gnathopods of both sexes subchelate, male gnathopod 2 larger than
1, article 5 produced between articles 4 and 6, female gnathopod 2
like gnathopod 1; uropod 3 lacking inner ramus; telson cleft. Species:
12, cosmopolitan, littoral. At least two kinds of telson occur in this
genus, one like that of Hyale, with subconical lobes fully split from
each other and set at an angle to each other forming a tent; and the
other a quadratiform flat plate with partial cleft. Reorganization of
this genus needs investigation.

MARINE GAMMARIDEAN AMPHIPODA 469

Austrochiltonia Hurley

Austrochiltonia Hurley, 1958.

Type-species: Hyalella australis Sayce, 1901 (original designation).

Maxilla 1 lacking palp; article 4 of maxillipedal palp short, conical;
gnathopods subchelate in both sexes, male gnathopod 2 much larger
than 1, article 5 not projecting between 4 and 6, female gnathopod
2 like 1; male pleopod 1 normal (Chiltonia); uropod 3 lacking inner
ramus; telson entire. Species: 2, Australia, freshwater.

Chiltonia Stebbing

Chiltonia Stebbing, 1899a.—Stebbing, 1906.

Type-species: Hyalella mihiwaka Chilton, 1898 (original desig-
nation). See Hurley, 1954a.

Maxilla 1 lacking palp; article 4 of maxillipedal palp short, conical;
genathopods subchelate in both sexes, male gnathopod 2 much larger
than 1, article 5 not projecting between articles 4 and 6, female
enathopod 2 like 1; male pleopod 1 uth inner ramus attenuated to a
backturned whip-like lash (Afrochiltonia and Austrochiltonia); uropod 3
lacking ramus; telson entire. Species: 3, New Zealand, freshwater.

Hyale Rathke

Hyale Rathke, 1837.—Stebbing, 1906.

Nicea Nicolet, 1849.

(Allorchestina) Brandt, 185la (valid subgenus). Type by present selection:
Amphithoe [sic] prevostiz Milne Edwards, 1830.

Galanthis Bate, 1857a.

Type-species: H. pontica Rathke, 1837 (monotypy). See Sars,
1895 (as H. lubbockiana).

Maxilla 1 with 1-articulate palp (but note Sars, 1895, pl. 11);
article 4 of maxillipedal palp unguiform; gnathopods subchelate in
both sexes, male gnathopod 2 larger than 1, article 5 not projecting
between articles 4 and 6, female gnathopod 2 like gnathopod 1;
uropod 3 lacking inner ramus; telson cleft. Species: 48, cosmopolitan,
especially tropics, littoral.

Hyalella Smith
Hyalella Smith, 1874.—Stebbing, 1906.

Type-species: Amphitoe aztecus Saussure, 1858 (monotypy and
subsequent synonymy). See Bulycheva, 1957.

Maxilla 1 with short l-articulate palp; article 4 of maxillipedal
palp unguiform; gnathopods of both sexes subchelate, male gnathopod
2 larger than 1, article 5 produced between articles 4 and 6; female

470 U.S. NATIONAL MUSEUM BULLETIN 271

enathopod 2 like gnathopod 1 or minutely chelate; uropod 3 lacking
inner ramus; telson entire. Species: 27, primarily S. America,
freshwater.

Insula Kunkel

Insula Kunkel, 1910.

Type-species: J. antennulella Kunkel, 1910 (montypy).

Maxilla 1 bearing a palp; article ?4 (Kunkel claims 3 and figures
only 3 articles) of maxillipedal palp unguiform; gnathopods subchelate
in male (female unknown), relatively alike in size, rather slender
(possibly juvenile), article 5 lobed between articles 4 and 6; uropod 3
lacking inner ramus; telson entire. Species: 1, Bermuda, littoral.

Najna Derzhavin

Najna Derzhavin, 1937.—Gurjanova, 1951.

Type-species: N. consiliorum Derzhavin, 1937 (monotypy).

Mandibular molar obsolete; maxilla 1 with vestigial palp; article 4 of
maxillipedal palp tiny, short, not unguiform; gnathopods similar in
both sexes, subchelate, gnathopod 2 not larger than 1, article 5 lobed
between articles 4 and 6; uropod 3 lacking inner ramus; telson entire
or slightly notched [probably requires erection of new family].
Species: 1, boreal, N. Pacific, littoral.

[Neobule Haswell]

Neobule Haswell, 1880b.—Stebbing, 1906.

Type-species: N. algicola Haswell, 1880b (monotypy).

A problematical genus, probably not a talitroid; uropod 3 biramous,
rami equal in size but small; gnathopods subchelate, gnathopod 2
larger than 1, palms nearly transverse. Species: 1, Australia, littoral.

Orchestia Leach

Orchestia Leach, 1814a.—Stebbing, 1906.
Scamballa White, 1847a.
Parorchestia Stebbing, 1899a.

Type-species: Oniscus gammarellus Pallas, 1766 (monotypy and
subsequent synonymy). See Sars, 1895 (as Orchestia littorea).

Maxilla 1 usually with palp, 2-articulate or absent; article 4 of
maxillipedal palp a vestigial bud or absent; gnathopods subchelate in
both sexes, male gnathopod 2 much larger than 1, female gnathopod 2
mitten-like; uropod 3 lacking inner ramus; telson entire. Species: 69,
cosmopolitan, primarily tropical, terrestrial (beachhoppers).

MARINE GAMMARIDEAN AMPHIPODA A471

Orchestoidea Nicolet

Orchestoidea Nicolet, 1849.—Stebbing, 1906.
Talitronus Dana, 1850, 1852a.
Megalorchestia Brandt, 1851b.

Type-species: O. tuberculata Nicolet, 1849 (monotypy).

Maxilla 1 with small 2-articulate palp; palp article 4 of maxilliped
a vestigial bud or absent; male gnathopod 1 subchelate but often
poorly, gnathopod 2 larger than 1, subchelate, article 5 not produced
between articles 4 and 6, female gnathopod 1 simple, gnathopod 2
mitten-like; uropod 3 lacking inner ramus; telson entire. Species:
12, primarily E. Pacific, terrestrial (beachhoppers).

Parallorchestes Shoemaker
Parallorchestes Shoemaker, 1941a.

Type-species: Allorchestes ochotensis Brandt, 1851c (original desig-
nation). See J. L. Barnard, 1962c.

Maxilla 1 with large 2-articulate palp; palp article 4 of maxilliped
unguiform; gnathopods subchelate, male gnathopod 2 much larger
than 1, article 5 produced between articles 4 and 6, female gnathopod
2 like 1; uropod 3 with small scale-like inner ramus; telson cleft.
Species: 1, boreal N. Pacific, littoral.

Parhyale Stebbing

Parhyale Stebbing, 1897; 1906.
Hyaloides Schellenberg, 1939.

Type-species: P. fasciger Stebbing, 1897 (monotypy). See Shoe-
maker, 1956b.

Maxilla 1 with 1-articulate palp; article 4 of maxillipedal palp
unguiform; gnathopods subchelate, male gnathopod 2 larger than 1,
article 5 produced between articles 4 and 6, female gnathopod 2 like
enathopod 1 and slightly larger; uropod 3 with small scale-like inner
ramus; telson cleft. Species: 2, circumtropical, littoral.

Parhyalella Kunkel

Parhyalella Kunkel, 1910.—Schellenberg, 1938.
Exhyalella Stebbing, 1917.

Type-species: P. batesoni Kunkel, 1910 (monotypy).

Maxilla 1 lacking palp; article 4 of maxillipedal palp unguiform;
gnathopods subchelate, male gnathopod 2 larger than 1, article 1
produced between articles 4 and 6, female gnathopod 2 like male but
smaller; uropod 3 uniramous; telson entire. Species: 7, pantropical,
freshwater and marine.

472 U.S. NATIONAL MUSEUM BULLETIN 271

Talitrus Latreille

Talitrus Latreille, in Bosc, 1802.—Latreille, 1802.

?(Talitrorchestia) Brandt, 185la (subgenus). Type-species: Orchestza cloqueti
Audouin, 1826 (obscure species) .

Talitroides Bonnier, 1898.—Stebbing, 1906. Type-species: 7. bonnieri Stebbing,
1906 (monotypy).

Talitriator Methuen, 1913.

Type-species: Cancer (Gammarus) saltator Montagu, 1808 (new
name for Oniscus locusta Pallas, 1766), selected by Boeck (1876).
This designation remains questionable. When Bosc (1802), in copying
Latreille’s manuscript, erected Talitrus (for which the latter deserves
nomenclatural credit in Bose), two species ‘‘Gammarus locusta Fab.”
and “Oniscus gammarellus Pallas” were assigned to it (vol. 1 of Bose,
p. 78). In vol. 2 of the same work, the assigned species were stated as
Talitrus grillus, a new species later attributed to Bose, but which, as
indicated in the description, should be attributed to Latreille. By
strict adherence to rules, and without either original description,
Bose’s second volume or Latreille’s work of a few months later in
1802, we would expect the genus Talitrus to be composed of two
species, one of which is precisely the type-species of the earlier genus
Gammarus, the other of which later came to be a species of Orchestia.
Only by reference to volume 2 do we assume Latreille was referring
earlier to the species described by Pallas and not that described by
Fabricius. They become homonyms when thrown together in Cancer
by other authors. Thus by elimination, now outmoded, Oniscus
gammarellus Pallas could yet be selected as the type-species of Talitrus
and Orchestia would fall as a synonym to Talitrus, a highly undesirable
occurrence at this late day in amphipodan taxonomy. And, G. locusta
Fabricius (identical with Cancer locusta Linnaeus) could also be
selected. If necessary, the case should be submitted to the ICZN in
the hope that they would fix Oniscus locusta Pallas as type-species,
and eliminate the ambiguity of Latreille (in Bosc) having used Fabri-
clus’ name as author of the species.

Maxilla 1 with small 2-articulate palp; article 3 of maxillipedal palp
a vestigial bud; gnathopod 1 of both sexes simple, gnathopod 2 of
both sexes small, feebly chelate, mitten-shaped; uropod 3 lacking inner
ramus; telson entire or slightly notched. Species: 11, (terrestrial,
circumtropical, biboreal).

Talorchestia Dana

(Talorchestia) Dana, 1852b (subgenus).—Stebbing, 1906.

Type-species: Talitrus gracilis Dana, 1852a (selected by Chevreux
and Fage, 1925).

MARINE GAMMARIDEAN AMPHIPODA 473

Maxilla 1 with small 2-articulate palp; article 4 of maxillipedal palp
a vestigial bud or absent; gnathopod 1 of both sexes simple or poorly
subchelate, male gnathopod 2 larger than 1, subchelate, article 5 not
produced between articles 4 and 6, female gnathopod 2 small, mitten-
shaped; uropod 3 lacking inner ramus; telson entire or slightly notched.
Species: 39, circumtropical, amphiboreal terrestrial (some beach-
hoppers).

Thaumatelsonidae
Fieures 170, 171

Diacnosis.—Accessory flagellum absent or vestigial; mandibular
molar evanescent; coxa 1 very small or absent, always partially
covered by following coxae; coxa 4 large, shield-like, not excavate
posteriorly; uropod 3 uniramous; outer lobes of maxilliped vestigial;
telson entire, greatly thickened dorsoventrally; urosomal segments
coalesced or partially so; article 2 of pereopod 3 (and 4-5) linear. See
Stenothoidae, Cressidae, Pagetinidae, Leucothoidae, Anamixidae,
Amphilochidae.

Description.—Rostrum inconspicuous; accessory flagellum absent
or vestigial; body smooth or pleon with dorsal carinae; coxa 1 small
or absent, hidden by following coxae; coxa 4 large, shield-like, not
excavate posteriorly; upper lip probably incised; mandible with
evanescent molar, palp present or absent; lower lips unknown; maxilla
1 with 2-articulate palp; maxilla 2 normal; maxilliped slender, outer
plates small or vestigial; gnathopods moderately powerful, subchelate
or chelate; uropod 3 uniramous, the ramus 2-articulate; telson greatly
thickened dorsoventrally; urosomal segments coalesced or partially
so, or urosomite 1 often with large dorsal process covering remainder
of urosome dorsally.

RELATIONSHIP.—T he Stenothoidae have distinct urosomal segments
and unthickened telsons.

The Cressidae have the unthickened telson fused with urosomite 3.

Parathaumatelson Gurjanova (1938) is synonymized with Pseudo-
thaumatelson Schellenberg (1931).

Key to the Genera of Thaumatelsonidae

1. Gnathopod 2 chelate (fig. 1717)... ..... +... . Prothaumatelson
Cnathopods2;subchelate (igseplaiee yen ne in ere cine fn) eee Ui nae

2. Palp of mandible 1-articulate or absent
Pseudothaumatelson (= Parathaumatelson)
Palprotimandiblers-articulabe: ys 0 awit) ee ee Thaumatelson

MARINE GAMMARIDEAN AMPHIPODA 475

Genera of Thaumatelsonidae

Prothaumatelson Schellenberg
Prothaumatelson Schellenberg, 1931.—Gurjanova, 1938.
Type-species: Thaumatelson nasutum Chevreux, 1912a; 1912b
(monotypy).
Species: 2, bipolar, littoral.
Pseudothaumatelson Schellenberg

Pseudothaumatelson Schellenberg, 1931.—Gurjanova, 1938.
Parathaumatelson Gurjanova, 1938.

Type-species: P. patagonicum Schellenberg, 1931 (present selec-
tion). :
Species: 3, subantarctic, littoral.

Ficure 171.—Thaumatelsonidae: Mouthparts, Thaumatelson herdmani Walker (1907):
a, mandible; b,c, maxillae 1, 2; d, maxilliped. Prothauwmatelson nasutum (Chevreux,
1912b): e, maxilliped. Gnathopod 1: f, Thawmatelson walkeri Chilton (1912b); g, Pro-
thaumatelson carinatum Shoemaker (1955). Gnathopod 2: h, Thaumatelson herdmani;
1, Thaumatelson walkeri; 7, Prothaumatelson carinatum. Uropod 3: k, Prothaumatelson
nasutum. Urosome and pleonite 3, urosomites coalesced, bearing 3 uropods and large
telson: 1, Thaumatelson walkeri.

285-135 O - 69 - 31

476 U.S. NATIONAL MUSEUM BULLETIN 271

Thaumatelson Walker

Thaumatelson Walker, 1906b, 1907.

Type-species: T. herdmani Walker, 1906b; 1907 (monotypy). See
Schellenberg, 1931.
Species: 3, subantarctic, littoral.

Vitjazianidae
Ficure 172

Diaanosis.—Accessory flagellum long, composed of a few long
articles; base of primary flagellum conjoint; gnathopod 1 simple,
enathopods feeble, anterior members intersimilar; coxae short. See
Synopiidae, Astyridae, Eusiridae, Liljeborgiidae, Hyperiopsidae,
Melphidippidae, Argissidae.

Description.—Accessory flagellum long, composed of only a few
articles; base of primary flagellum of antenna 1 conjoint, forming an
article nearly as long as peduncle; body smooth except for slight
carinations on urosome; rostrum slightly developed; coxae very short
to medium in length, scarcely touching; upper lip minutely incised;
lower lip with or without inner lobes; mouthparts otherwise basic;
enathopods feeble, gnathopod 1 simple, gnathopod 2 simple or sub-
chelate; uropods biramous; telson short, cleft.

RELATIONSHIP.—The Hyperiopsidae differ from Vitjazianidae by
the bent first maxillary palps, the poorly toothed primary cutting
edge and small molar of the mandible and the elongate fourth articles
of pereopods 1 and 2.

The Eusiridae have an elongate peduncle of antenna 1 and usually
large subchelate gnathopods. Liljeborgiidae have large subchelate
enathopods.

The Melphidippidae have an elongate uropod 3, and an elongate
peduncle of antenna 1.

The Astyridae have a 1-articulate accessory flagellum, a reduced
mandibular molar, and gaping lower lip.

The Stilipedidae lack an accessory flagellum, have foliaceous first
maxillae, short gnathopods, and no mandibular molar.

—

Ficure 172.—Vitjazianidae: a, Vitjaziana gurjanovae Birstein and Vinogradov (1955);
b, Vemana compressa J. L. Barnard (1964a). Lower lip: c, Vemana; d, Vitjaziana.
Telson: e, Vitjaziana. Maxilla 1: f, Vitjaziana; g, Vemana. Antenna 1, basal portion:
h, Vitjaziana. Maxilliped: 1, Vitjaziana. Uropod 3: 7, Vemana. Mandible: k, Vit-
jaziana; 1, Vemana. Upper lip: m, Vitjaziana. Gnathopod 1: n, Vitjaziana; 0, Vemana.
Gnathopod 2: p, Vitjaziana; gq, Vemana. Maxilla 2: 7, Vitjaziana.

