The Families and Genera

of Hyperiidea

(Crustacea: Amphipoda)

THOMAS E. BOWMAN

and

HANS-ECKHARD GRUNER

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 146
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 14 6

The Families and Genera of Hyperiidea

(Crustacea: Amphipoda)

Thomas E. Bowman

and Hans-Eckhard Gruner

JAN 8 1974

SMITHSONIAN INSTITUTION PRESS

City of Washington

1973
ABSTRACT

Bowman, Thomas E., and Hans-Eckhard Gruner. The Families and Genera of

Hyperiidea (Crustacea: Amphipoda). Smithsonian Contributions to Zoology,

number 146, 64 pages, 82 figures, 1973.—A synopsis of the amphipod suborder

Hyperiidea is presented, including diagnoses of its 2 infraorders, 6 superfamilies,

21 families, and 71 genera. Diagnostic keys are given for the families and genera,

and illustrations of significant characters are given for each genus. Several changes

are made in the classification of the Hyperiidea. Tribes are changed to infraorders

and subtribes to superfamilies, with the necessary nomenclatural changes being

made. A new superfamily, Lycaeopsoidea, for the family Lycaeopsidae, and a new

family, Anapronoidae, for Anapronoe, are proposed. A new genus, Spinoscina,

is proposed for Acanthoscina spinosa; and Vibilioides and Parascelus are reduced

to synonyms of Vibilia and Thyropus, respectively.

OFFICIAL PUBLICATION DATE is handstampcd in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 4776. SERIES COVER DESICN:

The coral Montastrea cax>ernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Bowman, Thomas E.

The families and genera of Hyperiidea.

(Smithsonian contributions to zoology, no. 146)

1. Amphipoda—Classification. I. Gruner, Hans-Eckhard, joint author. II. Title. III. Series:

Smithsonian Institution. Smithsonian contributions to zoology, no. 146.

QL1.S34 no. 146 [QL444.A5] 591'.08s [595'.371] 73-1579

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 - Price SI.35
Contents

Page

Introduction 1

Morphology 2

Natural History 6

Classification 7

Suborder Hyperiidea H. Milne Edwards, 1830 9

Infraorder Physosomata Pirlot, 1929 9

Superfamily Scinoidea, new name 10

Family Archaeoscinidae Stebbing, 1904 10

Archaeoscina Stebbing, 1904 10

Family Mimonectidae Bovallius, 1885 10

Mimonectes Bovallius, 1885 11

Pseudomimonectes Vinogradov, 1960 11

Family Proscinidae Pirlot, 1933 12

Proscina Stephensen and Pirlot, 1931 12

Mimoscina Pirlot, 1933 12

Family Scinidae Stebbing, 1888 12

Scina Prestandrea, 1833 14

Acanthoscina Vosseler, 1900 14

Ctenoscina Wagler, 1926 14

Spinoscina, new genus 16

Superfamily Lanceoloidea, new name 16

Family Microphasmidae Stephensen and Pirlot, 1931 16

Microphasma Woltereck, 1909 17

Microphasmoides Vinogradov, 1960 17

Mimonecteola Woltereck, 1909 18

Family Chuneolidae Woltereck, 1909 19

Chuneola Woltereck, 1909 19

Family Lanceolidae Bovallius, 1887 19

Lanceola Say, 1818 20

Megalanceola Pirlot, 1935 20

Metalanceola Pirlot, 1931 21

Paralanceola Barnard, 1930 22

Prolanceala Woltereck, 1907 22

Scypholanceola Woltereck, 1905 22

Infraorder Physocephalata, new name 23

Superfamily Vibilioidea, new name 24

Family Vibiliidae Dana, 1852 24

Vibilia H. Milne-Edwards, 1830 24

Cyllopus Dana, 1852 25

Family Cystisomatidae Willemoes-Suhm, 1875 25

Cystisoma Guerin-Mdneville, 1842 26

Family Paraphronimidae Bovallius, 1887 26

Paraphronima Claus, 1879 27

Superfamily Phronimoidca, new name 27

Family Hyperiidae Dana, 1852 27

Hyperia Latreille, in Desrnarest, 1823 28

Bougisia Laval, 1966 29

Hyperiella Bovallius, 1887 30

Hyperietta Bowman, 1973 30

Hyperioides Chevreux, 1900 31

Hyperionyx Bowman, 1973 32

iii
Page

Hyperoche Bovallius, 1887 32

Iulopis Bovallius, 1887 33

Lestrigonus Milne Edwards, 1830 33

Parathetnisto Boeck, 1870 34

Pegohyperia Barnard, 1932 34

Phronimopsis Claus, 1879 35

Themistella Bovallius, 1887 36

Family Dairellidae Bovallius, 1887 36

Dairella Bovallius, 1887 36

Family Phronimidae Dana, 1853 37

Phronima Latreille, 1802 37

Phronimella Claus, 1871 38

Family Phrosinidae Dana, 1853 38

Phrosina Risso, 1822 39

Anchylomera H. Milne-Edwards, 1830 39

Primno Guerin-Me'neville, 1836 40

Lycaeopsoidea, new superfamily 41

Family Lycaeopsidae Chevreux, 1913 41

Lycaeopsis Claus, 1879 41

Superfamily Platysceloidea, new name 41

Family Pronoidae Claus, 1879 42

Pronoe Guerin-Meneville, 1836 42

Eupronoe Claus, 1879 43

Paralycaea Claus, 1879 43

Parapronoe Claus, 1879 44

Sympronoe Stebbing, 1888 44

Anapronoidae, new family 45

A napronoe Stephensen, 1925 45

Family Lycaeidae Claus, 1879 46

Lycaea Dana, 1852 46

Brachyscelus Bate, 1861 47

Pseudolycaea Claus, 1879 47

Thamneus Bovallius, 1887 48

Tryphana Boeck, 1870 48

Family Oxycephalidae Bate, 1861 48

Oxycephalus H. Milne-Edwards, 1830 49

Calamorhynchus Streets, 1878 50

Cranoecephalus Bovallius, 1890 51

Glossocephalus Bovallius, 1887 51

Leptocotis Streets, 1877 52

Rhabdosoma White, 1847 52

Simorhyncfiotus Stebbing, 1888 53

Streetsia Stebbing. 1888 53

Tullbergella Bovallius, 1887 53

Family Platyscelidae Bate, 1862 54

Platyscehis Bate, 1862 55

Arnphithyrus Claus, 1879 55

Hemityphis Claus, 1879 55

Paratyphis Claus, 1879 55

Tetrathyrus Claus, 1879 56

Family Parascelidae Bovallius, 1887 57

Thyropus Dana, 1852 57

Euscelus Claus, 1879 57

Schizoscelus Claus, 1879 58

Sources of Illustrations 58

Literature Cited 60

IV
The Families and Genera of Hyperiidea

(Crustacea: Amphipoda)

Thomas E. Bowman

and Hans-Eckhard Gruner

The Hyperiidea are, therefore, those species

among the Amphipoda, in which nature indulges

in her widest diversities of development. . . .

—JAMES DWIGHT DANA

Introduction

The hyperiid amphipods are important marine

crustacean zooplankters, ranking third in abundance behind the Copepoda and Euphausiacea.

Contrary to the statement of Hurley (1969), identification of hyperiids often is a frustrating task, not

only because of the unsatisfactory state of our

knowledge of such genera as Eitpronoe and Lycaea

but especially because of the scattered condition of

the literature on hyperiid systematics. Much of the

essential literature is contained in works of the late

19th century and in reports of the major oceanographic expeditions, which often are not readily

available, especially at the smaller and newer

marine laboratories. No attempt has been made at

a summary of hyperiid systematics since the pioneer

monographs of Bovallius (1887a, 1887b, 1887c,

1889, 1890) and Claus (1879a, 1879b), although

many important works have since appeared. The

need for an up-to-date synopsis has been made more

Thomas E. Bowman, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Hans-Eckhard Gruner, Zoologisches Museum der Humboldt Universitdt zu Berlin,

Invalidenstrasse 43, 104 Berlin, Deutsche Demokratische

Republik.

urgent by the increased activity in plankton research in recent years.

A concurrent 2-month visit as consultants at the

Indian Ocean Biological Centre, Cochin, India,

afforded us the opportunity—free from the usual

interruptions at our respective museums—to work

closely together in preparing a large part of the

present synopsis. Our goal has been to prepare a

synopsis that will make it possible for a trained

biologist to identify any hyperiid amphipod to

genus without having to refer to other publications.

We have attempted to write clear and unequivocal

diagnoses and we include a section on morphology

so that the meanings of terms will be unmistakable.

All characters used in our diagnoses are illustrated.

We have used published illustrations of other authors freely, sometimes modifying them by eliminating unnecessary or confusing lines or by spreading

out overlapping appendages. When adequate illustrations were not available we prepared original

drawings.

Sources of the illustration are given in the appendix. The following abbreviations are used in

the illustrations:

II

A.I

A .2

head

antenna 1

antenna 2
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Md

Mx.l

Mx.2

Mxp

P.I to P.7

Up.l to Up.3

T

Us.l

Us.2 + 3

mandible

maxilla 1

maxilla 2

maxillipcd

pcrcopods 1-7

uropods 1-3

telson

urosomite 1

double urosomite

Our diagnoses are based in part on published

accounts, but we have examined numerous specimens in the collections of the Indian Ocean Biological Centre and our own museums in order to

verify, correct, or supplement these accounts. Synonymies are omitted, since they will be given fully in

a section on Hyperiidea in the Catalogns Cntstaceorum, now being prepared. For each family or

genus we list a few of the most useful taxonomic

works, but we have not attempted to provide an

extensive bibliography of the Hyperiidea.

This work is the result of equal effort by each

of us, and we should be considered coauthors rather

than junior and senior authors.

Morphology

Figure 1 shows a generalized hyperiid with the

principal parts labeled.

The body form is quite varied. In some hyperiid

amphipods, including the Physosomata and Cystisomatidae, the cuticle is thin and transparent and

the muscles are weakly developed. These hyperiids

often have rather globular bodies and are weak

swimmers. They are mainly bathypelagic. Others,

such as Pronoidae (Figures 5-1-59) and Lycaeidae

(Figures 60-64), have rather compact bodies with

strong musculature and can swim very rapidly. The

plump Platyscelidae and Parascelidae can undergo

conglobation (rolling into a ball) by bringing the

operculate bases of the left and right pereopods

5 and 6 together ventrally and fitting them against

the posteroventral margin of the head (Figure 74).

The telson and uropods are tucked under the body

and into a unnamed groove on the posterior margin of each pereopod 6 that we propose to call

the "telsonic groove." The function of this conglobation is unknown; possibly it aids these strong

swimmers to avoid predators by sinking.

The opposite extreme in body form from the conglobate families is found in the slender elongate

Oxycephalidae, culminating in the needle-shaped

Rhabdosoma (Figure 70).

The head is typically globular, with most of its

surface occupied by the compound eyes. The dorsal

midline is free from ommatidia and serves as a line

of attachment for a muscle that inserts on the dorsal

surface of the midgut. In the Physosomata the eyes

are small or absent and situated on the sides of the

rather short head. Among the Physocephalata the

eyes are small or medium-sized only in Vibilia and

Bougisia where they are located on the sides of

the cuboidal head. In the Oxycephalidae the anterior part of the head in front of the eyes is drawn

out into a short-to-long triangular rostrum.

The pereon comprises 7 segments (pereonites 1-7

= thoracic somites 2-8). Some of the anterior pereonites may be fused in certain genera. The coxae ( =

coxal plates), or the proximal segments of the

thoracic legs, may be fused with the pereonites or

separated from them by a suture; they are never

enlarged as in many Gammaridea.

The abdomen is formed of the pleon and the

urosome. The pleon contains 3 segments or pleonites. The urosome, which is composed of 3 segments

or urosomites in most gammarideans is 2-segmented

in hyperiids, urosomites 2 and 3 being fused to

form the double urosomite. The abdomen ends in a

thin triangular piece, the telson, which may be

fused with the double urosomite. The telson is

never divided or incised at its apex as it is in many

Gammaridea.

The 2 pairs of antenna, antenna 1 and antenna 2,

may be inserted on either the anterior or the ventral surface of the head. The site of insertion and

the structure of antennae 1 and 2 are the major

criteria by which the superfamilies of Physocephalata are distinguished (Figure 2). Each antenna consists of a basal part, the peduncle, followed by a

flagellum. In the Gammaridea the peduncle is 3-

segmented in antenna 1 and 5-segmented in antenna 2, but in the Hyperiidea the number of

peduncle segments is often fewer in antenna 1 and

always fewer in antenna 2. The second segment of

antenna 2, which bears the gland cone on which

the antennal gland opens, is fused to the head,

hence the peduncle consists of only 3 (or fewer)

segments.

The Hagellum may be filiform and multisegmented, composed of a few long segments that fold upon
NUMBER 146

-PEREON

elson

FIGURE 1.—Diagram of a hyperiid amphipod, based on $ Hyperia.

FIGURE 2.—Diagrams of $ heads of the 4 superfamilies of

Physocephalata: A, Vibilioidea; B, Phronimoidea; c, Lycaeopsoidea; D, Platysceloidea.

each other, or reduced to a single large segment

which may be followed by 1 or a few small or rudimentary additional segments. The distal segments

may be inserted at the tip of the first segment

(terminally) or proximally to the tip (subterminally). A secondary or accessory flagellum, present

in many Gammaridea, is never found in Hyperiidea. Both antennae are usually reduced in the

female, especially antenna 2, which may be rudimentary or absent. In a few instances antenna 2 is

absent in the male.

The mouthparts of the Hyperiidea (Figure 3)

are much reduced compared with those of the

Gammaridea. The mandible may or may not have

a 3-segmented palp. Frequently the palp is present

in the male and absent in the female of the same

species. Rarely it is reduced to 1 segment (Pseudomi monectes) or 2 segments ($ Tryphana). The

molar may be rudimentary or absent. Maxilla 1 has

an inner lobe in the Physosomata, but not in the

Physocephalata, and the palp is 1-segmented. Both

maxilla 1 and maxilla 2 may be rudimentary (Lycaeidae) or absent (Oxycephalidae). In the maxillipeds the palp is usually absent, although a 1-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

INNER LOBES

FIGURE 3. Mouthparts of some Hypcriidea: Md, Cyllopus magellanicus; Md\ Chuneola paradoxa (molar and palp absent); Mx.l, Mx.2, Cyllopus magellanicus; Mxp, Proscina magna

(inner lobes separate) ; Mxp', Parathemisto gaudichaudii (inner lobes fused) .

COXA

BASIS

DACTYL

FIGURE 4.—Diagram of an amiphipod pereopod.

segmented rudiment is present in a few instances.

The inner lobes are partly or completely separate

in most of the Physosomata, but are completely

fused into a single median lobe in other hyperiids.

FIGURE 5.—Distal segments of simple hyperiid pereopods:

A, Lanceola; B, Tryphana; c, Pronoe; D, Paratyphis; E, Thyropus; F, Vibilia; G, Dairella; H, Sympronoe. All drawings

are of pereopod 1, except c, which is of pereopod 2.

In a few species the inner lobe is rudimentary or

absent.

Each of pereonites 1-7 bears a pair of 7-segmented

appendages, the pereopods. The proximal segment,

or coxa, forms an immovable flat coxal plate fixed

to the lateral surface of the pereonite and sometimes fused to it. The remaining 6 segments forming the functional leg are the basis, ischium, merus,

carpus, propus, and dactyl (Figure 4). When a

pereopod is not prehensile it is said to be simple

(Figure 5). A prehensile pereopod may be either
NUMBER 146

FIGURE 6.—Distal segments of subchelate hyperiid pereopods:

A, Phrosina, pereopod 4; B, Anchylomera, pereopod 3; c,

Simorynchotus, pereopod 2; D, Eupronoe, pereopod 1; E,

Tetrathyrus, pereopod 2; F, Paraphronima, pereopod 1; c,

Brachyscelus, pereopod 2.

subchelate or chelate. In the Hyperiidea prehension

is usually effected by closure of the propus against

the expanded carpus, whereas in Gammaridea the

dactyl closes against the expanded propus. In a

subchelate pereopod (Figure 6) the prehensile surface of the carpus is its widened distal margin or its

convex anterior or posterior margin, but in a

chelate pereopod the carpus is produced distally

into a carpal process against which the propus

closes (Figure 7). When the carpal process is small

it is difficult to decide whether the pereopod is

chelate or subchelate. Here the term "weakly

chelate" may be used, but illustrations show the

condition far more clearly than words. Rarely, a

chela or subchela is formed between the dactyl and

propus (Figures 19, 35, 45) and a complex chela is

found in Amphithyrus (Figure 7<;).

