Revision of Corophiidae and

Related Families (Amphipoda)

t

J. LAURENS BAR

I

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 151
SERIAL PUBLICATIONS OF TH E SMITHSONIA N INSTITUTION

The emphasis upon publications as a means of diffusing knowledge was expressed

by the first Secretary of the Smithsonian Institution. In his formal plan for the Institution, Joseph Henry articulated a program that included the following statement:

"It is proposed to publish a series of reports, giving an account of the new discoveries

in science, and of the changes made from year to year in all branches of knowledge."

This keynote of basic research has been adhered to over the years in the issuance

of thousands of titles in serial publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with

the following active series:

Smithsonian Annals of Flight

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoology

Smithsonian Studies in History and Technology

In these series, the Institution publishes original articles and monographs dealing

with the research and collections of its several museums and offices and of professional

colleagues at other institutions of learning. These papers report newly acquired facts,

synoptic interpretations of data, or original theory in specialized fields. These publications are distributed by mailing lists to libraries, laboratories, and other interested

institutions and specialists throughout the world. Individual copies may be obtained

from the Smithsonian Institution Press as long as stocks are available.

S. DILLON RIPLEY

Secretary

Smithsonian Institution
SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 15 1

Revision of Corophiidae and

Related Families (Amphipoda)

J. Laurens Barnard

SMITHSONIAN INSTITUTION PRESS

City of Washington

1973
ABSTRACT

Barnard, J. Laurens. Revision of Corophiidae and Related Families (Amphipoda). Smithsonian Contributions to Zoology, number 151, 27 pages, 1 figure,

1973.—A superfamily Corophioidea is revived to include Corophiidae, Ampithoidae, lschyroceridae, Podoceridae, Biancolinidae, and Cheluridae. The Podoceridae are presumed ancestors of Caprogammaridae and the suborder Caprellidea. The Photidae, Aoridae, and Isaeidae are abolished and absorbed into

the Corophiidae. Axial reversal of dominance in gnathopods 1-2 is now overruled as a fundamental evolutionary feature of familial impact; instead, axial

reversal is seen to have occurred several times and possibly itself has been reversed at least once. Dorsoventral flattening of the urosome is also discarded as

of no conservative evolutionary content within the old Photidae and Corophiidae. The loss of web spinning glands has occurred at least thrice in the

superfamily within the Podoceridae, Cheluridae, and Isaeidae, but in the latter

family the loss is unconnected to any other morphological revolution and

therefore Isaeidae are synonymi/ed with Corophiidae. Podoceridae, Ampithoidae, and Cheluridae are strictly definable clusters of genera descended from

various corophiids, whereas lschyroteridae are possibly polyphyletic but can be

easily recognized and defined. Within the highly diverse Corophiidae at least

16 clusters within the 50 genera are loosely defined. Though few of the clusters

can be interconnected by visualizing a few simple morphological changes in

evolution, the nomenclature! recognition of these clusters as subfamilies is

pointless because of the multifactorial diagnoses and long discussion of exceptional cases necessarv for their definition.

OFFICIAL PUBLICATION DA IF is handstamped in a limited number of initial copies and is recorded in thr Institution's annual report, Smithsonian Year. SI PRESS NUMBER 4809. SERIES

COVER DESIGN: The coral Montastrea cuvernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Barnaul. Jerry Laurens. 1928-

Revision of Corophiidae and related families (Amphipoda) .

(Smithsonian contributions to zoology, no. 151)

I. Corophiidae. U. Amphipoda. I. Title. II. Series: Smithsonian Institution. Smithsonian

contributions to /oology, no. 151.

QLI.S'M no. 151 [Ql.444.A5| 591'.08s [595\S7I| 73--.4H

for sale by the Superintendent of Document*. U.S. Goverment Printing Office, Washington, D.C. 20402

Price: 55 cents, domestic postpaid; 40 cents, GPO Bookstore
Contents

Page

Introduction 1

Evolutionary Primitivity in Corophioidea 5

Pattern of Relationships in Corophiidae 6

Procedures 9

Corophioidea 9

Key to Families of Corophioidea 9

Corophiidae (=Aoridae, Isaeidae, Photidae) 9

Key I to the Genera of Corophiidae 10

Key II to the Genera of Corophiidae 12

Diagnoses of Genera 14

Aciiminodeutopus J. L. Barnard 15

Aloiloi J. L. Barnard 15

Atnpelisciphotis Pirlot, new synonymy 15

Amphideidopus }. L. Barnard 15

Aura Kr0yer 15

Aurcho J. L. Barnard 16

Aoroides Walker 16

Catnacho Stebbing 16

Cerapopsis Delia Valle 16

Cheiriphotis Walker 16

Chevalia Walker 16

Chevreuxius Bonnier (Ischyroceridae) 16

Concholestes Giles 16

Corophium Latreille 17

Dryopoides Stebbing 17

Gammaropsis Liljeborg 17

Goesia Boeck . 18

Grandidierella Coutiere 18

Haplocheira Haswell 19

Isaea Milne Edwards 19

Kamaka Derzhavin 19

Konatopus J. L. Barnard 19

Kuphocheira K. H. Barnard 19

Lemboides Stebbing . 19

Lembnpsis Pearse 20

Lcmbos Bate 20

Leptocheirus Zaddach 20

Microphotis Ruffo 20

Microdeutopus Costa 20

Microprotopus Norman 20

Neohela Smith 21

Neomegamphopus Shoemaker 21

Neomicrodeutopus Schellenberg 21

iii
Pige

Paracorophium Stebbing 21

Paraoroides Stebbing 21

Parunciola Chevreux 21

Photis Kx0yer and Cedrophotis J. L. Barnard 22

Protomedia Kr0yer and Cheirimedeia J. L. Barnard 22

Psrudomegamphopus Myers 22

Rildardanus J. L. Barnard, revised 22

Rudilemboides J. L. Barnard 22

Siphonoecetes Kr0yer 28

Uncinotarsus L'Hardy and Truchot 23

Unciola Say, revised 23

Unciolella Chevreux 23

Xenocheira Chevreux 23

Ischyroceridae 23

Key to the Genera of Ischyroceridae 24

Diagnoses of Genera 23

Bathyphotis Stephensen 23

Bogenfelsia J. L. Barnard 24

Bonnierella Chevreux 25

Cerapus Say 25

Chevreuxius Bonnier 25

Dryopoides Stebbing 25

Ericthonius Milne Edwards 25

Isaeopsis K. H. Barnard 25

Ischyrocerus Kr0yer 25

Jassa Leach 25

Microjassa Stebbing 26

Paradryope Stebbing . 26

Parajassa Stebbing 26

Pseudericthonius Schellenberg 26

Pseudischyrocerus Schellenberg 26

Runanga J. L. Barnard 26

Ventojassa J. L. Barnard 26

Ampithoidae 26

Biancolinidae 26

Podoceridae . 27

Cheluridae 27

Literature Cited 27

IV
Revision of Corophiidae and

Related Families (Amphipoda)

J. Laurens Barnard

Introduction

The families Corophiidae, Photidae, Isaeidae,

Aoridae, Ampithoidae, Ischyroceridae, Podoceridae, and Cheluridae were extant until about 1962,

when J. L. Barnard synonymized Isaeidae and Photidae. Recently, Barnard (1970) described Aloiloi,

a new genus of Aoridae appearing to confirm the

polyphyletic descent of aorids from various photids

and isaeids by reversal of dominance in gnathopods. In a compendium of genera, Barnard

(1969a) pointed out the unsatisfactory arrangement of Corophiidae, Aoridae, and Isaeidae and

offered a suggestion made by Dr. E. L. Bousfield

that Ericthonius of the Corophiidae belongs to the

Ischyroceridae. The former arrangements of these

families have been unsatisfactory not only because

certain pairs and triads of genera with apparent

affinities have been segregated in different families

but because the routine taxonomic practices of

identifying these genera were largely unworkable

or very tedious. Keys and identificatory schemes

of earlier days often focused on characters of quantitative value on which systematic judgments might

be highly erratic: for example, the degree of dorsoventral flattening of the urosome to mark Corophiidae from other families.

This paper attempts to reorganize the taxonomic

concepts of corophiids by reestablishing a superfamily Corophioidea and suppressing or extinguishing the old concepts of Corophiidae, Photidae,

Aoridae, and Isaeidae as based mainly on depressed

/. Laurens Barnard, Department of Invertebrate Zoology,

Smithsonian Institution, Washington, D.C. 20560.

urosome and axial reversal of dominance in gnathopods, or selective loss of tube-spinning glands in

pereopods, and removes several genera of those

four families to the Ischyroceridae, which is also

to be maintained in the Corophioidea. The Ampithoidae, Podoceridae, and Cheluridae remain

unmodified except for their assignment to the

Corophioidea; however, Amphitholina Ruffo is returned to the Ampithoidae from its former assignment to the Eophliantidae.

The redefined Corophiidae are to include the

Isaeidae, Photidae, and Aoridae but no subfamilies

aie recognized, though several flocks of genera are

arranged in Figure 1. For example, two genera of

aorids have merochelate first gnathopods, and several genera similar to Microdeutopus have carpochelate first gnathopods.

The Corophiidae, Photidae, and Aoridae are

three of the most familiar names in Amphipoda,

but as taxonomic and evolutionary devices they

have been highly inappropriate and unfortunate.

The establishment of these family names and their

original contents and the subsequent expansion of

their conceptual limits over the last century have

stretched good order to the breaking point. The

name Corophiidae, based on Corophium Latreille,

concerns one of the most specialized and advanced

genera in Amphipoda; Aoridae, based on Aora

Krrtyer, took another advanced but peculiar genus

as its type; and Photidae as based on Photis Kr0yer,

are poorly representative of generalized members

in the group, as hindsight now demonstrates. The

orderly development of classification and the naming of groups would probably have been enhanced

had a more generalized genus such as Gammaropsis

1
2

Liljeborg been the first type-genus of a family in

this group. If a family Corophiidae had subsequently been considered necessary, it would then

have been restricted to a very few genera rather

than having embraced a host of genera inextricably

confounded with the Photidae and Aoridae. A

greater stability in names would undoubtedly have

ensued, the conceptual limits these names encompassed would have been steadier, and, perhaps,

more attention would have been paid to fine structure and the evolutionary impact of characters in

the group. This might have permitted more rapid

biological investigation of an extremely important

group of amphipods of which only Corophinm has

had much attention.

One must appreciate the difficulty of the taxonomist in propounding a classification in which

Corophinm stands at one end of a family and

Gammaropsis at the other, but the vast array of

more than 50 genera displaying a wide variety of

character variables softens this extraordinary morphological distance. The special straining apparatus on gnathopod 2 of Corophinm and Paracorophinm Stebbing is apparently facilitated by a

merochelate development similar to that seen in

Aora Kr0yer and Aoroides Walker, but in Corophium the chelation is joined immovably together.

No straight line evolutionary pattern can be seen

in merochelation because Paracorophium appears

to be far more primitive than Corophium in terms

of uropod 3, coxae, and antenna 2. Hence there

is no value in retaining a subfamily Corophiinae

for a single genus since the unique combination of

corophiin characters is no more unusual than other

unique patterns within the Corophioidea.

The loss of web-spinning glands in the pereopods of Isaea Milne Edwards and the development

of prehensibility on pereopods 3-5, though unique

as a combination of characters in Corophiidae, is

probably matched in the Ischyroceridae by Isaeopv/.v K. H. Barnard. Presumably Isaea and Isaeopsis

are commensals of other orders of animals rather

than tube dwellers, but this remarkable change in

behavior and morphology is not always accorded

value at family level as one may see in the Dexaminidae and Lysianassidae where some genera are

commensals and others are free nestlers.

The general evolutionary progress in Corophiidae, as now constituted, includes pygidization

(rigidification or loss of structure on the urosome)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and coxal reduction, and some of the morphological, if not behavioristic, results of this evolution

are as highly valued as loss of spinning glands.

These glands have also been lost in the Podoceridae (as far as is known) and Cheluridae, both

families yet retained in the Corophioidea because

of the lleshy telson and the uniformly basic mouthparts. Pygidization and coxal reduction in podocerids apparently anticipate the evolution and

radiation ot the suborder Caprellidea; the connection between the two groups is well seen in the

Caprogammaridae (Kudrjaschov and Vassilenko,

1966) and it any more such links are discovered,

taxonomists may necessarily evaluate the Caprellidea as a superlamily and not a suborder, or make

some other noinendatural union between the two

groups.

Since pygidization is so generally characteristic

of amphipod advancement in many evolutionary

lines (Barnard, 1969a), one presumes that an early

mark of this process is the coalescence of the telsonic lobes into a solid flap. The main characteristic ol Coropliioidea is the inflated and fleshy telson. In practical taxonomy this character is the

main distinction of many corophiids (photids)

from generalized gammarids (Gammaridae). Barnard (1969a) has suggested that the basic gammaridean can be conceptualized as a primitive

corophiid such as Gammaropsis as easily as can

primitive members of Gammaridae, but the descent of the vast majority of other amphipods then

requires stages of evolution in which the telson

FIGURE 1 Pattern of relationships in Corophiidae. Genera

in upper half cany normal heads, genera in lower half carry

recessed heads. Column 1 is characterized by small coxae

(squares) and reduced uropod 3; column 7 by enlarged

carpochclate gnathopod 1 (males). Gammaropsis in midcolumn 4 is the basic genus but immediately connected to

Protomedeia, another basic genus. Parts of other columns

are characterized by symbols concerning coxae, gnathopod 1,

and uropod 3. Numbers in circles refer to generic clusters.