MARINE GAMMARIDEAN AMPHIPODA

Neveeee

le

Li a

Oo Mie .
ve

A478 U.S. NATIONAL MUSEUM BULLETIN 271

A few setae of pereopods 1-5 are elongate in some vitjazianids and
the term “fossorial’” might be applied to them but other fossorial
families do not have a basally conjoint primary flagellum on antenna 1.

Vitjazianids have definitive similarity to Gammaridae and Eusiri-
dae, from which they differ by the very short peduncle and the long
conjoint base of the primary flagellum on antenna 1. By evidence of
gnathopods and mandible, but not of telson and head the vitjazianids
are closely related to synopiids (tironids) and demonstrate again the
difficulty in defining families because of the centripetal radiation of
evolutionary lines around the Gammaridae.

Key to the Genera of Vitjazianidae

1. Gnathopod 2 subchelate; coxae of medium length; mandibular palp claviform.
Vemana
Gnathopod 2 simple; coxae minute; mandibular palp linear. . . Vitjaziana

Genera of Vitjazianidae

Vemana J. L. Barnard
Vemana J. L. Barnard, 1964a.
Type-species: V. compressa J. L. Barnard, 1964a (original
designation).
Species: 2, Caribbean, bathy-abyssopelagic.
Vitjaziana Birstein and Vinogradov
Vitjaziana Birstein and Vinogradov, 1955.

Type-species: V. gurjanovae Birstein and Vinogradov, 1955 (origi-
nal designation).
Species: 1, N. W. Pacific, abyssopelagic.

Incertae Sedis

FiGurRE 173

Didymocheila K. H. Barnard
Didymocheila K. H. Barnard, 1931; 1932.

Type-species: D. spongicola K. H. Barnard, 1931 (original desig-
nation) ; 1932.

Having all characteristics of Lysianassidae except article 3 of
gnathopod 2 not elongate and article 6 strongly elongate, gnathopods

MARINE GAMMARIDEAN AMPHIPODA 479

1 and 2 slender, subequal in size and weakly chelate; mouthparts from
lateral view conically grouped, upper lip substyliform, entire, mandible
substyliform, cutting edge slightly toothed, molar well developed,
?barrel-shaped or cup-shaped, unknown if triturative, 3-articulate
palp attached level with molar; lower lip with Trischizostoma appear-
ance; inner plate of maxilla 1 heavily setose medially, palp 2-articulate;
maxilla 2 and maxillipeds normal; uropod 3 lacking rami, composed of
a simple, ovate peduncle; telson broader than long, entire, apically
concave.

With strong resemblance to Sebidae in gnathopods but article 2 of
antenna 1 peduncle not elongate, upper lip elongate, mandibular
molar and palp strong, uropod 3 lacking any rami, telson very short,
etc. Species: 1, S. Georgia Islands, littoral.

i ¢

Figure 173.—Incertae Sedis: a, Sphaerophthalmus grobbeni Spandl (1924). Didymocheila
spongicola K. H. Barnard (1932): b, lateral aspect; c, telson ard third uropods; d,e, gnatho-
pods 1, 2; f, mandible; g, lower lip.

d

ASO U.S. NATIONAL MUSEUM BULLETIN 271

Metoediceros Schellenberg

Metoediceros Schellenberg, 1931.

Type-species: MM. fuegiensis Schellenberg, 1931 (monotypy).

Head poorly rostrate, not massive; eyes lateral; antennae short,
subequal in length, peduncle of antenna 1 slightly longer than head,
articles 1-3 successively shorter, male antenna 2 with calceoli, ac-
cessory flagellum absent; upper lip not incised; [epistome unknown];
mandible lacking palp, with large triturative molar and toothed incisor;
lower lip with inner lobes separate, outer lobes with mandibular
extensions; inner plate of maxilla 1 with one terminal seta, outer plate
with 10 spines, palp 4-articulate, article 4 unguiform; coxae long as in
normal gammaridean, coxa 4 not excavate posteriorly; gnathopods
1-2 weakly subchelate, of medium size, similar to each other, articles
5-6 subequal in length; pereopods generally like those of Oediceros
saginatus (see Sars, 1895, pl. 102), thus fossorial, pereopods i-4 with
vestigial dactyls; uropods 1-2 normal, apices of rami spinose; uropod
3 reduced to a small simple triangular peduncle; telson nearly circular,
entire. Affinities with Dogielinotidae but epistome unknown. Species:
1, Tierra del Fuego, brackish water.

Sphaerophthalmus Spandl
Sphaerophthalmus Spandl, 1923; 1924.

Type-species: S. grobbent Spandl, 1923; 1924 (monotypy).

Probably assignable to the Dexaminidae.

Head and thorax very tall, eye forming a large bead; apparently
urosomal segments 2-3 coalesced, very small and flat; metasome
dorsally carinate; accessory flagellum apparently absent, antenna 1
long, slender, article 2 longer than 1, flagellar articles numerous,
antenna 2 “3-articulate”’; maxillipeds apparently with medium sized
inner lobes, enormous subclavate outer lobes nearly exceeding the
moniliform 4- (?3-) articulate palp, the terminal article of which lacks
distinction; other mouthparts not described; gnathopod 1 stout, like
that of stegocephalids, gnathopod 2 more slender, elongate, article 6
expanded and bearing transverse palm as in gnathopod 1; pereopods
of medium length, second articles of pereopods 3-5 expanded; uropods
1-2 normal, rami interequal, apically spinose, uropod 3 with short
peduncle and leaf-lke rami; telson elongate, cleft nearly to base.
Species: 1, Red Sea, littoral.

MARINE) GAMMARIDEAN AMPHIPODA A81

Supplement

Recent studies by the writer in Hawai, New Zealand, and Australia
suggest the following changes and additions to the classificatory
scheme of Gammaridea.

The Eusiridae and Calliopiidae must be amalgamated.

The Aoridae must be amalgamated with the Isaeidae; most genera
of the Corophiidae should also be placed in the greater Isaeidae but
Corophium and Paracorophium might be retained within a subfamily.

Ceina is to be removed from the Phliantidae to form the type of a
new family with close affinities to the Hyalidae; the family will be
characterized by the loss of rami on uropod 3 and the rough skin
texture; it has affinities with various freshwater chiltonias that may
require removal from the Hyalellidae to form a group distinct from
the American hyalellas.

Biancolina is to be removed from the Eophliantidae to form the
type of a new family. The biramous uropod 3 of Biancolina is one
mark of its distinction from Eophliantidae. Amphitholina should also
be removed from Eophliantidae and returned to the Ampithoidae as
type of a new subfamily.

The freshwater genus Paracalliope is added to the list of marine
genera as one of its species, Oedicerus novizealandiae Dana (1853), is
definitely intertidal in New Zealand. That species is not a member
of Carolobatea as heretofore considered (Chilton, 1909). Stebbing
(1906) had already suggested Dana’s species to be identical with P.
fluviatilis (Thomson) but the two are definitely distinct.

Paracalliope Stebbing
Paracalliope Stebbing, 1899d; 1906.

Type-species: Calliope fluviatilis Thomson, 1879 (original desig-
nation).

Eusirid-calliopiid with incipient characters of Oedicerotidae, es-
pecially in the disproportionately elongate pereopod 5 with elongate,
terminally setose dactyl and elongate uropod 3.

Eyes paired, discontiguous; accessory flagellum vestigial; inner
plates of maxillae 1-2 heavily setose medially, maxilla 2 with addi-
tional submarginal row of medial setae; female gnathopods weak,
those of male stout, with erect lobes on fifth articles, hands twisted
mediad, palms bearing large spines; coxa 4 weakly or not postero-
dorsally excavate; telson ovoid, thin, uncleft; wrosomites 2-3 coalesced.
Species: 5 (2 undescribed), New Zealand, Australia, Philippine
Islands, India, marine littoral, brackish and freshwaters.

Literature Cited

[*=Not seen, reference copied from Stebbing, 1888.]

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Bruce, J. R.; Cotman, J. 8.; and Jonss, N. §., ed.
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Mar. Biol. C. mem. 36, Liverpool Univ. Press, pp. xii+307, plus
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486 U.S. NATIONAL MUSEUM BULLETIN 271

BriiacEen, E. v. p.

1905. Die Amphipoden des Katharinenhafens (Murmankiiste) und seiner
Umgebungen. Trav. Soc. Imp. Nat. St.-Pétersbourg, vol. 36,
livr. 1, no. 8, pp. 1-10, pl. 1.

Bruzeuius, R. M.

1859. Bidrag till kinnedomen om Skandinaviens Amphipoda Gammaridea.
Kongl. Svenska Vetenskaps-Akad. Handl., NF, vol. 3, pp. 1-104,
pls. 1-4.

BucuuHotiz, R.

1874. 8, Crustaceen: Die zweite deutsche Nordpolarfahrt in den Jahren
1869 und 1870 unter Fiihrung des Kapitan Karl Koldewey, vol. 2,
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Butycuetva, A. I.

1955. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz Japonskogo
Morja, II. Trudy Zool. Inst., Akad. Nauk SSSR, vol. 21, pp.
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1957. Morskie bloxi morej SSSR i sopredel’nyx vod (Amphipoda-
Talitroidea). Opred po Faune SSSR, Adad. Nauk SSSR, vol. 65,
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CaLMAN, W. T.

1910. On two new species of wood-boring Crustacea from Christmas Island.

Ann. Mag. Nat. Hist., ser. 8, vol. 5, pp. 181-186, pl. 5.
Catta, J.-D.

1875. Note pour servir 4 Vhistoire des amphipodes du Golfe de Marseille.

Rev. Sci. Nat., vol. 4, pp. 161-169.
CuHEvreux, E.

1887. Catalogue des crustacés amphipodes marins du sud-ouest de la
Bretagne, suivi d’un apercu de la distribution géographique des
amphipodes sur les cotes de France. Bull. Soc. Zool. France,
vol. 12, pp. 288-340, pl. 5.

1889. Amphipodes nouveaux provenant des campagnes de 1’ Hirondelle
1887-1888. Bull. Soc. Zool. France, vol. 14, pp. 284-289, figs.
1-5.

1891. Podoprion bolivari, amphipode nouveau de la famille des Lysz-
anassidae. Voyage de la goelette Melita aux Canaries et au Sénégal,
1889-1890. Mém. Soc. Zool. France, vol. 4, pp. 5-10, pl. 1.

1895. Sur un amphipode, Pseudotiron bouvieri, nov. gen. et sp., de la famille
des Syrrhoidae, nouvella pour la faune méditerranéenne. Bull.
Soc. Zool. France, vol. 20, pp. 165-170, figs. 1-14.

1899. Sur deux espéces géantes d’amphipodes provenant des campagnes
du yacht Princesse Alice. Bull. Soc. Zool. France, vol. 24, pp.
152-158, figs. 1-6.

1900. Amphipodes provenant des campagnes de I’ Hirondelle (1885-1888).
Rés. Campagnes Sci. Albert Ie, Monaco, vol. 16, pp. i-iv + 1-195,
pls.. 1-18.

1901la. Crustacés amphipodes: Mission scientifique de M. Ch. Alluaud aux
Iles Séchelles (Mars, Avril, Mai, 1892). Mém. Soe. Zool. France,
vol. 14, pp. 388-488, figs. 1-65.

1901b. Description d’un crustacé amphipode nouveau de la famille des
Stenothoidae (Parametopa Kervillei nov. gen. et sp.) capturé au
moyen d’une nasse par M. Henri Gadeau de Kerville, dans la
région d’Omonville-la-Rogue (Manche). Bull. Soe. Amis Sci.
Nat. Rouen, 1900, pp. 231-237, pl. 3.

1903.

1905a.

1905b.

1905c.

1906a.

1906b.

1906c.

1908a.

1908b.

1908c.

1909.

1910.

1911a.

1911b.

1911e.

1912a.

1912b.

MARINE GAMMARIDEAN AMPHIPODA 487

Note préliminaire sur les amphipodes de la famille des Lysianassidae
recueillis par la Princesse-Alice dans les eaux profondes de
V Atlantique et de la Méditerranée. Bull. Soc. Zool. France, vol.
28, pp. 81-97, figs. 1-7.

Paracyphocaris praedator type d’un nouveau genre de Lysianassidae.
Bull. Mus. Océanog., Monaco, no. 32, pp. 1-6, figs. 1-3.

Diagnoses d’amphipodes nouveaux provenant de l’expédition ant-
arctique du Francais. Bull. Soc. Zool. France, vol. 30, pp. 159-165.
figs. 1-3.

Description d’un amphipode (Katius obesus, nov. gen. et sp.), suivie
d’une liste des amphipodes de la tribu des Gammarina remenés
par le filet 4 grande ouverture pendant la derniére campagne de la
Princesse-Alice en 1904. Bull. Mus. Océanog., Monaco, no. 35,
pp. 1-5, figs. 1-7, list on p. 7.

Diagnoses d’amphipodes nouveaux provenant de l’expédition ant-
arctique du ‘Francais,’ II: Metopidae—Iphimedidae. Bull. Soc.
Zool. France, vol. 31, pp. 37-40, figs. 1-2.

Diagnoses d’amphipodes nouveaux provenant de l’expédition ant-
arctique du Francais, III: Oediceridae—Calliopidae, pp. 76-80,
figs. 1-2; IV: Atylidae, pp. 82-86, figs. 1-3; V: Phliadidae, pp.
87-89, figs. 1-2. Bull. Soc. Zool. France, vol. 31, pp. 76-89.

Crustacés amphipodes: Expédition Antarctique Francaise (1903-—
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pp. 1-100, figs. 1-56.

Diagnoses d’amphipodes nouveaux provenant des campagnes de la
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Monaco, no. 117, pp. 1-13, figs. 1-7.

Diagnoses d’amphipodes nouveaux provenant des campagnes de la
Princesse-Alice dans 1|’Atlantique nord. Bull. Inst. Océanog.,
Monaco, no. 121, pp. 1-15, figs. 1-8.

Diagnoses d’amphipodes nouveaux provenant des campagnes de la
Princesse-Alice dans ]’Atlantique nord. Bull. Inst. Océanog.,
Monaco, no. 129, pp. 1-12, figs. 1-6.

Diagnoses d’amphipodes nouveaux provenant des campagnes de la
Princesse-Alice dans ]’Atlantique nord. Bull. Inst. Océanog.,
Monaco, no. 150, pp. 1-7, figs. 1-3.

Diagnoses d’amphipodes nouveaux provenant des campagnes de la
Princesse-Alice dans 1’Atlantique nord. Bull. Inst. Océanog..
Monaco, no. 156, pp. 1-4, figs. 1-2.

Campagnes de la Melita: Les amphipodes d’Algerie et de Tunisie.
Mém. Soe. Zool]. France, vol. 23, pp. 145-285, pls. 6-20.

Diagnoses d’amphipodes nouveaux provenant des campagnes de la
Princesse-Alice dans l Atlantique nord. Bull. Inst. Océanog.,
Monaco, no. 204, pp. 1-3, figs. 1—6.

Sur les amphipodes des expédition antarctiques francaises. C.R.
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Deuxiéme expédition dans 1’antarctique dirigée par le Dr. Charcot,
1908-1910. Diagnoses d’amphipodes nouveaux. Bull. Mus. Hist.
Nat., Paris, 1912, no. 4, pp. 1-12.

Amphipodes: Deuxiéme Expédition Antarctique Francaise (1908-
1910) commandée par le Dr. Jean Charcot. Sci. Nat: Doce. Sci.,
pp. 79-186, figs. 1-62.