Pereopods 3 and 4 are usually slender and simple,

but in Parathemisto (Figure 43), Phronimopsis

(Figure 45), Anchylomera (Figure 6B), and Phrosina

(Figure 6.\) they are prehensile. Pereopod 5 has

a strong subchela in Phronimidae and Anchylomeridae. In pereopods 5 and 6 the basis is often

FIGURE 7.—Distal segments of chelate hyperiid pereopods:

A, Parapronoe; B, Platyscelus; c, Sympronoe; D, Vibilia; E,

Phronimopsis; F, Oxycephalus; c, Amphithyrus; H, Lestrigonus. \-c, pereopod 2; H, pereopod 1.

quite broad, especially in the Parascelidae and

Platyscelidae, where they form broad opercula

covering the other pereopods during conglobation.

In these families pereopod 6 is especially enlarged;

on its posterior margin is the telsonic groove, and

on its lateral surface is usually a pocket-shaped

fissure of unknown function (Figure 74). Pereopod

7 is often smaller than the other pereopods, sometimes consisting only of the basipod (Figure 74).

In the Lanceolidae and Chuneolidae the apex of

the propus in some pereopods is expanded into a

hood surrounding the base of the dactyl (Figure 8).

The dactyl can be flexed posteriad so that it lies

within the spoon-shaped cavity within the hood,

hence it has been called a "retractile dactyl," but

we propose the term "hooded dactyl," which more

accurately describes the structure.

The gills are hollow sacs arising from the medial

surfaces of the coxae. Typically there are 5 pairs,

on pereonites 2-fi, but they may be reduced to 2, 3,

or 4 pairs. The gills may have transverse pleats or

folds, which increase the respiratory surface.

Medial to the gills, on mature females, are the

oostegites, commonly on pereonites 2-5. Oostegites

are thin plates which overlap to form a pouch

beneath the pereon in which the eggs and young

are brooded. In the Hyperiidea the oostegites
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Pcrcopod 6 of Scypholanceola vanhoeffeni, showing hooded dactyl.

usually do not have marginal setae as in the

Gammaridea. In Rhabdosoma the oostegites are

reduced in size and their function is taken over by

the gills.

Each pleonite bears a pair of biramous swimming

appendages, the pleopods. Thus far they have not

been shown to be useful in taxonomy, but they have

not been closely studied.

The urosome bears 3 pairs of uropods, each consisting of a proximal segment, the protopod, bearing distally an exopod and an endopod (Figure 9).

The exopods and endopods are flattened, moderately to very broad, rarely quite narrow, but never

styliform as in some Gammaridea. One or more of

FIGURE 9.—Urosomes: A, Pronoe capito (telson not fused

with double urosomite; endopods not fused with protopods) ;

B, Oxycephalus piscator (telson fused with double urosomite;

endopods of uropods 2 and 3 fused with protopods).

the endopods, and less commonly the exopods, may

be fused with the protopod. In the Anchylomeridae

each uropod is a single leaflike segment, and in the

Cystisomatidae and the female Phronimella uropod

2 is entirely absent.

Natural History

The Hyperiidae are entirely pelagic, mostly

oceanic, although a few species are found in coastal

waters. The swimming ability, which we have

observed in living specimens, varies from feeble

movements of the limbs in the weakly muscled

Cystisoma to very rapid swimming in Parapronoe,

which has strongly developed pleonal muscles.

Feeding habits are largely unknown. The appendages lack the rows of plumose setae found in

filter-feeding Crustacea. Some species are definitely

known to be associated with medusae or salps, e.g.,

Hypcria (Hollowday, 1947; Pirlot, 1932; Dahl,

1959a, 1959b; Bowman, Meyers, and Hicks, 1963;

Laval, 1968a). Dahl (1959a, 1959b) identified
NUMBER 146

nematocysts from the host in the gut of Hyperia

galba and concluded that it feeds on the medusa.

But the observations of Pirlot (1932) and Bowman,

Meyers, and Hicks (1963) suggested that the host

tissues are not the only and perhaps not the

principal source of nourishment. Possibly the

medusa serves as a feeding platform from which

the amphipod makes short excursions to collect

food particles, including some from food captured

by the medusa. Certainly the well-developed grinding surface on the molars of Hyperia mandible is

fitted for chewing tougher material than the

gelatinous tissues of a medusa.

Observations on living hyperiids have been made

in recent years by Laval (1963, 1968a, 1968b, 1972),

who raised Lestrigonus schizogeneios and Bougisia

ornata with their host, the leptomedusa Phialidium.

As long as the medusa was adequately fed (in this

case on Artemia) the amphipod shared the food of

its host. If the medusa did not capture enough

food, the amphipod fed on the host tissues, beginning with the gonads.

The Phronimidae live in transparent "barrels,"

open at both ends, made by remodeling pyrosomes

or siphonophores. Laval (1968b) describes how a

nectophore of the siphonophore Abylopsis is fashioned into a barrel by Phronima curvipes. The

Phronima resides in the barrel, capturing prey

passing within reach with the 3rd pereopods. It

probably feeds mainly on slow-moving and softbodied forms. By holding the abdomen outside of

one of the barrel entrances and beating the pleopods,

the Phronima can propel the barrel through the

water.

In the genus Vibilia, ovigerous females attach

themselves to salps during the approximately 20

days that the eggs arc being incubated in the

marsupium. After the eggs hatch, the mother

reaches into her marsupium and grasps a larva

with the digitiform dactyls of her 7th pereopods

and places it on the surface of a salp. The larva,

which cannot swim since the pleopods and uropods

have not yet developed, enters the atrium of the

salp by way of one of the siphons and feeds on the

tissues of its host until it has developed to the

stage where it can become planktonic (Laval, 1963).

Members of the deep-water family Lanceolidae

are believed to live on deep-water medusae and

siphonophores and feed on their tissues. The

hooded dactyls of pereopods 5-7 (or of 6 and 7 in

Prolanceola) enable them to clasp the host tissue

firmly.

Vinogardov (1957) found the guts of Lanceolidae

full of a detritus-like mass containing many small

bodies that appeared to be nematocysts. In

Lanceolidae stomachs, Woltereck (1927) found

polychaete worms (Alciopidae), chaetognaths, and

even the anterior end of a rather large cephalopod

larva. He felt certain that such prey could not have

been captured by the slow-moving Lanceolidae and

must have been obtained secondarily from the

medusa or siphonophore host.

Doubtless many more associations between

hyperiids and their hosts, probably mostly gelatinous

animals such as coelenterates and thaliaceans, remain to be elucidated. In particular the Platysceloidea, with their reduced mouthparts, must feed

on soft tissues. However, some hyperiids appear to

be entirely free-living. Parathemisto is predaceous

on other plankters (Bigelow, 1926:107; Dunbar,

1946; Bowman, 1960); and unpublished studies of

gut contents indicate that Primno is also to a large

extent predaceous.

Classification

The classification used here is essentially that of

Pirlot (1929) but incorporates the additions of

Stephensen and Pirlot (1931). Our primary aim is

not to revise the classification of the Hyperiidea,

but certain nomenclatural changes have been necessary to bring the classification into harmony with

modern taxonomic practice and the International

Code of Zoological Nomenclature. Preparation of

the diagnoses forced us to examine in detail the

characters of all taxa of hyperiids at and above the

generic level. As a result we have made the changes

given below.

INFRAORDF.RS AND SUPERFAMILIES.—We consider

the names "Hyperiidea Gammaroidea" and "Hyperiidea Genuina" of Woltereck (1909) and

"Hyperiidea Physosomata" and "Hyperiidea Eugenuina" of Pirlot (1929) for their tribes (our

infraorders; see below) to be singularly inappropriate and meaningless. Moreover, they are

double words where single names are required.

This legal technicality gives us the opportunity,

which we gladly seize to propose the new infraorder
8

name Physocephalata (from "physos," swollen, plus

"cephalon," head), referring to the large head of

most of its members. The name has the advantage

of similarity in construction to "Physosomata," the

other infraorder, which is characterized by the

swollen pereon.

Tribes have been changed to infraorders and

subtribes to superfamilies. These changes were

made to conform with the usual rank of the category

"tribe," between family and genus. Since superfamily names are formed by adding a standard

ending (usually "oidea") to the stem of one of the

included families, we have replaced the old names

of subtribes. Thus, Lanceoliformata and Sciniformata become Lanceoloidea and Scinoidea, using

the original stems. We regret that it is necessary

to replace Bovallius' (1890) Recticornia, Filicornia,

and Curvicornia, which have been widely used since

1890, but they do not qualify as names of superfamilies. In place of Filicornia we have chosen

Phronimoidea rather than Hyperioidea, in order to

avoid confusion with Hyperiidea. For Curvicornia

we have selected Platysceloidea over other possibilities because Claus (1879b, 1887) included all the

families of Curvicornia in his "Platysceliden."

Although the family Lycaeopsidae until now has

been included in the Curvicornia, we are convinced

that it differs sufficiently from the other families to

warrant for it the establishment of a new superfamily, Lycaeopsoidea.

FAMILIES.—Stephensen (1925) assigned Anapronoe

to the family Pronoidae without giving his reasons

for placing it in that family rather than in the

Lycaeidae. The principal criteria by which the

two families may be distinguished are found in

antenna 2 of the female, mandibular palp, maxillae

1 and 2, and pereopod 7. Of these, only pereopod

7 has been described for Anapronoe, and in this

character Anapronoe agrees with the Lycaeidae

rather than the Pronoidae. We have been unable

to assign Anapronoe to either family, since it shares

some characters with each and some with neither.

Redefining either family to enable it to encompass

Anapronoe would drastically alter that family's

definition and increase the difficulty of distinguishing between the two families; hence, we have

erected a new family, Anapronoidae, for Stephensen's genus.

GENERA.—We have reduced 3 genera to synonyms

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of older genera because the characters used to

separate them seemed insufficient. Vibilioides

Chevreux (1905) differs from Vibilia Milne Edwards

(1830) only in the reduced number of segments on

pereopod 7. Since Hurley (1955) has shown that

this character is variable in Cyllopus, the only

other genus of Vibiliidae, we do not believe it to

have generic value. Vibilioides therefore becomes

a synonym of Vibilia, and its type and only species

becomes Vibilia alberti (Chevreux).

Similarly, as Stephensen (1925) pointed out,

Thyropus Dana, 1852, differs from Parascelus Claus,

1879, only in the longer pereopod 6, a difference of

less than generic significance. We are therefore

combining the 2 genera under the older name

Thyropus. The family name remains Parascelidae,

in accordance with Article 40 of the International

Code.

Metalycaea was proposed by Stephensen (1925)

for a new species, M. globosa, represented by 3

females in the collections of the Thor expedition.

We believe that M. globosa is the female of Lycaea

serrata Claus, described by Claus (1879b) from the

male. Stephensen had 5 males of L. serrata but did

not associate them with his M. globosa, perhaps

because they did not occur at any of the same

stations. Shoemaker's (1945) 2 males and 1 female

of L. serrata were obtained in 3 different net tows,

but he recognized that they were conspecific and

gave what he thought was the first description of a

female L. serrata. His drawing of the whole animal

does not resemble closely Stephensen's habit figures,

since the latter are of immature specimens (4 and

5 mm), whereas Shoemaker's ovigerous female

measured 10.5 mm. The appendages drawn by

Stephensen were from his 8-nim female and show

good agreement with those figured by Shoemaker.

Shoemaker's males agree closely with Claus' description and illustrations of L. serrata.

We propose 1 new genus, Spinoscina. Acanthoscina

spinosa Chevreux, 1914, placed in Ctenoscina by

Wagler (1926), has been shown by Vinogradov

(1957) to have characters of both Ctenoscina and

Acanthoscina. Rather than combine these genera,

we have elected to establish the new genus

Spinoscina for S. spinosa.

We have omitted from this synopsis the genera

Cyllias Bovallius, 1887, and Daira Milne Edwards,

1830 (synonyms: Dairilia Dana, 1953; Eudaira
NUMBER 140

Bovallius, 1889). Cyllias was proposed as a genus

of Cyllopodidae (now included in Vibiliidae) by

Bovallius (1887b) for Hyperia tricuspidata Streets

(1877). Both Dairilia and Eudaira were substituted

for Daira in the mistaken belief that the latter

name was preoccupied by Daira Leach, a genus of

crabs. In fact, no such genus was ever published

by Leach, and the crab genus Daira de Haan (1833)

is a junior homonym of Daira Milne Edwards.

However, the International Commission on Zoological Nomenclature (1957) has placed Daira

Milne Edwards on the Official Index, thereby

suppressing it, and has placed both Daira de Haan

and Dairilia Dana on the Official List, making them

the valid names for the crab and amphipod genera

respectively. Both Cyllias Bovallius and Daira

Milne Edwards were described in vague terms and

were not illustrated, and we are unable to assign

them even to families.

REVISED CLASSIFICATION.—With the above changes,

our classification is as follows:

Order Amphipoda

Suborder Hyperiidea

Infraorder Physosomata

Superfamily Lanceoloidea

Family Lanceolidae

Chuneolidae

Microphasmidae

Superfamily Scinoidea

Family Archacoscinidae

Scinidae

Mimonectidae

Proscinidae

Infraorder Physocephalata

Superfamily Vibilioidea

Family Cystisomatidae

Vibiliidae

Paraphronimidae

Superfamily Phronimoidea

Family Hyperiidae

Dariellidae

Anchylomeridae

Phronimidae

Superfamily Lycaeopsoidea

Family Lycaeopsidae

Superfamily Platysceloidea

Family Pronoidac

Anapronoidae

Lycaeidae

Oxycephalidae

Platyscelidae

Parascelidae

Suborder HYPERIIDEA H. Milne Edwards, 1830

Body form ranging from extremely slender and

elongate to nearly globular. Head and pereon

generally tumid and round in cross section or

somewhat flattened dorsoventrally. Eyes sometimes

small, more often enormously developed, frequently

covering most of surface of head. Pereonite 1 never

fused with head. Coxae small, often fused with

pereonites. Pleon usually powerfully developed;

urosomites 2 and 3 fused (=double urosomite).

Telson entire, never cleft or incised, without setae.

Antenna 1 without accessory flagellum. Antennae

1 and 2 often reduced in female. Maxilla 1 with or

without inner lobe; palp 1-segmented. Maxilliped

with inner lobes usually fused; palp absent.

Pereopods 1 and 2 infrequently simple, usually

subchelate or chelate or intermediate between

these 2 conditions; prehension almost always between carpus and propus rather than between

propus and dactyl as in Gammaridea. One or more

of pereopods 3-7 also sometimes prehensile. Pereopod 7 often reduced in size and number of segments. Uropods never with 2-segmented rami;

without long marginal setae. Gills varying in number from 5 pairs (on pereonites 2-6) to 2 pairs (on

pereonites 5 and 6). Gut with 1 pair of digestive

caecae and no rectal glands. Heart with 2 (rarely

3) pairs of ostia.

Infraorder PHYSOSOMATA Pirlot, 1929

Head small, usually shorter than pereonite 1.

Eyes small, composed of few facets, often completely absent. Mandible without molar. Maxilla

1 with inner lobe. Maxilliped almost always with

inner lobes separate. Pereopods 1 and 2 simple;

rarely, pereopod 1 chelate.

References: Stephensen and Pirlot, 1931:501;

Vinogradov, 1957:186.

The infaorder is divided into 2 superfamilies.

Since the characters used to separate the superfamilies are sometimes difficult to see, we give a

kev to the seven families of the infraorder.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Families of Physosomata

1. Mandible with 3-segmcnted palp 2

Mandible without palp, rarely with rudimentary 1-segmented palp 4

2. At least pereopods 6 and 7 with spoon-shaped propus and hooded dactyl VII. LANCEOI.IDAE

Pereopods without spoon-shaped propus and hooded dactyl 3

3. Carpus of pereopods 1 and 2 not broadened distally. Mandible with narrow serrate incisor

and well-developed lacinia mobilis I. ARCHAEOSCINIDAF.

Carpus of pereopods 1 and 2 broadened distally. Mandible with broad incisor, nearly as

broad as body of mandible; lacinia reduced V. MICROPHASMIDAK

4. Antenna somewhat shorter than head. Pereopods 3-7 with spoon-shaped propus and hooded

dactyl VI. CHUNEOI.IDAF.

Antenna 1 much longer than head. Pereopods 3-7 without spoon-shaped propus; pereopods

5—7 rarely with hooded dactyl 5

5. Pereopod 5 longer than other pereopods; basis usually with seriate margin (s) . Endopods

of all uropods fused with protopods. Inner lobes of maxilliped always fused; more or less

reduced, sometimes completely lacking IV. SCINIDAF

Pereopod 5 not longer than other pereopods; basis with smooth margins. Uropods with free

exopods and endopods. Inner lobes of maxilliped well developed and separate 6

6. Anterior pcreonites of female much inflated. Antenna 1 inserted in middle of anterior surface

of head. Antenna 2 of male at most one-third as long as antenna 1 II. MIMONECTIDAE

Body form of female like that of male; anterior pereonitcs not inflated. Antenna 1 inserted

on dorsal part of anterior margin of head. Antenna 2 of male as long as antenna 1

III. PROSCINIDAE

Superfamily SCINOIDEA, new name

Synonym: Sciniformata Stephensen and Pirlot,

1931.