C = gnathopod I enlarged, H = head, P = pereopods prehensile, R = urosomites coalesced, U = uropod. Arrows

suggi-st possible lines of descent. Genera in top and bottom

halves are fitted to 3 thermal regimes, tropical (T), warmtemperate (WT) and boreal-arctic, antiboreal-antarctic (B) .

Following genera omitted, similar to Gammaropsis (=

Eurystheus): Kermystheus, Podoceropsis, Segamphopus,

Audulla (valid subgenus) ; following omitted genus similar

to Microdeutopus: Hansenella.
NUMBER 151

"I

n

n

"1

D

CD

a
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

becomes flat and split into two lobes; in the most

advanced members of noncorophiid amphipods the

telson then returns to an unsplit condition though

it does not again become fleshy except in rare

cases (tliaumatelsonins). Though it may not be

truthful, the simplest pattern is to derive corophiids from hypothetical protogammarids already

progressing in an evolutionary sequence toward an

inflated and solid telson.

In the early and middle morphological evolution of corophiids, pygidization is presumably associated with the domicolous habit. But clearly the

opposite to pygidization also occurs in corophioideans, such as ampithoids, and, ultimately, in

the Podoceridae and Caprellidea, the domicolous

habit is surrendered in favor of nestling in anastomoses, the hypothetical primitive state. One can

visualize a sequence emerging from Gammaropsis,

which, morphologically, is scarcely distinct from the

basic gammaridean but which dwells in a tube

formed out of web-spinning glands. On one phyletic line through the Corophioidea the coxae become smaller, and the urosome becomes reduced

by flattening and by loss of various uropodal structures. But a side branch, the Ampithoidae, have

uropod 3 especially enlarged and provided with

hooks; ampithoids are also strong tube dwellers but

the coxae generally are larger than in the primitive

corophiid, Gammaropsis. In the Ischyroceridae,

;ilso tube dwellers, the coxae are variously sized and

uropod 3 remains fully formed but has a hook

morphology distinct from that of ampithoids.

Within the Corophiidae the tendency to evolve a

hooked uropod 3 is seen in Photis, so that one

judges that both evolutionary directions provide

adaptations lor domicoly. The Podoceridae can

scarcely lie cast out of the Corophioidea on their

loss of web-spinning glands as they have the fleshy

telson and stable mouthpart morphology of other

corophiids, and yet Podoceridae have more advanced cases of coxal reduction and loss of uropodal structure than do the majority of other corophioideans. The values of natural selection in the

functional morphology of these events are highlv

intriguing, as domicoly and nestling are both well

served by superficially similar advancements.

The fine structure of uropod 3 contains the presently most valuable set of characters distinguishing

families in Corophioidea. The Podoceridae are

discounted from this statement as they are recognized mainly through their elongate urosomite 1

and flexion of abdomen under the thorax (requiring more study as this situation is possibly not universal). Uropod 3 is often so degenerate that it can

not be used in a classificatory frame of reference.

In two genera of corophiids this is also true and

casts doubt on the true phylogeny of those two

genera (whether or not they are Corophiidae or

Ischyroceridae). Although several genera of Ischyroceridae have obvious phyletic connections, a high

probability exists that others now placed there according to morphological precepts have had diverse

origins from various primitive corophiids; eventually, Ischyroceridae may have to fall to Corophiidae

but such a move is not now supportable by any

direct evidence.

Certain indirect evidence in this problem involves similarities between pairs and triads of

genera, now partially in Corophiidae and partially

in Ischyroceridae, which share similarities in carpochelation of gnathopods and loss of structure on

uropod 2. For example, Ericthonius Milne Edwards

of Ischyroceridae and several genera of Corophiidae

in the Microdeutopus group have a similarly carpochelate gnathopod, though in Ericthonius the

gnathopod is number 2 and in the Microdeutopus

group the gnathopod is number 1. The reversal

of gnathopod dominance does not now seem to be

a character of high conservation in corophiids as

formerly believed, but, as evidenced by the genera

Aloiloi Barnard (1970) and perhaps by Pseudomegamphopus Myers (1968), this axial reversal is a

direct evolutionary step occurring diversely. The

return of dominance from gnathopod 1 to gnathopod 2 should therefore be of no greater moment

in the evolution of Ericthonius from a member of

Microdeutopus group than the step from Gammaropsis to Aloiloi.

With one exception, the relative elongation of

the peduncle of uropod 3 and the relative shortening of the rami have no obvious intergrades between corophiids and ischyrocerids or between

corophiids and ampithoids, but such evolutionary changes may also, like gnathopodal reversal,

have no long line of increments necessary or

even viable. The thin, nonsetose rami of ischyrocerids could have their origins in ampithoids

because the latter already have the elongate peduncle and short rami; again intergradation among

known genera is absent. This conclusion discounts
NUMBER 151

the reduction of rami in Dryopoides Stebbing, as

the outer ranuis of uropod 3 otherwise lacks uncination and the genus has other specializations

not characteristic- of ischyrocerids, such as an

evanescent pleonite 6, falcate article 3 of the mandibular palp, and short article 3 of antenna I.

Except for thin pereopods, Paradryupe Stebbing

is a good intergrade between corophiids and

ischyrocerids, as the outer ramus of uropod 3 is

slightly uncinate and gnathopod 2 has teeth resembling those in ischyrocerids. Uropod 3 of ampithoids is unmistakable and fairly uniform; it is

composed of a large, elongate, subcylindrical

peduncle bearing 2 short fleshy, ovate padlike rami,

the inner with apical setae, the outer with 1 or 2

very large spines pointing and curving dorsolaterally as hooks.

Uropotl 3 of ischyrocerids is less definable and,

indeed, 4-5 genera of this family have been placed

in Corophiidae for many decades. To some extent this error occurred because Corophiidae were

defined as a group containing members with

flattened urosome and therefore Ericthonius and

Cerapus Say were seen to be corophiids. Uropod

3 of ischyrocerids is now restricted to those having

an elongate, subcylindrical peduncle with a short

outer ramus bearing some kind of apical hook

dissimilar to that of Ampithoidae. The body of

the ramus may be apically hooked or may bear

one small hookspine; the ramus is rarely pad

shaped as in Ampithoidae but is usually thin and

tapering. An inner ramus is usually present, but

unlike uropod 3 of Ampithoidae it is thin and

tapering and lacks apical setae except perhaps for

one weak spinule.

The exceptional genera to be placed in Ischyroceridae are Cerapus, Runanga J. L. Barnard, and

Ericthonius, all of which have only one hooked

ramus on uropod 3, the inner ramus presumably

being lost. The hook on Ericthonius is unmistakable and unlike any member of Corophiidae. But

Cerapus and Runanga have a short, padlike ramus,

almost vestigial, bearing a strong apical hook (articulate or not). The padlike appearance of the

ramus recalls the Ampithoidae, but otherwise the

gnathopodal context of Cerapus connects it to

Ericthonius, and the general body shape of Runanga connects it to Cerapus so as to exclude both

from the Ampithoidae. In the latter family, Amp/iitholina represents a cylindrioid body form

suggesting a link between Ampithoidae and

Corophiidae (and by interpretation therefore to

Cerapus and Runanga).

EVOLUTIONARY PRIMITIVITY IN COROPHIOIDEA.—

The number of primitive genera in Corophioidea is

surprisingly low if one bases this estimate on logical

assumptions. Characteristics of the basic gammaridean (Barnard, 1969a) are used to define primitivity.

The basic (assume primitive) amphipod characters

in the corophioid context are: short article, 3 of

antenna 1 (or generally short antennae), long

accessory Hagellum, nonpedunculate ocular lobes,

3-articulate mandibular palp, large coxae, free and

unfiattened urosomites, large subchelate gnathopod 2 dominant and enlarged in males, simple

pereopods, long, equally sized rami of uropod 3,

and a telson as long as it is broad. Additional, less

important characters are: uniformity in extension

and size of coxae 1-5 (or 1-4), and thin and unornamented antenna 2. Because the vast majority

of Corophioidea are tube dwellers, one presumes

that the modification and specialization of attributes stated above have a relationship to domiciliary

habits. These are mainly visualized by assuming

that (1) elongate antennae have selective value to

a tube dweller in reaching out of the tube without

committing the entire body to predatory attack; (2)

reduction in coxae modifies the body toward a

cylindrical plan well adapted to a tube; the tube

may act as a partial substitute for any egg protection afforded by the coxae; (3) loss of uropodal

and telsonic structure and depression of the urosome (pygidization) are connected with ability

to grasp the inside of the tube and make quick

withdrawals into it; (4) the axial reversal of dominance from gnathopod 2 to gnathopod 1 carries

the prehensibility forward toward the tube opening; (5) the pedunculate ocular lobes increase the

ability of the animal to see by a slight margin

as the eyes project farther forward on the head

than in the basic amphipod.

These hypothetical examples illustrate, perhaps,

why few corophiid genera retain all of the attributes of the general gammaridean. Despite the

apparent advantages of domicolous specialization,

one of the most primitive genera in Corophioidea,

Gammaropsis, remains the most diversely speciated

genus confined to the open sea. It is approximated
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in diversity only by Corophium and Grandidierella

Coutiere, which occur not only in open seas but

mainly in brackish waters or in lagoons.

PATTERN OF RELATIONSHIPS IN COROPHIIDAE.—

The diversity and absence of intermediate stages

among certain corophiids are of such degree as to

thwart any clear understanding of evolutionary

lines in Corophiidae. The pattern of relationship

(Figure 1) duplicates the general pattern (Barnard,

1969a) found in Gammaridea as a whole, in which

evolution may be characterized as centrifugal. In

this concept one finds that basic corophiids, much

as protogammaruses, appear to have spun off evolutionary lines in various morphological directions.

So-called straight-line evolutionary sequences are

therefore very short.

In the pattern of Figure 1 the genera are arranged in two main groups: those above midline

with ordinary heads, and those below with the

head recessed for the basal attachment of antenna

2. In the Corophiidae this character alternative is

of reasonably broad scope and ease of definition. It

also provides a good basis on which to erect a

strawman, and provides a point of departure and

basis of discussion in attempting to deal with

possible parallels in evolution among the several

groups of genera. In Figure 1 each half of the

diagram is also subdivided into geographic zones:

tropical (T), warm-temperate (WT) and borealarctic (B)1

; the latter division also contains deepsea genera. Each genus on the diagram is fitted

closest to its center of geographic zonation.

Gammaropsis appears to have all of the theoretically basic or primitive conditions necessary for

a corophiid in terms of ordinary gnathopods, unmodified urosome and its appendages, mouthparts,

and pereopods. But Gammaropsis bears an elongate

article 3 of antenna 1 and a partially recessed head,

whicli in general terms within the Gammaridea

are atypical. One might however theorize that the

early tube-dwelling amphipods of the Corophioidea

would have necessarily had these mechanisms to

extend the first antenna and the eyes toward the

opening of the tube. The alternative conditions,

unrecessed head and/or unextended ocular lobes,

appear just as successful for survival because half

or more of the genera of Corophiidae have one or

both of these protrusive characteristics in condition

'Antiborcal=antarctic also included.

normal to nestling gammarideans. Hence the early

evolution of these characters may have been correlative with tube dwelling but may have been

selected against because of a relatively neutral

value to certain species in which evolved other

characteristics of far more value to the tube

dweller. In many of the genera with normal head

and antenna 1, the first pair of gnathopods (pereopods of thoracic segment 1) is enlarged and often

is prehensile, perhaps an adaptation to bring

grasping organs nearer to the tube opening. Because the second pair of gnathopods in these

"aorids," with enlarged gnathopod 1, appears similar to first gnathopods of related genera, one supposes that enlargement of gnathopod 1 is a simple

reversal mutation in the axial genetic systems of the

organism. The term "simple" is applied to this

case because an alternative development system

would call for gradual reduction of gnathopod 2

and gradual enlargement of gnathopod 1 in an

evolutionary sequence. There is virtually no evidence for this more complicated system as aorid

amphipods simply appear to have gnathopods 1-2

exchanged by direct axial reversal. Reversals appear

to have occurred in several diverse generic lines

independent from each other. Returned reversals

also appear to have occurred in which the enlarged

gnathopod 1 may have developed a distinctive

marker such as carpochelation and is then axially

reversed to segment 2 as appears to be the case in

Ericthonius (Ischyroceridae). Genera with enlarged

gnathopod 1 were formerly placed in the Aoridae,

but the polyphyly of this arrangement is now too

obvious to merit its continuation. The enlargement

of gnathopod 1 is poorly correlated with protrusive

cephalic features, as at least 7 genera with normal

head have enlarged gnathopod 1. The evolutionary

sequences are therefore multifactorial and undoubtedly contain assortive correlations with degrees

of pygidization, reduction of coxae, kinds of domiciles formed by the tube spinners of the pereopods,

and details of behavior of the several species. Great

assistance will be afforded to our understanding

of evolution in this group once functional morphologists explain some of these correlatives.