488 U.S. NATIONAL MUSEUM BULLETIN 271

CuEevrevx E.—Continued
1914. Diagnoses d’amphipodes nouveaux provenant des campagnes de la
Princesse-Alice dans Atlantique nord. Bull. Inst. Océanog.,
Monaco, no. 296, pp. 1—4, figs. 1-3.
1919. Note préliminaire sur les amphipodes recueillis par les expéditions
du Travailleur et du Talisman (1880-1883). Bull. Mus. Hist.
Nat., Paris, 1919, no. 7; 1920, no. 1, pp. 574-580, 7-12.
1920. Sur quelques amphipodes nouveaux ou peu connus provenant des
cotes de Bretagne. Bull. Soc. Zool. France, vol. 45, pp. 75-87,
figs. 1-9.
1925. Amphipodes I: Gammariens. Voyage de la Goélette Melita aux
Canaries et au Sénégal (1889-1890). Bull. Soc. Zool. France,
vol. 50, pp. 278-311, figs. 1-12.
1926. Diagnoses d’amphipodes nouveaux provenant des campagnes de la
‘‘Princesse-Alice’’ dans l’Atlantique et dans l’Océan Arctique.
Bull. Inst. Océanog., Monaco, no. 475, pp. 1-12, figs. 1-6.
1927. Crustacés amphipodes: Expéd. Sci. ‘‘Travailleur” et du “Talisman”
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CHEVREUX, E., and Bouvisr, E. -L.
1892. Perrierella crassipes, espéce et genre nouveaux d’amphipodes des
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CHEVREUX, E., and Faas, L.
1925. Amphipodes. Faune de France, vol. 9, pp. 1-488, figs. 1-438.
CHILTON, C.
1883. Further additions to our knowledge of the New Zealand Crustacea.
Trans. Proc. New Zealand Inst., vol. 15, pp. 69-86, pls. 1-38.
1884. Additions to the sessile-eyed Crustacea of New Zealand. Trans.
Proc. New Zealand Inst., vol. 16, pp. 249-265, pls. 17-21.
1898. A new freshwater amphipod from New Zealand. Ann. Mag. Nat.
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1900. A New Zealand species of the amphipodan genus Cyproidia. Ann.
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1909. The Crustacea of the subantarctic islands of New Zealand. Sub-
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1912. The Amphipoda of the Scottish National Antarctic Expedition.
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1914. A new amphipodan genus and species (family Dexaminidae) from
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1921. Report on the Amphipoda obtained by the F.I.S. “Endeavour” in
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1924. Some New Zealand Amphipoda: No. 5. Trans. Proc. New Zealand
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Costa, A.
*185la. Fauna del Regno di Napoli.
*1851b. Catalogo dei crostacei Italiani e di moltri altri del Mediterraneo per
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*1853a. Fauna del Regno di Napoli.

MARINE GAMMARIDEAN AMPHIPODA A489

*1853b. Descrizione di tre nuovi crostacei del Mediterraneo discoperti dal
Rev. G.F. Hope. Fauna del Regno di Napoli, fasc. 83, 10 pp.,
3 pls.

*1853c. Relazione sulla memoria del Dottor Achille Costa, di ricerche su’
crostacei anfipodi del regno di Napoli. Rend. Soc. Reale Borbonica,
Accad. Sci., Anno 1853, pp. 166-178.

1857. Ricerche sui crostacei anfipodi del regno di Napoli. Mem. Reale
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1867. Saggio della collezione de’crostacei del Mediterraneo del Museo
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CoutTibre, H.

1904. Sur un type nouveau d’amphipode Grandidierella Mahafalensis,
provenant de Madagascar. Bull. Soc. Philomath., 1904, vol. 6,
pp. 166-174, figs. 1-18.

CzERNIAVSKI, W.
*1868. Materialia ad zoographiam ponticam comparatum. Studiosi uni-
versitatis charcoviensis Voldemari Czerniavski.
Dau, E.

1945. Menigratopsis svennilssoni n. gen. et spec., a lysianassid amphipod
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vol. 15, no. 24, pp. 1-7, figs. 1-4.

1959. Amphipoda from depths exceeding 6000 meters. Galathea Rep.,
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Dana, J. D.

1849. Synopsis of the genera of Gammaracea. American Journ. Sci. Arts,
ser. 2, vol. 8, pp. 135-140.

1850. A new genus of Orchestidae. American Journ. Sci. Arts, ser. 2, vol
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1852a. Conspectus crustaceorum quae in orbis terrarum circumnavigatione,
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1852b. On the classification of the crustacean Choristopoda or Tetradecapoda.
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1853. Crustacea, Part II. U.S. Expl. Exped., vol. 14, pp. 689-1618, atlas
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DE.vLA VALLE, A.

1893. Gammarini del Golfo di Napoli. Fauna und Flora des Golfes von

Neapel . . . Monog. 20, pp. i-xi + 1-948, atlas of 61 pls.

DersHAVIN, A. N. [Derjavin, Derzhavin]

1923. Malacostraca der Siisswasser-gewdsser von Kamtschatka. {Russk.].
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1927. The Gammaridae of the Kamchatka Expedition 1908-1909. [Russk.].
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1937. Talitridae of the Soviet coast of the Japan Sea. Issledovanija
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A490 U.S. NATIONAL MUSEUM BULLETIN 271

Dysowsky, B. N.

1874. Beitrage zur naéheren Kenntniss der in dem Baikal-vorkommenden
niederen Krebse aus der Gruppe der Gammariden. Herausgegeben
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EneEauist, P.

1950. Studies on the soft-bottom amphipods of the Skagerrak. Zool.

Bidrag fran Uppsala, vol. 28, pp. 297—492, figs. 1-67, charts 1-6.
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*1775. Systema entomologiae, sistens insectorum classes, ordines, genera,
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*1779. Reise nach Norwegen, mit Bemerkungen aus der Naturhistorie und
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*1793. Entomologia systematica emendata et aucta: Secundum classes,
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Fasrictius, O.

*1780. Fauna Groenlandica, systematice sistens animalia Groenlandiae
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vernaculumque; synonyma auctorum plurium, descriptionem, lo-
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Gites, G. M.

1885. Natural history notes from H. M.’s Indian Marine Survey Steamer
‘Investigator,’ Commander Alfred Carpenter, R. N., Commanding,
No. 2: Description of a new species of the amphipod genus Melita
from the Bay of Bengal. Journ. Asiatic Soc. Bengal, vol. 54,
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1888. Natural History notes from H. M.’s Indian Marine Survey Steamer
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1890. Natural History notes from H.M.’s Indian Marine Survey Steamer
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Gois, A.

1866. Crustacea Amphipoda maris Spetsbergiam alluentis, cum speciebus
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GRUBB, A. E.

1861. Hin Ausflug nach Triest und dem Quarnero. Beitrige zur Kenntniss
der Thierwelt dieses Gebietes. Berlin, Nicolaische Verlagsbuch-
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1864a. Beschreibungen einiger Amphipoden der istrischen Fauna. Archiv
fir Naturgeschichte, Jahrg. 30, vol. 1, pp. 195-213, pl. 5.

1864b. Die Insel Lussin und ihre Meeresfauna. Nach einem sechswéchent-
lichen Aufenthalte geschildert von Dr. Adolph Eduard Grube.
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MARINE GAMMARIDEAN AMPHIPODA 491

GuERIN, F.-E.

1836. Description de quelques genres nouveaux de crustacés appartenant
a la famille des hypérines. Magasin de Zoologie Journal, Ann.
6, Cl. 7, pp. 1-10, + 2 unnumbered pp., pls. 17-18, 19.

GuRJANOVA, E.

1934. Neue Formen von Amphipoden des Karischen Meeres. Zool.
Anzeiger, vol. 108, pp. 122-130, figs. 1-6.

1938. Amphipoda, Gammaroidea of Siaukhu Bay and Sudzukhe Bay
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1948. Amphipoda Tixogo Okeana II: Stenothoidae dal’nevostochyx morei.
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1951. Bokoplavy morej SSSR i sopredel’nykh vod (Amphipoda-Gam-
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1952. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz dal’neve-
stochnykh morel. Akad. Nauk SSSR, Trudy Zool. Inst., vol.
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1953. Novye dopolnenija k dal’nevostochnoi fauna morskik bokoplavov.
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1-19 [in Russian].

1955. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz severnoi
chasti Tixogo Okeana. Trudy Zool. Inst., Akad. Nauk SSSR,
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1958. K fauna amfipod (Amphipoda) Ostrova Makkuori. Biol. Sovet.
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1962. Bokoplavy severnoi chasti Tixogo Okeana (Amphipoda-Gammaridea)
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Hansen, H. J.

1887. Malacostraca marina Groenlandiae occidentalis. Oversigt over
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L’Harpy, J. -P., and Trucuot, J. -P.

1964. Description d’Uncinotarsus pellucidus nov. gen., nov. sp., nouvel
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HASWELL, W. A.

1880a. Preliminary report on the Australian Amphipoda. Ann. Mag. Nat.
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1880b. On Australian Amphipoda. Proc. Linn. Soe. New South Wales,
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1880c. On some additional new genera and species of amphipodous crus-
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1880d. On some new amphipods from Australia and Tasmania. Proc. Linn.
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1882. Catalogue of the Australian stalk- and sessile-eyed Crustacea.
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285-135 O - 69 - 32

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HasweE.ui, W. A.—Continued

1885. Notes on the Australian Amphipoda. Proc. Linn. Soc. New South

Wales, vol. 10, pp. 95-114, pls. 10-18.
HELLuER, C.

1875. Die Crustacean, Pycnogoniden und Tunicaten der K.K. Osterr.-
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Herest, J. F. W.

1793. Garneelasseln. Onisci gammarelli. Part 6 in vol. 2 of “Versuch einer
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HeERpMAN, W. A.

1889. Second annual report of the Liverpool Marine Biological Station on

Puffin Island. Proc. Liverpool Biol. Soc., vol. 3, pp. 23-45, fig. 1.
Husss, E.

*1873. Mémoire sur les crustacés rares ou nouveaux des cétes de France,
(Vingt et uniéme article): Description de crustacés nouveaux
appartenant 4 la légion des Edriophtalmes, de l’ordre des Amphipodes,
de la famille des Piscicoles, de la tribu des Enoplopodes, nobis, du
genre des Ichthyomyzoques, nobis. Ann. Sci. Nat., ser. 5, vol. 17,
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Hoxrk, P. P. C:

*1879. Carcinologisches, grésstentheils gearbeitet in der zoologischen Station
der niederlindischen zoologischen Gesellschaft. Tijdschrift der
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1889. Crustacea Neerlandica, II: Nieuwe lijst van tot de fauna van Neder-
land behoorende Schaaldieren, met bijvoeging van enkele in de
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Houmss, S. J.

1903. Synopses of North-American invertebrates. American Nat., vol. 37,
pp. 267-292.

1905. The Amphipoda of southern New England. Bull. U.S. Bureau of
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1908. The Amphipoda collected by the U.S. Bureau of Fisheries steamer,
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Hurtey, D. EH.

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MARINE GAMMARIDEAN AMPHIPODA 493

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1954e. Studies on the New Zealand amphipodan fauna. No. 10: A new
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KossMann, R.

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*1802. Histoire naturelle, génerale et particuliére des crustacés et des
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LATREILLE P. A.—Continued
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*1825. Encyclopedie Methodique. Histoire Naturelle. Entomologie ou
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MEINERT, FR.

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496 U.S. NATIONAL MUSEUM BULLETIN 271

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1932. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les
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1933a. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les
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1934. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les
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1936. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les
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498 U.S. NATIONAL MUSEUM BULLETIN 271

Pirtot J. M.—Continued
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1824. Invertebrate Animals. [Marine Invertebrate Animals (per title page) ].
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Sars, G. O.
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1879. Crustacea et Pycnogonida nova in itinere 2do et 3tio expeditionis
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Sars, M.

1858. Oversigt over de i den norsk-arctiske Region forekommende Krebsdyr.

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500

U.S. NATIONAL MUSEUM BULLETIN 271

ScHELLENBERG A.—Continued

1926b.

1926c.

1927.

1929a.

1929b.

1930.

1931.

1935.

1937a.

SEXTON, E.
1909.

1911.

SEXTON, H.
1940.

SHEARD, K.
1936a.

Amphipoda 3: Die Gammariden der deutschen Tiefsee-Expedition.
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Report on the Amphipoda. Jn Zoological Research Cambridge
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Kritische Bemerkungen zur Systematik der Susswassergammariden.
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W.

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MARINE GAMMARIDEAN AMPHIPODA 501

1936b. Amphipoda from a South Australia reef, Part I. Ree. So. Australian
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1930. The Amphipoda of the Cheticamp Expedition of 1917. Contr.
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1933b. Amphipoda from Florida and the West Indies. American Mus.
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1935. The Amphipods of Porto Rico and the Virgin Islands. Sci. Survey,
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1945b. The amphipod genus Unciola on the east coast of America. American
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502 U.S. NATIONAL MUSEUM BULLETIN 271

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1894. The Amphipoda collected during the voyages of the William Barents
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1914. Crustacea from the Falkland Islands collected by Mr. Rupert Val-
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STEBBING, T. R. R., and Rosertson, D.

1891. On four new British Amphipoda. Trans. Zool. Soc. London, vol. 13,

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1925a. Crustacea Malacostraca, VI: (Amphipoda, II). Danish Ingolf-
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504 U.S. NATIONAL MUSEUM BULLETIN 271

STEPHENSEN K.—Continued

1925b. Danaella mimonectes (n. gen., n. sp.), a new bathypelagic gammarid
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STIMPson, W.

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MARINE GAMMARIDEAN AMPHIPODA 505

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506 U.S. NATIONAL MUSEUM BULLETIN 271

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Appendix I

Dissection of an Amphipod
For Right-handed Operators.

In a flat-bottomed syracuse dish, the amphipod is laid on its left
side with its legs projecting away from the operator, so that it appears
up-side down through the oculars of the stereoscope. The body is
covered to more than twice its depth with alcohol to prevent the
effects of surface tension during dissection and to ameliorate the
glare of reflected light from projecting legs.

A pair of jeweler’s forceps with very fine points, a fine dissecting
needle such as an insect pin mounted on a stick of wood, and a coarse,
standard dissecting needle are used. At least five standard glass
slides, one depression slide, and six thin cover slips are needed. A
small drop of glycerine is placed on two of the flat slides, a tiny drop
each on three of the flat slides, and the depression-slide concavity is
sparingly filled with glycerine (other media are used for permanent
mounts; permanent slides have the disadvantage of restricting the
manipulation of mounted parts for 3-dimensional observation; parts
from glycerine slides may be stored permanently in alcohol in a tiny
vial made of a bit of capillary tubing with one end closed by melting
in a fire, the other end stoppered with cotton, pith, or plastic foam).

One commences removing the pereopods (legs) of the amphipod
at either the fourth or fifth coxa (sideplate) depending on which of
these coxae is largest or would pull away from the body without
entangling other legs or coxae. The amphipod is up-side down on its
left side, being held with a coarse needle in the left hand through a
body segment or with forceps or a blunt stick, and the coxa is being
pulled and ripped gently at its base with the fine forceps. In most
cases the coxa can be pulled free of the body carrying some of its
proximal musculature. Occasionally the firmness of the attachment
dictates the use of a fine scalpel.

When the coxa is removed, the remainder of the leg and gill (and
if a female the brood lamella) will come with it. As the legs are excised
identifying marks are noted in order to record the leg sequence for
positioning on the slide. Particularly confusing are coxae 3 and 4
because they are often similar in size and shape, as are the last three
pereopods.

507

285-135 O - 69 - 33

508 U.S. NATIONAL MUSEUM BULLETIN 271

Pereopods and gnathopods are removed to one side of the dish
until all seven legs have been collected. Antennae 1 and 2 are dissected
at their bases (right side only). Care in removal of antenna 2 at its
juncture is needed because it often breaks easily at joint 2 or 3.

The seven coxae-legs and two antennae are removed in a group
from the dish of alcohol to the flat slide with the largest drop of glyc-
erine. When placed in the glycerine the parts will disperse the drop,
but a light breath of air will accelerate evaporation of the alcohol
and the amalgamation of the puddle. The legs must be fully immersed
in the glycerine to prevent drying and uptake of air bubbles. Do
not put on the cover slip.

The right uropods 1, 2, and 3, both lobes of the telson, and one
member of each pair of the pleopods are removed and placed on two
of the flat slides with tiny drops of glycerine; the parts are manip-
ulated, while the glycerine puddles coalesce, and arranged so that
their respective dorsal (uropods) and anterior (pleopods) sides are
up. A clean cover slip, gripped in the forceps, is lowered horizontally
over the glycerine until it can be dropped smartly onto the puddle
without engaging air bubbles. Glycerine is to be applied sparingly
so as to prevent excessive sliding of the cover slip. If the perimeter
of the cover slip lacks glycerine it may be added later by placing a
small drop at the edge.

Before removing mouthparts determine whether they are grouped
in a coniform or quadratiform bundle from lateral view.

Mouthparts are removed from the head, again with the amphipod
head pointing away from the observer so that motion to the right
with the forceps can be used to snap off the mouthparts. The maxil-
lipeds, which are the most posterior mouthparts, cover all the other
mouthparts and must be removed at their base first; both maxillipeds
will come off together. More anteriorly, a pair of bilobed second
maxillae is to be removed and then the first maxillae, each of which
appears to have three lobes (inner lobe, outer lobe, and palp but in
a few genera lacking a palp). The inner lobes are difficult to remove in
connection with the outer unless special care is taken and caution
must be exercised not to damage the lower lip. Mandibles are removed
next; they are usually brittle and easily broken; they are most easily
removed by rotating them to ascertain the basal muscular attach-
ment and snipping this with forceps. Sclerotic connections to upper
and lower lips also must be broken to avoid their damage. Usually
each mandible will have a palp. After maxilla 1 and the mandibles
are removed, a lower lip and an upper lip will remain; the lower lip
is extensive and for removal must be grabbed deeply in its muscular
and tendon attachments without separating the inner and outer lobes.
After practice one may desire to remove lower lips before dissecting

MARINE GAMMARIDEAN AMPHIPODA 009

mandibles as the two mouthparts often are closely connected with
tissues and the mandibles will tear the lower lip when being removed.