Male always with body of normal form, not

inflated. Mandible with well-developed lacinia

mobilis. Maxilla 2 with only a lew distal spines

on inner and outer lobes. Maxilliped with inner

lobes either completely separate or completely

fused. Pereopods 5-7 very rarely with hooded

dactyls. Gills on pereonite 2, when present, as large

as those of following pereonites; gills rarely absent

on pereonite 2 or 3. Oostegites on pereonites 2-5,

usually with marginal setae.

Four families (Archaeoscinidae, Mimonectidae,

Proscinidae, Scinidae).

1. Family ARCHAEOSCINIDAE Stebbing, 1904

Body form very different in male and female.

Male slender, similar to Scina; total length close to

2 mm. Female with pereonites 1-5 inflated into a

spherical form; head immersed in pereonite 1;

pereonites 6 and 7 and pleon narrow; total length

up to 6 mm. Cuticle transparent. Coxae distinct

from pereonites. Telson very small, triangular, not

fused with double urosomite. Eyes very small or

absent. Antenna 1 with large, densely setose 1st

flagellar segment; distal segments short, slender.

Antenna 2 about half as long as antenna 1, 5- or

6-segmented. Mandible with long, 3-segmented

palp; incisor very narrow; lacinia mobilis well

developed. Maxilliped with large outer lobes;

inner lobes small, fused proximally, separate distally. All pereopods simple, without chela or

subchela; dactyl not hooded; carpus of pereopods

1 and 2 not broadened distally. Uropods long and

very narrow; all ranii free. Gills on pereonites 3-6.

One genus.

1. Archaeoscina Stebbing, 1904

FIGURE 10

Type-species: Archaeoscina bonnieri Stebbing,

1904.

With the characters of the family.

References: Bovallius, 1889:70; Woltereck, 1906:

190; Stephensen and Pirlot, 1931:534; Vinogradov,

1956:200.

II. Family MIMONECTIDAE Bovallius, 1885

Body form different in male and female. Male

slender; head as high as pereonite 1. Female with

pereonites 1-6 strongly inflated; pereonite 7 and
NUMBER 146 11

FIGURE 10 Archaeoscina.

pleon slender; head much smaller than pereonite 1.

Total length 6-24 mm. Eyes very small and composed of few facets or completely lacking. Coxae

distinct from pereonites. Telson small, triangular,

not fused with double urosomite. Antenna 1 much

longer than head, inserted in middle of anterior

margin of head; with short peduncle, large 1st

flagellar segment and a few short distal flagellar

segments. Antenna 2 in*male composed of few

segments, not more than one-third as long as

antenna 1; in female rudimentary, with inflated

proximal segment and 1 or 2 stubby distal segments. Mandible without molar or palp, rarely

with rudimentary 1-segmented palp; lacinia mobilis

well developed. Maxilliped with large rounded

outer lobes (sometimes with 1-segmented rudimentary palp); inner lobes small, separate. Pereopods all simple, without chela or subchela; dactyls

not hooded. Uropods long, narrow; exopods and endopods free. Gills on pereonites 2-6.

Two genera.

Key to the Genera of Mimonectidae

Mandible without palp 2. Mimonectes

Mandible with 1-segmented rudimentary palp 5. Pseudomimonectes

2. Mimonectes Bovallius, 1885

FIGURE 11

Type-species: Mimonectes loveni Bovallius, 1885

(by present designation).

Mandible without palp. Pereopods 6 and 7,

propus slender; dactyl at most weakly curved.

References: Stephensen and Pirlot, 1931:503;

Vinogradov, 1957:205.

3. Pseudomimonectes Vinogradov, 1960

FIGURE 12

Type-species: Pseudomimonectes robustus VinoFIGURE 11.—Mimonectes.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gradov, 1960a. (The genus is based on a single,

sexually immature specimen.)

FIGURE 12.—Pseudomimonectes.

Mandible with rudimentary 1-segmented palp.

Pereopods 6 and 7 with broad propus and strong,

claw-shaped dactyl forming a prehensile organ.

Reference: Vinogradov, 1960a:219.

III. Family PROSCINIDAE Pirlot, 1933

Body in both male and female normal, not at all

or only slightly inflated; total length 4.5-18 mm.

Coxae separate from pereonites. Telson small, not

fused with double urosomite. Eyes absent. Antenna

1 much longer than head, inserted on dorsal part of

anterior surface of head, with large 1st flagellar

segment and a few short distal segments. Antenna

2 in male slender, about as long as antenna 1; in

female short or rudimentary. Mandible without

palp or molar; incisor serrate. Maxilliped with

large outer lobes; inner lobes separate. Pereopods

1 and 2 simple; carpus not broadened. Pereopods

5-7 sometimes with hooded dactyls. Uropods long

and slender, with free exopods and endopods. Gills

on pereonites 2-6.

Two genera.

Reference: Pirlot, 1939:24.

Key to the Genera of Proscinidae

Antenna 1 with 1st flagellar segment conical. Antenna 2 of female reduced to unsegmented

knob. Pereopods 5-7 without hooded dactyls 4. Proscina

Antenna 1 with 1st flagellar segment having nearly parallel margins. Antenna 2 of female 4-

segmented. Pereopods 5-7 with hooded dactyls 5. Mimoscina

4. Proscina Stephensen and Pirlot, 1931

FIGURE 13

Type-species: Parascina stephenseni Pirlot, 1929.

Both sexes with normal body, not inflated.

Antenna 1 with 1st flagellar segment long and

conical. Antenna 2 of female reduced to an unsegmented knob. Lacinia mobilis of left mandible

as broad as incisor. Pereopods 5-7 without hooded

dactyls.

References: Pirlot, 1939:25; Vinogradov, 1956:

202; 1957:208; 1960a:222; 1964:127.

5. Mimoscina Pirlot, 1933

FIGURE 14

Type-species: Mimoscina gracilipes Pirlot, 1933.

Body of female weakly inflated. Antenna 1 with

1st flagellar segment very long, with nearly parallel

margins. Antenna 2 of female composed of 4 free

segments. Lacinia mobilis of left mandible not as

broad as incisor. Pereopods 5-7 with sharply

curved, hooded dactyls.

References: Pirlot, 1933:2; 1939:29; Vinogradov,

1960a:222; 1962:13.

IV. Family SCINIDAE Stebbing, 1888

Body slender, usually flattened dorsoventrally;

total length including antennae and uropods, 5-30

mm. Head small, wider than long. Eyes small,

sometimes absent. Pereon always broader than

pleon, both sometimes with middorsal spines.

Pereonites 1 and 2 occasionally fused. Coxae dis-
NUMBER 146 13

FICURE 13.—Proscina.

FIGURE 14.—Mimoscina.

tinct. Telson quite small. Antennae 1 as long as

pereon or as pereon and pleon combined, rodshaped, inserted on dorsal anterior margin of head,

2- or 3-segmented, with very long 2nd segment, 3rd

segment minute or absent. Antenna 2 inserted on

ventral anterior margin of head; in male long,

slender, and folded beneath body, or sometimes

rudimentary; in female always rudimentary. IVfandible without palp or molar. Maxilliped with outer

lobes well developed, rarely reduced; inner lobes

completely fused, sometimes absent. Pereopods

slender, without spoon-shaped propus and hooded

dactyls; pereopod 1 simple, rarely chelate; pereopod

2 always simple. Pereopod 5 the longest; basipod of

pereopod 5 often broadened and armed with marginal teeth. All uropods with endopods fused with

protopods; exopods short, spiniform. Gills on pereonites 2-6, 3-6, or 4-6. Oostegites with marginal

setae.

References: Wagler, 1926:319; 1927:85; Vinogradov, 1960a: 224.

Four genera.

Key to the Genera of Scinidae

1. Body segments without middorsal spines 6. Scina

Body segments with middorsal spines 2

2. Pereonites 1 and 2 fused; pereon with 6 middorsal spines 3

Perconites 1 and 2 separate; pereon with 5 or 7 middorsal spines 8. Ctenoscina

3. Pereopod 1 simple 7. Acanthoscina

Pereopod 1 chelate 9. Spinoscina
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6. Scina Prestandrea, 1833

FIGURE 15

Type-species: Scina ensicorne Prestandrea, 1833.

Body segments without middorsal spines. Pereonites 1 and 2 separate. Antenna 2 of male well

developed, folded beneath body. Maxilliped with

inner lobe present; outer lobes well developed.

Pereopod 1 simple. Pereopod 5 without fusion of

segments. Gills on pereonites 2-6 or 3-6.

References: Wagler, 1926:324; 1927:85; Hurley,

1956:4; Vinogradov, 1960a:224.

FIGURE 15 Scina.

7. Acanthoscina Vosseler, 1900

FIGURE 16

Type-species: Scina acanthodes Stebbing, 1895.

Body segments with middorsal spines, 6 on

pereon, 3 on pleon. Pereonites 1 and 2 fused.

Second segment of male antenna 1 expanded at

base. Antenna 2 of male and female represented

by a pair of long sharp spines. Maxilliped without

inner lobe; outer lobes reduced to two small oval

plates. Pereopod 1 simple. Pereopod 5 with carpus,

merus, and propus fused. Gills on pereonites 2-6.

Reference: Wagler, 1926:424.

8. Ctenoscina Wagler, 1926

FIGURE 17

Type-species: Ctenocina tennis Wagler, 1926 (by

present designation).

Body segments with middorsal spines, 5 or 7 on

pereon, 3 or 4 on pleon. Pereonites 1 and 2

separate. Antenna 2 of male and female represented
NUMBER 146 15

FIGURE 16.—Acanthoscina.

FIGURE 17.—Ctenoscina.
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

by a pair of long sharp spines, except in the male

C. macrocarpa, which has antenna 2 composed of 5

short segments. Maxilliped without inner lobe;

outer lobes well developed. Pereopod 1 chelate.

Pereopod 5 without fusion of segments. Gills on

pereonites 4-6.

References: Wagler, 1926:430; Vinogradov, 1964:

140.

Body segments with middorsal spines, 6 on#

pereon, 4 on pleon. Pereonites 1 and 2 fused.

Antenna 2 of female (male unknown) represented

by a pair of long sharp spines. Maxilliped with

inner lobe present; outer lobes well developed.

Pereopod 1 chelate. Pereopod 5 without fusion of

segments. Gills on pereonites 3-6.

Reference: Vinogradov, 1957:219.

9. Spinoscina, new genus

FIGURE 18

Type-species: Acanthoscina spinosa Chevreux,

1914.

Superfamily LANCEOLOIDEA, new name

Synonym: Lanceoliformata Stephensen and Pirlot, 1931.

Body in both sexes more or less inflated. Mandible

FIGURE 18.—Spinoscina.

usually with weakly developed lacinia mobilis.

Maxilla 2 with numerous, usually very large spines

on both lobes. Maxilliped with inner lobes fused

proximally, separate distally. Pereopods 6 and 7

with spoon-shaped propus and hooded dactyl, or

some of posterior pereopods subchelate; rarely all

pereopods simple. Gills on pereonites 2-6 or 4-6,

those of pereonite 2 very small. Oostegites on

pereonites 2-5, usually without marginal setae.

Three families (Microphasmidae, Chuneolidae,

Lanceolidae).

V. Family MICROPHASMIDAE Stephensen

and Pirlot, 1931

Pereonites 2-4 strongly inflated; pereonites 5-7
NUMBER 146 17

and pleon slender; total length 3-25 mm. Cuticle

transparent. Coxae separate from pereonites. Telson

small, triangular, not fused with double urosomite.

Head about as long as pereonite 1, with massive

mouthpart region. Eye very small or absent.

Antenna 1 fiagellum with large 1st segment and a

few very small distal segments, the latter sometimes

missing. Antenna 2 5- or 6-segmented, shorter or

slightly longer than antenna 1. Mandible with

long, 3-segmented palp; incisor broad, lacinia

mobilis very narrow. Maxilliped with large outer

lobes; inner lobes fused proximally, separate distally. Pereopods 1 and 2, carpus broadened

distally; propus conical. Pereopods 3-5 and 7 often

prehensile. Pereopods never with hooded dactyls.

Uropods narrow, with free exopods and endopods.

Three genera.

Key to the Genera of Microphasmidae

1. Antennae 1 and 2 very long, both reaching far beyond anterior margin of head; antenna 2

somewhat longer than antenna 1. All pereopods with carpus as long as or almost as long

as propus 12. Mimonecteola

Antennae 1 and 2 short; antenna 2 distinctly shorter than antenna 1. All pereopods with

carpus distinctly shorter than propus - 2

2. Antenna 1 reaching beyond anterior margin of head. Pereopods 3-5 subchelate

10. Microphasma

Antenna 1 not reaching beyond anterior margin of head. Only pereopod 5 subchelate

11. Microphasmoides

10. Microphasma Woltereck, 1909

FIGURE 19

Type-species: Microphasma agassizi Woltereck,

1909.

Eye very small. Antenna 1 short, inserted on and

extending beyond anterior margin of head. Antenna

2 much shorter than antenna 1; proximal segment

much inflated; gland cone very large. All pereopods

with carpus distinctly shorter than propus. Pereopods 3-5 subchelate; propus broad, distal part of

closing margin with row of strong spines; dactyl

curved, on pereopods 3 and 4 almost half as long as

propus, on pereopod 5 somewhat shorter.

References: Woltereck, 1909:153; Stephensen and

Pirlot, 1931:539; Shoemaker, 1945:218.

FIGURE 19.—Microphasma.

11. Microphasmoides Vinogradov, 1960

FlCURE 20

Type-species: Microphasmoides vitjazi Vinogradov, 1960a.

Eye absent. Antenna 1 small, inserted on lateral

surface of head and not reaching anterior margin

of head. Antenna 2 shorter than antenna 1; proximal segment much inflated; gland cone very large.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

All pereopods with carpus distinctly shorter than

propus. Pereopods 3 and 4 with long narrow

propus. Pereopod 5 subchelate; propus with subparallel margins; distal margin with row of strong

spines; dactyl short, hook-shaped. Pereopods 6 and

7 with propus narrowing distally.

Reference: Vinogradov, 1960a:213.

FIGURE 20 Microphasmoides.

12. Mimonecteola Woltereck, 1909

FIGURE 21

Type-species: Mimonecteola diomedeae Woltereck, 1909.

Eye lacking or very small. Antennae 1 and 2

very long, inserted on and extending far beyond

anterior surface of head. Antenna 2 somewhat

longer than antenna 1; proximal segment not

inflated; gland cone small. All pereopods with

carpus as long as or nearly as long as propus. All

pereopods usually simple; pereopods 5 and 7 rarely

subchelate, but propus only slightly broadened.

References: Woltereck, 1909:153; Shoemaker,

1945:224; Vinogradov, 1964:118.

FIGURE 21.—Mimonecteola.
NUMBER 146

19

VI. Family CHUNEOLIDAE Woltereck, 1909

Body weakly arched, somewhat flattened dorsoventrally, slightly broader than high; total length

5-41 mm. Coxae not fused with pereonites. Telson

short, triangular, not fused with double urosomite.

Head with blunt to triangular rostrum, somewhat

longer thaa pereonite 1. Eyes very small or absent.

Antenna 1 somewhat shorter than head, inserted

in lateral depression of head beneath rostrum;

peduncle 3-segmented; flagellum with very large

proximal segment and sometimes a few minute

distal segments. Antenna 2 shorter than antenna

1, with large inflated proximal segment (containing

opening of antennal gland) and minute distal

segment. Mandible without palp or molar; lacinia

mobilis broad. Maxilliped with large, rounded

outer lobes; inner lobes small and completely

separate. All pereopods with hooded dactyls;

pereopods 3-7 with spoon-shaped propus. Uropods

with free exop'ods and endopods. Gills on pereonites 2-6. Oostegites on pereonites 2-5.

One genus.

13. Chuneola Woltereck, 1909

FICURE 22

Type-species: Chuneola paradoxa Woltereck,

1909.

With the characters of.the family.

References: Pirlot, 1930:3; Vinogradov, 1956:196;

1957:201; 1960b:251.

FIGURE 22.—Chuneola.

VII. Family LANCEOLIDAE Bovallius, 1887

Body greatly inflated in larva, only slightly so in

adult, total length 6-70 mm. Eyes small, rudimentary, or absent. Coxae separate from pereonites.

Antenna 1 with 1st flagellar segment large, robust,

nearly straight or slightly curved; distal segments

Key to the Genera of Lanceolidae

(Modified from Vinogradov, 1957)

1. Pereopods 5 and 6 with spoon-shaped propus and hooded dactyl 2

Only pereopods 6 and 7 with spoon-shaped propus and hooded dactyl. Pereopod 1 with

propus widening distally 18. Prolanceola

2. Gills on pereonites 4-6. Maxilla 1 with palp shorter than outer lobe 16. Metalanceola

Gills on pereonites 2-6. Maxilla 1 with palp longer than outer lobe 3

3. Pereopod 1 with propus produced into lobe above insertion of dactyl so that dactyl is inserted subterminally 17. Paralanceola

Pereopod 1 with propus not produced anteriorly above insertion of dactyl; dactyl inserted

terminally 4

i. Pereopod 1 with carpus subquadrate, only slightly wider at distal end than at proximal end.