Because Protomedeia is the least specialized

genus in the upper half of Figure 1, I have connected Gammaropsis and Protomedeia with reverse

arrows to mark Protomedeia as a better example of

primitivity in head than Gammaropsis and to sug-
NUMBER 151

gest Protomeileia as a possible precursor. This

would place the hypothetical geographic origins

of corophiids in the north boreal zone near the

most primitive living members of the Gammaridae,

a not unreasonable situation.

There is no conclusive evidence that a single reversible step between normal head and recessed

head occurs in the evolutionary sequences between closely related genera but two cases suggest that this is possible. In Figure 1 the two

columns at far right contain the Microdeutopus

and Lemboides groups in which the upper members have normal heads, the lower members recessed heads. The consistent presence of an enlarged

and carpochelate gnathopod 1 in the Microdeutopus line appears to dominate the group so strongly

as to negate the probability that the gnathopod

evolved twice (in upper and lower genera). Another kind of easily recognizable gnathopod 1

occurs in the Lemboides group. If a normal head

is considered more primitive and the recessed head

more advanced, one can point to another case, in

Microdeutopus, where the basic genus is geographically centered in the north boreal, whereas many

of its advanced descendants are found in the tropics

or farther south. These considerations suggest that

the basic corophiid did not have protrusive cephalic features but that these have evolved independently several times.

Arrows drawn between genera in Figure 1 imply

possible descent from species similar to the

ancestral genus. No other decisions can be implied

from these arrows except that the evolutionary

sequence could not be reversed because structurally

extinct parts of the morphology would have to be

reconstructed.

According to Figure 1 at least 16 evolutionary

clusters of genera are present in the Corophiidae

alone. Some of the clusters are poorly cohesive

(e.g., numbers 3 and 7) and some are poorly segregated from others (e.g., numbers 1 and 2; numbers 13 or 14 as derived from Gammaropsis). The

origins of clusters 5, 6, and 8 are almost completely

unknown and the origins of 3, 7, 8, and 10 are

very speculative. Only the sequences between Gammaropsis and clusters 4, 11, 12, 13, 14, and possibly

1 and 2 are highly reasonable.

The internal cohesiveness of cluster 1 is based

on equality of enlarged gnathopods with progress

from Aorcho to Camacho and Neohela by a reduction and loss of the inner ramus of uropod 3 and a

reduction in size of coxae. Parunciola in cluster 2

could be descendants of Aorcho by loss of the inner

ramus on uropod 3 and recessment of the head, and

Parunciola could then progress to its descendants

by way of reduction in coxae.

Cluster 3 is based on Protomedeia, a possible

ancestor or descendant of Gammaropsis, but which

is cold-adapted and may have radiated into the

"cheira" genera with heavily setose gnathopod 2

(and often gnathopod 1). Paracorophium and

Corophium are also remotely connected to cluster

3 in their heavily setose gnathopod 2 but, in Paracorophium, article 4 of gnathopod 2 is formed

into a setose lobe which in Corophium is joined

to article 5. Corophium is highly remote from

Paracorophium because of the reduction in or loss

ot structure in coxae, mandibular palp, and uropod

3.

Cluster 4, characterized by Photis, appears close

to Gammaropsis, but all genera within cluster 4

have a weak to strong degree of reduction in the

inner ramus of uropod 1, while otherwise retaining

large coxae and dominance of gnathopod 2.

Cluster 5, lacking any visible ancestors, is characterized by axial reversal of gnathopods 1-2, the

enlarged gnathopod 1 being carpochelate. High

setosity of gnathopod 2 and loss of the inner ramus

of uropod 3 are internal advancements within the

cluster.

The Aora cluster (6) contains two genera with

merochelate and enlarged gnathopod 1 from unknown ancestors. The merochelate gnathopod 1

resembles the merochelate and setose gnathopod 2

of Paracorophium, but little else between the genera

has similarity.

Lembos (cluster 7) can be characterized as a

Protomedeia with axial reversal of gnathopods.

Lembopsis has a tooth on article 5 of gnathopod 1

but is not to be considered truly carpochelate.

Lemboides and Rudilemboides have article 5 of

gnathopod 1 dominant over article 6, the reverse

of conditions in Lembos. Because this situation

occurs in gnathopod 2 of Protomedeia, Lemboides

appears as a better descendant of Protomedeia than

of Lembos, but no other genus with normal head,

other than Protomedeia, can be found as a better

ancestor of Lembos. A dashed arrow from Gammaropsis is therefore indicated as a remote possibility of relationship, but Lembos differs from
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gammaropsis in the short article 3 of antenna 1,

the normal head, and the axially reversed

gnathopods.

The Dryopoides cluster (8) is enigmatic. Paradryope differs from Protomedeia in the short rami

of uropod 3 (in resemblance to ischyrocerids) and

in the elongate article 3 of antenna 1, while Dryopoides has the short rami of uropod 3, short

article 3 of antenna 1, and axially reversed

gnathopods.

Isaea (9) resembles Protomedeia but has prehensile pereopods 3-5 and apparently has no webspinning glands. The species of Isaea are

commensals on decapods.

Chevalia (10), with elongate article 3 of antenna

1, small coxae and fused pleonites 4-5, has no close

relative.

Clusters 11-14, composed of Pseudomegamphopus, Aloiloi, Megamphopus, and Psendeurystheus,

appear simply to be products of Gammaropsis, the

first two characterized by axially reversed gnathopods from distinct ancestors within the Gammaropsis i>ool, Megamphopus characterized by diversity

in anterior coxae, and Pseudeurystheus by elongation of article 5 on gnathopod 1. Such unlisted

genera as Segamphopus, Kermystheus, and Podoceropsis are advanced products of the Gammaropsis

pool as based on reduction or loss of accessory

flagella or elongation of article 5 on gnathopod 1.

These three genera plus Megamphopus and

Pseudeurystheus actually appear to belong to what

should be called a supergenus Gammaropsis. Their

character differentiations are either so minor or

so well-connected by intergradations to good

species of Gammaropsis that they should be recognized only as subgenera or as marks of species

groups within Gammaropsis. Much investigation

and revision of species assigned to the Gammaropsis

genera await specialists throughout the world

(Barnard, 1962).

Neither Uncinotarsus (15) nor Concholestes (16)

can be integrated with those groups characterized

by Siphonoecetes or Camacho, but Concholestes

appears closest to Siphonoecetes, whereas Uncinotarsus may be remotely related to Unciola.

Other families of the Corophioidea are not

shown on Figure I but their relationships can be

approximated. The Ischyroceridae all have the

head recessed and article 3 of antenna 1 elongate.

The normal members appear to be a single evolutionary sequence with ancestors near Gammaropsis. The aberrant Ericthonius and Cerapus have

gnathopod 2 carpochelate in a fashion similar to

gnathopod 1 of the Microdeutopus line, but in

head and antenna 1 do not resemble the Grandidierella group (within the Microdeutopus line)

to which they correspond in the loss of the inner

ranius of uropod 3 or the full reduction of rami

on uropod 3, whereas they resemble the Neomegamphopus line in head and antenna 1 but have

a modified uropod 3. Gnathopod 2 of Erichthonius could be an axial reversal of gnathopod 1 in

Neomegamphopus. The slightly hooked single

ramus of uropod 3 in Ericthonius, with the pedunc le elongate, resembles certain Ischyroceridae

sufficiently to suggest that Ericthonius is an ischyrocerid lacking an inner ramus of uropod 3 but this

condition may have evolved independent of

ischyrocerids. In any corophiid with highly reduced

or missing rami of uropod 3, the probable ancestry

is obfuscated. Though Cerapus and Ericthonius can

be defined as ischyrocerids, their probable ancestry

in the Microdeutopus line should be revalued

continuously.

The Ampithoidae are characterized by poorly

recessed head and short article 3 of antenna 1 and

therefore are closest to Protomedeia in the Corophiidae. Some species of ampithoids have developed prehensile pereopods as in Isaea of the

Corophiidae but apparently they retain tubespinning glands.

Podoceridae probably have recessed heads, but

the heads and eyes of many species are so oddly

modified that a decision is difficult to make on

this point. Article 3 of antenna I is elongate and

thus relates this group to Gammaropsis ancestors.

The tendency to loss of rami on uropod 3 and

reduction in coxae places Podoceridae close to

Unciola and Unciolella of the Corophiidae or to

Neohela in the group with normal head. Neohela

is a poor hypothetical ancestor to podocerids because of its incipient blindness and slightly enInrgetl gnathopod 1 (or equally sized gnathopods

1-2), but its extremely slender body is good basic

material for the modifications found in Podoceridae. Urosomite 1 of podocerids is elongate and the

ulxlomen is usually heavily flexed beneath the

thorax, and in some podocerids the thorax is

broadened laterally. Podocerids clearly appear to

IK." ancestors of Caprogammaridae and the Caprel-
NUMBER 151

lidea, as podocerids show a tendency to reduction

of pereopods 1-2, a reduction and pygidization of

the urosome, and, in Dulichia, a shortening and

splitting away from the thorax of pereonite 1,

which in caprellids becomes fused with the head.

PROCEDURES.—This paper is written in condensed

form; references to old genera and illustrations may

be consulted in Barnard (1969a). Two distinct

keys offer a choice of characteristics for identification. New diagnoses to genera in the Corophiidae

and Ischyroceridae are presented. Diagnoses in the

other corophioidean families remain identical to

those in Barnard (1969a).

COROPHIOIDEA

DIAGNOSIS.—Telson fleshy, often pillow shaped

but always depressed dorsoventrally, fleshiness not

confined to ventral keel only, telson entire though

occasionally with weak lobation.

DESCRIPTION.—Except for Biancolinidae mouthparts like those of basic gammaridean with weak

exceptions; mandible always bearing incisor, triturative molar, spine row, and palp, latter rarely

reduced in number of articles; lower lip with inner

lobes, outer lobes occasionally subdivided; palp

of maxilla 1 large; inner and outer plates of

maxilliped large, palp large but rarely article 4

vestigial; coxa 1 generally as large as coxa 2 except

in two genera; coxae 2-3 well developed, or equal

to coxa 1, coxa 4 never posteriorly excavate;

rostrum rarely present, never hood shaped; peduncle of antenna 1 with elongate articles 1-2, never

with telescoped articles 2-3; article 3 of gnathopod

2 not elongate and hand not like Lysianassidae;

pereopod 5 longer than 4 but never vastly elongate

and fossorial as in Oedicerotidae; telson not

elongate: urosomites 2-3 not selectively coalesced

together, if fusion occurring either urosomites 1-2

together or 1-3 together.

Key to Families of Corophioidea

1. Urosome formed into large, rectangular, fiat rigid box much longer than pleonites 1-3

together, uropod 2 heavily flabellatc CHELURIDAE

Urosome not boxlike or if certain segments coalesced not significantly longer than pleonites

1-3 together, uropod 2 not fiabellate 2

2. Urosomite 1 more than twice as long as urosomite 2 and always free PODOCOUDAE

Urosomite 1 not more than 1.5 times as long as urosomite 2 even if coalesced with that

segment S

3. Uropod 3 lacking rami COROPHIIDAE (in part)

Uropod 3 with 1-2 rami 4

4. Ramus or rami of uropod 3 half or less as long as peduncle 5

Ramus or rami of uropod 3 as long as or longer than peduncle 6

5. At least one ramus of uropod 3 heavily setose terminally AMPITHOIDAE

No ramus of uropod 3 heavily setose, maximum of 3 wirelike setae on outer ramus

laterally ISCHYROCERIDAE

6. Mandibular palp absent, molar absent or vestigial, palp of maxilla 1 vestigial or absent,

palp of maxilliped 3-articulate BIANCOLINIDAE

Palps of mouthparts present and normal except palp of mandible occasionally 1-2

articulate COROPHIIDAE (in part)

COROPHIIDAE

(=AORIDAE, ISAEIDAE, PHOTIDAE)

DIAGNOSIS.—Corophioidean with ordinary urosomite 1, if rarely coalesced to other urosomites not

elongate; uropod 3 in two configurations: (1) lacking rami, peduncle setose; (2) bearing ramus or

rami as long as or longer than peduncle (except

Dryopoides and Paradryope), at least one ramus

with terminal setae, or bare, no hooks; pereopods

glandular with rare exception.