The upper lip and epistome are not removed from the head at
this time. Their interrelationship from lateral view must be pre-
served. The observer should note the condition of the ventral margin
of the upper lip from anterior view (rounded, incised, truncated)
before mounting the carcass on the depression slide.

Mouthparts are transferred to a tiny drop of glycerine on a flat
slide, arranged in sequence and fitted with a cover slip. Preferably
the mouthparts are arranged so that the following parts project
upward or are on top: mandibular molars and the inner lobes of
lower lip, maxillae, and maxillipeds. If the base of the maxillipeds
curves upward, it may be cut off so that the cover slip will set firmly.
The mandibles should be arranged with the molars projecting
obliquely toward the observer or directly lateral, if the center of
gravity so permits. Often mandibles are placed on a separate slide
with supports for the cover slip to prevent crushing. Supports may
be made of wire or sand grains.

Because a unilateral dissection has been made, the remaining
amphipod carcass has a complete set of pereonal and pleonal parts
remaining on one side (the left if done by a right-handed operator).
Coxa 1 and any other (left) legs are removed which would obscure
the head and pleon from lateral view. The carcass is mounted right
side down in the glycerine of the depression slide and a cover slip
firmly set. If the amphipod is so large that it will be crushed by
the cover slip or lie in a tilted position, two pieces of wire of appro-
priate thickness (or variously thick insect needles, pins, paper clips
cut with nipping pliers) are placed on each side of the amphipod,
which is covered with sufficient glycerine to fill the area between
the two wires, and the cover slip set on the supports. The top glass
should fit the carcass snugly so as to hold it in place but not to crush
it. Glycerine has sufficient surface tension so that it will not leak
_out from under the elevated coverslip as long as the slide is kept
in a horizontal position.

One now returns to the first flat slide on which were placed the
seven right pereopods (including gnathopods) and the two right
antennae. They are arranged in order from anterior to posterior in
two rows. Gills from legs 2 to 7 (or 2-6 or 2-5) are removed and
placed in sequence on the fourth flat slide of glycerine. If the animal
is a female, the brood lamellae are removed in sequence and placed
in a row on another slide. At this stage one is working from glycerine
to glycerine without the effects of a change in surface tension so
that it is easy to keep the parts in order, making notes of charac-
teristics that will permit proper orientation. Cover slips are set
in place.

510 U.S. NATIONAL MUSEUM BULLETIN 271

If the amphipod species is dimorphic, a slide of antennae, gnath-
opods, and uropod 3 of the other sex should be prepared for rapid
identification; but a full dissection should be prepared for descriptive
purposes.

Parts of greatly differing thickness should not be intermingled
on the same slide as the thinner parts will not be properly fitted
by the cover slip. Delicate parts may need artificial support of the
cover slip as noted above in discussion of the mandibles. Dirt may
be removed from heavily setose appendages by use of a fine camel’s-
hair brush.

Eventually the student will gain sufficient experience for examination
of most parts without dissection. Even mouthparts can be partially
to fully examined by careful manipulation under a fine stereoscope
with adequate 2-directional light sources. Mandibles often can be
rotated for viewing molars without their complete removal. This
protects unique specimens from unnecessary damage or loss of parts,
or the need to mount parts permanently.

The taxonomist anticipating a need to illustrate the organism will
leave the telson and the left first coxa attached to the carcass so that
a full lateral view of the amphipod is preserved. The telson can be
removed for flat mounting after the lateral view is drawn. Usually
the left legs distal to their coxae are removed and mounted. The
lateral in toto drawing represents a composite reconstruction of body
and coxae drawn first, with legs superimposed on the drawing by use
of a microprojector or camera lucida in which degree of magnification
can be replicated. In this way legs are attached to the body drawing
in perfectly flat but somewhat unnatural condition. One must deter-
mine accurately the attachment loci of the legs to their coxae by
study of the proposed slides 5 and 6 noted below. Generally, it is
preferable to make slides of the following composition if illustrations
are to be made.

1. Depression slide with carcass.

2. Mandible, maxilliped, lower lip, with support for cover slip.

3. Maxillae, 1, 2.

4. Antennae 1, 2, with support for cover slip to allow rotation and

examination of all surfaces by movement of cover slip.

5. Gnathopod 2, if especially thick.

6. Gnathopod 1 and pereopods 1-5.

7. Pleopods 1-3.

8. Uropods 1-3, with support for cover slip to allow rotation of parts.
9. Telson (removed from carcass after lateral illustration).

10. Upper lip and epistome (removed from carcass after lateral

illustration).
11. Left legs, except for their coxae.

MARINE GAMMARIDEAN AMPHIPODA 511

12. Gills, with support for cover slip.
13. Brood lamellae.

Very frequently, preserved amphipods have broken appendages.
Sometimes the loss of uropod 3 is a consistent occurrence especially
in gammarids and oedicerotids. So few Gammaridea lack a third
uropod that the first assumption always should be that the part has
been removed accidently and close examination should be made for
sockets and musculature indicating the loss.

Antennae are often broken and such specimens should be avoided
until experience is sufficient to recognize amphipods by other means.
In the photid-corophiid complex, legs (except gnathopods) and
antennae are frequently autotomized when the animals are preserved,
and specialists usually have found other means of identification in
those families.

The ecologist making a study of a single species should be prepared
to take special care in preservation of his material to ensure com-
pleteness of the specimens. He may find slow dilution of seawater or
special anesthetics suitable to kill the organisms slowly and to prevent
autotomy.

Appendix II

Amphipod Analytical Sheet
A Checklist of Characters to be Examined Before Using Keys

Before the keys are utilized for the identification of an amphipod,
the morphological characteristics of the organism should be deter-
mined. This requires a complete dissection, mounting, and examina-
tion of parts on microscope slides as discussed in Appendix I. The
following checklist may be useful as a guide to characters that should
be determined. This procedure is very laborious at first but pays
dividends in the final result and makes familiar, more rapidly, those
characters which ultimately are most important. Principal characters
are italicized.

The checklist is designed to indicate the characters present by
circling the pertinent items in the sentences, or adding percentages
to blanks or zeros to the ends of inappropriate sentences. Descriptive
phrases applied to various shapes are minimal in number; the observer
may have to compose further refinements. But descriptive refine-
ments can be overextended to the unnecessary wastage of time, for
this checklist applies to initial analysis of the generic position and not
to fine details necessary for specific identity.

One may find it useful to replicate copies of this checklist for the
keeping of records.

BODY

(Pereon and Pleon [metasome and urosome))

General, segments:
Abnormal disproportion or enlargement (example, Danaella).
Cylindricalization (examples, Colomastigidae, Eophliantidae).
Dorsal depression with or without splaying of coxae.
Ornamentation:
Teeth, dorsal and lateral; pereon 1, 2, 3, 4, 5, 6, 7; pleon 1, 2, 3, 4, 5, 6.
Spine groups, dorsal, especially on pleon 1, 2, 3, 4, 5, 6.
Elongation of metasome: example, is it as long as 5 or 6 pereonites? (for dis-
tinguishing a few Gammaridae and Eusiridae).
Urosome:
Coalescence of segments: 1-2, 1-3, 2-3 (examples: Ampeliscidae, Kuriidae
Podoceridae).
Dorsal depression (examples, Corophiidae, Cheluridae).
Elongation of wrosomite 1 (example, Podoceridae).
Pleonal epimera:
Shape and ornamentation from lateral view, especially of epimeron 3 (primarily
for identification to species).
512

MARIND GAMMARIDEAN AMPHIPODA 013

COXAE
Size:

Normal: like basic gammaridean; 1-4 of medium length, subquadrate, 5-7
shorter, successively smaller, 5-6 slightly lobed.

Elongation (example, Stegocephalidae).

Reduction (example, Podoceridae).

Disproportional sizes of 1-4: 1 long, 2 shorter, 3 shorter, 4 long (example,
Argissidae).

Coza 1 absent or vestigial (rare examples, Bateidae, Anamizis).

Coxa 1 reduced in size and partially to fully covered by following coxae (example,
Stenothoidae). (Caution: keep amphipod specimen in unflexed condition
for proper observation.)

Coxae 1-2 or 1-3 reduced in size and covered by following coxae (primarily
genera of Lysianassidae).

Coxa 3 larger than 4 (example, Synopiidae).

Coxa 5 as long as 4 (rare).

Position:

Serial contiguity: coxae contiguous or overlapping.

Serial discontiguity (example, Podoceridae, Eophliantidae).

Concealment of one coxa by another (see above).

Lateral splaying (Phliantidae).

Shape:

Coxae 1-4 subquadrate.

Acumination of coxae 1, 2, 3, 4.

Excavation posteriorly of coxa 4: posterior margin straight, concave or deeply
excavate and/or bearing posterior lobe (contrast Stegocephalidae and
Isaeidae).

Coxa 1 tapered, expanded, oval, semicircular, quadrate, conical, acutely lobed.

Ventral serrations.

Special patterns:
Disproportional sizes of 1-4 (Argissidae, see above).
Crescentic curve formed ventrally by coxae 4-5 together (example Epimeria).

HEAD
Size:

Length as a function of one or more pereonites (1, 1-2, 1-3), head measured
on horizontal axis from front of lateral lobe to perpendicular line from
posterior cephalic extent.

Massive (see definition in glossary).

Shape:

Normal gammaridean: head cuboidal, with lateral lobes.

Globular: subspheroid; neck cylindrical.

Galeate (see definition in glossary).

Rostrum:
Length in relation to head ( %) and to article 1 of antennal ( %).
Shape: acute, spatulate; horizontal, deflexed.

Lateral lobe:

Shape and extent of projection.

Notch or ornamentation.

Ocular bulge on side of head.

Marginal details of anteroventral corner of head near insertion of antenna 2.

514 U.S. NATIONAL MUSEUM BULLETIN 271

EYES
Composition:
Presence or absence (careful examination required because eyes often lose
pigment in preservative).
Paired ommatidial mass below cephalic cuticle (common).
Cuticular lenses in lateral pairs (Ampeliscidae almost exclusively). Occasionally
lenses occupy anterior surface of head.
Diffused pigment or stain.
Quadrigeminous lenticular bodies (example, Argissidae).
Bright pigmentary masses enveloping brain (especially Ampeliscidae).
Shape: ovoid, flask-like, reniform.
Position:
Near lateral cephalic surface.
In lateral lobes.
In rostrum (especially Oedicerotidae).
Dorsally confluent (especially Oedicerotidae).
Accessory detached ommatidia (pattern often confused by preservational
accidents).
Occupying cephalic extent almost fully (Hyperiopsidae).

ANTENNA 1

Length:
As percent of total body (front of head to base of telson ( %).
In relation to antenna 2 and to its peduncle ( %).
Flagellum:
Proportion to peduncle ( %).
Elongation of basal article (often conjoint, composed of incompletely seg-
mented articles).
Proportion to peduncular article 3, especially when latter elongated ( %).
Number of articles: 0, 1, 2, 3, 4, 5, 6-10, 11-15, 16++.
Peduncle: Proportion to head.
Relative lengths of all three articles (value of 100 assigned to length of article
1): 1=100%; 2=—( %);3=( %).
Ornamentation on any article, all sides.
Distinctive spines or setal bundles.
Possible geniculation between articles (example, several Haustoriidae).
Accessory flagellum (attached to article 3 of peduncle, medial):
Number of articles: 0, 1, 2, 3, 4, 5-10, 11+.
A fused scale.
Special shape.
Elongation of basal article.
Accessory organs:
Calceoli.
Aesthetases, if especially enlarged or in dense bundles.

ANTENNA 2

Length as percent of body length (including head) ( %).
Peduncle: proportion to head ( %).
Relative proportions of articles: 4=100%;5=( %).
Tumidity of articles 3, 4, or 5; article 1 large and subspherical.
Gland cone and/or ensiform process on articles 2 and 1; extreme enlargement
and shape. (Ensiform process generically important primarily in Phoxo-
cephalidae. Gland cone often medial and hidden from lateral view.)

MARINE GAMMARIDEAN AMPHIPODA 515

Flagellum: proportion to peduncle ( %) or article5( %).
Number of articles: 0, 1, 2, 3, 4, 5, 6-10, 11-15, 16+.
Ornaments:
Aesthetases, calceoli on peduncle and/or flagellum.
Distinctive spine groups.
“Fossorial’’ condition, with long plumose setae.

MOUTHPARTS

From lateral view forming a conical bundle below head (rare) or a quadrate bundle
(common). To be examined before dissection.
Amalgamation of mandibles and mazillae into ventral keel (Anamixidae).

EPISTOMAL-LABRAL COMPLEX (prebuccal)

Lateral view (primarily of generic importance in Lysianassidae) :

Epistome and labrum separated by notch (common) or coalesced.

Epistome formed as lobe dominating labrum, vice versa, or produced together.
Shape of lobes:

Epistome: flat, rounded, acute.

Labrum: flat, rounded, acute.
Prebueccal mass inconspicuous and of normal gammaridean proportions.

UPPER LIP (anterior view)

Ventral margin: rounded, truncate, incised, lobed asymmetrically or symmetri-
cally.

MANDIBLE

Shape and size of body: bulky (Synopiidae), styliform (Acanthonotozomatidae) ,
elytriform (Stegocephalidae, Pardaliscidae), OR normal (Gammaridae).
Incisor: normal, extremely broadened; needle-like, toothed, untoothed, teeth
separated by flat margin.
Lacinia mobilis, if present on either right or left mandible: toothed, special
shape such as vermiform.
Spines proximal to lacinia mobilis: 1-2; 3-6; 7+.
Molar: Absent.
Size: small (fig. 1267), medium (fig. 126e), large (fig. 126g), fully dominating
mandible (fig. 166d).
Shape: cylindrical, cuboidal, laminate, conical, tuberous.
- Texture: triturative (rasp-like), spinose (spines articulate), setulose, minutely
fuzzy, striate, smooth.
Accessory seta or spine on triturative molar.
Palp:
Number of articles: 0, 1, 2, 3.
Attachment position relative to molar: over (level with), distal to, proximal
to (variation from “‘level”’ primarily in Lysianassidae).
Relative length of articles: 1=( %);2=100%; 3=( %).
Shape:
Article 8: cylindrical, falconiform, tuberculiform.
Article 2: occasionally curved strongly.
Article 1: rarely with distal cusp.
Setation: article 3: distal only, medial.
Disymmetry of right and left members (especially lacinia mobilis, incisor, spine-
row).

516 U.S. NATIONAL MUSEUM BULLETIN 271

LOWER LIP

Normal gammaridean (fig. 2).

Inner lobes: weak (fig. 71d), absent.

Mandibular projection of outer lobes: pointed, obtuse, absent.

Outer lobes: distally notched; medially excavate (Ampithoidae, figs. 50c,d).
Special shapes: Pleustidae (fig. 37a), Trischizostoma (fig. 125v).

MAXILLA 1
Inner plate:
Size: absent (very rare), small (fig. 127f), medium (fig. 127e), as large (broad
and bulky) as outer plate (rare).
Setation: terminal, medial, or both.
Number of setae: 1, 2, 3, 4-6, 6-12.
Structure of setae: normal; sickle-shaped or strongly constricted.
Outer plate:
Number of spines: 1-4, 5-6, 7-8, 9-11+.
Shape of spines: normally slender, extremely stout, some bifid, some serrate,
in two distinct groups by position or structure.
Palp:
Number of articles: 0, 1, 2.
Normal structure: article 1 short, article 2 long.
Article 1 long, article 2 short.
Modifications: strongly bent (geniculate), foliaceows, bearing scales (examples:
Stilipedidae, H yperiopsidae).

MAXILLA 2

Normal gammaridean (fig. 1270).

Abnormally small, plates partially coalesced, setae very sparse.
Breadth of lobes: subequal, inner broader, outer broader.

Axial divergence of lobes (fig. 127)). |

Extension of outer plate on basal article (fig. 161a).

Specialized spines (rare).

Extent of medial setation on inner plate: strong, sparse, absent.

MAXILLIPED
Inner lobes (proximal):
Size: vestigial (fig. 128e), normal (fig. 128a).
Abnormal shape: foliaceous (rare), styliform (note: inner lobes often appear
styliform if not fully depressed by cover slip).
Outer lobes:
Size relative to inner: usually larger, vestigial (fig. 114d), foliaceous (rare).
Spination: absent, medial, distal.
Palp:
Extension in relation to outer plate: shorter, equal, longer.
Number of articles: 0, 2, 3, 4.
Medial or terminal extensions of articles, e.g., lobes, cusps: articles 1, 2, 3.
Elongation of articles 1, 2, 3. ©
Terminal palp article (usually 4): claw-like (normal); barrel-shaped; vestigial;
bearing distal nail, spine, or setae.