Left mandible, lacinia nearly as wide as incisor 15. Megalanceola

Periopod 1 with carpus subtriangular, markedly wider at distal end than at proximal end.

Left mandible with lacinia minute 5

5. Lateral surface of head with 2 concavities separated by an oblique ridge .19. Scypholanceola

Lateral surface of head without concavities 14. Lanceola
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

small, few in number. Antenna 2 usually longer

and more slender than antenna 1. Mandible with

3-segmented palp, without molar. Pereopod 1 with

carpus usually more or less expanded distally.

Pereopods 6 and 7 and usually pereopod 5 with

spoon-shaped propus and hooded dactyl.

Six genera.

14. Lanceola Say, 1818

FIGURE 23

Type-species: Lanceola pelagica Say, 1818.

Head usually with well-developed rostrum; interantennal lobe rounded, rudimentary to well developed. Eyes small, round to oval. Mandible with

3rd segment of palp shorter than 2nd. Maxilla 1

with well-developed palp and broad inner lobe.

Pereopod 1 with carpus expanded distally. Pereopods 5-7 with spoon-shaped propus and hooded

dactyl.

References: Bovallius, 1887c:28; Stebbing, 1888:

1301; Stephensen, 1918:8; Woltereck, 1927:60; Barnard, 1932:254; Shoemaker, 1945:206; Vinogradov,

1957:189; 1960a:200; 1926:6; 1961:109.

c?

FIGURE 23.—Lanceola.

15. Megalanceola Pirlot, 1935

FIGURE 24

Type-species: Lanceola stephenseni Chevreux,

1920.

Large amphipods, up to 70 mm in length. Head

short; rostrum very small; interantennal lobe rather narrow, with concave lateral surface, its distal

margin flaring laterad to form a rim. Eyes oval,

without pigment. Mandible with lacinia mobilis

nearly as broad as incisor. Maxilla 1 with narrow

inner lobe. Pereopod 1 with carpus only slightly
NUMBER 146 21

expanded distally. Pereopods 5-7 with spoonshaped propus and hooded dactyl.

References: Pirlot, 1935:passim; 1939:8; Shoemaker, 1945:212; Vinogradov, 1964:114.

FIGURE 24 Megalanceola.

16. Metalanceola Pirlot, 1931

FIGURE 25

Type-species: Metalanceola chevreuxi Pirlot,

1931.

Head with inconspicuous rostrum. Eyes absent.

Antenna 1 with well-developed distal segments.

Antenna 2 short, only slightly longer than peduncle of antenna 1. Mandible with 3rd segment of

palp slightly longer and not much narrower than

FIGURE 25.—Metalanceola.
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2nd segment. Maxilla 1 with palp shorter than

outer lobe. Maxilliped with very small inner lobes.

Pereopod 1 with carpus not expanded. Pereopods

3 and 4 with strong, recurved dactyls. Pereopods

5—7 with spoon-shaped propus and hooded dactyls.

Gills on pereonites 4-6.

References: Pirlot, 1931:passim; 1939:12; Vinogradov, 1960a: 210.

17. Paralanceola Barnard, 1930

FIGURE 26

Type-species: Paralanceola anomala Barnard,

1930.

Head with inconspicuous rostrum. Eyes minute,

composed of only 4 facets. Antenna 1 very short

and stout. Antenna 2 with last segment of peduncle

FIGURE 26.—Paralanceola.

elongate and slightly enlarged distally; flagellum

very long, filiform, without visible sutures. Mandibular palp with long 3rd segment. Pereopod 1 with

carpus not expanded; propus produced into lobe

above insertion of dactyl so that dactyl is inserted

subterminally. Pereopods 5-7 with spoon-shaped

propus and hooded dactyl.

Rejerence: Barnard, 1930:398.

narrow incisor and well-developed lacinia mobilis;

palp with long 3rd segment. Pereopod 1 subchelate,

with triangular carpus; propus produced distally

into lobes above and below insertion of dactyl.

Pereopods 6 and 7, but not pereopod 5, with hooded

dactyls and spoon-shaped propus.

References: Woltereck, 1909:157; Vinogradov,

1957:198.

18. Prolanceola Woltereck, 1907

FIGURE 27

Type-species: Prolanceola vibiliformis Woltereck,

1907.

Body not tumid. Head only slightly shorter than

pereonite 1, with short rostrum. A row of 4 eyespots extending dorsally from usual eye on side of

head. Antenna 1 with 1st flagellar segment tapering

and becoming very slender distally. Antenna 2

much longer than antenna 1; flagellum 6-segmented, longer than peduncle. Mandible with rather

19. Scypholanceola Woltereck, 1905

FIGURE 28

Type-species: Scypholanceola vanhoeffeni Woltereck, 1909.

Large amphipods, body length up to about 60 mm.

Head with well-developed rostrum; lateral surface

with 2 concavities separated by ridge running

obliquely posteroventrad from interantennal lobe.

Eyes without refracting elements or pigment, in

form of band with ends lying in concavities and

connected across ridge. Coxae 2-5 sharply pointed
NUMBER 146 23

FIGURE 28 Scypholanceola.

anteriorly. Mandible with 3rd segment of palp

shorter than 2nd. Maxilla 1 with well-developed

palp and broad inner lobe. Pereopod 1 with carpus

expanded dis tally. Pereopods 5-7 with spoonshaped propus and hooded dactyl.

References: Wokereck, 1909:161; 1927:65; Chevreux, 1920:7; Shoemaker, 1945:215; Vinogradov,

1956:197; 1960a:207.

Infraorder PHYSOCEPHALATA, new name

Head large, longer than pereonite 1. Eyes large

(small to moderately large in Vibilia), occupying

most of head surface (excluding rostrum in Oxycephalidae). Maxilla 1 without inner lobe. Maxilliped

with inner lobes completely fused. Pereopods 1 and

2 usually chelate or subchelate, less commonly simple.

Four superfamilies.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Superfamilies of Physocephalata

1. Antenna 1 inserted on ventral surface of head; 1st segment of flagellum of male curved.

Antenna 2 of male folded in zigzag PLATYSCFLOIDEA

Antenna 1 inserted on anterior surface of head. Antenna 2 of male not folded 2

2. Antenna 1 of male with flagellum multisegmented PHRONIMIMOIDEA

Antenna 1 of male with flagellum composed of large first segment and sometimes a few

rudimentary distal segments 3

3. Antenna 1 not inserted in groove of head; first segment of flagellum of male lanceolate or

conical; distal segments inserted terminally or absent VIBILIOIDEA

Antenna 1 inserted in groove of head; first segment of flagellum of male conical, distal segments inserted subterminally LYCAEOPSOIDEA

Superfamily VIBILIOIDEA, new name

Antenna 1 inserted on anterior surface of head;

flagellum with large straight 1st segment and one

or a few small or rudimentary additional segments

inserted terminally. Antenna 2 inserted on anterior

or ventral surface of head, short, composed of a

few segments or rudimentary. Pereopod 5 never

with large subchela.

Three families (Vibiliidae, Cystisomatidae, Paraphronimidae).

Key to the Families of Vibilioidea

1. Head very large, rounded dorsally; ventral surface flat with dentate border. Uropod 2 absent;

uropods 1 and 3, endopod fused with protopod IX. CYSTISOMATIDAE

Head subquadrangular or rounded, without dentate border. Uropods all present, endopods

not fused with protopods 2

2. Body not transparent. Mandible with palp. Pereopod 2 chelate VIII. VIBILIIDAE

Body transparent. Mandible with palp in male, not in female. Pereopod 2 simple

X . PARAI'HRONIMIDAt

VIII. Family VIBILIIDAE Dana, 1852

Body compact, moderately slender; total length

from about 5 to 20 mm. Head small to moderately

large, subquadrangular or rounded. Eyes small to

moderate-sized and separated, or large and occupying most of head surface. Pereonites all separate.

Coxae separate from pereonites. Telson short, triangular or semicircular. Antenna 1 with short

3-segmented peduncle; 1st segment of flagellum

large, straight, spatuliform or conical; remaining

segments rudimentary. Antenna 2 filiform, subequal to antenna 1. Mandible with palp and molar

process. Maxilliped with short rounded inner lobe;

outer lobes rounded at apices, with straight medial

margins and rounded lateral margins. Pereopod 1

simple; pereopod 2 chelate. Pereopods 5 and 6 the

longest; pereopod 7 short, segments sometimes reduced in number, distal segment always digitiform.

Uropods with free exopods and endopods. Gills on

pereonites 2-6. Oostegites on pereonites 2-5.

Two genera.

Key to the Genera of Vibiliidae

Eyes small to moderately large, but never occupying most of head surface. Antenna 1 with

spatuliform flagellum. Antenna 2 inserted anteriorly 20. Vibilia

Eyes occupying most of head surface. Antenna 1 with conical flagellum. Antenna 2 inserted

ventrally 21. Cyllopus

20. Vibilia H. Milne-Edwards, 1830

FICURE 29

Type species: Vibilia peronii H. Milne-Edwards

1830.

Head rather small, subquadrangular. Eyes small

to moderately large but never occupying most of

head surface. Antenna 1 with broad, laterally

flattened flagellum. Antenna 2 inserted on anterior

surface of head. Pereopod 7 usually with complete
NUMBER 146 25

number of segments, rarely with only 3 segments

distal to basipod. Protopod of uropod 3 shorter

than that of uropod 1.

References: Behning, 1913, 1925; Stephensen,

1918:33; Chevreux and Fage, 1925:382; Pirlot,

1929:91; Laval, 1963.

FIGURE 29.—Vibilia.

21. Cyllopus Dana, 1852

FIGURE 30

Type-species: Cyllopus magellanicus Dana, 1852.

Head large, almost globular. Eyes large, occupying most of head surface. Antenna 1 with slender

conical flagellum. Antenna 2 inserted on ventral

surface of head. Pereopod 7 with basipod longe

than remaining segments combined, the latter com

prising 5 or 3 segments. Protopod of uropod '

longer or shorter than that of uropod 1.

References: Barnard, 1930:405; Hurley, 1955

129.

FIGURE 30.—Cyllopus.

IX. Family CYSTISOMATIDAE

Willemoes-Suhm, 1875

Body large, cuticle thin, almost completely transparent; musculature very weak. Total length up to

120 mm. Head very large, rounded dorsally; ventra

surface flat or slightly concave, with dentate bordei

Eyes comprising 2 oval areas on dorsal surface c

head. Short middorsal spines often present o

pereonites and pleonites. Pereonites 1 and 2 o
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

even 1-4 fused. Coxae fused with pereonites. Telson

very small. Antenna 1 with 1 or 2 poorly defined

peduncle segments; flagellum composed of a slender

lanceolate proximal segment and a minute distal

segment. Antenna 2 represented by a pair of small

spines on ventral surface of head. Mandible with

molar; palp absent. Maxilla 2 without inner lobe.

Maxilliped with long inner lobe, widening distally;

outer lobes pointed at apices. Pereopod segments

with serrate margins. Pereopods 1 and 2 distinctly

chelate. Pereopod 5 the longest. Pereopod 7 sometimes prehensile, with dactyl closing against distal

end of propus. Uropod 2 absent. Endopod of uropods 1 and 3 fused with protopod. Gills on pereonites 2-6 or 4-6. Oostegites on pereonites 2-5, those

on pereonites 4 and 5 rudimentary.

One genus.

22. Cystisoma Gu£rin-M£neville, 1842

FIGURES 31,32

Type-species: Cystisoma neptunus Gue"rin-Meneville, 1842.

With the characters of the family.

References: Stephensen, 1918:56; Barnard, 1932:

268; Woltereck, 1903:passim; 1904:passim; Willemoes-Suhm, 1874: passim.

X. Family PARAPHRONIMIDAE

Bovallius, 1887

Body rather slender, transparent; total length

6-15 mm. Head large, almost cubical. Eyes large,

occupying most of head surface. Pereonites all

separate. Coxae fused with pereonites. Telson very

short. Antenna 1 shorter than head; peduncle 3-

segmented; first segment of flagellum enlarged, the

others rudimentary. Antenna 2 inserted on ventral

margin of head, rudimentary in female, slightly

MARGINAL SPII«S

FIGURE 31.—Cystisoma: diagram of head, ventral.

FIGURE 32.—Cystisoma.
NUMBER 146

longer than antenna 1 in male, composed of a few

segments. Mandible without palp and without

molar. Maxilliped with inner and outer lobes fused

into a single broad plate. Pereopods 1 and 2 much

shorter than following pereopods. Pereopod 1 weakly subchelate, merus produced into short spoonshaped lobe beneath carpus. Pereopod 2 with

propus produced into a thin hollowed process on

either side of dactyl. Pereopods 3-6 slender, subequal; pereopod 7 shorter. Uropods with free exo27

pods and endopods. Gills on pereonites 3-6. Oostegites on pereonites 2-5.

One genus.

23. Paraphronima Claus, 1879

FIGURE 33

Type-species: Paraphronima gracilis Claus, 1879a.

With the characters of the family.

References: Claus, 1879a:64; Bovallius, 1889:22;

Stebbing, 1888:1335; Vosseler, 1901:94.

FIGURE 33.—Paraphronima.

Superfamily PHRONIMOIDEA, new name

Antenna 1 inserted on anterior surface of head;

flagellum of male with long straight 1st segment

and shorter, terminally inserted distal segments;

flagellum of female 1-segmented. Antenna 2 inserted on anterior surface of head; flagellum of male

long, multisegmented; flagellum of female short,

composed of a few segments, sometimes rudimentary or absent. Pereopod 5 sometimes with large

subchela.

Four families (Hyperiidae, Dairellidae, Phronimidae, Phrosinidae).

Key to the Families of Phronimoidea

1. Pereopod 5 without large subchela 2

Pereopod 5 with large subchela 3

2. Pereopod 2 chelate XI. HYPERIIDAE

Pereopod 2 simple XII. DAIRELLIDAE

3. Uropods with free exopods and endopods. Head subconical. narrowing and prolonged

ventrally XIII. PHROMMIDAE

Uropods composed each of a single leaflike segment only. Head globular XIV. PHROSINIDAE

XI. Family HYPERIIDAE Dana, 1852

Body compact, form rather variable, most often

subglobular in female, more slender in male; total

length, 2-30 mm. Head usually large, globular. Eyes

usually large and occupying most of head surface.

Pereonites all separate, or up to 5 anterior pereon-
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ites fused. Coxae separate or fused with pereonites.

Telson small to moderate-sized, rounded or triangular. Antenna 1 with long filiform flagellum in

male, with short 1-segmented flagellum in female.

Antenna 2 similar to antenna 1, but often more

reduced in female. Mandible with molar; palp

present in both sexes or absent in female. Maxilliped quite variable; outer lobes usually well developed, sometimes fused medially; inner lobe well

developed, reduced, or even absent. Pereopod 1

simple, subchelate, or chelate. Pereopod 2 chelate.

Pereopods 3-7 varying in relative lengths, but

pereopod 7 not reduced. Uropods with free exopods

and endopods. Gills on pereonites 2-6. Oostegites

on pereonites 2-5.

Thirteen genera.

Key to the Genera of Hyperiidae

1. Pereonites all separate. Coxae separated from pereonites by sutures 2

At least pereonites 1 and 2 fused. Coxae fused with pereonites (except in Bougisia) 7

2. Eyes without facets. Head produced anteriorly into acute process between antenna 1 and

antenna 2 34. Pegohyperia

Eyes normal, with facets. Head not produced between antenna 1 and antenna 2 3

3. Anterior pereonites raised into transverse folds. Body covered with fine pubescence

31. lulopis

Pereonites not raised into folds. Body not covered with fine pubescence 4

4. Pereopods 1 and 2 with carpal process laterally compressed, knife-shaped, without spines

30. Hyperoche

Pereopods 1 and 2 with carpal process spoon-shaped, with somewhat concave center and

raised margins armed with spines 5

5. Pereopods 3 and 4 prehensile; propus closing against the dilated carpus 33. Parathemisto

Pereopods 3 and 4 with carpus not dilated 6

6. Pereopod 5 or pereopods 5 and 6 longer than pereopods 3 and 4 26. Hyperiella

Pereopods 3 and 4 longer than pereopods 5 and 6 24. Hyperia

7. Eyes small. Head produced anteriorly into acute process between antenna 1 and antenna 2.

Coxae separated from pereonites by sutures 25. Bougisia

Eyes covering most of head surface. Head not produced between antenna 1 and antenna 2.