Species without rami on uropod 3 can be distinguished from Ampithoidae in the undivided

outer lobes of the lower lip but cannot be distinguished from Ischyroceridae. In other words, an

ischyrocerid without rami on uropod 3 could not

be distinguished from Corophiidae and therefore

the two genera of corophioideans in this circumstance must be placed in the Corophiidae.
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key I to the Genera of Corophiidae

1. Rami of uropod 3 absent Key ^

One or two rami of uropotl 3 present

2. Uropod 3 with one ramus Key **

1

 Q

Iropod 3 with two rami °

3. Inner ramus of uropod 3 much shorter than outer ramus Key C

Rami of uropod 3 subequal in length to each other Key ^

KEY A (a triplet)

la. Gnathopod 2 larger than gnathopod I. antenna 2 of medium stoutness, uncuspidate,

longer than antenna 1, article 3 of antenna 1 subequal to article 1, coxae 5-7 not acuminate, rami of uropod 1 equal Concholestes

b. Gnathopod 1 larger than gnathopod 2. antenna 2 slender, uncuspidate, shorter than antenna 1, article 3 of antenna I shorter than article 1, coxae 5-7 acuminate, rami of

uropod I equal Uttcinotarsus

c. Gnathopod 1 larger than gnathopod 2, antenna 2 of medium stoutness, cuspidate, shorter

than antenna 1, article 3 of antenna 1 shorter than article 1, coxae 5-6 acuminate,

inner ramus of uropod 1 vestigial Rildardanus

KEY B

1. (•nathopod 2 with articles 4-"> contiguous in corophiid fashion (see text)... Corophium

(•nathopod 2 with articles 4-5 attached in ordinary fashion 2

2. Pleonites 4—5 coalesced together Katnaka

Pleonites 4-5 separate 3

3. Coxae serially discontiguous and very small 4

Coxae serially contiguous and usually large 10

4. I'ropotl 2 with one ramus 5

Iropod 2 with 2 rami 6

">. Accessory flagellum absent, mantlibular palp 1-articulate, gnathopods weakly subchelate

and not subequal to each other, uropod 3 with peduncle medially expanded

Siphonoecetes

Accessory llagellum 2-articulate, mandibular palp 3-articulate, gnathopod 1 enlarged in

male, carpochelate, uropod 3 with normal peduncle Chevreuxius (Ischyroceridae)

6. Coxae angular 7

Coxao softly rounded 8

7. Article 6 of gnathopod 2 an elongate Iectangle, paiachelate, or palm transverse, antennae

usually much shorter than body, flagellum of antenna 2 scarcely longer than peduncular article ">. peduncle of uropod 3 often expanded medially very strongly, antenna 2

generally considered stout, often strongly so t nciola

Article t> of gnathopod 2 ovate or trapezoidal, expanded, palm oblique, antennae usually

as long as or longer than body, flagillum of antenna 2 nearly as long as all articles of

peduncle, peduncle of uropod 3 rarely expanded medially and usually poorly so, antenna 2 \ery slender Neohela

8. Accessory flagellum only I-articulate, male gnathopod 1 carpochelate

Neomicrodeutopus and Grandidierella

Accessory fiagellmn iarel\ I-articulate, usually multiarticulate, male gnathopod 1 not

caipochelate 9

!>. Inner plate of maxilla I with one s.ta. ramus of uropod 3 about 1.5 times as long as

peduncle Vnciolella

Inner plate of maxilla I with about 10 setae lining medial edge, ramus of uropod 3 about

twin- as long as peduncle Parunciola

Ml. Kycs on extended cephalic peduncles 11

Eyes on side of head in normal gainmaridean fashion 13

11. Ramus of uropod 3 basally almost as broad as apex of peduncle .. Microphotis

Ramus of uropod 3 basally half or less as broad as apex of peduncle 12

12. Male gnathopod 2 propodochclate Cerapopsis

Male gnathopod 2 subchelate Ampelisciphotis (—Gaviota)
NUMBER 151 H

13. (•nathopods 1-2 simple, both small, gnathopod 2 with vestigial setose dactyl Kuphocheira

(.nathopods 1-2 subchtiutc, one or the other enlarged, dactyls of gnathopod 2 normal 14

14. (.nathopod I slightly to greatly larger than gnathopod 2 15

(.nathopod 2 slightly to greatly smaller than gnathopod 1 17

I"). Kaiiius of tiropod 3 twice as long as peduncle, male gnathopod 1 carpochelatc 16

Raimis of moped 3 only as long as peduncle, male gnathopod 1 not carpochelatc

Paraoroidcs

16. Mandibular palp article 1 elongate Grandidieretla

Mandibular palp article 1 not elongate Neomicrodeutopus

17. Accessory flagellum absent Micropkotis

Accessory flagellum present 18

IK. Male coxae 1-4 similar to each other in size and shape Microprotopus

Male coxae 1-4 of varying sizes and shapes Chehiphotis

KEY C

1. Gnathopod 2 with articles 4-5 contiguous in corophiid fashion (see text), but article 4

free and projecting during flexion Paracorophium

Gnathopod 2 with articles 4-5 attached to each other normally 2

2. Coxae very small and discontiguous serially S

Coxae large and touching serially 4

3. Gnathopod 1 scarcely larger than and similar to gnathopod 2, palm smooth, inner ramus

of uropod 3 distinct, pleonal cpimcron 3 smoothly quadrate Camacho

Gnathopod 1 much larger than gnathopod 2, palm sculptured, inner ramus of uropod 3

usually represented by large peduncular process, when distinct scarcely shorter than

outer ramus, pleonal epimeron 3 bearing postcroventral tooth Vnciola

4. Coxa 1 smaller than coxa 2 and partially or fully hidden by coxa 2 Leptocheirus

Coxa 1 not grossly smaller than coxa 2 and not substantially hidden by coxa 2 5

3. Gnathopods 1-2 simple and small Haplechehra

Gnathopods subchclatc and either gnathopod 1 or gnathopod 2 enlarged in both sexes . 6

6. Gnathopod 1 larger than gnathopod 2. in male carpochelate, gnathopod 2 small and nearly

simple Acuminodeutopus

Gnathopod 2 larger than gnathopod 1, in male grossly subchelate, gnathopod 1 small and

normally subchelate 7

7. Accessory flagellum absent or formed as vestigial scale Photis

Accessory flagellum multiarticulate 8

8. Uropod 3 scalelike, peduncle platelike Cheiriphotis

Uropod 3 cylindrical Protomedeia (subgenus Cheirhnedeia)

KEY D

1. Anterior coxae small and slightly discontiguous, pleonites 4-5 coalesced together

Chevatia

Anterior coxae small or large but touching serially, pleonites 4-5 free 2

2. Gnathopod 2 distinctly larger than gnathopod 1 in both sexes (or value of this character

undecided) 3

Gnathopod 1 distinctly larger than gnathopod 2 8

3. Article 3 of antenna 1 equal to or slightly longer than article 1 4

Article 3 of antenna 1 shorter than article 1 6

4. Gnathopods 1-2 of medium size and equal to each other, pereopod 3 shorter than articles

1-5 of pereopod 5 together Aorcho

Gnathopod 2 distinctly larger than gnathopod 1 5

5. Coxae longer than height of pereonitcs, pereopods 3-5 subprehensile lsaea

Coxae equal to or much shorter than height of pcreonites, pereopods 3-5 simple

P$eudeury%theus, Segamphopus, Audulta,

Bonnierella (Ischyroceridae), Kermystheus, Podoceropsis, Megamphopus, Gammaropsis

6. Coxa 1 significantly smaller than and covered by coxa 2 Leptocheirus

Coxa 1 not significantly smaller than nor covered by coxa 2 7

7. Accessory flagellum absent, inner plate of maxilla 2 heavily setose basomedially, inner

plate of maxilla 1 as long as palp, article 4 of maxillipedal palp vestigial Goesia
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Accessory flagellum 3 +articulate, inner plate of maxilla 2 not setose basomedially. inner

plate of maxilla 1 half as long as palp, article 4 of maxillipedal palp ordinary

Protomedeia

8. Male gnathopod 1 incrochelate 9

Male gnathopod 1 not merochelate 10

9. Accessory flagclluin absent ... Aoroides

Accessory flagellum 3 + aiticulate Aora

10. Male gnathopod 1 with article 5 broader and/or longer than article 6 11

Male gnathopod 1 with article 6 as broad and as long as article 5 18

11. Male gnathopod I not carpochelate 12

Male gnathopod 1 carpochelate 14

12. Both rami of uropod 3 much shorter than peduncle, rami basally disjunct from each

other liryopoides

One ramus of uropod 3 as long as or longer than peduncle, rami attached to peduncle

contiguously 13

13. Gnathopods fully subchelate Lemboides

Gnathopods scarcely subchelate Rudilemboides

14. Gnathopod 2 of male heavily setose on full anterior margin of article 5 15

Gnathopod 2 of male sparsely setose on anterior margin of article 5 or setae compacted

into distal bundle 17

15. Article 5 of gnathopod 2 longest, female gnathopod 1 simple Neomegamphopus

Article 6 of gnathopod 2 longest, female gnathopod 1 subchelate 16

16. Head elongate, antenna 2 inserted in head behind level of eyes, gnathopod 2 of male as

stout as gnathopod 1, of female weakly setose anteriorly Amphideutopus

Head not elongate, antenna 2 inserted in head on level with eyes or in front of them,

gnathopod 2 of male much more slender than gnathopod 1, of female densely setose

anteriorly Microdeutopus (=Coremapus)

17. Antenna 2 inserted in head behind level of eye, inner plate of maxilla 1 with medial

setae, article 3 of mandibular palp stout and clavate Konatopus

Antenna 2 inserted in head level with eye, inner plate of maxilla 1 lacking medial setae

(one apical only) , article 3 of mandibular palp elongate and linear but falcate

Microdeutopus and Hansenella

18. Article 3 of antenna 1 as long as article 1 19

Article 3 of antenna 1 shorter than article 1 20

19. Gnathopod 2 stout, article 6 two-thirds as broad as article 6 of gnathopod 1 Aloiloi

Gnathopod 2 slender, article 6 one-third as broad as article 6 of gnathopod 1

Pseudomegamphopus

20. Male gnathopod I with long tooth on article 5 Lembopsis

Male gnathopod I with posterior lobe but not distinct tooth on article 5 21

21. Gnathopod 2 simple, article 5 of gnathopod 1 very bulbous Xenocheira

Gnathopod 2 subchelate. article 5 of gnathopod 1 not bulbous Lembos

Key II to the Genera of Corophiidae

1. Articles 4-5 of gnathopod 2 contiguous in corophiid fashion (see text) 2

Articles 4-5 of gnathopod 2 attached to each other normally 3

2. Uropod 3 with 2 unequal rami, niatidibiilar palp 3-articulate, article 4 of gnathopod 2

detached from article 5 along contiguous margin Paracorophium

L'ropod 3 with one ramus. mandibulai palp 2-articulate, article 4 of gnathopod 2 fully

joined to article 5 Corophhtm

3. Pleonites 4-5 coalesced together 4

IMeonites 4-5 free 5

4. L'ropod 3 biramous, antenna I longer than antenna 2, anterior coxae all subequal to each

other in size, small, discontiguous Chevalia

I ropod 3 with one ramus, antenna 1 less than half as long as antenna 2, anterior coxae

large, touching, coxa 1 enlarged Kamaka

5. Coxa 1 much shorter, smaller than, and covered by coxa 2 Leptocheirus

Coxa 1 subequal in size and not covered by coxa 2 6
NUMBER 151 13

6. Pcreopods 8-5 prehensile Isaea

Pcreopods 3-5 simple 7

7. Anterior coxae serially discontiguous, not deeper than broad 8

Anterior coxae serially overlapping, either long or short 16

H. Rami of uropod 3 absent . Key A of Key I

One or two rami of uropod 3 present 9

!>. Accessory flagcllum absent, mandibular palp 1-articulate Siphonoecetes

Accessory Hagellum present, mandibular palp 3-articulate 10

10. Uropod 3 with 2 rami, inner ramus small Camacho

Uropod 3 with one ramus 11

11. Male gnathopod 1 carpochelate 12

Male gnathopod 1 not carpochelate 13

12. Uropod 2 with one ramus Chevreuxius

Uropod 2 with 2 rami Neomicrodeutopus and Grandidierella

13. Peduncle of uropod 3 medially expanded I in tola

Peduncle of uropod 3 not medially expanded 14

14. Accessory Hagellum 1 articulate, article 3 of antenna 1 equal to or shorter than article 1

UnciolcUa

Accessory Hagellum inuliiarticulate, article 3 of antenna 1 longer than article 1 . 15

I.'). Coxae angular, inner plate of maxilla 1 with 5 setae Xeohela

Coxae softly rounded, inner plate of maxilla 1 with 10 setae (coxae actually touching

serially) .- Parunciola

16. Male gnathopod 1 larger than gnathopod 2, article 5 longer and stouter than shortened and

slender article 6 Key E

Male gnathopod 1, if larger than gnathopod 2, with article 6 equal in size to or dominating

article 5 17

17. Article 3 of antenna J equal to or longer than article 1 18

Article 3 of antenna 1 shorter than article 1 27

18. Uropod 3 with one ramus 19

Uropod 3 with 2 rami 22

19. Gnathopods simple, dactyl of gnathopod 2 vestigial and setose Kuphocheira*

Gnathopods subchelate, dactyl of gnathopod 2 normal 20

20. Ramus of uropod 3 basally almost as broad as apex of peduncle Microphotis

Ramus of uropod 3 basally half or less as broad as apex of peduncle 21

21. Male gnalhopod 2 propodochelate Cerapopsis

Male gnathopod 2 subchelate Ampelisciphotis (=Gaviota)

22. Inner ramus of uropod 3 very short Photis

Rami of uropod 3 subequal to each other 23

23. Gnathopod 1 much larger than gnathopod 2 24

Gnathopod 1 equal to or smaller than gnathopod 2 25

24. Gnathopod 2 stout, article 6 over two-thirds as broad as article 6 of gnathopod I Aloiloi

Gnalhopod 2 slender, article 6 one-third as broad as article 6 of gnathopod 1

Pseudomegatnphoptts

25. Gnathopods 1-2 equal to each other, pereopod 3 shorter than articles 1-4 of pereopod 5

Aorcho

Gnathopod 2 larger than gnathopod 1, pereopod 3 longer than articles 1-5 of pereopod 5