MARINE GAMMARIDEAN AMPHIPODA 517

GNATHOPODS

Judgment of gnathopods 1-2 as: feeble together (fig. 165a), normal (basic gam-
maridean with gnathopod 2 powerful), powerful together (fig. 115a).

GNATHOPOD 1 (excluding coxa 1)

Present, vestigial, or absent (Bateidae and Paranamizis only).
Size (or length) relative to gnathopod 2: smaller, equal, larger.
Sexual dimorphism: similar or different in male and female.
Articles:
2: Length in relation to coxa 1 (rarely important except when coxa | abnormal).
3: Length normal; or elongate (like fig. 3a).
4: Merochelation: with strong thumb-like extension (rare).
5: Length relative to article6( %).
Posterior lobe: present, weak, absent.
Carpochelation: with strong distoposterior tooth or teeth forming thumb
or guarding article 6.
6: Breadth: relative to article 5; wider, equally wide, narrower.
Shape: ovate, pyriform, quadrangular, rectangular, linearly rectangular,
styliform.
Palm:
Present, absent, undecided.
Slope: transverse, oblique: slight; moderate; extreme.
Chela if present: parachelate (describe if strongly chelate).
Definition of proximoposterior corner of palm: spines, protuberance, tooth,
change in slope only.
Ornamentation: special spines, teeth.
7: Fit of the dactyl to palm: congruent, overlapping, not fitting.
Shape and ornaments: claw-like (normal); vestigial; absent (rare); with
special setae or spines; hidden in setae or cirri; flagelliform.
Distal articles especially scaly or with small stiff setae (Lysianassidae).

GNATHOPOD 2
Articles:
3: Length normal; elongate (fig. 3a).
4: Merochelation: with strong thumb-like extension.
5: Length relative to article6( %).
Posterior lobe: present, weak, absent.
Carpochelation: with strong distoposterior tooth or teeth forming thumb or
guarding article 6.
Seales or stiff setae (pineapple cushion of Lysianassidae).
General shape: cup-like, elongate rectangle, ovate.
6: Breadth relative to article 5: wider, equal, narrower.
Shape: ovate, pyriform, rectangular, linearly rectangular, mitten-like (Lysi-
anassidae, Talitroidea), quadrangular.
Palm:
Present, absent, undecided.
Slope: transverse; oblique: slight, moderate, extreme.
Chela if present: parachelate (describe if strongly chelate).
Definition of proximoposterior corner of palm: spines, protuberance, tooth,
change in slope only.
Ornamentation: special spines, teeth.

518 U.S. NATIONAL MUSEUM BULLETIN 271

7: Fit of this dactyl to palm: congruent, overlapping, not fitting. Shape and
ornaments: claw-like (normal); vestigial; absent (rare); with special setae,
spines, or processes.

Sexual dimorphism in female: gnathopod 2 like male but much smaller, like gnath-

opod 1 and of similar size, like gnathopod 1 but larger.

PEREOPODS 1-2

Internal glands present or absent.

Orientation of pereopod 2 like that of pereopod 3 (Hohaustorius).
Chelate or prehensile.

Articles 4—5, 4-6, or 4, 5, 6 inflated strongly (rare).

Article 4 extraordinarily elongate (Ampeliscidae, Hyperiopsidae).
Special spines on article 6 near claw: spines striate, hooked.
Article 7 absent (Haustoriidae).

PEREOPODS 3-5

Relative lengths: pereopod3( %);4=100%;5=( %).

General stucture:
All similar in structure and slightly longer successively (normal).
Article 2 ecpanded: pereopod3( ),4( ),5(_).
Expanded lobe of article 2 of pereopod 5 different from pereopods 3 and 4.
Chelate, subchelate, or prehensile: pereopods 38, 4, 5.
Fossorial setation (see glossary) present, absent.
Article 7 absent (Haustoriidae, Stegocephalidae): pereopds 3, 4, 5.
Pereopod 5 reduced to fewer than 6 articles.

PLEOPODS

(Rarely significant in marine Gammaridea [but see Phliantidae especially]).
Relative length (size) of each pair: 1=100%;2=( %);3=( %).
Length of longest ramus relative to peduncle ( %).
Length of inner ramus to outer ( %) (note absence of rami or low number
of articles).
Shape of coupling hooks on peduncles.
Lobation of peduncles.
UROPODS 1 and 2

Absence (rare) or presence.

Projection along following uropods.
Uropod 1 reaching ( %) along uropod 2;( %) along uropod 3.
Uropod 2 reaching ( %) along uropod 3.

Relative length of rami: outer or inner shortened (occasional), inner absent or
vestigial (rare).

Spination density of peduncle and rami (usually of specific value only).

Incision of inner ramus (example Anonyz).

UROPOD 3
Absence (rare) or presence.
Rami absent (rare).
Length relative to other uropods; extension beyond longest of other uropods ( % of
its own length).

MARINE GAMMARIDEAN AMPHIPODA 519

Length of peduncle relative to wrosomal segment 3 ( %), to peduncles of other
uropods ( % of peduncle of uropod 1), or to telson ( %).

Length of longest ramus relative to peduncle ( %).

Length of inner ramus to outer ( %).

Shape of rami: styliform, lanceolate, barrel-shaped, foliaceous.

Articles of outer ramus (1 or 2).

Minute ornamentation and hooks on rami (especially Ischyroceridae and Am-
pithoidae).

Special peduncular processes.

TELSON

Absence (rare) or presence.
Fused to urosomite 3 (rare).
General shape and length (make sketch).
Length in relation to urosomite 3( %) or uropod3( %).
Degree of cleft between lobes ( %), emarginate only.
Ornamentation: apically pointed, notched, trifid, truncate, rounded, concave.
Greatly enlarged and with ventral keel (rare), forming dorsoventral plate.
Dorsoventrally thickened (‘‘fleshy’’) (example, Isaeidae), bearing lateral nobs,
scales, hooks.
SEXUAL DIMORPHISM

(primarily for identification to species)

Especially:
Antennae:
Eyes:
Gnathopods:
Coxae:
Pleonal epimera 1-3:
Uropod 3:
Urosomal teeth:

Appendix III
Glossary

accessory flagellum. Thesecondary ramus of antenna 1, often absent or vestigial
(fig. 1), and attached medially to peduncular article 3.

aesthetasc, aesthete. Sensory setae of antennae, flattened and nontapering.

article. The segment of an appendage (fig. 1).

calceolus. A small globular or helmet-shaped, articulate sense organ on the
antennae; presumably a modified aesthetasc; of rare occurrence in Gammaridea
and most often seen in Eusiridae (fig. 89)).

carpochelate. Immovable finger of prehensile appendage occurring on carpus
(article 5); examples: Lewcothoe, Microdeutopus.

chela. Immovable finger of prehensile appendage.

chelate. Descriptive of the palm of a gnathopod protruding as an immovable
finger on which the dactyl closes (fig. 129h). See parachelate, carpochelate,
propodochelate, merochelate, complexly chelate.

claw, claw-like. Descriptive of a talon or simple, tapering nail. [Not descriptive
of chelae as used in decapod terminology.]

compressed. Flattened from side to side.

conjoint. Describing the basal amalgamation of flagellar articles on antennae.

corneal lens. A biconvex cuticular body occurring directly in or on the chitinous
cephalic surface in Ampeliscidae; contrasted with subcuticular ommatidia.

coxa, coxal plate. [Terms used synonymously herein.] Article 1 of a pereonal
appendage, expanded into a lateral lamella (fig. 1). [Terms for other articles of
the appendages such as basis, ischium, merus carpus, propodus, and dactyl
are frequently but not universally used in Gammaridea; instead, the articles
are simply numbered. ]

dactyl. Talon-like terminal article of pereopods (article 7) or maxillipeds (articles
3 or 4).

depressed. Flattened dorsoventrally.

emarginate. Descriptive of the concave posterior end of an uncleft telson (fig.
42¢e).

entire. Descriptive of an uncleft telson (fig. 38f).

epimeron. A lateral pleuron of pleonites 1-3; the ventrolateral plate-like extension
of the body segment (fig. 1).

epistome. The anterior surface of the head above the labrum; this area is often
extended ventrally to appear as a part of the labrum and may be anteriorly
produced as a cusp or lobe (fig. 4a).

flagellum. The distal portion of either antenna 1 or 2; on antenna 1 it commences
with article 4, on antenna 2 with article 6; because basal peduncular articles
of antenna 2 are often difficult to resolve, the juncture may be recognized
between the elongated final peduncular article and the shortened first flagellar
article which is followed by similar short articles; on antenna 1, however, article
1 of the flagellum is occasionally elongate and apparently composed of non-
segregated (thus conjoint) daughter articles (fig. 1).

520

MARINE GAMMARIDEAN AMPHIPODA 521

fossorial. Associated with the habit of burrowing, often referring to the exces-
sively spinose or setose condition of appendages used for burrowing by Gam-
maridea; especially applicable to Haustoriidae, Oedicerotidae, Phoxocephalidae
with some setae of articles 4-6 of pereopods 3-5 more than half as long as
those articles; and some spines in groups forming submarginal rows perpendicu-
lar to margins; long setae also occur on “‘filter’’ feeders such as Ampeliscidae.

galeate. Descriptive of the helmet-shaped heads of various oedicerotids and
synopiids.

gamopod. A gnathopod; referring to the use of gnathopods for grasping members
of the opposite sex during amplexus.

geniculate. Permanently bent, usually in reference to the flexeda ntennae of some
haustoriids, or the outer lobes of maxilla 2 in some stegocephalids in which the
bend occurs between articles; or applicable to bent palps of maxilla in Hyperiop-
sidae in which the bend occurs on one article.

gnathopod. One member of the first two pairs of free thoracic appendages; these
appendages differ in function and usually in appearance from following pereo-
pods; often called pereopods.

incisor. The apical portion of the mandible usually formed into a toothed chewing
edge or untoothed chopping plate.

joint. The juncture between two articles of an appendage.

labrum. (See upper lip.) ’

lacinia mobilis. An articulated accessory plate proximal to the mandibular
incisor, often absent or missing on either left or right mandibles, occasionally
indistinguishable from a spine of the spine-row.

low lip (labium). A fleshy complex posterior to the mandibles, always composed
of at least one pair of lobes (outer), often with a medioproximal pair of inner
lobes; the lateroproximal ends of the outer lobes are often attentuated as alae
and are denoted as mandibular lobes (fig. 2g).

mandible. The anterior movable appendage of the buccal group; usually composed
of a body bearing a distal incisor, a lacinia mobilis, spine row, molar, and
d-artculate palp (fig. 2f).

massive. A term applied to the heads of Synopiidae and Oedicerotidae; head
as long as pereonites 1-3 combined and as tall as or taller than long (length
not including rostrum). Heads of Ampeliscidae and Phoxocephalidae are
elongate but not massive; heads of Acanthonotozomatidae are as tall as long
but are not as long as pereonites 1-3 combined.

maxilla 1. A pair of cephalic appendages posterior to the lower lip; for taxonomic
purposes only three portions of each member are named: the medial lobe
(plate) usually bearing marginal setae, the lateral and larger lobe (plate)
bearing terminal spines, and, attached to the outer lobe, a palp usually com-
posed of two articles but occasionally absent (fig. 29).

maxilla 2. A pair of cephalic appendages posterior to maxilla 1; for taxonomic
purposes each member recognized as a pair of lobes (plates) medial and lateral,
usually strongly setose (fig. 272).

maxillipeds. The posteriormost pair of ‘‘cephalic’’ appendages, representing
the primitive first thoracic segment now amalgamated with the head but in
amphipod taxonomy not included in the sequential numbering of thoracic
appendages; for taxonomic purposes recognized as a pair of basally amalga-
mated appendages, each member composed of a proximal (inner) plate, a distal
(outer) plate, and a palp of four articles, rarely reduced to 3 or 2 articles or
absent (fig. 21).

merochelate. Immovable finger of prehensile appendage occurring on merus
(article 4); example: gnathopod 1 of Aora.

22 U.S. NATIONAL MUSEUM BULLETIN 271

mesosome. The pereon or thorax. [Term rarely used.]

metasome. Pleonites 1-3. [Term rarely used.]

molar. A process of the mandible, located on the midmedial margin; when
completely developed it is a large, massive, subcylindrical body with a surface
of ridges and teeth used for grinding (triturative) (fig. 2f).

ommatidium (singular), ommatidia (plural), ommatidial (adjective). Terms
applying to the parts of the subintegumentary compound eye, not to be confused
with the corneal lenses of the integument of Ampeliscidae.

palm. A posterior surface or margin of article 6 of a gnathopod or pereopod on
which article 7 (dactyl) closes for the purpose of prehension; usually recog-
nizable because of expansion of article 6 or by occurrence of special spines or
ornamentation and usually with a proximal defining limit marked by a change
in marginal slope or occurrence of special spines (fig. 2e).

palp. Terminal articles of a buccal appendage, in Amphipoda occurring only
on mandibles, first maxillae, and maxillipeds as the stenopodous terminal
articles distal to the expanded outer plates or main body.

parachelate. A rarely used term in Amphipoda applied to propodochelate
gnathopods and pereopods in which the immovable finger is distinct, but
article 6 is otherwise unexpanded or nonpalmate and the dactyl strongly
overlaps the apex of the immovable finger; gnathopods of various Eophliantidae
are good examples but the term may also be applied to numerous other cases,
such as those linear, chelate gnathopods of Sebidae, Didymocheila, and various
second gnathopods of Lysianassidae.

peduncle. The basal articles of a fundamentally biramous appendage; in
Amphipoda applied to antennae, pleopods, and uropods; antenna 1 with three
peduncular articles, antenna 2 with five peduncular articles (but appendage
not biramous); pleopods with one definitive ,-eduncular article but remnants
of others occurring proximally; uropods each with one peduncular article
(fig. 1).

pereon. The complex of seven free thoracic segments bearing gnathopods and
pereopods, not including the maxillipeds (fig. 1).

pereonite. A segment of the pereon.

pereopod. A walking, grasping, standing, or feeding appendage attached to a
pereonite; normally composed of seven articles, including coxa; in Amphipoda
the first two pairs are often termed gnathopods and only the last five pairs of
thoracic legs are called pereopods (fig. 1).

plate. A flattened lobe on an article of a maxilla or maxilliped.

pleopod. A biramous swimming appendage on pleonites 1-3, one pair for each
pleonite (fig. 1).

pleon. The abdomen (of six free segments in Gammaridea, rarely with some
segments coalesced). See metasome and urosome.

pleonite. A segment of the pleon.

prebuccal complex. The labrum and epistome together.

prehensile. Adapted for seizing or grasping; applicable to but rarely used for
gammaridean gnathopods; especially useful in denoting pereopods of cyphocarids
Isaea, Pleonexes, etc., which either are subchelate or chelate, or have distinct,
spinose palms or nonskid surfaces indicating their use in grasping.

propodochelate. Synonymous with chelate.

propodus. ‘The sixth article of a thoracic appendage (especially used to denote
the palmar article of a gnathopod).

scale, scale-like. Terms applied to the accessory flagellum when forming a
small lamella immovably fused to article 3 of antenna 1; and to the inner ramus
of uropod 3 when strongly reduced and plate-like.

MARINE GAMMARIDEAN AMPHIPODA 523

simple. Used in amphipod taxonomy to denote the absence of spines or setae
on appendages; or the occurrence of buta single article in the ramus of a uropod;
or especially to the absence of a palm ona gnathopod or pereopod. Distinction
between subchelate and simple is often weak.

splayed. Descriptive of the lateral spreading of coxae, especially in Phliantidae.

subchelate. Article 6 of a gnathopod or pereopod having a distal palm against
which article 7 closes; a prehensile condition in which the palm is not produced
to form a finger; intermediate in condition between chelate and simple. Com-
plexly subchelate or complexly chelate are terms referring to the formation
of a false chela by protrusion of teeth, cusps, or lobes from articles other than
the sixth and upon which article 7 impinges to form a prehensile condition;
occurring especially in Aoridae, Corophiidae, and Leucothoidae (fig. 2e).
See “simple.”

telson. A flap dorsal to the anus attached to pleonite 6, primitively bilobed
but usually in Amphipoda with bases coalesced and often with lobes completely
coalesced to form a single plate (figs. 1, 2d).

triturative. Descriptive of the rasp-like surface of a mandibular molar, composed
of teeth, ridges, and cusps.

upper lip (labrum). A fleshy lobe attached to the anterior cephalic margin in
front of the mandibles; occasionally the anterior surface of the labrum _ pro-
trudes as a lobe or cusp; often the cephalic area to which the labrum is attached
is recognizable as an “‘epistome’”’ and may also be lobed; or both labrum and
epistome may be indistinguishable and produced together as a single lobe.

uropod. Oue member of the three pairs of terminal pleonal appendages, each
formed of a peduncle and two rami (fig. 1) (occasionally rami of uropod 3 reduced
or absent, rarely rami of uropods 1—2 absent or reduced).

urosome. The complex of pleonites 4, 5, 6, carrying uropods, and telson. Often
numbered as urosomites 1, 2, 3 (fig. 1).

urosomite. A segment of the urosome.