Coxae fused with pereonites 8

8. First 2 pereonites fused in adult male and female 11

First 3, 4, or 5 pereonites fused in adult female; first 2, 3, or 4 pereonites fused in adult

male 9

9. Pereopods 5-7 subequal in length; pereopod 6 only slightly longer than pereopods 5 and 7.

Telson of moderate size 32. Lestrigonus

Pereopod 5 much shorter or distinctly longer than pereopods 6 and 7. Telson very short 10

10. Pereonites 1-3 fused. Pereopod 5 much shorter than pereopods 6 and 7. 29. Hyperionyx

Pereopods 1-5 fused. Pereopod 5 much longer than pereopods 6 and 7 36. Themistella

11. Pereopod 2 chelate; dactyl closing against process of robust propus. Pereopods 3-7 prehensile 35. Phronimopsis

Pereopod 2 chelate; propus closing against carpal process. Pereopods 3-7 not prehensile 12

12. Antenna 2 of female moderately long; shape sinuous in profile. Mandible incisor with serrate apex. Maxilliped with well-developed inner lobe. Exopods of uropods with notched

lateral margins 28. Hyperioides

Antenna 2 of female rudimentary. Mandible incisor with smooth apex. Maxilliped with

rudimentary inner lobe. Exopods of uropods without notches 27. Hyperietta

24. Hyperia Latreille, in Desmarest, 1823

FICURE 34

Type-species: Cancer medusarum O. F. Miiller,

1776.

Large species, 10-30 mm in length. Head deeper

than long. Pereon more or less dilated in female;

all pereonites free. Coxae not fused with pereonites.

Antenna 1 with 3-segmented peduncle. Mandible

with serrate incisor; palp present in both sexes.

Maxilla 1 with outer lobe having 5 apical spines;

palp with robust spine at mediodistal corner. Maxilliped with inner lobe well developed, usually with

2 terminal spines. Pereopod 1 subchelate or barely
NUMBER 146 29

chelate; carpal process only slightly developed.

Pereopod 2 chelate; carpal process spoon-shaped,

bearing spines along edge of spoon. Pereopods 1

and 2, carpus, and propus with many spines. Pereopods 5-7 shorter than pereopods 3 and 4. Uropod

3 of male with broad endopod.

References: Bovallius, 1889:129; Sars, 1890:6;

Bowman, 1972:5.

FIGURE 34.—Hyperia.

25. Bougisia Laval, 1966

FIGURE 35

Type-species: Bougisia ornata Laval, 1966.

Head with well-developed rostrum and acute

lateral lobes. Eyes small, lateral. Pereonites 1 and

2 fused. Coxal plates separate from pereonites.

Antenna 2 of female composed of a single segment.

Mandible with palp in both sexes. Maxilla 1 with

rather broad palp. Maxilliped with very short inner

lobe. Pereopod 1 subchelate; pereopod 2 chelate,

with short spoon-shaped carpal process. Pereopods

3-7 subequal; dactyl of pereopod 5 prehensile, closing against expanded distal margin of propus.

Reference: Laval, 1966.

FIGURE 35 Bougisia.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

26. Hyperiella Bovallius, 1887

FIGURE 36

Type-species: Hyperiella antarctica Bovallius,

1887b.

Body small to moderate-sized; pereon rather

plump; pereonites all separate. Antenna 1 and antenna 2 of female 4-segmented. Mandible with

palp in both sexes; incisor with serrate margin.

Maxilla 1 with inner lobe having 5 terminal spines.

Maxilla 2 with 1 terminal spine on inner lobe,

2 or 3 on outer lobe. Maxilliped with inner lobe

well developed, with 2 terminal spines; outer lobes

separate. Pereopod 1 subchelate; pereopod 2 chelate;

posterior margin of propus serrate in pereopods 1

and 2. Pereopods 3 and 4 with posterior margins of

carpus and propus serrulate; posterior margins of

carpus bearing a few slender spines. Pereopod 5

(or pereopods 5 and 6) much longer than pereopods

3 and 4; pereopod 7 (or pereopods 6 and 7) subequal to pereopods 3 and 4. Dactyls of pereopods

3-5 very long. Uropod 3 elongate, except in H.

macronyx.

References: Bovallius, 1889:242; Stebbing, 1888:

1403; Walker, 1907:7; Bowman, 1973:26.

FIGURE 36.—Hyperiella.

27. Hyperietta Bowman, 1973

FIGURE 37

Type-species: Hyperia luzoni Stebbing, 1888.

Small species, with body rather compressed laterally. Head rather short anteroposteriorly; eyes occupying most of its surface. Pereonites 1 and 2 fused

in both sexes. Coxae fused with pereonites. Telson

inserted distinctly anterior to insertion of protopod

of uropod 3. Female telson at least half as long as

FIGURE 37 Hyperieita.
NUMBER 14(5 31

protopod of uropod 3. Epistome small, inconspicuous. Antenna 1 of female 2-segmented. Antenna 2

of female 1-segmented, rudimentary; gland cones

rather inconspicuous, in anterior view converging

medially. Mandible with smooth incisor and dentate lacinia; molar narrow; with palp in male, without in female. Maxilla 1 with outer lobe having 3

terminal spines. Maxilliped with outer lobes fused

medially; inner lobe rudimentary. Pereopod 1

simple, weakly subchelate or barely chelate. Pereopod 2 chelate, with spoon-shaped carpal process

bearing marginal spines. Pereopods 3 and 4 of

female with 1 conspicuous spine on posterior margin of merus, 2 spines on posterior margin of carpus; pereopods 3 and 4 of male with shorter spines

in same positions. Distal margin of propus of pereopods 6 and 7 and sometimes of pereopod 5 produced into spinose lobe medial to base of dactyl;

dactyl unarmed. Uropods slender, margins of exopods and endopods smooth or with extremely fine

serrations.

References: Stebbing, 1888:1382; Vosseler, 1901:

64; Stephensen, 1924:83; Yang, 1960:33; Bowman,

1973:55.

28. Hyperioides Chevreux, 1900

FIGURE 38

Type-species: Hyperioides longipes Chevreux,

1900.

Small species with body rather compressed laterally. Head globular, produced anteriorly above

insertion of antenna 1; eyes occupying most of its

surface or limited entirely to dorsal surface. Pereonites 1 and 2 fused in both sexes. Coxae fused with

pereonites. Posterior elevation of pleonite 1 of

male pronounced. Telson rather short. Antenna 1

of female 2- or 3-segmented. Antenna 2 of female

1-segmented, moderately long, with characteristic

sinuous shape in lateral view; gland cone distally

rounded. Mandible with dentate incisor; with palp

in male, without in female. Maxilla 1 with outer

lobe having 3 large terminal spines and a smaller

subterminal spine. Maxilla 2 with outer lobe having 1 terminal and 1 subterminal spine; inner lobe

with 1 terminal spine. Maxilliped having inner lobe

well developed, with 2 terminal spines. Pereopods 1

and 2 chelate, with spoon-shaped carpal process

bearing marginal spines. Pereopods 5 and 6 disFICURE 38.—Hyperioides.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tinctly longer than pereopods 3 and 4 and pereopod

7; dactyls of pereopods 5-7 long, with comb of fine

setules on proximal 3rd of anterior margin. Uropods, exopods notched on lateral margins.

References: Chevreux, 1900:143; Stebbing, 1888:

1379; Vosseler, 1901:60; Bowman, 1973:30.

29. Hyperionyx Bowman, 1973

FICURE 39

Type-species: Hyperia macrodactyla Stephensen,

1924.

Small species. Head globular, eyes occupying

most of its surface. Pereonites 1-3 fused in both

sexes. Coxae fused with pereonites. Telson very

short. Antenna 1 of female 2-segmented. Antenna 2

of female 1-segmented, slender, moderately long;

gland cone small, triangular. Mandible reduced

[?]; with palp in male, without palp in female.

Maxilla 1 with outer lobe having 1 terminal spine;

palp with very few marginal setae. Maxilla 2 with

few setae; outer lobe with 2 terminal spines; inner

lobe very short. Maxilliped with very few setae;

inner lobe slender; outer lobes obovate. Pereopod

1 subchelate. Pereopod 2 chelate, with spinose,

spoon-shaped carpal process. Pereopods 3-7 with

strong, curved, unarmed dactyls. Pereopods 3 and 4

and pereopod 6 subequal; pereopod 5 much shorter; pereopod 7 somewhat longer.

References: Stephensen, 1924:90; Yang, 1960:35;

Bowman, 1973:71.

Al

FIGURE 39.—Hyperionyx.

30. Hyperoche Bovallius, 1887

FIGURE 40

Type-species: Metoecus medusarum Kr0yer, 1838.

Pereonites all separate. Coxae separate from pereonites. Antenna 1 with 3-segmented peduncle. Mandible with palp in both sexes; molar laminate,

triangular, nearly smooth. Maxilla 1 with broad

palp. Maxilliped with inner lobe well developed;

outer lobes separate. Pereopods 1 and 2 distinctly

chelate; carpal process laterally compressed, knifeshaped. Pereopods 3-7 subequal; pereopods 3 and

FIGURE 40.—Hyperoche.
NUMBER 146

4 with posterior margin of carpus produced into

thin ridge which may end distally in a tooth.

References: Bovallius, 1889:83; Senna, 1906:154;

Steuer, 1911:674.

31. Iulopis Bovallius, 1887

FIGURE 41

Type-species: Iulopis loveni Bovallius, 1887b (by

present designation).

Body and pereopods covered with fine setae, giving the species a furry appearance. Pereonites all

free. Pereonites 2-7 raised into transverse rounded

folds. Coxae not fused with pereonites. Antenna 2

much reduced in female. Mandible with palp in

male, without palp in female. Maxilliped with

obsolete inner lobe and sparsely spinose outer lobes.

Pereopod 1 subchelate; pereopod 2 chelate; carpal

process narrowly hollowed in both. Pereopods 3-7

subequal; pereopods 3 and 4 may be slightly shorter or slightly longer than pereopods 5-7.

References: Bovallius, 1889:116; Spandl, 1927:

159; Shoemaker, 1945:238.

FIGURE 41.—Iulopis.

32. Lestrigonus Milne Edwards, 1830

FIGURE 42

Type-species: Lestrigonus jabrei Milne Edwards,

1830.

Small species with rather plump pereon. Head

globular; eyes occupying most of its surface. Some

of anterior pereonites fused, first 2 to first 5

in female, first 2 to first 4 in male, always more

in female. Coxae fused with pereonites. Telson of

moderate size. Antenna 1 of female 2-segmented.

Antenna 2 of female 1-segmented, usually very

small; gland cone conspicuous, with rounded or

pointed apex. Epistome prominent, strongly convex

anteriorly. Mandible with dentate incisor, with

palp in male, without palp in female. Maxilla 1

with outer lobe having 3 large terminal spines and

usually a smaller subterminal spine. Maxilliped

with outer lobes separate, tapering distally; inner

lobe well developed. Pereopod 1 subchelate or

barely chelate. Pereopod 2 distinctly chelate; carpal

process spoon-shaped with spines along margin of

spoon. Pereopods 5-7 usually longer than pereopods 3 and 4; pereopods 5 and 7 subequal, slightly

shorter than pereopod 6.

References: Vosseler, 1901:66; Yang, 1960:15;

Laval, 1968a:passim; Bowman, 1973:33.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 42.—Lestrigonus.

33. Parathemisto Boeck, 1870

FIGURE 43

Type-species: Parathemisto abyssorum Boeck,

1870.

Body rather compressed laterally. Pereonites all

free. Coxae not fused with pereonites. Antenna

1 with 3-segmented peduncle. Mandible with

palp in both sexes. Pereopod 1 simple. Pereopod 2

chelate, with spoon-shaped carpal process. Pereopods 3 and 4 with dilated carpus, forming prehensile organ with propus. Pereopods 5—7 longer than

pereopods 3 and 4.

References: Stephensen, 1924:94; Barnard, 1930:

419; Hurley, 1955:151; Bowman, 1960:passim;

Kane, 1966:passim.

FIGURE 43.—Parathemisto.

34. Pegohyperia Barnard, 1932

FIGURE 44

Type-species: Pegohyperia princeps Barnard,

1932.

Body rather elongate. Head short; eyes without

facets. Pereonites all free, with rounded transverse

folds. Coxae not fused with pereonites; coxae 3-5

much deeper than others. Telson nearly as long as

protopod of uropod 3. Antenna 1 of female with

large laminate flagellum. Mandible with palp in

both sexes; molar laminate, triangular. Pereopods
NUMBER 146

1 and 2 distinctly chelate; carpal process laterally

compressed, knife-shaped. Pereopods 5 and 6 some35

what longer than pereopods 3, 4, and 7. Protopods

of uropods broad.

References: Barnard, 1932:277; Hurley, 1960:112.

FIGURE 44.—Pegohyperia.

35. Phronimopsis Claus, 1879 Body slender in male; pereon somewhat inflated

FIGURE 45 in female. Head higher than long. Pereonites 1 and

Type-species: Phronimopsis spinifera Claus, 2 fused. Coxae fused with pereonites. Telson very

1879a. short. Antenna 1 of female with 1-segmented peFIGURE 45.—Phronimopsis.
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

duncle produced into long spine beneath flagellum.

Mandible with palp in male, without palp in

female. Pereopod 1 simple; dactyl covered with

short setae. Pereopod 2 chelate; dactyl closing

against anteroventral process of strongly swollen

propus. Pereopods 3-7 prehensile; dactyl closing

against expanded distal margin of propus. Pereopods 5-7 subequal, longer than pereopods 3 and

4. Uropods slender.

References: Claus, 1879a:63; Bovallius, 1889:318;

Vester, 1900:passim; Vosseler, 1901:51.

36. Themistella Bovallius, 1887

FIGURE 46

Type-species: Themistelln stcenstrupi Bovallius,

1887b.

Small species with rather broad pereon. Head

rather broad; eyes occupying most of its surface.

Pereonites 1-5 fused in both sexes. Coxae fused

with pereonites. Telson very short. Antenna 1 of

female 2-segmented, rather long. Antenna 2 of

female 1-segmented, rudimentary; gland cone small.

Mandible with serrate incisor; with palp in male,

without palp in female. Maxilliped, outer lobes

narrow, tapering distally; inner lobe almost completely absent, apparently represented by slight

unarmed bulge on basal segment. Pereopods with

rather broad segments. Pereopods 1 and 2 chelate;

carpal process gauge-shaped, with marginal spines.

Pereopod 5 much longer than pereopods 6 and 7.

Dactyls of pereopods 6 and 7 with flexure slightly

distal to midlength.

References: Bovallius, 1889:312; Bowman, 1973:

63.

P.6

FICURE 46.—Themistella.

XII. Family DAIRELLIDAE Bovallius, 1887

Body moderately slender, but pereon of female

somewhat expanded, transparent; total length 6-10

mm. Head large, globular in female, somewhat

smaller in male. Eyes large, occupying most of

head surface. Pereonites 1 and 2 fused dorsally.

Coxae fused with pereonites. Telson rounded, very

short. Antenna 1 with short 3-segmented peduncle;

flagellum of male long, filiform, multisegmented;

flagellum of female composed of a single cylindrical segment. Antenna 2 of male long, multisegmented; absent in female. Mandible without palp,

with molar. Maxilliped with inner and outer lobes

fused into a single subquadrate plate. Pereopods 1

and 2 slender, simple, only slightly shorter than

pereopods 3-7, which are subequal. Uropods with

free exopods and endopods. Gills on pereonites 2-6.

Oostegites on pereonites 2-5.

One genus.

37. Dairella Bovallius, 1887

FICURE 47

Type-species: Paraphronima californica Bovallius, 1885.

With the characters of the family.

The mouthparts of Dairella have been described

by Stebbing (1888) and Bovallius (1889). The small

size makes dissection of these mouthparts difficult,

and it is not surprising that both authors misinterpreted maxilla 2 and maxilliped. The maxilliped is

a single subquadrate plate with a slightly concave
NUMBER 146 37

distal margin bearing a few spines at the distolateral corners. Stebbing considered this plate to be

the base only of the maxilliped. The pyriform

maxilla 2 lies directly in front of and close to the

maxilliped. Both Bovallius and Stebbing believed

it to be the outer lobe of the maxilliped, but

careful dissection showed it to be completely

separate from the latter throughout its entire

length. The reason for Stebbing's failure to find

maxilla 2 is now obvious, but the nature of the

structure that Bovallius described as maxilla 2 is

not clear.

References: Stebbing, 1888:1342; Bovallius, 1889:

332.

FIGURE 47.—Dairetta.

XIII. Family PHRONIMIDAE Dana, 1853

Body slender, elongate; total length 5-40 mm.

Head subconical, widest dorsally, narrowing and

prolonged ventrally. Eyes composed of a larger

dorsal part and a smaller bulging ventrolateral

part. Pereonites all separate or pereonites 1 and 2

fused. Coxae fused with pereonites. Telson very

small. Antenna 1 in male long, multisegmented,

first segment of flagellum distinctly enlarged; antenna 1 of female very short, 2-segmented. Antenna

2 of male long, multisegmented, sometimes absent;

antenna 2 of female reduced to a tubercle. Mandible without palp in both sexes. Maxilliped with

slender outer lobes and short inner lobe. Pereopods

1 and 2 simple, subchelate, or chelate depending

on development of carpal process; propus produced

into thin triangular process on either side of

dactyl. Pereopods 3, 4, 6, and 7 simple, slender;

pereopod 7 not reduced. Pereopod 5 ending in subchelate claw. Uropods slender, with free exopods

and endopods. Gills on pereonites 4-fi. Oostegites

on pereonites 2-5.