26

26. Accessory Hagellum 3+ articulate Pseudeurystheus, Gammaropsis^

Accessory flagellum 0-2 articulate

Bonnierella, Segamphopus, Kermystheus, Podoceropsis, Audulla, Megamphopusf

27. I'ropod 3 with 2 rami equal to each other in size 28

Uropod 3 with less than 2 full rami 34

28. Gnathopod 2 larger than gnathopod 1 29

Gnathopod 1 larger than gnathopod 2 30

29. Accessory flagellum absent, inner plate of maxilla 2 heavily setose basomedially, inner plate

of maxilla 1 as long as palp, article 4 of maxillipedal palp vestigial C.oesia

• Antenna 1 unknown.

f These genera difficult to distinguish at piesent.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Accessory flagellum 3 + -articulate, inner plate of maxilla 2 not setose basomedially, inner

plate of maxilla 1 half as long as palp, article 4 of maxillipedal palp ordinary

Protomedeia

30. Article 5 of gnathopod 2 strongly bulbous Xenocheira

Article 5 of gnathopod 2 not bulbous 31

31. Male gnathopod 1 merochelate 32

Male gnathopod 1 not merochelate 33

32. Accessory flagellum absent Aoroides

Accessory flagellum present Aora

33. Article 5 of male gnathopod 1 with long posterior tooth, anterior margin of article 5 on

male gnathopod 2 heavily setose Lembopsis

Article 5 of male gnathopod 1 lobate but lacking tooth, anterior margin of article 5 on

male gnathopod 2 usually poorly setose Lembos

34. Uropod 3 with 2 unequally sized rami 35

Uropod 3 with one ramus or none 36

35. Cnathopods simple Haplocheira

Gnathopods subchelate Cheiriphoth

36. Gnathopod 1 larger than gnathopod 2 Paraoroides

Gnathopod 2 larger than gnathopod 1 37

37. Gnathopods simple, dactyl of gnathopod 2 vestigial and setose Kuphocheira

Gnathopods subchelate, dactyl of gnathopod 2 normal 38

38. Coxae 1-4 of varying sizes and shapes in male Cheiriphoth

Coxae 1-4 of similar sizes and shapes among themselves in male Microprotopus

KEY E (part of Key II)

1. Article 5 of male gnathopod 1 lacking processes, softly lobate 2

Article 5 of male gnathopod I carpochelate or bearing sharp process 3

2. Gnathopods fully subchelate Lemboides

Gnathopods poorly subchelate Rudilemboides

3. Antenna 2 attached to head level with eye 4

Antenna 2 attached to head posterior to level of eye 8

4. Uropod 3 with one ramus 5

Uropod 3 with 2 rami 6

5. Mandibular palp article 1 elongate Grandidierella

Mandibular palp article 1 not elongate Neomicrodeutopus

6. Rami of uropod 3 much shorter than peduncle Dryopoides

Rami of uropod 3 as long as or longer than peduncle 7

7. Articles 4-6 of gnathopod 2 poorly setose anteriorly, article 4 short

Hansenella and Microdeutopus

Articles 4-6 of gnathopod 2 heavily setose anteriorly, article 4 elongate

Microdeutopus (—Coremapus)

8. Inner ramus of uropod 3 shortened Acuminodeutopus

Rami of uropod 3 subequal to each other in length 9

9. Articles 5-6 of male gnathopod 2 poorly setose anteriorly Konatopus

Articles 5-6 of male gnathopod 2 strongly setose anteriorly 10

10. Male gnathopod 2 with well-defined palm, rami of uropod 3 nearly twice as long as

peduncle Amphideutopus

Male gnathopod 2 scarcely subchelate, rami of uropod 3 subequally as long as peduncle

Neomegamphopus

Diagnoses of Genera ing coxae 1-4) longer than broad and touching or

PROCEDURES.—Synonymies, type-species, number overlapping; (2) gnathopods: small and subchelate;

of species, and distributions are found in Barnard (3) uropod 3: peduncle short, about as long as

(1969a), except for those genera described since telson, rami longer than peduncle, thin, moderately

1965. setose, extending subequally; (4) telson: subThe term "ordinary" is used for brevity to des- circular, trapezoidal, rectangular, width and length

cribe the following conditions: (1) coxae: (mean- dimensions not extraordinary, telson generally less
NUMBER 151 15

than 1.5 times as broad as long or less than 1.5

times as long as broad, posteriorly truncate or

weakly concave or convex, with sets of parasagittal

(weakly apicolateral) setae, setules, spinules, or

nasiform humps.

Extraordinary characters are cited in a second

paragraph of pertinent diagnoses. These deviations

from a normal condition are of low frequency

sufficient to eliminate them from description in

every diagnosis, thus contributing to the brevity

of the arrangement. The deviant characters are

described in the following points: (1) head extremely elongate or eyes placed on pedunculate

ocular lobes; (2) mandibular palp article 1

elongate; (3) mandibular palp article 3 clavate;

(4) inner plate of maxilla 1 with medial setae;

(5) inner plate of maxilla 2 lacking medial setae;

(6) palp article 4 of maxilliped not unguiform and

bearing single apical nail; (7) anterior male coxae

of grossly variable shapes, if of significant taxonomic value; (8) pereopods prehensile; (9) one

or more rami of uropods 1-2 shortened or absent;

(10) any urosomites coalesced; (11) epimeron 3

bearing large tooth; (12) second antenna especially

thickened or toothed.

Certain genera of Ischyroceridae are included in

keys and diagnoses because of their similarity to

corophiids.

Acuminodeutopus J. L. Barnard

Article 3 of antenna 1 shorter than article 1;

accessory flagellum about 2-articulate; head deeply

recessed; coxae short but contiguous; gnathopod

1 enlarged, ordinary on female, on male carpochelate, article 6 thin, simple, shorter and narrower

than article 5, dactyl elongate, overlapping chela

of article 5, gnathopod 2 small but article 5 elongate, scarcely subchelate, poorly setose anteriorly;

uropod 3 with short peduncle, outer ramus longer

than peduncle, inner ramus short and scalelike oi

less than half as long as outer ramus; telson ordinary but armaments obsolescent.

Mandibular palp article 1 slightly elongate.

Aloiloi J. L. Barnard

Aloiloi J. L. Barnard, 1970:65-65.

TYPE-SPECIES.—A. nenue J. L. Barnard, 1970.

Article 3 of antenna 1 as long as article 1; accessory flagellum present; head deeply recessed:

coxae short but contiguous; gnathopod 1 enlarged,

in male subchelate, article 6 strongly broadened

and dominating article 5, dactyl fitting palm,

gnathopod 2 also enlarged slightly, subchelate,

article 6 dominating article 5; uropod 3 ordinary

but stout, telson ordinary.

Mandibular palp article 3 clavate; inner plate

of maxilla 1 medially setose; palp article 4 of maxilliped subclavate but with long nail; male coxa 1

slightly enlarged.

Ampelisciphotis Pirlot, new synonymy

Gaviota J. L. Barnard, 1958:87.

Article 3 of antenna 1 as long as or longer than

article 1; accessory flagellum absent; head deeply

recessed; coxae ordinary; gnathopods 1-2 ordinary;

uropod 3 uniramous, peduncle elongate, ramus

shorter than peduncle; telson ordinary.

Ocular lobes pedunculate.

Amphideutopus J. L. Barnard

[Article 3 of antenna 1 and accessory flagellum

unknown]; head deeply recessed; coxae short but

contiguous; gnathopods 1-2 of female of approximately equal size to each other, ordinary, poorly

setose anteriorly; gnathopods 1-2 of male both enlarged, gnathopod 1 carpochelate, article 6 narrower and much shorter than article 5, poorly

subchelate but dactyl elongate and overlapping

chela of article 5, posterior margins of articles 4-6

heavily setose, gnathopod 2 with articles 4—5

inflated, article 6 slightly dominating article 5,

subchelate, anterior margins of articles 5-6 heavily

setose; uropod 3 ordinary; telson ordinary.

Mandibular palp article 3 slightly clavate; inner

plate of maxilla 1 with 1 medial seta.

A or a Kr0yer

Article 3 of antenna 1 shorter than article 1;

accessory flagellum present; head poorly recessed;

coxae ordinary or slightly shortened; gnathopod

1 enlarged, ordinary or scarcely subchelate in female, merochelate in male, articles 5-6 linear,

palm absent, dactyl elongate, gnathopod 2 subchelate, moderately setose anteriorly; uropod 3

ordinary; telson ordinary.

Male coxa 1 enlarged or projecting askew.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Aorcho J. L. Barnard

Article 3 of antenna 1 as long as article 1;

accessory flagellum multiarticulate; head poorly to

moderately recessed; coxae very short but contiguous; gnathopods 1-2 slightly enlarged together,

equal to each other in size, ordinary and subchelate, article 6 weakly dominating article 5; uropod

3 ordinary but peduncle slightly flattened; telson

ordinary.

Urosome depressed.

A oroides Walker

Like Aora but accessory flagellum absent.

Camacho Stebbing

Article 3 of antenna 1 much shorter than article

1; accessory flagellum multiarticulate; head poorly

recessed; coxae very small, discontiguous; gnathopods 1-2 small, subequal in size to each other but

gnathopod 1 slightly the larger, ordinary, subchelate; peduncle of uropod 3 short, outer ramus

elongate and slender, inner ramus very short and

scalelike; telson ordinary.

Cerapopsis Delia Valle

Article 3 of antenna 1 as long as article 1; accessory flagellum absent; head deeply recessed;

coxae ordinary to elongate; female gnathopod 1

ordinary but poorly subchelate, dactyl elongate,

female gnathopod 2 ordinary and slightly enlarged,

article 6 dominating article 5; male gnathopod 1

highly elongate, almost simple, gnathopod 2 enlarged, weakly to strongly propodochelate, chela

nmiltifid, article 6 dominating article 5, dactyl

slightly overlapping protuberant palm; uropod 3

uniramous, peduncle elongate, ramus as long as

peduncle; telson ordinary.

Mandibular palp article 3 weakly clavate; ocular

lobes elongate; urosome depressed.

Cheiriphotis Walker

Article 3 of antenna 1 shorter than article 1;

accessory flagellum multiarticulate; head deeply

recessed; coxae ordinary but anterior coxae of

male of varying sizes and shapes; gnathopod 1

ordinary, gnathopod 2 enlarged especially in male,

article 6 heavily dominating article 5, palm sculptured, dactyl fitting palm, female gnathopod 2

scarcely enlarged, palm distinct; uropod 3 short,

peduncle flattened, rami shorter than peduncle,

inner ramus vestigial or absent; telson ordinary.

Mandibular palp article 3 weakly clavate; inner

plate of maxilla 1 medially setose; palp article 4

of maxilliped subclavate.

Chevalia Walker

Article 3 of antenna 1 slightly shorter than article

1; accessory flagellum 2-articulate; head poorly

recessed; coxae short, discontiguous; gnathopod 1

ordinary, gnathopod 2 enlarged in both sexes, subchelate, article 6 enlarged and dominating article

5, dactyl scarcely fitting palm, latter transverse to

oblique; uropod 3 short, rami scarcely as long as

peduncle but extending equally; telson ordinary.

Urosomites 1-2 coalesced; inner plate of maxilla

1 medially setose; palp article 4 of maxilliped

clavate.

Chevreuxius Bonnier (Ischyroceridae)

Article 3 of antenna 1 elongate but slightly

shorter than article 1; accessory flagellum 2-articulate; head poorly recessed; coxae very short and

discontiguous; male gnathopod 1 enlarged and

carpochelate, article 6 very short and narrower

than article 5, sculptured but subchelate, dactyl

overlapping palm; gnathopod 2 ordinary, small,

moderately setose; uropod 3 uniramous, peduncle

flattened and expanded, ramus subequal to peduncle in length; telson ordinary.

Palp article 4 of maxilliped slightly shortened;

uropod 2 uniramous.

Concholestes Giles

Article 3 of antenna 1 subequally as long as

article 1; accessory flagellum absent; head poorly

recessed; coxae very short, discontiguous; gnathopod

1 ordinary, poorly subchelate, gnathopod 2 slightly

enlarged, article 6 dominating article 5, subchelate,

palm sculptured; uropod 3 composed only of

setose peduncle; telson apparently coalesced with

urosomite 3, thus telson enlarged, armaments obso-
NUMBER 151 17

lete [but Giles describes 6 pleonites and telson as

short, squamous and semilunar].

PAntenna 2 slightly thickened; Puropod 2 uniramous; body subcylindrical, urosome depressed.

Corophium Latreille

Article 3 of antenna 1 shorter than article 1;

accessory flagellum absent; head deeply recessed;

coxae very short, discontiguous; gnathopods alike

between the sexes, gnathopod 1 generally ordinary

but heavily setose, article 5 elongate, palm transverse, gnathopod 2 scarcely enlarged, article 4 enlarged, inflated, incipiently merochelate but extended and fused distally along posterior edge of

article 5, heavily setose posteriorly, article 6 longer

than article 5, slender, simple, dactyl elongate;

uropod 3 uniramous, short, peduncle and ramus

flattened and broad, ramus heavily setose; telson

softly triangular, slightly enlarged, armaments

obsolescent.