285-135 O - 69 - 34

Index and Familial Assignment of Gammaridean Genera

[Valid genera in capitals; subgenera in capitals and parentheses;
dubiously valid genera in brackets. See Stebbing (1888, 1906) for
additional nomina dubia and nuda not assignable to valid genera.
Page numbers of primary citations of genera in boldface.|

Acanthechinus, to ACTINACANTHUS,
Paramphithoidae, 393
Acanthogrubia, to CYMADUSA,
Ampithoidae, 144
ACANTHOHAUSTORIUS, Haustori-
idae, 254
ACANTHONOTOZOMA, Acanthono-
tozomatidae, 120
ACANTHONOTOZOMELLA,
thonotozomatidae, 122
ACANTHONOTOZOMOIDES, Acan-
thononotozomatidae, 122
Acanthonotus, to ACANTHONOTO-
ZOMA, Acanthonotozomatidae,
120
Acanthopleustes, to AMATHILLOP-
SIS, Paramphithoidae, 394
Acanthosoma, to PARAMPHITHOH,
Paramphithoidae, 396
ACANTHOSTEPHEIA, Oedicerotidae,
378
Acanthozone, to PARAMPHITHOE,
Paramphithoidae, 396
ACCEDOMOERA, Eusiridae, 219
ACEROIDES, Oedicerotidae, 378
Aceropsis, to ACEROIDES, Oedicero-
tidae, 378
Aceros, to ARRHIS, Oedicerotidae, 381
ACIDOSTOMA, Lysianassidae, 317
Acidostomella, to SOCARNOIDES,
Lysianassidae, 363
ACONTIOSTOMA, Lysianassidae, 317
ACTINACANTHUS, Paramphithoi-
dae, 393
ACUMINODEUTOPUS, Aoridae, 150
ADELIELLA, Lysianassidae, 320
AFROCHILTONIA, Hyalellidae,
Talitroidea, 468
ALEXANDRELLA, Stilipedidae, 161,
263, 453

524

Acan-

see

ALIBROTUS, dubious genus, Lysianas-
sidae (not discussed, see Stebbing,
1906)

ALICELLA, Lysianassidae, 320

ALLOGAUSSIA, Lysianassidae [also
assigned to ? Orchomene], 320, 353

ALLORCHESTES, Hyalidae, see Tali-
troidea, 468

(Allorchestina), subgenus to H YALE,
Hyalidae, see Talitroidea, 469

AMAR YLLIS, Lysianassidae, 320

Amathia, to GAMMARELLUS, Gam-
maridae, 242

Amathilla, to GAMMARELLUS,
Gammaridae, 242

Amathillopleustes, to CLEON A RDOP-
SIS, Eusiridae, 223, 392, 393

AMATHILLOPSIS, Paramphithoidae,

394

AMBASIA, Lysianassidae, 321

AMBASIELLA, Lysianassidae, 321

AMBASIOPSIS, Lysianassidae, 321

AMPELISCA, Ampeliscidae, 8, 130

AMPELISCIPHOTIS, Isaeidae, 271

AMPHIDEUTOPUS, Aoridae, 150,
267 , 269

AMPHILOCHELLA, Amphilochidae,
136

AMPHILOCHOIDES, Amphilochidae,
136, 421

AMPHILOCHOPSIS, Amphilochidae,
136

AMPHILOCHUS, Amphilochidae, 136

AMPHIPOREIA, MHaustoriidae, 254

AMPHITHOIDES, Ampithoidae, 143

AMPHITHOLINA, Eophliantidae, 142,
211, 481

Amphithonotus, to DEXAMINE, Dex-
aminidae, 203

MARINE GAMMARIDEAN AMPHIPODA

AMPHITHOPSIS,
174

AMPITHOE, Ampithoidae, 143

ANAMIXIS, Anamixidae, 146

ANATYLUS, Dexaminidae, 202

ANCHIPHIMEDIA, Acanthonotozo-
matidae, 122

Andania, to ANDANIEXIS, Stego-
cephalidae, 440

ANDANIELLA, Stegocephalidae, 440

ANDANIEXIS, Stegocephalidae, 440

ANDANIOPSIS, Stegocephalidae, 441

ANDANIOTES, Stegocephalidae, 441

ANELASMOPUS, Gammaridae, 217,
236

ANISOGAMMARUS, Gammaridae, 42
237

Anisopus, to PLEONEXES, Ampi-
thoidae, 145

ANOEDICEROS, Ocedicerotidae,

ANONYX, Lysianassidae, 324

AORA, Aoridae, 152

AORCHO, Aoridae, 152, 184, 186, 264,
267

AOROIDES, Aoridae, 152

APHERUSA, Calliopiidae, 4, 174

Araneops, to AMPELISCA, Ampelis-
cidae, 130

ARCULFIA, Pardaliscidae, 401

ARGISSA, Argissidae, 159

ARISTIAS, Lysianassidae, 324

ARISTIOPSIS, Lysianassidae, 324

AROUI, Lysianassidae, 328

ARRHINOPSIS, Oedicerotidae, 381

ARRHIS, Oedicerotidae, 381

Aruga, to LYSIANASSA, Lysianas-
sidae, 348

Arugella, to LYSIANASSA, Lysianas-
sidae, 348

Aspidophoreia, to ALLORCHESTES,
Hyalidae, see Talitroidea, 468

Aspidopleurus, to PHIPPSIA, Stego-
cephalidae, 442

ASTYRA, Astyridae, 161, 392

Astyroides, to ALEXANDRELLA,
Stilipedidae, 161, 453

ATYLOELLA, Eusiridae, 222

Atyloides, to SCHRADERIA, Eusiri-
dae, 230

ATYLOPSIS, Calliopiidae, 175, 218

ATYLUS, Atylidae, 163

Aucklandia, to PARAMOERA, Eusiri-
dae, 227

Calliopiidae, 26,

380

y45)

Audouinia, to COROPHIUM, Coro-
phiidae, 190

(AUDULLA), subgenus of GAMMA-
ROPSIS, Isaeidae, 271

AUST ROCHILTONIA, Hyalellidae,see
Talitroidea, 469

Austronisimus, to PARAL YSIANOP-
SIS, Lysianassidae, 356

AUSTROPLEUSTES, Pleustidae, 115,
ZOOM A 22) ey

AUSTROSYRRHOE, Synopiidae, 458

Autonoe, to LEMBOS, Aoridae, 154

AZOTOSTOMA, Lysianassidae, 328

Barentsia, to GULBARENTSIA, Oedi-
cerotidae, 382

BATEA, Bateidae, 164

BATH YAMARYLLIS, Lysianassidae,
328

BATHCALLISOMA, Lysianassidae [also
assigned to SCOPELOCHEIRUS],
328, 362

BATH YCERADOCUS, Gammaridae,
238

BATHYMEDON, Oedicerotidae,
381

BATH YPANOPLOEA, Acanthonoto-
zomatidae (provisional) [seeParam-
phithoidae], 122, 389

BATH YPHOTIS, Ischyroceridae, 109,
231, 267, 276

BATH YPOREIA, Haustoriidae, 254

BATH YPOREIAPUS, Oedicerotidae,
381

BATH YSCHRADERIA, Eusiridae, 222

BATH YSTEGOCEPHALUS, Stegoce-
phalidae, 441

BEAUDETTIA, Beaudettiidae, 37, 167

Bellia, to HAUSTORIUS, Haustoriidae,
257

Bemlos, to LEMBOS, Aoridae, 154

BIANCOLINA, Eophliantidae, 39, 211,
433, 481

BIRCENNA. Eophliantidae, 212, 433

Boeckia, to LEPTOCHEIRUS, Aoridae
154

BOECKOSIMUS, Lysianassidae, 330

BOGENFELSIA, Ischyroceridae, 186,
276

BONNIERELLA, Ischyroceridae (and
subgenus to MEGAMPHOPUS,
Isaeidae), 267, 273, 276

Boscia, to MELITA, Gammaridae, 245

BOUVIERELLA, Calliopiidae, 175

13,

526

BOVALLIA, Eusiridae, 222
BRUZELIA, Synopiidae, 459
Bruzeliclla, to JASSA, Ischyroceridae,
279
BRUZELIOPSIS, Synopiidae, 82, 460
BYBLIS, Ampeliscidae, 130
BYBLISOIDES, Ampeliscidae, 132
Cacao, to STILIPES, Stilipedidae, 454
Caliniphargus, to MELITA, Gammari-
dae, 245 f
Callimerus, to AMPHILOCHUS, Am-
philochidae, 136
Calliope, to CALLIOPIUS, Calliopiidae,
176
CALLIOPIELLA, Calliopiidae, 175
CALLIOPIUS, Calliopiidae, 4, 176
Callisoma, to SCOPELOCHEIRUS,
Lysianassidae, 362
CAMACHO, Corophiidae, 188, 267
CARANGOLIA, Haustoriidae, 108, 256
CARDENIO, Haustoriidae, 256
CARINOBATEA. Bateidae, 164
‘“Carinogammar us’’, see ‘‘Marine Car-

inate Gammarus,’’ Gammaridae,
244

CAROLOBATEA, Oedicerotidae, 382,
481

CASCO, Gammaridae, 107, 238

CEBOCARIS, Lysianassidae, 331

CEINA, Phliantidae, 39, 410, 432, 433,
481

CEININA, Eophliantidae, 213, 433

CENTROMEDON, Lysianassidae, 333

CERADOCOIDES, Gammaridae, 238

CERADOCOPSIS, Gammaridae, 239

CERADOCUS, Gammaridae, 28, 42,
239

Cerapodina, to CERAPUS, Corophiidae,
188

CERAPOPSIS, Corophiidae, 188

CERAPUS, Corophiidae, 188

Chagosia, to ASTYRA, Astyridae, 161

Charcotia, to WALDECKIA, Lysian-
assidae, 368

(CHEIRIMEDEIA), subgenus
PROTOMEDHEIA, Isaeidae, 274

CHEIRIMEDON. Lysianassidae, 333

CHEIRIPHOTIS, Isaeidae, 271

CHEIROCRATELLA, Gammaridae,
239

CHEIROCRATUS, Gammaridae, 32,
239

CHELURA, Cheluridae, 39, 180

of

U.S. NATIONAL MUSEUM BULLETIN 271

CHEVALIA, Isaeidae, 106, 271

CHEVREUXIELLA, Lysianassidae,
335, 373

CHEVREUXIUS, Corophiidae, 190

CHILTONIA, Hyalellidae, see Tali-
troidea, 43, 469

Chimaeropsis, to ARGISSA, Argissidae,
159

Chironesimus, to ANONYX, Lysian-
assidae, 324

Chloris, to PARAPHERUSA, Gammar-
idae, 247

CHOSROKHES, Calliopiidae, 25, 39, 176,
217,219,408, 409

CLARENCIA, Calliopiidae, 176

CLEIPPIDES, Calliopiidae, 176, 283,
392

CLEONARDO, Eusiridae, 222

CLEONARDOPSIS, Eusiridae,
392,393

CLEPIDECRELLA, Lysianassidae,
338

COLOMASTIX, Colomastigidae, 184

CONCHOLESTES, Corophiidae, 156,
190

223,

COREMAPUS, Aoridae, 152

COROPHIUM, Corophiidae, 102, 190,
481

Cratippus, to COLOMASTIX, Colo-

mastigidae, 184

Cratophium, to JASSA, Ischyroceridae,
279

CRESSA, Cressidae, 200

CRESSINA, Cressidae, 200

CRYBELOCEPHALUS, Lysianassi-
dae, 338

CRYBELOCYPHOCARIS, Lysianas-
sidae, 338

Cuviera, to LEUCOTHOE, Leucothoi-
dae, 289

CYCLOCARIS, Lysianassidae, 338
CYCLOTELSON, Amphilochidae, 138
CYLINDRYLLIOIDES, Eophlianti-
dae, 213, 433
CYMADUSA, Ampithoidae, 144
CYPHOCARIS, Lysianassidae, 339
CYPROIDEA, Amphilochidae, 139
Cyproidia, to CYPROIDEA, Amphilo-
chidae, 139
CYPSIPHIMEDIA,
matidae, 123
CYRTOPHIUM, Podoceridae, 427
DANAELLA, Lysianassidae, 339, 373

Acanthonotozo-

MARINE GAMMARIDEAN AMPHIPODA

Danaia, to CRESSA, Cressidae, 200

Darwinia, to LAF YSTIUS, Lafystiidae,
283

DAUTZENBERGIA, Eusiridae, 223

Dercothoe, to ?7MICROPROTOPUS,
Isaeidae, 274

DERJUGIANA, Lysianassidae, 339

Dermophilus, to ?7LAFYSTIUS, Lafy-
stiidae, 283

DEXAMINE, Dexaminidae, 203

DEXAMINELLA, Dexaminidae, 203

DEXAMINOIDES, Dexaminidae, 203

DEXAMONICA, Dexaminidae, 203

Dexiocerella, to PODOCERUS, Podo-
ceridae, 431

DID YMOCHEILA, see Incertae Sedis,
39, 110,294,478

DJERBOA, Eusiridae, 167, 223

DOGIELINOTUS, Dogielinotidae, 208

Dorbaneila, to LEPECHINELLA,
Lepechinellidae, 286

Dryope, Driope, to UNCIOLA, Coro-
phiidae, 197

DRYOPOIDES, Aoridae (2), Coro-
phiidae (<7), 153, 190, 275, 426

DULICHIA, Podoceridae, 427

Dulichiella, to MELITA, Gammaridae,
245

Dyopedos, to DULICHIA, Podoceridae,
427

ECHINIPHIMEDIA, Acanthonotozo-
matidae, 124

“Eechinogamma rus’’, se ANISOGAM-
MARUS, Gammaridae, 238

ECLYSIS, Paramphithoidae, 115, 160,
161, 392, 394

Egidia, to UROTHOE, Haustoriidae,
260

Eiscladus, to PHOTIS, Isaeidae, 274

ELASMOPOIDES, Gammaridae, 240

ELASMOPUS, Gammaridae, 28, 37,
42, 166, 217, 240

Elimedon, to 7HIPPOMEDON, Lysi-
anassidae [also assigned to PARA-
CENTROMEDON], 345, 355