Two genera.

Key to the Genera of Phronimidae

Pereonites 1 and 2 not fused. Carpus of pereopod 5 with smooth anterior margin and pointed

process at anterodistal angle 38. Phronima

Pereonites 1 and 2 fused. Carpus of pereopod 5 with dentate anterior margin, without clearly

defined anterodistal angle 39. Phronimella

38. Phronima Latreille, 1802

FIGURE 48

Type-species: Cancer sedentarius Forskal, 1775.

Body moderately slender. Pereonites all free.

Pereopod 5 with carpus markedly widened distally;

anterodistal angle forming strong pointed process;

anterior margin smooth. Uropod 2 present in both

sexes; endopod sometimes reduced but never missing.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 48.—Phronima.

References: Claus, 1879a:passim; Vosseler, 1901:

1; Stephensen, 1924:112; Shih and Dunbar, 1963:

passim; Shih, 1969:passim.

39. Phronimella Claus, 1871

FICURE 49

Type-species: Phronima elongata Claus, 1862.

Body and pereopods extremely slender. Pereonites 1 and 2 fused. Pereopod 5 with carpus elongate,

only slightly widened distally, without clearly defined anterodistal angle; anterior margin dentate.

Uropod 2 absent in female, rudimentary in male,

sometimes with small exopod.

References: Same as for Phronima.

FICURE 49.—Phronimella.

XIV. Family PHROSINIDAE Dana, 1853

Body rather compact; total length 4-30 mm.

Head large, globular; eyes occupying most of head

surface. Pereonites all separate or pereonites 1 and

2 fused. Coxae separate from pereonites. Telson

well developed, oval or triangular. Antenna 1 with
NUMBER 14fi 39

long filiform flagellum in male; with 1-segmented

flagellum in female, sometimes reduced. Antenna 2

similar to antenna 1 in male, rudimentary in female. Mandible with palp in male, without palp in

female. Maxilliped with slender outer lobes and

well-developed inner lobe.

Pereopods 1 and 2 simple; one or more of following pereopods prehensile, subchelate; pereopod

5 always the longest, with large subchela; pereopod

7 reduced in length or number of segments. Uropods composed each of a single leaflike segment

only. Gills on pereonites 2-6. Oostegites on pereonites 2-5.

Three genera.

Key to the Genera of Phrosinidae

1. Pereonites 1 and 2 separate. Pereopods 3 and 4 simple 42. Primno

Pereonites 1 and 2 fused. Pereopods 3 and 4 subchelate 2

2. Head produced into 2 rostral points. Pereopod 7 composed of broad basis and sometimes a

rudimentary second segment 40. Phrosina

Head rounded, without rostral points. Pereopod 7 with all segments present 41. Anchylomera

40. Phrosina Risso, 1822

FIGURE 50

Type-species: Phrosina semilitnata Risso, 1822.

Head produced into 2 rostral points. Pereonites 1

and 2 fused. Telson round-triangular, nearly as

wide as urosome. Antenna 1 of female very short.

Antenna 2 absent in female. Pereopods 3 and 4

subchelate, with sharp carpal process; distal margin

of carpus dentate, with larger teeth in pereopod 4.

Pereopods 5 and 6 subchelate; carpus relatively

short and broad, with dentate distal margin; propus

much longer than carpus. Pereopod 7 composed of

broad basis and sometimes a rudimentary second

segment. Uropods rounded, with smooth margins.

Gills without folds.

References: Stebbing, 1888:1424; Vosseler, 1901:

89.

FIGURE 50.—Phrosina.

41. Anchylomera H. Milne-Edwards, 1830

FIGURE 51

Type-species: Anchylomera blossevillii H\ MilneEdwards, 1830.

Head globular, without rostrum. Pereonites 1

and 2 fused. Telson round-triangular, nearly as

wide as urosome. Antenna 1 of female very short.

Antenna 2 of female reduced to a tubercle. Pereopods 3 and 4 subchelate, with sharp carpal process;

distal margin of carpus smooth. Pereopod 5 with

broad basis, having greatest width proximally; carpus with broad distal margin with rounded teeth.

Pereopod 6 prehensile, with swollen and rounded

carpus. Pereopod 7 with all segments present;

dactyl digitiform. Uropods rounded, with smooth

margins. Gills with folds.

References: Bovallius, 1889:408; Vosseler, 1901:
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 51.—Anchylomera.

42. Primno Guerin-Meneville, 1836

FICURE 52

Type-species: Primno macropa Guerin-Meneville,

1836.

Head produced into single, very short rostrum.

Pereonites 1 and 2 separate. Telson much narrower

than urosome. Antenna 1 of female longer than

head. Antenna 2 of female reduced to a tubercle.

Pereopods 3, 4, and 6 simple, with some teeth on

margin of carpus. Pereopod 5 prehensile, entire

anterior margin of carpus dentate; propus shorter

than carpus. Pereopod 7 with all segments present,

dactyl digitiform. Uropods pointed; uropods 2 and

3 with teeth on outer margins. Gills without folds.

References: Stebbing, 1888:1440; Bovallius, 1889:

397; Vosseler, 1901:87.

FIGURE 52.—Primno.
NUMBER 14« 41

LYCAEOPSOIDEA, new superfamily

Antenna 1 inserted on anterior surface of head;

flagellum of male with large triangular 1st segment

and a few small distal segments inserted subterminally; flagellum of female short, composed of a few

segments. Antenna 2 absent in female; in male

inserted on ventral surface of head, very short,

curved, composed of a few segments. Pereopod 5

never with large subchela. Pereopod 7 reduced in

size, but all segments present.

One family (Lycaeopsidae).

XV. Family LYCAEOPSIDAE Chevreux, 1913

Body slender, somewhat compressed laterally;

total length 3-6 mm. Head globular, higher than

long; eyes large, occupying most of head surface.

Coxae not fused with pereonites. Telson usually

rather narrow, not fused with double urosomite.

Mandible with palp in male, without palp in

female. Maxilliped with outer lobes pointed, only

slightly longer than inner lobe.

Pereopods 1 and 2 simple. Pereopod 5 long and

very slender in male, normal in female and young

male. Pereopod 6 long, with broad segments, especially in male. Uropods 1-3 having exopods and

endopods not fused with protopods. Uropod 3 with

endopod modified in male. Gills on pereonites 5

and 6, without folds. Oostegites on pereonites 2-5.

One genus.

43. Lycaeopsis Claus, 1879

FIGURE 53

Type-species: Lycaeopsis themistoides Claus,

1879b.

With the characters of the family.

References: Pirlot, 1930:27; 1939;42; Spandl,

1927:213.

Superfamily PLATYSCELOIDEA, new name

Antenna 1 inserted on ventral surface of head;

flagellum of male with large curved 1st segment

and a few small distal segments inserted terminally,

or more commonly subterminally, on 1st segment;

flagellum of female usually curved, composed of a

few segments. Antenna 2 inserted on ventral surface of head; in male composed of 5 usually long

segments (3 peduncular, 2 flagellar) that fold on

one another in zigzag fashion; in female short,

composed of a few segments, or absent. Pereopod

5 never with large subchela. Pereopod 7 reduced in

size and sometimes in number of segments.

Six families (Pronoidae, Anapronoidae, Lycaeidae, Oxycephalidae, Platyscelidae, Parascelidae).

References: Claus, 1887; Spandl, 1927.

FICURE 53.—Lycaeopsis.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Families of Platysceloidea

1. Body not conglobate. Pereopods 5 and 6 with basis somewhat broadened, but never operculate. Pereopod 6 with distal segments inserted on margin of basis or on medial surface a

short distance back from margin 2

Body conglobate. Pereopods 5 and 6 with basis transformed into broad operculum. Pereopod

6 with distal segments inserted on medial surface far back from margin 6

2. Body elongate; head produced into distinct rostrum anterior to eyes

XIX. OXYCEPHAUDAE (except Simorhyncholus)

Body rather compact; head globular 3

3. Pereopod 7 consisting only of basis and 1 or 2 rudimentary segments. Mandible with palp

in both sexes XVI. PRONOIDAF.

Pereopod 7 having all segments present. Mandible without palp in female 4

4. Antenna 2 of female short, composed of a few segments. Mandible without palp in male

XVII. ANAPRONOIDAE

Antenna 2 of female rudimentary or absent. Mandible with palp in male 5

5. Maxillae 1 and 2 absent. Uropod 2 with endopod fused with protopod XVIII. LYCAF.IDAK

Maxillae 1 and 2 reduced but present. Uropod 2 with endopod not fused with protopod

XIX. OXYCEPHALIDAE (Simorhynchotus)

6. Mouthparts in form of a broad short cylinder. Pereopod 7 consisting only of basis and sometimes 1 or 2 rudimentary additional segments XX. PLATYSCELIDAE

Mouthparts pointed, in form of a cone. Pereopod 7 with all segments present

XXI. PARASCELIDAE

XVI. Family PRONOIDAE Claus, 1879

Body compact, more or less laterally compressed;

total length 3-16 mm. Head globular; eyes large,

occupying most of head surface. Coxae not fused

with pereonites. Telson triangular, separate from

or very rarely fused with double urosomite. Antenna 2 of female short, with few segments. Mandible

with palp in both sexes. Maxillae 1 and 2 well developed. Pereopod 1 simple, sometimes subchelate.

Pereopod 2 simple or chelate. Pereopod 5 longer

than pereopod 6; basis somewhat broadened. Pereopod 6 having basis much broader than that of

pereopod 5, but not operculate; distal segments

inserted subterminally. Pereopod 7 with welldeveloped basis followed by 1 or 2 rudimentary

segments; all segments may be present in juveniles.

Uropods usually with free rami; endopods of uropods 2 and 3 rarely fused with protopods. Gills on

pereonites 2—6, with or without folds. Oostegites on

pereonites 2-5.

Five genera.

References: Claus, 1887:47-55; Spandl, 1927:216-

227.

Key to the Genera of Pronoidae

1. Pereopod 2 simple 2

Pereopod 2 chelate 3

2. Pereopod 1 with basis nearly as broad as long. Uropod 3 much longer than telson 44. Pronoe

Pereopod 1 with basis narrow. Uropod 3 only slightly longer than telson AQ.Paralycaea

3. Double urosomite distinctly longer than wide 4

Double urosomite about as long as wide 45. Eupronoe

4. Pereopod 2 with carpal process pointed, nearly as long as propus. Uropod 3 slightly longer

than telson 47. Parapronoe

Pereopod 2 with carpal process rounded, about half as long as propus. Uropod 3 much

longer than telson 48. Sympronoe

44. Pronoe Guerin-Meneville, 1836

FIGURE 54

Type-species: Pronoe capilo Guerin-Meneville,

1836.

Body rather slender; double urosomite about as

wide as long. Telson very short, not fused with

double urosomite. Antenna 2 of male short, folded

only once or twice. Pereopod 1 simple; basis oval,
NUMBER 146 43

almost as broad as long. Pereopod 2 simple. Pereopod 5 distinctly longer than pereopod 6. Pereopod

7 composed of basis and very small second segment.

Uropod 1 with exopod and endopod pointed. Uropod 2 with exopod pointed, endopod rounded.

Uropod 3 much longer than telson. Gills without

folds.

References: Claus, 1887:48; Stebbing, 1888:1507.

FIGURE 54.—Pronoe.

45. Eupronoe Claus, 1879

FIGURE 55

Type-species: Eupronoe maculata Claus, 1879b.

Body rather stout; anterior pereonites short, appearing as though compressed together longitudinally; double urosomite about as wide as long. Telson moderately long, not fused with double urosomite. Antenna 2 of male long, with very short terminal segment. Pereopod 1 simple or subchelate;

basis twisted in male. Pereopod 2 chelate; carpal

process pointed or rounded. Pereopod 5 distinctly

longer than pereopod 6. Pereopod 7 with pearshaped basis and small elliptical second segment.

Uropod 1 with exopod and endopod pointed. Uropods 2 and 3 with exopod and endopods rounded,

very delicate; uropod 3 much longer than telson.

Gills with folds.

References: Claus, 1887:50; Spandl, 1927:222;

Hurley, 1955:175.

FIGURE 55.—Eupronoe.

46. Paralycaea Claus, 1879

FIGURE 56

Type-species: Paralycaea gracilis Claus, 1879b.

Body rather stout; double urosomite somewhat

broader than long; telson long, fused with or separate from double urosomite. Antenna 2 of male

with long terminal segment. Pereopods 1 and 2

simple. Pereopod 5 distinctly longer than pereopod

6. Pereopod 7 with narrow basis; distal segments
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sometimes separate, usually fused into short recurved segment. Uropod 2 having endopod fused

with or separate from protopod. Uropod 3 having

endopod fused with protopod. Uropods 1 and 2

having exopods and endopods pointed. Uropod 3

slightly longer than telson; exopod pointed; endopod obtusely rounded. Gills without folds.

References: Glaus, 1887:63; Pirlot, 1930:30.

FIGURE 56.—Paralycaea.

47. Parapronoe Claus, 1879

FICURE 57

Type-species: Parapronoe crustulum Claus, 1879b.

Body rather slender; double urosomite distinctly

longer than wide; telson long, not fused with double urosomite. Antenna 2 of male moderately long,

with very short terminal segment. Pereopod 1 simple. Pereopod 2 chelate; carpal process pointed,

about as long as propus. Pereopod 5 much longer

than pereopod 6; basis of pereopod 6 with concave

distal margin. Pereopod 7 with broad basis and

very small additional segments. Uropods with

pointed endopods and exopods; uropod 3 only

slightly longer than telson. Gills with folds.

References: Claus, 1887:53; Stebbing, 1888:1521;

Barnard, 1930:427; Pirlot, 1939:52.

FIGURE 57.—Parapronoe.

48. Sympronoe Stebbing, 1888

FIGURE 38

Type-species: Parapronoe parva Claus, 1879b.

Body rather slender; double urosomite about

twice as long as wide; telson very short, not fused

with double urosomite. Antenna 2 of male long,

with very short terminal segment. Pereopod 1 simple. Pereopod 2 chelate; carpal process rounded,

about half as long as propus. Pereopod 5 much

longer than pereopod 6; basis of pereopod 6 with

rather narrow distal margin produced into broadly

rounded anterior lobe and narrow, pointed posterior process. Pereopod 7 with pear-shaped basis and
NUMBER 146 45

2 very small additional segments. Uropods 1 and 2

wtih pointed endopods and exopods; uropod 3

much longer than telson, endopod rounded, exopod

bluntly pointed. Gills with folds.

References: Stebbing, 1888:1533; Spandl, 1927:

225.

FIGURE 58.—Sympronoe.

XVII. ANAPRONOIDAE, new family

Body rather plump, not laterally compressed.

Head globular; eyes large, occupying most of head

surface. Double urosomite shorter than wide. Telson triangular, not fused with double urosomite.

Antenna 2 of female short, with few segments.

Mandible without palp in both sexes. Maxillae 1

and 2 moderately well developed. Pereopod 1 subchelate; pereopod 2 chelate. Pereopod 5 longer

than pereopod 6; basis of both broad, but not operculate. Pereopod 6 with distal segments inserted on

medial surface of basis proximal to its distal end,

their combined length subequal to that of basis.

Pereopod 7 with all segments present.

One genus.

49. Anapronoe Stephensen, 1925

FIGURE 59

Type-species: Anapronoe reinhardti Stephensen,

1925.

With the characters of the family.

References: Stephensen, 1925:163; Spandl, 1927:

217.

FIGURE 59.—Anapronoe.
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

XVIII. Family LYCAEIDAE Claus, 1879

Body rather compact, laterally compressed or

somewhat flattened dorsoventrally; total length 3-

22 mm. Head globular or somewhat flattened dorsoventrally; eyes large, occupying most of head

surface. Coxae not fused with pereonites. Telson

triangular, separate from or fused with double

urosomite. Antenna 2 rudimentary or absent in

female. Mandible with palp in male, without palp

in female (except 2-segmented palp in female Tryphana). Maxillae 1 and 2 reduced but present.

Pereopods 1 and 2 simple or subchelate. Pereopod

5 slightly or distinctly longer than pereopod 6;

basis of both somewhat broadened but not operculate. Pereopod 6 with distal segments inserted on

margin of basis or on medial surface proximal to

margin. Pereopod 7 with all segments present.

Uropod 3 having endopod sometimes fused with

protopod. Gills on pereonites 2-6, with folds.

Oostegites on pereonites 2-5.

Five genera.

References: Claus, 1887:55-65; Chevreux and

Fage, 1925:426.