Antenna 2 thickened and elongate but sexually

dimorphic, male with weak teeth on articles 4-5;

mandibular palp 2-articulate, geniculate between

articles 1-2; urosomites depressed, articulate or

coalesced; body weakly subcylindrical; ocular lobes

weakly pedunculate.

Dryopoides Stebbing

Article 3 of antenna 1 shorter than article 1;

accessory flagellum 2-articulate; head poorly recessed; coxae ordinary; gnathopod 1 of female

slightly enlarged, gnathopods 1-2 subchelate,

gnathopod 2 weakly setose anteriorly, male gnathopods 1-2 enlarged, gnathopod 2 larger and longer

than gnathopod 1, both subchelate; uropod 3 with

weakly elongate and broadened peduncle, rami

extending equally but tiny, half or less as peduncle,

rami disjunct from each other on attaching to

peduncle; telson enlarged, corresponding with dorsal evanescence of urosomite 3.

Note asynchrony of male and female gnathopods.

Gammaropsis Liljeborg

(Including Audulla Chevreux, Bonnierella Chevreux (also

Ischyroceridac), Kermystheus J. L. Barnard, Megamphopus

Norman, Podoceropsis Boeck, Pseudeurystheus Schellenberg,

Segamphopus J. L. Barnard, and unnamed subdivision)

Article 3 of antenna 1 as long as article 1; accessory flagellum multiarticulate (typical) to absent;

head deeply recessed but occasionally recessment

diminished; coxae ordinary to sh6rt but contiguous; gnathopod 1 ordinary or article 5 elongate

(Pseudeurystheus and Segamphopus); gnathopod 2

enlarged especially in male, ordinary, with article

6 dominating article 5 or latter also elongate and

enlarged (Megamphopus), article 6 occasionally

elongate, palm extended along posterior margin

and dactyl strongly elongate (Podoceropsis); uropod 3 ordinary or with peduncle elongate and rami

subequal to peduncle or slightly shorter than

peduncle; telson ordinary, but occasional species

with deep posterior recessment.

Mandibular palp article 3 often clavate; inner

plate of maxilla 1 variable, often with medial

setae; palp article 4 of maxilliped variable, ordinary, or in various degrees of clavateness, usually

setose apically.

This is the most diverse and perhaps the most

primitive genus of Corophiidae. The several genera

and subgenera appended to Gammaropsis are

joined by intergrading species or are characterized

by apparently insignificant anomalies. Where one

good character might be used to define a subgenus

in one cluster of species, this character is found

to be variable in a second cluster based on other

criteria. The accessory flagellum occurs in all stages

from multiarticulate to 3-articulate to 2-articulate,

1-articulate, and absent. Clusters based on the alternatives of accessory flagellum appear definitely

to be polyphyletic as based on gnathopods and

uropod 3. The elongation of article 5 on gnathopods of Megamphopus, of article 5 on gnathopod

1 of Pseudeurystheus, or the enlargement and elongation of palm and dactyl on Podoceropsis are connected to the central concept of Gammaropsis by

various intergrades.

The named subdivisions of Gammaropsis are

briefly diagnosed below but the diagnoses only encompass the central species of those clusters and

serve to identify the concepts as stated in earlier

years. The writer is impressed, without adequate

study of any evolutionary sequence, that the trends

to loss of accessory flagellum and modifications of

gnathopods occur in parallel within several world

biomes and so asynchronously as to defy any application of Linnean didactyly to the several polyphyletic systems.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

AmiuIhi Chevreux: Like Gammaropsis but flagellum of male antenna 2 stout, male gnathopod 2

with thumb on transverse palm.

Bonnierella Chevreux: Assignable to Ischyroceriilae because of elongate peduncle of uropod 3 and

slightly shortened rami but not highly distinct

from various species ot Gammaropsis; accessory

Hagellum 2-articulate, article 2 of pereopods 3-5

linear, anterior and posterior margins parallel;

deep-sea.

Kermystheus J. L. Barnard: Like Gammaropsis

but accessory Hagellum composed only of short

scale; peduncle of uropod 3 elongate; gnathopods

of 2 known species highly diverse, these species

probably not closely related. Podoceropsis dubia

Shoemaker and A', ociosus (J. L. Barnard) more

closely related to each other than K. ociosus and

A', kermadeci, yet P.d. and K.o. presently segregated subgenerically.

Mcgamphopus Norman: Like Gammaropsis but

accessory Hagellum usually 1-2 articulate; male anterior coxae often of varying sizes and shapes but

occasionally static; male gnathopod 1 often extraordinary, enlarged nearly to size of gnathopod 2

and bearing other similarities to gnathopod 2; or

male gnathopod 2 relatively reduced in size to fit

enlarged gnathopod 1; or male gnathopod 2 with

article 5 dominating article 6 in length or length

plus width. New list of species, others rejected:

cor mi tus Norman, longicornis Chevreux; (note

that pachypus Schellenberg keys to Protomedcial).

Podoceropsis Boetk: Like Gammaropsis but accessory Ragellum absent (typical) or 1-2 articulate; article (*> of male gnathopod 2 dominating

short article 5, and palm usually extended along

posterior margin of article f> and dactyl elongate;

peduncle of uropod 3 usually elongate. New list

of species, other rejected: angulosa Chevreux, dubia

Shoemaker, elephantis K. H. Barnard, inaequistylis Shoemaker, insignis Chilton, lindahli Hansen,

longidactylus Chevreux, nitida Stimpson, pusilla

Chevreux, sophiae Boeck. Latter two species with

reasonably typical gnathopod of Gammaropsis.

Pseudeurystheus Schellenberg and Segamphopus

|. L. Barnard: Like Gammaropsis but accessory

Ragellum 3-articulate or 1-2 articulate, article 5

of male gnathopod 1 at least 1.5 times as long as

article 6, article 5 of male gnathopod 2 also elongate.

Unnamed division: Like Gammaropsis but telson deeply excavate posteriorly, gnathopod 1 like

Pseudeurystheus, gnathopod 2 like Podoceropsis:

chilloni (Thomson), haswelli (Thomson), and possibly anomalus Chevreux.

Examples of ordinary Gammaropsis with 0-2

articulate accessory Hagellum: ctenurus Schellenberg, georgianus Schellenberg, kcrguelcni Schellenberg, longicornis Walker, martesia J. L. Barnard,

palmatus Stebbing and Robertson, similis Schellenberg.

Example of ordinary Gammaropsis with gnathopod 2 like Podoceropsis: lobatus Chevreux.

Examples of elongate peduncle on uropod 3:

exsertipes Stebbing, spinosus Shoemaker.

Unnamed division: Megamphopus effrenus J. L.

Barnard; keying to vicinity of Aloiloi or Pseudomegamphopus because of slight enlargement of

gnathopod 1, but otherwise with gnathopods resembling certain species of Megamphopus; possibly a case of axial reversal of gnathopods from

megamphopus ancestor but anterior male coxae

static.

Goesia Boeck

Article 3 of antenna 1 shorter than article 1;

accessory Hagellum absent; head deeply recessed

[but genus placed near Protomedeia in Figure 1];

coxae ordinary; gnathopod 1 elongate, thin, heavily setose, article 6 slightly dominating article 5 or

these articles of equal length, palm transverse and

short; gnathopod 2 slightly enlarged, heavily setose,

article 5 dominating article 6, palm weak, bearing

one enlarged spine; uropod 3 short, rami as long

as peduncle, extending equally, heavily setose; telson ordinary.

Palp article 1 of maxilliped very small, short,

apically setose; epimeron 3 with posteroventral

tooth.

Grandidierella Coutiere

Article 3 of antenna 1 shorter than article 1;

accessory Hagellum 1-2 articulate; head poorly recessed; coxae short, scarcely contiguous or slightly

discontiguous; female gnathopods small, ordinary

but moderately setose, gnathopod 1 slightly the

larger, male gnathopod 1 enlarged, carpochelate,

article 6 slender, shorter and narrower than article

5, simple or weakly subchelate and sculptured,
NUMBER 151 19

dactyl elongate, gnathopod 2 ordinary, often heavily setose; uropod 3 uniramous, peduncle usually

platelike, ramus much longer than peduncle; telson

ordinary but occasionally short, armaments possibly

obsolescent.

Mandibular palp article 1 elongate on typespecies but not necessarily on other species; palp

article 4 of maxilliped short, with 2 or more apical

nails or setae; urosome depressed.

See Neomicrodeutopus.

subchelate, article 6 dominating article 5, palm

with large excavation and defining tooth; uropod

3 uniramous, peduncle slightly elongate and moderately expanded, ramus equal to or shorter than

peduncle, heavily setose apically; telson and urosomite 3 apparently coalesced.

Urosomites 1-2 coalesced but free from telson

and urosomite 3; mandibular palp article 1 elongate or not; inner plate of maxilla 2 poorly to

strongly setose medially; 3 species, east Asia.

Haplocheira Haswell

Article 3 of antenna 1 shorter than article 1; accessory flagelliim 2-f articulate; recessment of head

apparently variable; coxae ordinary; gnathopods

simple or with tiny transverse palm, slender, elongate, apparently similar between the sexes, gnathopod 1 slightly the stouter, type-species with weakly

falcate article 6, gnathopod 2 the longer, gnathopods heavily setose, dactyls of gnathopods elongate or not, setose or not; uropod 3 short, outer

ramus as long as peduncle, inner ramus very short

or vestigial; telson ordinary.

Inner plate of maxilla 1 heavily setose medially;

palp article 4 of maxilliped blunt or thick and

apically setose.

Isaea Milne Edwards

Article 3 of antenna 1 as long as article 1; accessory flagellum multiarticulate; head of moderate

recessment (neither deep nor shallow); coxae ordinary; gnathopod 1 ordinary, gnathopod 2 enlarged,

article 6 especially enlarged and dominating article

5, palm oblique, weakly sculptured, dactyl fitting

palm, female gnathopod 2 ordinary but less enlarged; uropod 3 ordinary but peduncle slightly

elongate; telson ordinary.

Mandibular palp article 3 weakly clavate or not;

palp article 4 of maxilliped elongate but apically

blunt and setose; pereopods 3-5 prehensile; tubespinning glands of pereopods apparently absent.

Kamaka Derzhavin

Article 3 of antenna 1 slightly shorter than article 1; accessory flagellum absent; head deeply

recessed; coxae ordinary but coxa 1 enlarged;

gnathopod 1 ordinary, male gnathopod 2 enlarged,

Konatopus J. L. Barnard

Konalopus J. L. Barnard, 1970:70-72.

TYPE-SPECIES.—K. paao J. L. Barnard, 1970.

Article 3 of antenna 1 as long as article 1; short

accessory Hagellum present; head deeply recessed;

coxae short but contiguous; gnathopod 1 enlarged,

of female ordinary, of male carpochelate and

weakly propodochelate, article 6 shorter and narrower than article 5, dactyl elongate, overlapping

chela of article 5; gnathopod 2 ordinary, poorly setose anteriorly; uropod 3 ordinary, rami extending

subequally but outer slightly the shorter; telson

ordinary.

Mandibular palp article 3 clavate; inner plate of

maxilla 1 with medial setae; palp article 4 of maxilliped subclavate, apically setose.

Kuphocheira K. H. Barnard

[Article 3 of antenna 1 unknown]; accessory

flagellum absent; head apparently poorly recessed;

coxae short but contiguous; gnathopods 1-2 simple, similar between the sexes, heavily setose, small,

slender, article 5 longer to much longer than article

(), dactyl of gnathopod 2 reduced and setose; uropod 3 uniramous, ramus shorter than elongate

peduncle; telson circular, armaments obsolete.

Inner plate of maxilla 1 medially setose; epimeron 3 with tooth.

Lemboides Stebbing

Article 3 of antenna 1 shorter than article 1;

accessory flagellum 2-4 articulate; head poorly recessed; coxae short but contiguous; typical gnathopod 1 of female slightly enlarged, ordinary, of male

enlarged, article 5 tumid and dominating article 6,
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

latter narrower and shorter than article 5, subchelate, dactyl fitting palm; other species with article 6 as long as 5, bearing large thumb and dactyl

overlapping palm or gnathopod 1 scarcely larger

than gnathopod 2; gnathopod 2 ordinary, strongly

setose, in male article 5 usually dominating article

6, in female article 6 elongate and dominating

article 5; uropod 3 ordinary, rami elongate, outer

slightly the shorter; telson ordinary.

South African species possibly generically distinct because of odd gnathopods.

Lembopsis Pearse

Article 3 of antenna 1 shorter than article 1;

accessory flagellum multiarticulate; head poorly

recessed; coxae short but contiguous; gnathopod 1

enlarged, weakly carpochelate, thus article 5 with

1 or more posterior teeth but article 6 enlarged

and dominating article 5, subchelate, palm sculptured, dactyl slightly overlapping palm; gnathopod

2 of medium size, subchelate, heavily setose; female

gnathopod 1 slightly enlarged, ordinary; uropod 3

ordinary; telson ordinary.

Mandibular palp article 1 elongate.

Lembos Bate

Article 3 of antenna 1 shorter than article 1;

accessory Hagellum present; head poorly recessed;

coxae short but contiguous; gnathopod 1 enlarged,

weakly in female, generally article 6 inflated and

dominating article 5, subchelate, palm occasionally sculptured, dactyl fitting or overlapping palm;

gnathopod 2 small, subchelate, weakly to strongly

setose anteriorly; uropod 3 ordinary; telson ordinary.