ENDEVOURA, Lysianassidae, 342

ENSAYARA, Lysainassidae, 342

(EOGAMMARUS), subgenus of ANI-
SOGAMMARUS, Gammaridae,
237

EOHAUSTORIUS, Haustoriidae, 256

EOPHLIANTIS, Eophliantidae, 213,
433

527

Ephippiphora, to WALDECKIA, Ly-
sianassidae, 368

Epidesura, to ATYLUS, Atylidae, 163

EPIMERIA, Paramphithoidae, 394

EPIMERIELLA, Paramphithoidae,
115, 160, 161, 392, 395

Epimeriopsis, void ab initio, see BATH -
YPANOPLOEA, Acanthonotozo-
matidae, 122

ERICTHONIUS, Corophiidae, 39, 192,
275

EKriopis, to ERIOPISA, Gammaridae,
240

ERIOPISA, Gammaridae, 240

ERIOPISELLA, Gammaridae, 241

EUANDANIA, Stegocephalidae, 441

EUCALLISOMA, Lysianassidae, 342

EUON YX, Lysianassidae, 342

EURYMERA, Eusiridae, 223

Euryporeia, to KHURYTHENES, Lysi-
anassidae, 343

Eurystheus, to GAMMAROPSIS, Is-
aeidae, 271

Eurytenes, to EURYTHENES, Lysi-
anassidae, 343

EURYTHENES, Lysianassidae, 343

EUSIRELLA, Eusiridae, 90, 114, 225,
292

EUSIROGENES, Eusiridae, 225, 233

EUSIROIDES, Eusiridae, 173, 225

EUSIROPSIS, Eusiridae, 114, 225, 292

EUSIRUS, Eusiridae, 173, 226

Euxinia, to PONTOGAMMARUS
(freshwater), Gammaridae, 248

EXAMPITHOE, Ampithoidae, 144

Exhyalella, to PARH YALELLA, Hya-
lellidae, see Talitroidea, 471

EXOEDICEROPSIS, Oe¢edicerotidae,
382

EXOEDICEROS, Oedicerotidae, 382

Exungia, to COLOMASTIX, Colo-
mastigidae, 184

FALKLANDELLA, Gammaridae, 97,
116, 170, 241

FIGORELLA, Lysianassidae, 343

Fimbriella, to GAMMAROPSIS, Is-
aeidae, 271

GAINELLA, Lysianassidae, 343

Galanthis, to HYALE, Hyalidae, see
Talitroidea, 469

Gallea, to CYPROIDEA, Amphilochi-
dae, 139

528

GAMMARACANTHUS, Gammaridae,
241

GAMMARELLA, Gammaridae, 241

GAMMARELLUS, Gammaridae, 242

GAMMAROPSIS, Isaeidae, 28, 38, 271

GAMMARUS, Gammaridae, 242

GAROSYRRHOE, Synopiidae, 460

GAVIOTA, Corophiidae, 192

GITNAN, Amphilochidae, 138

GITANOGEITON, Amphilochidae, 138

GITANOPSIS, Amphilochidae, 138

Glauconeme, to UNCIOLA, Corophii-
dae, 197

Glycera, to GLYCERINA, Lysianassi-
dae, 345

GLYCERINA, Lysianassidae, 345

GNATHIPHIMEDIA, Acanthonoto-

_ zomatidae, 124
GOESIA, Isaeidae, 272
Gossea, to APHERUSA, Calliopiidae,

174
Gracilipes, to RHACHOTROPIS,
EKusiridae, 229
GRANDIDIERELLA, — Corophiidae,
147, 155, 192

Grayia, to GAMMARELLUS, Gam-
maridae, 142

Grimaldia, to SEBA, Sebidae, 436

Grubia, to CYMADUSA, Ampithoidae,
144

Guerina, to TRISCHIZOSTOMA,
Lysianassidae, 365

Guerinia, to TRISCHIZOSTOMA,
Lysianassidae, 365

GUERNEA, Dexaminidae, 203

GULBARENTSIA, Oedicerotidae, 382

HALICE, Pardaliscidae, 401

HALICELLA, Pardaliscidae, 114, 402

HALICOIDES, Pardaliscidae, 402, 453

HALICREION, Oedicerotidae, 383

Halimedon, to WESTWOODILLA,
Oedicerotidae, 387

HALIRAGES, Calliopiidae, 108, 177,
393

HALIRAGOIDES, Calliopiidae, 177

HANSENELLA, Aoridae, 153

HAPLOCHEIRA, Isaeidae, 272

HAPLOOPS, Ampeliscidae, 132

HARCLEDO, Eusiridae, 226

Harmomia, Harmonia, to PARA-
PHERUSA, Gammaridae, 247

Harpina, to HARPINIA, Phoxo-
cephalidae, 415 —

U.S. NATIONAL MUSEUM BULLETIN

271

HARPINIA, Phoxocephalidae, 415

HARPINIOIDELLA, Eusiridae, 167,
226
HARPINIOIDES, Calliopiidae, 177,

218, 219, 421, 422
HARPINIOPSIS, Phoxocephalidae, 415
Haustoriella, to PHOXOCEPHALOP-

SIS, Haustoriidae, 257
HAUSTORIOIDES, Dogielinotidae, 3,

209
HAUSTORIOPSIS,

251, 434
HAUSTORIUS, Haustoriidae, 256
Hela, to NEOHELA, Corophiidae, 194
Helella, to NEOHELA, Corophiidae,

Prophliantidae,

194

Helleria, to GUERNEA, Dexaminidae,
203

Hemijassa, to JASSA, Ischyroceridae,
279

HETEROCRESSA, Pagetinidae, 389

HETEROPHILIAS, Phliantidae, 410

HETEROPHOXUS, Phoxocephalidae,
417

HIPPOMEDON, Lysianassidae, 345

HIRONDELLEA. Lysianassidae, 345

Hoplonyx, to TMETONYX, Lysian-
assidae, 365

HOPLOPHEONOIDES,
dae, 139

HOPLOPLEON, Amphilochidae,

HORNELLIA, Gammaridae, 242

HYALE, Hyalidae, see Talitroidea,
28, 42, 207, 469

HYALELLA, Hyalelliade,
troidea, 42, 469

Hyaloides, to PARH YALE, Hyalidae,
see Talitroidea, 471

HYPERIOPSIS, Hyperiopsidae,

ICHNOPUS, Lysianassidae, 346

Ichthyomyzocus, to 7LAFYSTIUS, La-
fystiidae, 283

ICILIUS, Podoceridae, 427

Icridium, to PEREIONOTUS, Phli-
antidae, 411

Iduna, to LILJEBORGIA, Liljebor-
giidae, 294

IDUNELLA, Liljeborgiidae, 293

ILERA USTROE, Synopiidae, 460

Incisocalliope, to PARAPLEUSTES,
Pleustidae, 425

INSULA, Hyalid ae, see Talitroidea, 470

Amphilochi-

140

see Tali-

263

MARINE GAMMARIDEAN AMPHIPODA

Iphigenia, to IPHINOTUS, Phliantidae,
410

IPHIMEDIA, Acanthonotozomatidae,
124

IPHIMEDIELLA, Acanthonotozoma-
tidae, 124

Iphimediopsis Della Valle, to PANOP-
LOEA, Acanthonotozomatidae, 126

Imphimediopsis Schellenberg, to BATH-
YPANOPLOEA, Acanthomotozo-
matidae, 122

IPHINOTUS, Phliantidae, 410

IPHIPLATHIA, Phliantidae, 410

ISAEA, Isaeidae, 38, 272

ISAEOPSIS, Ischyroceridae, 278

ISCH Y ROCERUS, Ischyroceridae, 279

Janassa, to PARAJASSA, Ischyroceri-
dae, 279

JASSA, Ischyroceridae, 6, 116, 141, 279

JEDDO, Synopiidae, 113, 159, 461

JOUBINELLA, Phoxocephalidae, 171,
418

KAMAKA, Corophiidae, 106, 194

Katius, to KURYTHENES, Lysianas-
sidae, 343

KERGUELENIA, Lysianassidae, 346

KERM YSTHEUS, Isaeidae, 272

KOROGA, Lysianassidae, 346

Kroyera, to MONOCULODES, Oedi-
cerotidae, 383

KUPHOCHEIRA, Isaeidae, 273

KURIA, Kuriidae, 281, 433

KYSKA, Lysianassidae, 346

LABRIPHIMEDIA, Acanthonotozom-
atidae, 125

LAETMATOPHILUS, Podoceridae, 430

LAF YSTIUS, Lafystiidae, 283

Lakota, to ANONYX, Lysianassidae,
324

Lalaria, to AORA, Aoridae, 152

Lampra, to TRITAETA, Dexaminidae,

206
LAOTHOES, Calliopiidae, 177
LAPHYSTIOPSIS, Laphystiopsidae,
86, 115, 285

LEIPSUROPUS, Podoceridae, 430
LEMBOIDES, Aoridae, 153
LEMBOPSIS, Aoridae, 153

LEMBOS, Aoridae, 28, 154

LEPECHINELLA, Lepechinellidae,
163, 286

Lepidactylis, to ?HAUSTORIUS,

Haustoriidae, 256

029

LEPIDEPECREELLA, Lysianassidae,
347

LEPIDEPECREOIDES,
dae, 347

LEPIDEPECREOPSIS, Lysianassidae,
347

LEPIDEPECREUM, Lysianassiade,
348

LEPTAMPHOPUS, Calliopiidae, 178

LEPTOCHEIRUS, Aoridae, 154, 267,
269

Leptochela, to EHUON YX, Lysianassi-
dae, 342

LEPTOPHOXOIDES, Phoxocephali-
dae, 418

LEPTOPHOXUS,
419

Leptothoe, to MAERA, Gammaridae,
243

LEUCOTHOE, Leucothoidae, 28, 289

LEUCOTHOELLA, Leucothoidae, 290

LEUCOTHOIDES, Leucothoidae, 290

LIAGOCERADOCUS, Gammaridae,
217, 243

LILJEBORGIA, Liljeborgiidae, 294

Lilljeborgiella, to LILJEBORGIA
Liljeborgiidae, 294

Linguimaera, to MAERA, Gammaridae,
243

LIOUVILLEA, Eusiridae, 226

LISTRIELLA, Liljeborgiidae, 29, 36,
294

Lonchomerus, to AORA, Aoridae, 152

(LOPICEROS), subgenus of OEDI-
CEROIDES, Oedicerotidae, 384

Lusyta, to JASSA, Ischyroceridae, 279

Lycesta, to LEUCOTHOE, Leucothoi-
dae, 289

LYSIANASSA, Lysianassidae, 348

Lysianassina, to LYSIANASSA, Lysi-
anassidae, 348

Lysianax, to LYSIANASSA, Lysianas-
sidae, 348

LYSIANELLA, Lysianassidae, 349

Lysianopsis, to LYSIANASSA, Lysi-
anassidae, 348

Macleayia, to JASSA, Ischyroceridae,
279

MACROPISTHOPUS, Ampithoidae,
144

MAERA, Gammaridae, 28, 42, 243

MAERACUNHA, Gammaridae, 243

MAERELLA, Gammaridae, 243

Lysianassi-

Phoxocephalidae,

530

Maeroides, to GAMMAROPSIS, Isaei-

dae, 271

MAEROPSIS, Gammaridae, 244

MANDIBULOPHOXUS, Phoxocepha-

lidae, 419

MAORIPHIMEDIA, Acanthonotozo-
matidae, 125

‘““Marine Carinate, Gammarus,” Gam-
maridae, 244

(MARINOGAMMARUS), Gammari-
dae, see GAMMARUS, 242

MAXILLIPHIMEDIA, Acanthonoto-
zomatidae, 125, 451

Megalonoura, to MEGALUROPUS,
Gammaridae, 244

Megalorchestia, to ORCHESTOIDEA,

Talitridae, see Talitroidea, 471

MEGALUROPUS, Gammaridae,

94, 107, 157, 158, 244, 369

Megamoera, to MELITA, Gammaridae,

245

MEGAMPHOPUS, Isaeidae, 273

MELITA, Gammaridae, 245

MELITOIDES, Gammaridae, 245

MELPHIDIPPA, Melphidippidae, 371

MELPHIDIPPELLA, Melphidippidae,

371

MELPHISANA, Melphidippidae, 172,
371

MENIGRATES, Lysianassidae, 349

MENIGRATOPSIS, Lysianassidae, 349

MESOCYCLOCARIS, Lysianassidae,
349

MESOCYPHOCARIS, Lysianassidae,
350

MESOGAMMARUS, Gammaridae
245

MESOMETOPA, Stenothoidae, 448

MESOPLEUSTES, Pleustidae, 88, 167,

30,

392, 422

MESOPROBOLOIDES, Stenothoidae,
448

Mesostenothoides, to STENOTHOI-

DES, Stenothoidae, 451

METACERADOCOIDES, Gammari-

dae, 245

METACERADOCUS, Gammaridae
246

METACYCLOCARIS, Lysianassidae,

350

METACYPHOCARIS, Lysianassidae,
350

U.S. NATIONAL MUSEUM

BULLETIN 271

METALEPTAMPHOPUS,
piidae, 108, 178, 369
METAMBASIA, Lysianassidae, 350
Metandania, to ANDANIOTES, Ste-
gocephalidae, 441
METAPHOXUS, Phoxocephalidae, 419
METEPIMERIA, Paramphithoidae

Callio-

396

METEUSIROIDES, Eusiridae, 167,
227

METHALIMEDON, Oedicerotidae
383

Metharpinia, to PARAPHOXUS,
Phoxocephalidae, 420
METOEDICEROS, [Oedicerotidae], see
Incertae Sedis, 37, 113, 207, 376,
377, 480
METOPA, Stenothoidae, 448
METOPELLA, Stenothoidae, 449
METOPELLOIDHES, Stenothoidae, 449
Metopina, to METOPA, Stenothoidae,
448
METOPOIDHES, ’Stenothoidae, 449
Microcheles, to IPHIMEDIA, Acantho-
notozomatidae, 124
MICRODEUTOPUS, Aoridae, 154
MICROJASSA, Ischyroceridae, 279
MICROLYSIAS, Lysianassidae, 351
MICROPHOTIS, Isaeidae, 273
MICROPHOXUS, Phoxocephalidae,
419
Microplax, to LILJEBORGIA, Lilje-
borgiidae, 294
MICROPROTOPUS, Isaeidae, 110, 274
MICROSTENOTHOEH, Stenothoidae,
449
MONOCULODES, Oedicerotidae, 383
MONOCULOPSIS, Oedicerotidae, 383
Montagua, to STENOTHOEH, Steno-
thoidae, 450
Montaguana, to STENOTHOE, Steno-
thoidae, 450
Mulleria, to MAERA, Gammaridae,
243
Naenia, to PODOCEROPSIS, Isaeidae,
274
NAJNA, see Talitroidea (provisional),
37, 59, 113, 209, 406, 470
NANNON YX, Lysianassidae, 351
NECOCHEA, Pardaliscidae, 402
NEOAMBASIA, Lysianassidae; 351
NEOBULE, Talitroidea (provisional),
470

MARINE GAMMARIDEAN AMPHIPODA

NEOCYPROIDEA, Amphilochidae,
140
Neogammaropsis, to HLASMOPUS,

Gammaridae, 240
NEOHAUSTORIUS, Haustoriidae, 257
NEOHELA, Corophiidae, 194
NEOMEGAMPHOPUS, Aoridae, 155,

267, 269
NEOMICRODEUTOPUS, Aoridae,
155, 184, 194
Neophotis, to CHEVALIA, Isaeidae,

271
NEOPLEUSTES, Pleustidae, 424
NEOXENODICEH, Podoceridae, 431
NETAMELITA, Gammaridae, 246
Nicea, to HYALE, Hyalidae, see Tali-
troidea, 469
NICIPPE, Synopiidae, 403
NIPPOCHELURA, Cheluridae, 181
Normania, to NORMANION, Lysi-
anassidae, 351
NORMANION, Lysianassidae, 351
Nototropis, to ATYLUS, Atylidae, 163
OCHLESIS, Ochlesidae, 373
OCOSIN GO, Lysianassidae, 352
ODIUS, Acanthonotozomatidae, 125
OEDICERINA, Oedicerotidae, 384
OEDICEROIDES, Oedicerotidae, 384
OEDICEROPSIS, Oedicerotidae, 384
Oediceropsoides, to OEDICEROIDES,
Oedicerotidae, 384
OEDICEROS, Oedicerotidae, 385
ONESIMOIDES, Lysianassidae, 352
ONISIMUS, Lysianassidae, 352
“ONISIMUS” auct., to BOECKO-
SIMUS, Lysianassidae, 330
Opis, to OPISA, Lysianassidae, 353
OPISA, Lysianassidae, 353
ORADAREA, Calliopiidae,
393
ORCHESTIA,
Talitroidae, 470
ORCHESTOIDEA, Talitridae,
Talitroidae, 12, 471
ORCHOMENT, Lysianassidae, 40, 353
Orchomenella, to ORCHOMENE.
Lysianassidae, 365
“Orchomenella”’ greenlandica, Lysian-
assidae, see keys, 306, 311, 313
Orchomenopsis, to ORCHOMENE,
Lysianassidae, 353
Orthopalame, to MICROPROTOPUS,
Isaeidae, 274

178, 392,
Talitridae, see

see

531
Otus, to ODIUS, Acanthonotozo-
matidae, 125
PACH YCHELIUM, Lysianassidae, 354
PACH YNUS, Lysianassidae, 354
PAGETINA, Pagetinidae, 389
PALINNOTUS, Phliantidae, 411
PANOPLOEA, Acanthonotozomatidae,
126
PANOPLOEOPSIS,
matidae, 126
PARACALLIOPE (freshwater), Cal-
liopiidae, see supplement, 172, 481
PARACALLISOMA, Lysianassidae, 89,
111, 354
PARACALLISOMOPSIS,
sidae, 355
[PARACENT ROMDEON],
sidae [also assigned to
POMEDON|], 345, 355
PARACERADOCUS, Gammaridae,
246
PARACOROPHIUM,
194, 481
Paracyclocaris, to LEPIDEPECRE-
ELLA, Lysianassidae, 347
PARACYPHOCARIS, Lysianassidae,
355
PARACYPROIDEA, Amphilochidae,
140
PARADEXAMINE, Dexaminidae, 206
PARADRYOPEH, Aoridae, 155, 275
PARADULICHIA, Podoceridae, 431
PARAGRUBIA, Ampithoidae, 144
PARAHALICE, Pardaliscidae, 114, 403
PARAHAUSTORIUS, Haustoriidae,
257
PARAJASSA, Ischyroceridae, 279
PARALEPECHINELLA, Lepechinel-