Key to the Genera of Lycaeidae

1. Pereopods 1 and 2 simple 2

Pereopods 1 and 2 subchelate 3

2. Pereopod 1 with basis nearly as broad as long. Uropod 3 much longer than telson; endopod

not fused with protopod 54. Tryphana

Pereopod 1 with basis about half as broad as long. Uropod 3 slightly longer than telson;

endopod fused with protopod 52. Pseudolycaea

3. Pereopods 1 and 2 with margins of carpus and propus smooth or finely serrulate. Uropod 1

with protopod much longer than endopod 50. Lycaea

Pereopods 1 and 2 with margins of carpus and propus dentate. Uropod 1 with protopod and

endopod subequal in length 4

4. Body somewhat flattened dorsoventrally. Distal segments of pereopods covered with

numerous setae 53. Thamneus

Body laterally compressed. Distal segments of pereopods without setae 51. Brachyscelus

50. Lycaea Dana, 1852

FIGURE 60

Type-species: Lycaea ochracea Dana, 1853.

Body rather plump, especially in female. Telson

fused with double urosomite. Antenna 2 of male

with very short terminal segment. Pereopods 1 and

2 subchelate, margins of carpus and propus smooth

or finely serrulate; basis of pereopod 1 not markedly broadened. Pereopod 5 distinctly longer than

pereopod 6. Uropod 1 with protopod much longer

than endopod. Uropod 3 slightly longer than telson; endopod not fused with protopod.

References: Claus, 1887:61; Barnard, 1930:428;

Shoemaker, 1945:243.

FIGURE 60.—Lycaea.
NUMBER 146 47

51. Brachyscelus Bate, 1861

FIGURE 61

Type-species: Brachyscelus cruscuhim Bate, 1861.

Body compact, moderately slender. Telson not

fused with double urosomite. Antenna 2 of male

with short terminal segment. Pereopods 1 and 2

subchelate, margins of carpus and propus dentate;

basis of pereopod 1 not markedly broadened. Pereopod 5 slightly longer than pereopod 6. Uropod 1

with protopod and endopod subequal in length.

Uropod 3 slightly longer than telson; endopod not

fused with protopod.

References: Claus, 1887:56; Stephensen, 1925:172.

FIGURE 61.—Brachyscelus.

52. Pseudolycaea Claus, 1879

FIGURE 62

Type-species: Pseudolycaea pachypoda Claus,

1879b.

Body moderately plump. Telson fused with double urosomite. Antenna 2 of male with very short

terminal segment. Pereopods 1 and 2 simple; basis

of pereopod 1 not markedly broadened. Pereopods

5 and 6 subequal in length. Uropod 1 with protopod much longer than endopod. Uropod 3 slightly

longer than telson; endopod fused with protopod.

References: Claus, 1887:64; Chevreux and Fage,

1925:430; Spandl, 1927:215.

FIGURE 62.—Pseudolycaea.
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

53. Thamneus Bovallius, 1887

FIGURE 63

Type-species: Thamneus rostratus Bovallius,

1887b.

Body moderately plump, flattened dorsoventrally.

Head somewhat flattened, with short, broad, upturned rostrum in male. Telson not fused with

double urosomite. Antenna 2 of male much shorter

than head, 4-segmented. Pereopods 1-6 covered

with numerous setae on distal segments. Pereopods

1 and 2 subchelate, margins of carpus and propus

dentate; basis of pereopod 1 not markedly broadened. Pereopods 5 and 6 subequal in length. Uropod 1 with protopod and endopod subequal in

length. Uropod 3 distinctly longer than telson;

endopod not fused with protopod.

Reference: Stebbing, 1888:1558.

FIGURE 63.—Thamneus.

54. Tryphana Boeck, 1870

FIGURE 64

Type-species: Tryphana malmi Boeck, 1870.

Body robust. Telson not fused with double urosomite. Antenna 2 of male with long terminal segment. Mandible of female with 2-segmented palp.

Pereopods 1 and 2 simple; basis of pereopod 1 oval,

nearly as broad as long; pereopod 2 with slender

tlistal segments, propus and dactyl covered with

short setae. Pereopods 3-6 stout, with prominent

glands in carpus, distally prehensile with dactyl

closing against produced distal margin of propus.

Pereopod 5 distinctly longer than pereopod 6. Uropod 1 with protopod and endopod subequal in

length. Uropod 3 much longer than telson; endopod not fused with protopod.

Reference: Sars. 1890:16.

FIGURE 64.—Tryphana.

XIX. Family OXYCEPHALIDAE Bate, 1861

Body form variable, sometimes compact, usually

elongate, sometimes extremely long and slender;

total length 3-120 mm. Head sometimes globular,

usually produced anteriorly into long rostrum,

sometimes with distinct neck; eyes large, occupying
NUMBER 146 49

most of head surface except rostrum and neck.

Coxae usually separate from pereonites, sometimes

fused. Telson fused with or separate from double

urosomite. Antenna 2 absent in female. Mandible

with palp in male, without palp in female. Maxillae 1 and 2 absent. Pereopod 1 rarely simple,

usually subchelate or chelate. Pereopod 2 subchelate or chelate. Pereopod 5 subequal to or distinctly

longer than pereopod 6; basis of both rarely slender, usually broadened but not operculate. Pereopod 6 with distal segments inserted terminally or

rarely subterminally on basis, their combined

length greater than that of basis. Pereopod 7 usually with all segments present; distal segments

sometimes reduced in number or absent. Uropods

2 and 3 having endopods sometimes fused with

protopods. Gills on pereonites 2-6, sometimes rudimentary or absent on one or more of pereonites

2-4. Oostegites on pereonites 2-5, sometimes on

pereonites 3-5, sometimes reduced in size.

Nine genera.

References: Bovallius, 1890; Fage, 1960; Pillai,

1966a.

Key to the Genera of Oxycephalidae

1. Double urosomite 3 or more times as long as wide 2

Double urosomite less than 3 times as long as wide 4

2. Body extremely slender. Pereopod 7 reduced to basis and sometimes one or more additional

rudimentary segments 60. Rabdosoma

Body only moderately slender. Pereopod 7 with all segments present 3

3. Rostrum with broad lateral flanges. Uropods 1 and 3 with well-developed exopods and

endopods 56. Calamorhynchus

Rostrum without lateral flanges. Uropod 1 with endopod very small; uropod 3 with

exopod very small 59. Leptocotis

4. Rostrum rounded or absent 5

Rostrum pointed 6

5. Body very slender. Pereopods 1 and 2 chelate; pereopods 5 and 6 paddle-like

58. Glossocephalus

Body compact. Pereopod 1 simple; pereopod 2 subchelate; pereopods 5 and 6 not paddle-like

61. Simorhynchotus

6. Uropods 2 and 3 having endopods not fused with prolopods 7

Uropods 2 and 3 or uropod 3 having endopods fused with protopod 8

7. Pcreonite 5 produced into backward-pointed processes on either side. Pereopods 5-7 having

basis with row of 3 or 4 pitlike glands 57. Cranocephalus

Perconitc ") without processes. Pereopods 5-7 having basis without pitlike glands 62. Streetsia

8. Pereopods 1 and 2 subchelate. Pereopod 6 with distal posterior corner of basis produced into

triangular process. Endopod fused with protopod in uropod 3, not fused in uropods

1 and 2 63. Tullbergella

Pereopods 1 and 2 chelate. Pereopod 6 having basis without process. Endopod fused with

protopod in uropods 2 and 3 55. Oxycephalus

55. Oxycephalus H. Milne-Edwards, 1830

FIGURE 65

Type-species: Oxycephalus piscatoris H. MilneEdwards, 1830.

Body elongate. Head somewhat shorter than

pereon, produced into long, pointed rostrum. Coxae

fused with pereonites. Double urosomite longer

than wide, sometimes twice as long, fused with

telson. Pereopods 1 and 2 chelate. Pereopod 7 with

all segments present. Uropods 2 and 3 having endopods fused with protopods. Uropod 3 slightly

shorter, equal to, or slightly longer than telson.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 65.—Oxycephalus.

56. Calamorhynchus Streets, 1878

FIGURE 66

Type-species: Calamorhynchus pellucidus Streets,

1878.

Body elongate. Head longer than pereon, produced into long, pointed rostrum expanded into

broad lateral flanges. Coxae fused with pereonites.

Double urosomite more than 3 times as long as

wide, fused with telson. Pereopods 1 and 2 chelate.

Pereopod 7 with all segments present. Uropod 3

shorter than telson; endopod fused with protopod.

FIGURE 66.—Calamorhynchus.
NUMBER 146 51

57. Cranocephalus Bovallius, 1890

FIGURE 67

Type-species: Cranocephalus goesi Bovallius,

1890.

Body rather compact, only moderately elongate.

Head somewhat shorter than pereon, produced into

pointed rostrum. Coxae fused with pereonites but

demarcated by indentations. Pereonite 5 produced

into backward-pointed processes on either side.

Double urosomite somewhat longer than wide,

fused with telson. Pereopods 1 and 2 subchelate;

pereopod 2 sometimes almost chelate. Pereopods

5-7 having broad basis, with row of 3 or 4 pitlike

glands. Pereopod 7 with all segments present. Uropod 3 subequal to or shorter than telson.

FIGURE 67.—Cranocephalus.

58. Glossocephalus Bovallius, 1887

FIGURE 68

Type-species: Glossocephalus milne-edwardsi Bovallius, 1887b.

Body very slender, elongate. Head much shorter

than pereon, produced into short rounded rostrum.

Coxae separate on pereonites 1-6, fused with pereonite 7. Double urosomite about as long as wide,

fused with telson. Pereopods 1 and 2 chelate. PereUp.3

FIGURE 68.—Glossocephalus.
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

opods 3 and 4 very slender. Pereopods 5 and 6

paddle-like, with broad segments; dactyl very small.

Pereopod 7 with all segments present. Uropod 3

somewhat longer than telson. Gills on pereonite 2

absent in male, rudimentary in female; on pereonite 3 rudimentary in both sexes. Oostegites on

pereonites 3-5.

59. Leptocotis Streets, 1877

FIGURE 69

Type-species: Leptocotis spinifera Streets, 1877.

Body elongate. Head longer than pereon, produced into long, pointed rostrum. Coxae not fused

with pereonites. Double urosomite more than 3

times as long as wide, fused with telson. Pereopods

1 and 2 chelate. Pereopod 7 with all segments present. Uropod 1 having endopod very small. Uropod

2 having endopod fused with protopod in female,

separate in male. Uropod 3 shorter than telson;

endopod fused with protopod; exopod very small.

FIGURE 69.—Leptocotis.

60. Rhabdosoma White, 1847

FIGURE 70

Type-species: Oxycephalus armatus H. Milne-Edwards, 1840.

Body extremely slender and elongate. Head much

f

longer than pereon, with long neck and very long,

needle-shaped rostrum. Coxae fused with pereonites. Double urosomite more than 3 times as long

as wide, not fused with telson. Pereopods 1 and 2

very short, chelate. Pereopods 3-6 very slender. Pereopod 7 very small, usually consisting only of basis,

FICURE 70.—Rhabdosoma.
NUMBER 14€

sometimes with one or more additional rudimentary segments. Uropods very slender; uropods 2 and

3 having endopod fused with protopod, exopod

very short. Gills on pereonites 2-6 or 5 and 6 in

female, on pereonites 4-6 or 5 and 6 in male. Oostegites reduced in size, on pereonites 2-6.

61. Simorhynchotus Stebbing, 1888

FIGURE 71

Type-species: Simorhynchus antennarius Claus,

1871.

Body compact. Head much shorter than pereon,

globular, with short, rounded rostrum in male.

Coxae separate. Double urosomite slightly shorter

than wide, fused with telson. Pereopod 1 simple.

Pereopod 2 subchelate. Pereopod 7 with all segments present. Uropods 2 and 3 having endopod

fused with protopod. Uropod 3 slightly longer than

telson.

FIGURE 71 Simorhynchotus.

62. Streetsia Stebbing, 1888

FIGURE 72

Type-species: Streetsia challengeri Stebbing, 1888.

Body elongate. Head variable in length, from

somewhat shorter to distinctly longer than pereon,

produced into pointed rostrum. Coxae not fused

with pereonites. Double urosomite from slightly

longer to 2.5 times as long as wide, fused with telson. Pereopod 1 subchelate; pereopod 2 subchelate

or chelate. Pereopod 7 with all segments present.

Uropod 3 shorter than telson.

FICURE 72.—Streetsia.

63. Tullbergella Bovallius, 1887

FICURE 73

Type-species: Tullbergella cuspidata Bovallius,

1887b.

Body rather compact, only moderately elongate.

Head much shorter than pereon, produced into

short, pointed rostrum. Coxae not fused with pereonites. Pleonite 3 produced into long, backwardpointing spines on either side. Double urosomite
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

slightly shorter than wide, fused with telson. Pereopods 1 and 2 subchelate. Pereopod 6 with distal

posterior corner of basis produced into triangular

process. Pereopod 7 composed of basis and 3 small

distal segments. Uropod 3 as long as telson; endopod fused with protopod.

FK;URE 73.—Tullbergella.

XX. Family PLATYSCELIDAE Bate, 1862

FIGURE 74

Body wide, stout, conglobate, usually somewhat

depressed dorsoventrally. Pleon distinctly narrower

than pereon and capable of folding under pereon.

Total length 2-24 mm. Head short, wider than

pereon, ventral margin (viewed from front) produced into median round-triangular process. Eyes

large, occupying most of head surface. Coxae not

fused with pereonites. Telson broadly triangular,

fused with double urosomite. Antenna 2 of female

short, of few segments. Mouthparts in the form of

a broad, short cylinder. Mandible short, with broad

incisor; with palp in male, wtihout palp in female.

Maxillae 1 and 2 with large broad lobes.

Pereopods 1 and 2 simple, subchelate, or chelate.

Pereopods 5 and 6 with bases transformed into

broad opercula, always longer in pereopod 6; distal

segments inserted subterminally in basis of pereopod 5, much farther proximally in pereopod 6;

posterior margin of basis of pereopod 6 with longitudinal groove (telsonic groove) into which telson

and uropods fit during conglobation; outer surface

of basis of pereopod 6 often with fissure. Pereopod

7 with narrow, curved basis and sometimes one or

more rudimentary additional segments. Uropods 2

and 3 having endopods sometimes fused with protopods. Gills on pereonites 2-6. Oostegites on pereonites 2-5.

Five genera.

telsonic

groove

ventral margin

FIGURE 74.—Platyscelidae.
NUMBER 146 55

1. Pereoj

Peieoj

fissi

Pereor.

Key to the Genera of Platyscelidae

ods 1 and 2 having dactyl forming subchela with concave distal margin of propus

ods 1 and 2 without subchela between dactyl and propus

2. Pereopods 1 and 2 with pointed carpal process. Pereopod G having basis with semicircular

65. Amphithyrus

ods 1 and 2 without carpal process. Pereopod 6 having basis without fissure

67. Tetrathyrus

3. Pereopod 1 simple. Pereopod 2 with short carpal process 68. Paratyphis

Pereopods 1 and 2 chelate. Pereopod 2 with carpal process about as long as propus 4

1. Antenna 2 of male with two distal segments much shorter than half the length of preceding segment 64. Platyscelus

Antenna 2 of male with two distal segments more than half the length of preceding segment

66. Hemityphis

64. Platyscelus Bate, 1862

FICURE 75

Type-species: Typhis ovoides Risso, 1816.

Antenna 2 of male having 2 distal segments much

shorter than half the length of preceding segment

(third segment from end). Pereopods 1 and 2 chelate, with pointed carpal process. Pereopod 6 having basis with nearly straight fissure.

References: Claus, 1887:31; Stebbing, 1888:1480;

Spandl, 1927:240; Shoemaker, 1945:255.

FIGURE 75 Platyscelus.

65. Amphithyrus Claus, 1879

FICURE 76

Type-species: Amphithyrus bispinosus Claus,

1879b (by present designation).

Cuticle with polygonal pattern of surface sculpturing. Antenna 2 of male with two distal segments

subequal in length to preceding segment. Pereopods 1 and 2 complexly chelate; carpus with pointed process forming chela; propus with concave

distal margin forming subchela. Pereopod 6 having basis relatively short and wide, with semicircular fissure.

References: Claus, 1887:41; Pirlot, 1930:43;

Spandl, 1927:246.

66. Hemityphis Claus, 1879

FIGURE 77

Type-species: Hemityphis tennimnnus Claus,

1879b.

Antenna 2 of male having two distal segments

more than half the length of preceding segment.

Pereopods 1 and 2 chelate, with pointed carpal

process. Pereopod 6 having basis with short, curved

fissure.

References: Claus, 1887:38; Stebbing, 1888:1471;

Spandl, 1927:233; Pirlot, 1930:37; Shoemaker, 1945:

259.

67. Paratyphis Claus, 1879

FIGURE 78

Type-species: Paratyphis maculatus Claus, 1879b.

Antenna 2 of male having 2 distal segments

slightly less than half or somewhat more than half
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 76.—Amphithyrus.

FIGURE 77.—Hemtiyphis.

the length of preceding segment. Pereopod 1 simple.

Pereopod 2 weakly chelate, with short carpal process. Pereopod f> having basis with nearly straight

fissure.