Male coxa 1 often enlarged or askew.

Leptocheirus Zaddach

Article 3 of antenna 1 shorter than article 1;

accessory Hagellum multiarticulate; head moderately or occasionally deeply recessed [but placed

near Protnmedeia in Figure 1]; coxae 2-5 very long

and overlapping, coxa 1 variable, usually very small

and hidden by enlarged coxa 2 but occasionally

coxa 1 slightly enlarged and partially visible;

gnathopods 1-2 subequal to each other in size, not

enlarged but heavily setose, gnathopod 1 ordinary

but with inflated article 3 and heavily setose posteriorly, gnathopod 2 slender, article 5 elongate

and dominating article 6, latter narrow but short,

palm obsolescent, dactyl short; uropod 3 short, peduncle flattened, rami longer than peduncle and

extending equally; telson ordinary.

Mandibular palp article 1 slightly elongate.

Microphotis Ruffo

Article 3 of antenna 1 as long as article 1; accessory flagellum absent; head probably deeply recessed; coxae ordinary to shortened; gnathopod 1

ordinary; male gnathopod 2 slightly enlarged, article 6 dominating article 5, palm with small thumb,

dactyl overlapping thumb; uropod 3 uniramous,

peduncle slightly elongate, ramus shorter than peduncle, apically setose; telson ordinary.

Mandibular palp article 3 clavate.

Microdeutopus Costa

(Including Hansenella Chevreux and Coremapus Norman—

see Myers, 1969:96)

Article 3 of antenna 1 shorter than article 1;

accessory flagellum present; head poorly recessed;

coxae short but contiguous, male or female coxa 1

often elongate and pointed; gnathopod 1 enlarged,

male member strongly carpochelate, article 5 thin

to inflated and with one or more teeth, article 6

narrow and shorter than article 5, poorly subchelate

or simple, occasionally sculptured, dactyl elongate,

overlapping chela of article 5, gnathopod 2 ordinary or occasionally enlarged and thickened, subchelate or palm sculptured, weakly to moderately

setose anteriorly, female gnathopod 1 ordinary but

slightly enlarged; uropod 3 ordinary but peduncle

slightly elongate; telson ordinary.

Hansenella Chevreux: Like Microdeutopus but

female gnathopods like those of male. Coremapus

Norman: Synonym of Microdeutopus, article 4 of

gnathopod 2 highly elongate and heavily setose.

Microprotopus Norman

Article 3 of antenna 1 shorter than article 1;

accessory flagellum 2-articulate; head deeply to

moderately recessed; coxae elongate; gnathopod 1
NUMBER 151 21

ordinary; gnathopod 2 enlarged, ordinary in female, especially enlarged in male, article 6 strongly

dominating article 5, palm and posterior margin

of article 6 largely confluent, sculptured, dactyl

elongate but fitting enlarged palmer margin; uropod 3 uniramous, peduncle of medium elongation,

ramus longer than peduncle; telson ordinary.

Mandibular palp article 3 subclavate; inner plate

of maxilla 1 medially setose.

erately setose, article 5 dominating article 6, palm

transverse, dactyl fitting palm; uropod 3 uniramous, peduncle expanded, ramus much longer than

peduncle; telson ordinary.

Palp article 4 of maxilliped stubby and apically

setose.

Probably synonymous with Grandidierella as

some species of latter bearing short mandibular

palp article 1.

Neohela Smith

Article 3 of antenna 1 as long as article 1; accessory ilagellum multiarticulate; head poorly recessed; coxae very short, discontiguous; gnathopods

similar between the sexes, both pairs enlarged,

gnathopod 1 slightly the larger, both ordinary, subchelatc, palm of gnathopod 1 weakly sculptured;

uropod 3 uniramous, peduncle short, ramus elongate, weakly setose; telson enlarged, irregularly

triangular, armaments obsolete.

Body thin, cylindrical, urosome depressed.

Neomegamphopus Shoemaker

Article 3 of antenna 1 as long as article 1; accessory flagellum 2-articulate; head deeply recessed;

coxae ordinary; female gnathopods 1-2 small, ordinary, similar to each other; male gnathopod 1 enlarged, heavily setose, carpochelate, article 6 thin,

shorter and narrower than article 5, simple but

sculptured, dactyl elongate, overlapping chela of

article 5, gnathopod 2 heavily setose, highly elongate and slender, simple; uropod 3 ordinary but

subequally extending rami scarcely longer than

peduncle; telson ordinary.

Mandibular palp article 3 subclavate; inner plate

of maxilla 1 with 2 medial setae; palp article 4 of

maxilliped apically setose.

Neomicrodeutopus Schellenberg

Article 3 of antenna 1 shorter than article 1;

accessory flagellum 1- (?2-) articulate; head poorly

recessed; coxae short, discontiguous; [female unknown]; male gnathopod 1 enlarged, carpochelate,

article (i slender, shorter and narrower than article

5, weakly sculptured, dactyl elongate and reaching

base of chela on article 5, gnathopod 2 stout, modParacorophium Stebbing

Article 3 of antenna 1 shorter than article 1;

accessory flagellum absent; head deeply recessed;

coxae ordinary; gnathopod 1 ordinary but heavily

setose, gnathopod 2 of male enlarged slightly, freely

merochelate but article 6 also enlarged and either

chelate or subchelate, appendage heavily setose,

female article 6 narrow, poorly subchelate; uropod

3 short, outer ramus subequally as long as peduncle, inner ramus half as long as outer ramus, both

apically setose; telson small, broad, smooth.

Male antenna 2 slightly thickened basally, occasionally with weak distal peduncular tooth; mandibular palp article 1 slightly elongate, article 3

weakly clavate; inner plate of maxilla 1 finely

setose distally; palp article 4 of maxilliped long or

short, apically blunt and setose; urosome depressed.

Paraoroides Stebbing

Article 3 of antenna 1 shorter than article 1;

accessory flagellum vestigial; head recessment unknown; coxae ordinary; gnathopods 1-2 of female

equal to each other in size, weakly subchelate,

poorly setose anteriorly, male gnathopods 1-2

slightly enlarged, gnathopod 1 the larger, subchelate, palm with thumb, dactyl fitting palm, gnathopod 2 subchelate, moderately setose anteriorly; uropod 3 uniramous, short, ramus subequally as long

as peduncle; telson ordinary.

Parunciola Chevreux

Article 3 of antenna 1 longer than article 1;

accessory flagellum multiarticulate; head deeply recessed; coxae short but contiguous; male unknown;

female gnathopods 1-2 subequal in size to each

other, gnathopod 1 slightly the larger, otherwise
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ordinary; uropod 3 uniramous, ramus much longer

than peduncle; telson ordinary.

Mandibular palp article 3 ciavate; inner plate of

maxilla 1 medially setose, urosome depressed.

Photis Kr0yer and Cedrophotis J. L. Barnard

Article 3 of antenna 1 as long as article 1; accessory flagellum absent or vestigial; head deeply recessed; coxae ordinary, often strongly elongate;

gnathopod 1 ordinary; gnathopod 2 enlarged,

weakly so in female and with weakly differentiated

palm, strongly so in male, article 6 large and dominating article 5, palm often heavily sculptured,

transverse or oblique, dactyl fitting or weakly overlapping palm; uropod 3 with elongate peduncle,

outer ramus as long as peduncle, inner ramus half

or less as long as peduncle; telson ordinary.

Mandibular palp article 3 ciavate.

Cedrophotis Barnard, 1967: Like Photis but inner ramus of uropod 3 half (as against one-third

or less) as long as outer ramus; inner lobes of lower

lip partially fused together; upper lip with deep

incision.

Protomedeia Kroyer and Cheirimedeia

J. L. Barnard

Article 3 of antenna 1 shorter than article 1;

accessory flagellum multiarticulate; head poorly to

moderately recessed; coxae ordinary to short, occasionally partly discontiguous; gnathopod 1 ordinary, gnathopod 2 of female ordinary, subequal in

size to gnathopod 1, of male enlarged, article 5

usually as long or longer than and dominating

article (>, or articles 5-6 inflated, palm ordinary or

sculptured and narrowed or transverse, dactyl fitting palm or strongly overlapping; uropod 3 ordinary but peduncle slightly elongate and rami equal

to or longer than peduncle; telson ordinary.

Inner plate of maxilla 1 with medial setae; inner plate of maxilla 2 lacking medial setae.

Cheirimedeia J. L. Barnard: Like Protomedeia

but inner ramus of uropod 3 half as long as outer

ramus.

Pseudomegamphopus Myers

Pseudomegamphopus Myers, 1968:527.

TYPE-SFF.CIES.—P. barnardi Myers, 1968.

Article 3 of antenna 1 as long as article 1; accessory flagellum 2-articulate; head deeply recessed;

coxae ordinary; gnathopod 1 enlarged, article 6

heavily dominating article 5, palm oblique, dactyl

fitting palm, gnathopod 2 ordinary but slender and

poorly subchelate; female unknown; peduncle of

uropod 3 slightly elongate, rami extending equally

and about as long as or slightly longer than peduncle; telson ordinary.

Rildardanus J. L. Barnard, revised

Rildardanus Barnard, 1969b: 197.

TYPE-SPECIES.—R. tros Barnard, 1969b.

Article 3 of antenna 1 shorter than article 1;

accessory llagellum 1-articulate, scalelike; head

deeply recessed; coxae very short, discontiguous;

female unknown; male gnathopod 1 enlarged, article 6 dominating article 5, palm very oblique and

defined by thumb, dactyl overlapping thumb, gnathopod 2 slender, parachelate, moderately setose

anteriorly; uropod 3 composed only of setose peduncle, or bearing vestigial ramus, peduncle unexpanded; telson enlarged, suborbicular, armaments ordinary.

Antenna 2 thickened and heavily sculptured; inner rami of uropods 1-2 vestigial; epimeron 3 with

tooth; urosome depressed.

Diagnosis revised to include Unciola laminosa

Pearse, uropod 3 bearing vestigial ramus; this species bridges Rildardanus and Unciola, differing

from Unciola in vestigial accessory flagellum, unexpanded peduncle of uropod 3, vestigial ramus,

heavily toothed antenna 2, shortened inner rami

of uropods 1-2.

Rudilemboides J. L. Barnard

Article 3 of antenna 1 shorter than article 1;

accessory flagellum multiarticulate; head deeply recessed; coxae short but contiguous; gnathopods 1-2

of female approximately equal to each other in

size, gnathopod 1 ordinary, gnathopod 2 scarcely

subchelate, moderately setose anteriorly, male gnathopod 1 enlarged, article 5 elongate and inflated,

article 6 shorter and more slender than article 5,

scarcely subchelate, dactyl short but overlapping

imaginary palm, gnathopod 2 slightly thicker than

but similar to gnathopod 2 of female, anterior

setae poorly developed; uropod 3 ordinary; telson

ordinary.

Inner plate of maxilla 1 medially setose.
NUMBER 151 23

Siphonoecetes Kr0yer

Article 3 of antenna 1 as long as article 1; accessory flagellum absent; head deeply recessed; coxae

short and discontiguous; gnathopod 1 ordinary,

gnathopod 2 ordinary, slightly enlarged, article 5

dominating article 6, palm very oblique, dactyl

fitting or slightly overlapping palm; uropod 3

short, uniramous, peduncle flattened, ramus shorter

than peduncle, apically setose; telson enlarged, subcircular, armaments obsolescent.

Ocular lobes weakly pedunculate; antenna 2

slightly or strongly thickened in male; mandibular

palp 1-articulate; uropod 2 with 1-2 rami; body

thin and cylindrical, urosome depressed.

Uncinotarsus L'Hardy and Truchot

Article 3 of antenna 1 shorter than article 1;

accessory flagellum 2-articulate; head poorly recessed; coxae very short, discontiguous; gnathopod

1 enlarged, article 6 heavily dominating article 5,

subchelate, dactyl fitting palm, gnathopod 2 smaller

but similar, both gnathopods poorly setose anteriorly; uropod 3 lacking rami; telson ordinary.

Mandibular palp article 1 elongate; outer plates

of lower lip weakly incised; inner plate of maxilla

2 naked medially; palp article 4 of maxilliped

short, blunt, apically setose; uropod 2 with one

ramus; epimeron 3 with tooth.

Unciola Say, revised

Article 3 of antenna 1 as long as article 1; accessory flagellum multiarticulate; head deeply recessed; coxae very short, discontiguous; gnathopods

similar between the sexes, gnathopod 1 enlarged,

article 6 inflated and dominating article 5, subchelate, palm strongly oblique, often with thumb, dactyl fitting palm; gnathopod 2 of ordinary small

form, moderately setose anteriorly, subchelate or

simple or weakly parachelate, occasionally article

5 elongate; uropod 3 uniramous, small, peduncle

flattened and expanded medially and extending

distally alongside ramus, latter shorter than peduncle, apically setose; telson enlarged, subcircular,

armaments generally present.

Body thin, subcylindrical, urosome depressed;

antenna 2 slightly to strongly thickened, epimeron

3 with posteroven^al tooth.

Unciola laminosa Pearse transferred to Rildarclanus.