Acanthonotozo-

Lysianas-

Lysianas-
?HIP-

Corophiidae,

lidae, 288
PARALEPTAMPHOPUS, (fresh-
water), Calliopiidae, 172
PARALEUCOTHOE, Leucothoidae,
291

PARALIBROTUS, Lysianassidae, 356
PARALICELLA, Lysianassidae, 356
PARALLORCHESTES, Hyalidae, see
Talitroidae, 471
PARALYSIANOPSIS,
356
PARAMBASIA, Lysianassidae, 356
PARAMETOPA, Stenothoidae, 449
PARAMETOPELLA, Stenothoidae,
450

Lysianassidae,

032

PARAMOERA, Eusiridae, 227
PARAMPHITHOHE, Paramphithoidae,

119, 176, 396

Paranaenia, to GAMMAROPSIS,
Isaeidae, 271

PARANAMIXIS, Anamixidae, 146,
164

PARANCHIPHIMEDIA, Acantho-

notozomatidae, 126
PARANDANIA, Stegocephalidae, 442
Parandaniexis Nicholls, to PPEUDAN -

DANIEXIS, Stegocephalidae, 443
PARANDANIEXIS Schellenberg,

Stegocephalidae, 442
PARAOROIDES, Aoridae, 155
PARAPANOPLOEA, Acanthonotozo-

matidae, 127
PARAPERIOCULODES,

tidae, 385
PARAPHERUSA, Gammaridae, 107,

116, 170, 247, 275
PARAPHOXUS, Phoxocephalidae, 420
PARAPLEUSTES, Pleustidae, 425
PARARGISSA, Hyperiopsidae, 32, 112,

158, 264
Pararistias, to PERRIERELLA, Ly-

sianassidae, 358
Parastyra, to ASTYRA, Astyridae, 161
Parathaumatelson, to PSE UDOTHA U-

MATELSON, Thaumatelsonidae,

475
Paratryphosites, to HIPPOMEDON,

Lysianassidae, 345
Paratylus, to ATYLUS, Atylidae, 163
Paravalettia, to SEBA, Sebidae, 436
PARAWALDECKIA, _Lysianassidae,

357
“Parawaldeckia kidderi,’’ see PARA-

WALDECKIA, Lysianassidae, 357
PARDALISCA, Pardaliscidae, 403
PARDALISCELLA, Pardaliscidae, 251,

403
PARDALISCOIDES, Pardaliscidae,404
PARDALISCOPSIS, Pardaliscidae, 404
PARDISYNOPIA, Pardaliscidae, 404
PARELASMOPUS, Gammaridae, 247,

Oedicero-

291
PAREPIMERIA, Paramphithoidae,
396
PAREPIMERIELLA, Pleustidae, 392,
393, 425

PAREUSIROGENKHES, Eusiridae, 227
PARHALIMEDON, Oedicerotidae, 385

U.S. NATIONAL MUSEUM BULLETIN 271

Parharpinia, to PARAPHOX US,
Phoxocephalidae, 420

PARH YALH, Hyalidae, seeTalitroidea,
471

PARH YALELLA,
Talitroidea, 471

PARIPHIMEDIA, Acanthonotozoma-
tidae, 127

PARIPHIMEDIELLA, Acanthonoto-
zomatidae, 127

PARIPHINOTUS, Phliantidae, 411

PAROEDICEROIDES, Oedicerotidae,

Hyalellidae, see

385
PAROEDICEROS, Oedicerotidae, 386
PARONESIMOIDES, Lysianassidae,
357

PARONESIMUS, Lysianassidae, 357

Paropisa, to SOPHROSYNE, Lysian-
assidae, 363

Parorchestia, to ORCHESTIA, Tali-
tridae, see Talitroidea, 470

PARPANO, Pardaliscidae, 404

PARUNCIOLA, Corophiidae, 196, 267

(PATOIDES), subgenus, to ACE RO-
IDES, Oedicerotidae, 378

(PECTENOGAMMARUS), subgenus
of GAMMARUS, Gammaridae,
242

PELTOCOXA, Amphilochidae, 140

PELTOPES, Amphilochidae, 141

PEREIONOTUS, Phliantidae, 411

PERIOCULODES, Oedicerotidae, 386

PERIOCULOPSIS, Oedicerotidae, 386

Periphlias, to CEINA, Phliantidae, 410

PERRIERELLA, Lysianassidae, 358

Phaedra, removed from APHERUSA,
Calliopiedae, to dubious status, 174

Pherusa, to GAMMARELLA, Gam-
maridae, 241

Pherusana, to GAMMARELLA, Gam-
maridae, 241

PHIPPSIA, Stegocephalidae, 442

PHIPPSIELLA, Stegocephalidae, 442

PHLIAS, Phliantidae, 61, 411

PHOT IS, Isaeidae, 8, 274

PHOXOCEPHALOPSIS, Haustoriidae,
108, 257

PHOXOCEPHALUS, Phoxocephalidae,
420

PHOXOSTOMA, Lysianassidae, 358

Phoxus, to PHOXOCEPHALUS,
Phococephalidae, 420

MARINE GAMMARIDEAN AMPHIPODA

Phylluropus, to MEGALUROPUS,
Gammaridae, 158, 244

Platamon, to HIPPOMEDON, Lysian-
assidae, 345

Platophium, to PODOCERUS, Podo-
ceridae, 431

PLAT YISCHNOPUS,
258, 414, 436

PLEONEXHES, Ampithoidae, 145

PLEUSTES, Pleustidae, 425

PLEUSYMTES, Pleustidae, 425

PLIOPLATEIA, Phliantidae, 412

PODOCEROPSIS, Isaeidae, 274

PODOCERUS, Podoceridae, 22, 28, 39,
409, 431

PODOPRION, Lysianassidae, 358

PODOPRIONELLA, Lysianassidae,
358

PODOPRIONIDES, Lysianassidae,
359

POLYCHEIRA, Dexaminidae, 26, 30,
206

Pontharpinia, to PARAPHOXUS,
Phoxocephalidae, 420

PONTOCRATES, Oedicerotidae, 386

PONTOGENEIA, Eusiridae, 172, 228

PONTOGENEIELLA, Eusiridae, 228

PONTOGENEOIDES, Eusiridae, 167,
173, 228

PONTOPOREIA, Haustoriidae,
258

PRACHYNELLA, Lysianassidae, 359

Prinassus, to GUERNEA, Dex-
aminidae, 203

PRINCAXELIA, Pardaliscidae, 405

Priscilla, to PRISCILLINA, Haustori-
idae, 258

PRISCILLINA, Haustoriidae, 258

Proboliella, to PROBOLOIDES, Sten-
othoidae, 450

PROBOLISCA, Stenothoidae, 450

Probolium, to STENOTHOE,
othoidae, 450

PROBOLOIDES, Stenothoidae, 450

PROCYPHOCARIS, Lysianassidae,
359

PROHARPINIA, Phoxocephalidae, 420

PROLAPHYSTIOPSIS, Laphystiopsi-
dae, 115, 283, 285

PROLAPHYSTIUS, Laphystiopsidae,
86, 170, 174, 285

PROMETOPA, Stenothoidae, 450

Haustoriidae,

108,

Sten-

033

[PRONANNON YX], Lysianassidae
[also assigned to 7LYSIANASSA],
348, 359

PROPHLIAS, Prophliantidae, 434

PROSTEBBINGIA, Eusiridae, 228

PROSTENOTHOE, Stenothoidae, 450

PROTHAUMATELSON, Thaumatel-
sonidae, 475

PROTOHAUSTORIUS, Haustoriidae,
259

Protohyperiopsis, to PARARGISSA,
Hyperiopsidae, 264

PROTOMEDEHIA, Isaeidae, 274

Protophoxus, to PARAPHOXUS,
Phoxocephalidae, 420

Pseudalibrotus, to ONISIMUS, Lysi-
anassidae, 352

PSEUDAMBASIA, Lysianassidae, 360

PSEUDAMPHILOCHUS, Amphilo-
chidae, 36, 37, 115, 139, 217, 292,
421

PSEUDANDANIEXIS, Stegocephali-
dae, 443

PSEUDEPIMERIA, Paramphithoidae,
396

PSEUDERICTHONIUS, Corophiidae,
196

(PSEUDEURYSTHEUS), subgenus of
GAMMAROPSIS, Isaeidae, 271

PSEUDHARPINIA, Phoxocephalidae,
420

PSEUDIPHIMEDIELLA, Acanthono-
tozomatidae, 127

Pseudiphimediopsis, to BATH YPAN-
OPLOEA, Acanthonotozomatidae,
122

PSEHUDISCH YROCERUS, Ischyro-
ceridae, 280

PSEUDOANON YX, Lysianassidae,
360

Pseudoceradocus, to QUADRIVISIO,
Gammaridae, 247

PSEUDOHAUSTORIUS, Haustori-
idae, 259

PSEUDOKOROGA, Lysianassidae, 360

PSEUDOMOERA, Eusiridae, 229

Pseudonesimus, to SCHISTURELLA,
Lysianassidae, 361

Pseudophthalmus, to
Ampeliscidae, 130

PSEUDOPONTOGENEIA, Eusiridae,
172, 229

AMPELISCA,

534

PSEUDORCHOMENEH, Lysianassidae,
360

PSEUDOTHAUMATELSON, Thau-
matelsonidae, 475

PSEUDOTIRON, Synepiidae, 109, 397,
461

Pseudotryphosa, to ?7?URISTES, Lys-
ianassidae, 367

Pterygocerus, to
Haustoriidae, 256

Ptilocheirus, to LEPTOCHEIRUS,
Aoridae, 154

Pyctilus, to ERICTHONIUS, Coro-
phiidae, 192

QUADRIVISIO, Gammaridae, 247

QUASIMODIA, Philantidae, 412

REGALIA, Calliopiidae, 178, 283

RHACHOTROPIS, Eusiridae, 33, 173,
229, 392

RIFCUS, Lysianassidae, 361

RONCO, Eusiridae, 229

ROZINANTE, Etsiridae, 230

RUDILEMBOIDES, Aoridae, 156

RUNANGA, Corophiidae, 196, 426

SANCHO, Calliopiidae, 25, 26, 39,
108, 179, 215, 408, 409

Scamballa, to ORCHESTIA, Talitridae,
see Talitroidae, 470

SCHISTURELLA, Lysianassidae, 361

SCHRADERIA, Eusiridae, 230

SCOPELOCHEIROPSIS, Lysianas-

HAUSTORIUS,

sidae, 361
SCOPELOCHEIRUS, Lysianassidae,
362
SEBA, Sebidae, 436
(SEGAMPHOPUS), subgenus of

MEGAMPHOPUS, Isaeidae, 273
Sextonia, to IDUNELLA, Liljeborgi-
idae, 293
SHACKLETONIA, Lysianassidae, 362
Shoemakerella, to LYSIANASSA, Lysi-
anassidae, 348
SIPHONOECETES, Corophiidae, 197
SOCARNESLLA, Lysianassidae, 362
SOCARNEI, Lysianassidae, 362
SOCARNODES, Lysianassidae, 363
SOCARNOPSIS, Lysianassidae, 363
SOPH ROSHNE, Lysianassidae, 363
SPHAEROPHTHALMUS, © see
certae Sedis, 202, 480
Spinifer, to PHOXOCEPHALUS,
Phoxocephalidae, 420

In-

U.S. NATIONAL MUSEUM BULLETIN

271

Stebbingia, to
siridae, 227
STEGOCEPHALINA, Stegocephalidae,
443
STEGOCEPHALOIDES, Stegocepha-
lidae, 443
STEGOCEPHALOPSIS,
lidae, 443
STEGOCEPHALUS, Stegocephalidae,
444
STEGOPLAX, Amphilochidae, 141
STELEUTHERA, Stegocephalidae, 444
Stenia, dubious genus, Lysianassidae
(not discussed, see Stebbing, 1906)
STENOPLEURA, Calliopiidae, 179
STENOPLEUROIDES, Calliopiidae,
179
STENOPLEUSTHS, Pleustidae, 425
STENOTHOE, Stenothoidae, 28, 450
STENOTHOIDES, Stenothoidae, 451
STEN ULA, Stenothoidae, 451
STEPHENSENTA, Lysianassidae, 364
STEPHOBRUZELIA, Synopiidae, 461
Sthenometopa, to METOPA, Steno-
thoidae, 448
STILIPES, Stilipedidae, 161, 454
Stimpsonella, to MICRODEUTOPUS,
Aoridae, 154
Stimpsonia, to
Aoridae, 154
STOMACONTION, Lysianassidae, 364
Suleator, to HAUSTORIUS, Haus-
toriidae, 257
SUNAMPHITOE, Ampithoidae,
145
Sympleustes, to STENOPLEUSTHES,
Pleustidae, 425
SYNCHELIDIUM, Oedicerotidae, 387
SYNDEXAMINE, Dexaminidae, 206
SYNOPIA, Synopiidae, 34, 107, 115,

PARAMOERA, Eu-

Stegocepha-

MICRODEUTOPUS,

113,

116, 462
Synopioides, to HALICEH, Pardalisci-
dae, 401

SYRRHOEH, Synopiidae, 462

SYRRHOITES, Synopiidae, 462

Talitriator, to TALITRUS, Talitridae,
see Talitroidea, 472

Talitroides, to TALITRUS, Talitridae,
see Talitroidea, 472

Talitronus, to ORCHESTOIDEA, Tal-
itridae, see Talitroidea, 471

MARINE GAMMARIDEAN AMPHIPODA

(Talitrorchestia), subgenus, to ?TALI-
TRUS, Talitridae, see Talitroidea,
472

TALITRUS Talitridae, see Talitroidea,
472

TALORCHESTIA, Talitridae, see Tal-
itroidea, 472

TEMNOPHLIAS, Phliantidae, 21, 22,
26, 412

Teraticum, to SEBA, Sebidae, 436

Tessarops, to TIRON, Synopiidae, 463

TETRADEION, Stegocefhalidae, 444

Tetromatus, to AMPELISCA, Ampel-
iscidae, 130

Tetronychia, to HIRONDELLEA,
Lysianassidae, 345

THAUMATELSON, Thaumatelsoni-
dae, 476

Thersites, to BATHYPOREIA, Haus-
toriidae, 254

Thoelaos, to LAOTHOES, Calliopiidae,
177

THORIELLA, Lysianassidae, 364, 373

TIRON, Synopiidae, 463

TMETON YX, Lysianassidae, 365

“Tmetonyx’’ auct., to TRYPHOSEL-
LA, Lysianassidae, 365

Trichophoxus, to PARAPHOXUS,
Phoxocephalidae, 420

TRIODOS, Ampeliscidae, 132

TRISCHIZOSTOMA, Lysianassidae,
365

TRITAETA, Dexaminidae, 206

Tritropis, to RHACHOTROPIS, Eusi-
ridae, 229

TROPICHELURA, Cheluridae, 182

Tryphosa, to ORCHOMENEH, Lysian-
assidae, 353, 365

O30

“Tryphosa”’ auct., to TR YPHOSELLA,
Lysianassidae, 365
TRYPHOSELLA, Lysianassidae, 365
TRYPHOSITES, Lysianassidae, 366
TRYPHOSOIDES, Lysianassidae, 367
UNCINOTARSUS, Aoridae, 156, 190,
427
UNCIOLA, Corophiidae, 197
UNCIOLELLA, Corophiidae, 197
URISTES, Lysianassidae, 367

Uristoides, to URISTES, Lysianas-
sidae, 367

UROHAUSTORIUS, Haustoriidae,
259

UROTHOE, Haustoriidae, 108, 260
UROTHOIDES, Haustoriidae, 108, 260
USCHAKOVIELLA, Paramphithoidae,
397
VALETTIA, Lysianassidae, 367
VALETTIOPSIS, Lysianassidae,
VEMANA, Vitjazianidae, 478
Vertumnus, to 7EPIMERIA, Paramph-
ithoidae, 394
VIJAYA, Lysianassidae, 368
VITJAZIANA, Vitjazianidae, 478
WALDECKIA, Lysianassidae, 368
WANDELIA, Eophliantidae, 213, 433
Westwoodia, to WESTWOODILLA,
Oedicerotidae, 387
WESTWOODILLA, Oedicerotidae, 387
WEYPRECHTIA, Gammaridae, 170,
247, 392, 393
Wyvillea, to JASSA, Ischyroceridae, 279
XENOCHEIRA, Aoridae, 156
Xenoclea, to PODOCEROPSIS, Isaei-
dae, 274
XENODICH, Podoceridae, 431
ZARAMILLA, Eusiridae, 230
ZOBRACHO, Haustoriidae, 260

368

U. S. GOVERNMENT PRINTING OFFICE : 1969 O - 285-135

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