FIGURE 78.—Paratyphis.

References: Claus, 1887:39; Stebbing, 1888:1476;

Spandl, 1927:213; Pirlot, 1930:39.

68. Tetrathyrus Claus, 1879

FICURE 79

Type-species: Tetrathyrus forcipatus Claus,

1879b.

Antenna 2 of male with two distal segments more

than half the length of preceding segment. Pereopods 1 and 2 subchelate; dactyl closing against concave distal margin of propus. Pereopod 6 having

basis with long ventral margin, without fissure.

References: Claus, 1887:40; Stebbing, 1888:1180;

Spandl, 1927:240; Barnard, 1930:439.
NUMBER 146 57

FIGURE 79.—Tetrathyrus.

XXI. Family PARASCELIDAE Bovallius, 1887

Body wide, stout, conglobate, usually somewhat

depressed dorsoventrally. Pleon distinctly narrower

than pereon and capable of folding under pereon.

Total length 2.5-7 mm. Head short, wider than

pereon, ventral margin (viewed from front) broadly rounded. Eyes large, occupying most of head

surface. Coxae not fused with pereonites. Telson

broadly triangular, fused with double urosomite.

Antenna 2 of female short, of few segments. Mouthparts pointed, in the form of a cone. Mandible

elongate, with narrow incisor; with palp in male,

without palp in female. Pereopods 1 and 2 simple or

chelate. Pereopods 5 and 6 with basis transformed

into broad operculum, always longer in pereopod 6; distal segments inserted subterminally on

basis of pereopod 5, much farther proximally in

pereopod 6; posterior margin of pereopod 6 with

longitudinal groove (telsonic groove) into which

telson and uropods fit during conglobation; outer

surface of pereopod 6 with or without fissure. Pereopod 7 with all segments present. Uropods 2 and 3

having endopods sometimes fused with protopods;

exopods much shorter than endopods. Gills on pereonites 2-6. Oostegites on pereonites 2-5.

Three genera.

References: Claus, 1887:42; Stebbing, 1888:1491;

Spandl, 1927:250.

Key to the Genera of Parascelidae

1. Fcreopods 1 and 2 simple 69. Thyropus

Pereopod 2 chelate 2

2. Pereopod 1 simple. Pereopod 6, basis with long fissure 71. Schizoscelus

Pereopod 1 chelate. Pereopod 6, basis without fissure 70. Euscelus

69. Thyropus Dana, 1852

FIGURE 80

Synonym: Parascelus Claus, 1879.

Type-species: Thyropus diaphanus Dana, 1852.

Pereopods 1 and 2 simple. Pereopod 6 having

basis very long, narrowed in distal half, with or

without short fissure. Uropods 1-3 having exopods

and endopods not fused with protopod.

References: Claus, 1887:45; Stebbing, 1888:1492;

Stephensen, 1925:208; Spandl, 1927:258; Shoemaker, 1945:260.

70. Euscelus Claus, 1879

FIGURE 81

Type-species: Euscelus robustus Claus, 1879b.

Pereopods 1 and 2 chelate; carpal process narrow,

curved, with truncate and dentate distal margin.

Pereopod 6 having basis without fissure. Uropods

1-3 with exopods oval, much shorter than endopods; exopods and endopods not fused with protopods.

References: Claus, 1887:43; Spandl, 1927:251.
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 81 Euscelus.

71. Schizoscelus Claus, 1879

FIGURE 82

Type-species: Schizoscelus ornatus Claus, 1879b.

Pereopod 1 simple. Pereopod 2 chelate, with

pointed carpal process. Pereopod 6 having basis

subrectangular, with long, slightly curved fissure.

Uropods 2 and 3 having endopods fused with protopods.

References: Claus, 1887:43; Stebbing, 1888:1503;

Spandl, 1927:225.

FIGURE 82.-rSchizoscelus.

Sources of Illustrations

Figure 1: Original.

Figure 2: Original.

Figure 3: Md, original. Md', Pirlot, 1930, fig. 3. Mx.l,

Mx.2, original. Mxp, Stephensen and Pirlot, 1931, fig. 17.

Mxp', original.

Figure 4: Original.

Figure 5: Original.

Figure 6: Original.

Figure 7: Original.

Figure 8: Shoemaker, 1945, fig. 22j.

Figure 9: A, original, B, Fage, 1960, fig. 8.

Figure 10: $, Woltereck, 1909, pi. 5, fig. 13; $, fig. 12. A.I,

A.2, Md, P.I, P.2, Vinogradov, 1956, fig. 2. Us, Stephensen

and Pirlot, 1931, fig. 13.

Figure 11: £, Woltereck, 1909, pi. 3, fig. 8. $, Stephensen

and Pirlot, 1931, fig. 7; Md, Mxp, fig. 8; P.I, P.2, P.7, Us,

fig. 9.

Figure 12: Whole animal, Vinogradov, 1960a, fig. 12; Md,

P.7, fig. 13.

Figure 13: A.I, Vinogradov, 1956, fig. 4. A.2, Md, P.I, P.2,

Stephensen and Pirlot, 1931, fig. 17; Us, fig. 18.

Figure 14: 9, Pirlot, 1933, fig. 1. A.I, A.2, Vinogradov, 1960a,

fig. 1 I. Md, Mxp, Pirlot, 1933, fig. 2. P.I, P.2, P.7, Vinogradov, 1960a, fig. 14. Us, Pirlot, 1933, fig. 3.
NUMBER 146 59

Figure 15: $ , Chevreux and Fage, 1925, fig. 387. A.2, P.I, P.5,

Scina borealis, off California, original. Md, Mxp, Scina

curvidactyla, off Washington, original. Us, Scina tullbergi,

off California, original.

Figure 16: $, Vosseler, 1901, pi. 10, fig. 1. A.I, Wagler, 1926,

fig. 51b; Mxp, fig. 52d; P.I, P.5, fig. 52a.

Figure 17: $, Wagler, 1926, fig. 53; H (ventral), fig. 54; Mxp,

fig. 56d; P.I, P.5, fig. 56a. P.5 (dactyl, enlarged), original.

Figure 18: $, Mxp, Vinogradov, 1957, fig. 18. P.I, P.5, Chevreux, 1914, fig. 4.

Figure 19: Whole animal, Woltereck, 1909, pi. 4, fig. 11. A.I,

A.2, P.I, P.3, P.5, P.6, Us, Stephensen and Pirlot, 1931, fig.

16.

Figure 20: Whole animal, Vinogradov, 1960a, fig. 9; A.I, A.2,

Md, Mxp, Us, fig. 10; P.I, P.3, P.5, P.6, fig. 11.

Figure 21: Whole animal, Woltereck, 1909, fig. 10. Head, P.I,

P.2, Shoemaker, 1945, fig. 29; P.5, P.7 (left), fig. 30. P.5,

P.7 (right) , Vinogradov, 1964, fig. 10.

Figure 22: $, Pirlot, 1930, fig. 1; A.I, A.2, Md, Mxp, fig. 3;

P.I, P.2, P.3, P.7, fig. 4; Us, fig. 5. H (dorsal), Vinogradov,

1960b, fig. 2.

Figure 23: $, Stephensen, 1918, fig. 4. H, Md, Mxp, Shoemaker, 1945, fig. 15; Mx.l, fig. 20; P.I, P.2, fig. 19. Md

incisor and lacinia, Lanceola sayana, off New Jersey,

original.

Figure 24: 9, original. A.I, A.2, Pirlot, 1935, fig. 3; Md, Mx.l,

fig. 1. P.I, P.2, Shoemaker, 1945, fig. 21.

Figure 25: $, Pirlot, 1931, fig. 1; Md, Mxp, P.3, fig. 2. A.I,

A.2, Mx.l, P.I, P.2, Vinogradov, 1960a, fig. 6.

Figure 26: Barnard, 1930, fig. 52.

Figure 27: Whole animal, Woltereck, 1909, fig. 14. A.I, A.2,

Md, Mx.l, Vinogradov, 1957, fig. 7; P.I, P.2, P.5, fig. 8.

Figure 28: $, Woltereck, 1909, fig. 21. H, original. Md, Mx.l,

Mxp, P.I, P.2, Shoemaker, 1945, fig. 22.

Figure 29: $, Vosseler, 1901, pi. 11, fig. 6. Md, Mxp, Us,

Vibilia viatrix, off California, original. P.7, V. cultripes,

off California, original.

Figure 30: $, Hurley, 1955, fig. 23. P.7, Us, Cyllopus magellanicus, off New Zealand, original.

Figure 31: Original.

Figure 32: $, Spandl, 1927, fig. 8. Md, Mxp, P.I, P.2, original. Us, Stephensen, 1918, fig. 21.

Figure 33: $, Vosseler, 1901, pi. 8, fig. 22. Md, Mxp, P.I,

P.2, Us, Paraphronima gracilis, off California, original.

Figure 34: $, A.I, A.2, Bowman, 1973, fig. 18a-c; Md, Us,

fig. 2h,o; Mxp, fig. 15e,f; P.I, P.2, fig. 4a,b.

Figure 35: Laval, 1966.

Figure 36: $, Bowman, 1973, fig. 22a; Mxp, fig. 20i; P.I, P.2,

fig. 23a,b; Us, fig. 21a.

Figure 37: 9, Vosseler, 1901 pi. 5, fig. 16. A.I, A.2, Mxp, P.3,

Bowman, 1973, fig. 46d,e,i,j,m; Md, P.I, P.2, fig. 43d,i,j;

P.6, fig. 41o; Us, fig. 41 q.

Figure 38: $, Vosseler, 1901, pi. 7, fig. 6. Md, original. Mxp,

Bowman, 1973, fig. 24j and original. P.I, P.2, Bowman,

1973, fig. 24k,l. Us, original.

Figure 39: 9, P.I, P.2, P.5, Yang, 1960, fig. 9. Mx.l, Mxp,

P.5 dactyl, Bowman, 1973, fig. 52b,d-f. Us, Stephensen, 1924,

fig. 35.

Figure 40: 9, Md, Hyperoche martinizii, Gulf of Guinea,

original. Mx.l, Mxp, H. medusarum, Antarctic, original.

P.I, P.2, P.3, H medusarum, off California, original.

Figure 41: 9, Bovallius, 1889, pi. 8, fig. 1. Mxp, P.I, P.2,

Iulopis mirabilis, off California, original.

Figure 42. 9, P.I, P.2, Bowman, 1973, fig. 32a,o,p; A.2, fig.

37j; Md, Mxp, fig. 36c,f. A.I, original.

Figure 43: 9, $, Chevreux and Fage, 1925, figs. 407, 408.

Md, Bowman, 1960, fig. 1; P.I, P.4, P.5, fig. 16; P.2, fig. 2.

Figure 44: 9, original. A.I, P.I, P.2, Barnard, 1932, fig.

162b,d,e; Md, Mxp, fig. 163a,f; Us, fig. 164c.

Figure 45: 9, Chevreux and Fage, 1925, fig. 406. $, Us,

Bovallius, 1889, pi. 14. Md, Mxp, P.I, P.2, P.6, original.

Figure 46: Bowman, 1973, fig. 51a,e,f,g,j,k,p,r,s.

Figure 47: J, H9 , Bovallius, 1889, pi. 15. Md, Mx.l, Mx.2,

Mxp, P.I, P.2, Us, original.

Figure 48: 9,3 , Vosseler, 1901, pi. 2. Md, Mxp, Us, Phronima sedentaria, original.

Figure 49: 9, P.5, Shih and Dunbar, 1963, fig. 9. Us <$, Us 9,

original.

Figure 50: 9, Bovallius, 1889, pi. 18, fig. 3; P.3, fig. 6;

P.4, fig. 7; P.5, fig. 8; P.6, fig. 9; P.7, fig. 10 (distal segment

added); Us, fig. 11.

Figure 51: $, Stebbing, 1888, pi. 177. Md, Mxp. P.I, P.2,

P.3, P.5, P.6, P.7, Us, original.

Figure 52: Original.

Figure 53: $, Stebbing, 1888, pi. 181. A.I, P.I, P.2, P.5, P.6,

P.7, Us 9, Pirlot, 1930, fig. 9; Us $, fig. 9 (but Up.3 added

from Stebbing, 1888, pi. 180).

Figure 54: $, A.2, Stebbing, 1888, pi. 186. P.I, P.2, P.7,

Us, original.

Figure 55: $, Claus, 1887, pi. 13, fig. 7. A.2, P.I $ , P.I 9,

P.2, P.7, original. Us, Stebbing, 1888, pi. 187.

Figure 56: 9, P.I, P.2, P.7, Us, original. A.2<$, Claus, 1887,

pi. 20, fig. 2.

Figure 57: 9, Stebbing, 1888, pi. 191. A.2 $, P.I, P.2, P.6, P.7,

Us, original.

Figure 58: $, Stebbing, 1888, pi. 192. A.2, P.I, P.2, P.6, P.7.

Us, original.

Figure 59: 9, P.I, P.2, P.6, P.7, Anapronoe sp., off California,

original. A.I 9, A.2 9, A.2<$, Md, Mx.l, Us, A. reinhardti,

Gulf of Guinea, original.

Figure 60: 9, Us, Shoemaker, 1945, fig. 38; P.I, P.2, fig. 39.

A.2 $, P.I $ , original.

Figure 61: $ , Stebbing, 1888, pi. 195. A.2, P.I, P.2, Us,

original.

Figure 62: $, P.I, P.2, Chevreux and Fage, 1925, fig. 420.

P.7, Us, Stephensen, 1925, fig. 64. A.2 $, original.

Figure 63: $, <$ H, dorsal, A.2$, original. P.I, P.6, Pillai,

1966b, fig. 16. Us, Stebbing, 1888, pi. 198.

Figure 64: $, A.2.J, Stebbing, 1888, pi. 194. P.I, P.2, P.5,

Us, original.

Figure 65: 9, Fage, 1960, fig. 5a. P.I, P.2, P.7, Us, original.

Figure 66: 9, Pillai, 1966a, pi. 1; P.I, P.2, P.7, fig. 8. Us,

Fage, 1960, fig. 19.

Figure 67: 9, Shoemaker, 1945, fig. 44A. US<$, US9 . Fage.

1960, fig. 51. P.I, P.2, original.

Figure 68: 9, P.I, P.2, Up.3, Us, Shoemaker, 1945, fig. 45.

Figure 69: $, Pillai, 1966a, pi. 1; P.7, fig. 7. Up.29, Fage,

1960, fig. 22; Us $, fig. 21. P.I, P.2, original.
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Figure 70: 9, Bovallius, 1890, pi. 7, fig. 20. P.I, P.2, Pillai,

1966a, fig. 15; P.7, fig. 16. Us, Fage, 1960, fig. 63.

Figure 71: $, Pillai, 1966a, pi. 1. P.I, P.2, P.7, Us, original.

Figure 72: 9, Pillai, 1966a, pi. 1; Us, fig. 12. P.I, P.2, P.7,

original.

Figure 73: 9, Pillai, 1966a, pi. I; P.I, P.2, P.6, P.7, fig. 6. Us,

original.

Figure 74: Platyscelus ovoides, original.

Figure 75: A.2<$, Claus, 1887, pi. 2, fig. 6. P.I, P.2, P.6,

Spandl, 1927, fig. 45.

Figure 76: Amphithyrus bispinosus. Head and pereon, dorsal, original. A.2, P.I, P.2, P.6, Claus, 1887, pi. 6.

Figure 77: A.2^, Claus, 1887, pi. 4. P.I, P.2, P.6, Spandl,

1927, fig. 47.

Figure 78: A.2 <J , Claus, 1887, pi. 5; P.I, P.2, P.6, pi. 7.

Figure 79: A2$, Claus, 1887, pi. 5. P.I, P.2, P.6, Spandl,

1927, fig. 48.

Figure 80: $, A.2 $, P.7, Claus, 1887, pi. 8. P.I, P.2, P.6,

Us, Spandl, 1927, fig. 53. Mouth cone (labrum, Md, Mxp),

original.

Figure 81: Spandl, 1927, fig. 51.

Figure 82. Spandl, 1927, fig. 52.

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the author on the first page of text with an unnumbered footnote that includes his professional

mailing address. A table of contents is optional. An index, if required, may be supplied by the

author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further

headings may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet,

omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date:page) with a full reference at the

end of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent lines indented about three spaces. Within an entry, use a period-dash (.—) to separate

each reference. Enclose with square brackets any annotation in, or at the end of, the entry.

For references within the text, use the author-date system: "(Jones, 1910)" and "Jones

(1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

Use the metric system instead of, or in addition to, the English system.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively; however, if a group of figures is treated as a single figure, the components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on

pages separate from the text and not attached to the artwork. An instruction booklet for the

preparation of illustrations is available from the Press on request.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and

article titles, using initial caps with all words except minor terms such as "and, of, the." For

capitalization of titles in foreign languages, follow the national practice of each language.

Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume,

number, and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," "pages."

For free copies of his own paper, a Smithsonian author should indicate his requirements

on "Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will

receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.
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