Unciolella Chevreux

Article 3 of antenna 1 as long as article 1; accessory flagellum 2-articulate; head deeply recessed;

coxae very short, discontiguous; gnathopod 1

slightly enlarged, ordinary, gnathopod 2 ordinary

[subadult]; uropod 3 uniramous, peduncle slightly

flattened, ramus slightly longer than peduncle; telson ordinary.

Palp article 4 of maxilliped slightly blunted and

apically setose.

Xenocheira Chevreux

Article 3 of antenna 1 shorter than article 1;

accessory flagellum 2-articulate; head moderately

recessed; coxae ordinary; gnathopod 1 slightly enlarged, ordinary but heavily setose, gnathopod 2

with articles 1-5 inflated and heavily setose, article

(i slender, elongate, poorly subchelate; uropod 3

ordinary but peduncle slightly elongate; telson ordinary.

ISCHYROCERIDAE

DIAGNOSIS.—Corophioidean with ordinary urosomite 1; peduncle of uropod 3 elongate relative to

shortened rami, inner ramus, if present, not apically setose, bearing one small apical spine or not,

when uropod 3 bearing only one ramus that ramus

also not setose; outer ramus lacking terminal setae,

bearing apical spine or not, apicolateral margin

often with small denticles; pereopods glandular.

Diagnoses of Genera

The term "ordinary" is used for gnathopod 2

when this appendage is larger than gnathopod 1,

subchelate, with article 6 dominating article 5.

Bathyphotis Stephensen

Accessory flagellum multiarticulate; article 5 of

gnathopod 1 longer than article 6; coxa 1 more

than three-fourths as long as coxa 2, coxa 5 subequal in length to coxa 6, coxa 1 excavate pos-
2 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Genera of Ischyroceridae

(Note that Dryopoides and Paradryope are included here for comparisons)

1. Uropod 3 with one ramus or rami highly vestigial 2

Uropod 3 with 2 rami 6

2. Uropod 2 with one ramus 3

Uropod 2 with 2 rami 5

3. Male gnathopod 1 carpochelate, rsmui of uropod 3 as long as peduncle Chevreuxius

Male gnathopod 1 subchelate, ramus or rami of uropod 3 much shorter than peduncle . 4

4. Pleopod 3 with one ramus Runanga

Pleopod 3 with 2 rami Cerapus

5. Inner rami of uropods 1-2 less than half as long as outer rami Pseudericthonius

Rami of uropods 1-2 subequal to each other Ericthonius

6. Coxae minute, serially discontiguous Bogenfelsia

Coxae sufficiently large to be serially contiguous 7

7. Gnathopod 1 larger than gnathopod 2 8

Gnathopod 2 larger than gnathopod 1 or gnathopods of similar size 9

8. Pleonite 6 evanescent dorsally, article 3 of antenna 1 shorter than article 1, accessory tlagellum vestigial Dryopoides

Pleonite 6 visible dorsally, article 3 of antenna 1 longer than article 1, accessory flagellum

2 articulate Paradryope

9. Article 2 of pereopods 3-5 linear, article 4 maxillipedal palp clawlike, longer than

article 3 Bonnierella

Article 2 of pereopods 3-5 subovate or broadly rectangular, article 4 of maxillipedal palp

shorter than article 3, blunt or subconical (setose) 10

10. Article 5 of gnathopod 1 much longer than article 6 11

Article 5 of gnathopod 1 as long as or shorter than article 6 13

11. Spines on outer plate of maxilla 1 reduced to four, article 4 of pereopod 3 twice as wide

as on pereopods 4-5 Bathypkotis

Spines on outer plate of maxilla 1 nine or more, article 4 of pereopod 3 not wider than

on pereopods 4-5 12

12. Outer ramus of uropod 3 with 1-3 wirelike marginal setae Ventojassa

Outer ramus of uropod 3 lacking marginal setae Pseudischyrocerus

13. Accessory flagellum a small scale, articulate or not Parajassa

Accessory flagellum elongate 14

14. Coxa 5 subet]ual to coxa 6 in length, coxa 1 half as long as coxa 2 Microjassa*

Coxa 5 much longer than coxa 6, coxa 1 three-fourths as long as coxa 2 15

15. Pereopods subchelate Isaeopsis

Pereopods simple 16

16. Outer raraus of uropod 3 with basally immersed, hooked terminal spine, distolateral margin

of ramus with one to three large, reverted, sharp scales or several denticles, male gnathopod 2 usually with large tooth on proximal part of palm and with tooth on distal

part of palm Jassa

Outer ramus of uropod 3 with or without basally immersed terminal spine, never hooked,

distolateral margin of ramus with 0-7+ perpendicular, blunt, usually extremely small

denticles, male gnathopod 2 lacking proximal palmar tooth but often with distal palmar

process Ischyrocerus

'Dryopoides male of Stebbing (1910) his near this couplet but various characters are unknown. Male gnathopod 2 is not like that of ischyrocerids.

teriorly; uropod 3 with elongate peduncle, rami as broad as on pereopods 4-5; urosomite 1 with

short, weakly spinose on margins, outer ramus dorsal teeth,

hooked apically.

Outer plate of maxilla 1 with 4 spines (compare Bogenfelsia J. L. Barnard

Pseudischyrocnus); article 4 of pereopod 3 twice [Accessory flagellum unknown]; article 5 of gna-
NUMBER 151 25

thopod 1 longer than article 6; gnathopod 2 ordinary; coxae minute, serially discontiguous; rami of

uropod 3 much shorter than peduncle [apices unknown microscopically].

Bonnierella Chevreux

Accessory flagellum 2-articulate, elongate; article

6 of gnathopod 1 equal to or longer than article

5; gnathopod 2 ordinary; coxa 1 as long as coxa

2, coxa 5 slightly longer than coxa 6; rami of

uropod 3 shorter than peduncle, each with weak,

immersed apical spinule.

Article 2 of pereopods 3-5 narrow and linear;

article 4 of maxillipedal palp unguiform, longer

than article 3.

Cerapus Say

Accessory flagellum absent; article 6 of gnathopod 1 equal to or shorter than article 5; gnathopod 2 very large in male, carpochelate, article 5

multicuspidate, article 6 simple, shorter and more

slender than article 5, dactyl large and overlapping

half length of article 5; coxae short, discontiguous

serially; uropod 3 small, with one tiny vestigial

ramus bearing hook, inner ramus possibly present

on one species.

Rostrum thornlike; uropod 2 with 1 ramus; telson very broad and short, often with 2 humps bearing thorn spines or short scales; body cylindrical,

urosome depressed.

Chevreuxius Bonnier

Accessory flagellum 2-articulate; male gnathopod

1 enlarged, carpochelate, article 6 stout and subchelate or sculptured but narrower and shorter

than article 5; gnathopod 2 small, article 6 longer

than article 5; coxae short, discontiguous serially;

uropod 3 short, bearing 1 ramus as long as peduncle and with 2 apical setae.

Uropod 2 with 1 ramus; body subcylindrical,

urosome depressed.

Dryopoides Stebbing

See Corophiidae.

Ericthonius Milne Edwards

Accessory flagellum absent; article 6 of gnathopod 1 shorter than article 5; male gnathopod 2 enlarged, carpochelate, article 6 simple or sculptured

but narrower and shorter than article 5, dactyl

overlapping carpal thumb; coxae short, scarcely or

fully discontiguous serially; uropod 3 elongate,

single ramus shorter than peduncle and weakly

hooked apically.

Telson broad, short, bearing 2 knobs covered

with thorn spines or scales; body subcylindrical,

urosome depressed.

Isaeopsis K. H. Barnard

Accessory flagellum 1-articulate; article 6 of

gnathopod 1 longer than article 5; gnathopod 2

ordinary; coxa 1 "small," coxa 5 longer than coxa

6; uropod 3 with elongate peduncle, rami very

short, thin, outer apically hooked and weakly

toothed.

Pereopods prehensile.

Ischyrocerus Krpyer

Accessory Hagellum 2-articulate, elongate; article

6 of gnathopod 1 longer than article 5; gnathopod

2 ordinary, article 6 of male lacking large posteroproximal tooth (compare Jassa) but often with

distal tooth near base of dactyl; coxa 1 more than

three-fourths as long as coxa 2, coxa 5 longer than

coxa (>; uropod 3 slightly to fully elongate, rami

slightly shorter than peduncle, outer with apical

hook or not, small lateral denticles, no apical immersed spine.

Jassa Leach

Accessory flagellum 2-articulate, elongate; article

(> of gnathopod 1 longer than article 5; gnathopod

2 ordinary, article 6 of male bearing large posteroproximal tooth (compare Ischyrocerus); coxa 1

more than three-fourths as long as coxa 2, coxa 5

longer than coxa 6; uropod 3 slightly to fully

elongate, outer ramus with immersed and hooked

apical spine, distolateral margin with 1-3 large,

sharp reverted scales or denticles.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Microjassa Stebbing

Accessory Hagellum 1-articulate, very small; article 6 of gnathopod 1 longer than article 5; gnathopod 2 ordinary or propodochelate in male; coxa

1 half or less as long as coxa 2, coxa 5 variable,

longer or not longer than coxa 6; uropod 3 elongate, rami short, inner ramus occasionally shorter

than outer, latter with weak apical hook and tiny

apicolateral denticles.

Paradryope Stebbing

Accessory Hagellum 2-articulate; gnathopod 1

larger than gnathopod 2, otherwise ordinary; gnathopod 2 subchelate, articles 5-6 subequal in

length; coxae short but contiguous, coxa 1 as long

as coxa 2, coxa 6 as long as coxa 5; uropod 3 small,

rami minute but extending equally, outer weakly

hooked.

Parajassa Stebbing

Accessory Hagellum composed of small tubercle;

article 6 of gnathopod 1 longer than or subequal

to article 5; gnathopod 2 ordinary or in male with

large posteroproximal tooth on article 6; coxa 1 as

long as coxa 2, coxa 5 longer than coxa 6; uropod 3

small, stubby, with shortened outer ramus (type)

or ordinarily elongate with short rami, outer with

hooked apical spine.

Uropods 1-2 often with large lateral setae on

peduncles. Parajassa geurgiuna Schellenberg transferred to Ventojassa.

Pseudericthonius Schellenberg

Accessory flagellum uniarticulate, forming a

scale; article 6 of gnathopod 1 longer than article

5; male unknown, female gnathopod 2 similar to

and scarcely larger than gnathopod 1; coxae short,

serially discontiguous; uropod 3 with single ramus

slightly shorter than elongate peduncle, apically

uncinate.

Uropods 1-2 with inner rami much shorter than

outer rami; body cylindrical, urosome depressed.

Pseudischyrocerus Schellenberg

Accessory flagellum multiarticulate, elongate;

article 5 of gnathopod 1 longer than article 6;

gnathopod 2 ordinary; coxa 1 as long as coxa 2,

coxa 5 probably longer than coxa 6; peduncle of

uropod 3 elongate, rami elongate but slightly

shorter than peduncle, outer with 2-3 accessory

teetli near apex and several marginal spines.

Outer plate of maxilla 1 with 9 spines (compare

Bathyphotis)', article 4 of pereopod 3 wider than

on pereopods 1-5 in male.

Runanga J. L. Barnard

Accessory Hagellum formed of uniarticulate scale;

article 5 of gnathopod 1 slightly longer than article 6; gnathopod 2 ordinary; coxae short, discontiguous serially, coxa 5 foliaceous and setose;

uropod 3 with 2 vestigial rami, one ramus with

hook.

Uropod 2 with 1 ramus; pleopod 3 with 1 ramus;

body cylindrical, urosome depressed.

Ventojassa J. L. Barnard

Ventojassa Barnard, 1970: 204-20").

TYPE-SPECIES.—Eurysiheus ventosa J. L. Barnard,

1962.

Accessory Hagellum vestigial (type) or 2-articuIatc (\\ georgiana [Schellenberg]); article 5 of gnathopod 1 longer than article 6; gnathopod 2 ordinary; coxa 1 as long as coxa 2, coxa 5 longer than

coxa 6; uropod 3 slightly elongate, rami shorter

than peduncle, outer with apical hook and several

spimiles or setules, lateral margin with 1-3 wirelike setae (unique in family).

AMPITHOIDAE

DIAGNOSIS.—Corophioidean with ordinary urosomite 1; peduncle of uropod 3 elongate relative to

shortened padlike rami, inner ramus apically setose, outer ramus with one or two large reverted

hook-spines, often with facial scales and other setae; pereopods glandular; outer lobes of lower lip

usually subdivided partially (unique to Ampithoidae). See |. I.. Barnard (1969a) lor key, and diagnoses.

Amphitholina Ruffo is returned to this family

from the Eophliantidae.

BIANCOLINIDAE

DIAGNOSIS.—Corophioidean with ordinary uroso-
NUMBER 151 27

mite 1; uropod .'5 short, rami equally long and as

long as peduncle, outer ramus not uncinate, with 2

straight apical spines, inner ramus with 1-2 spines

or setae apically, never heavily setose; outer lobes

of lower lip undivided; mouthparts partially degenerate, palps of mandible and maxilla 1 absent

or vestigial, palp of maxilliped reduced to 3 articles, mandihular molar evanescent or absent; pereopods probably lacking spinning glands.

The attribution of this family to Corophioidea is

problematical. The genus Biancolina formerly was

placed in Eophliantidae but uropod 3 of BiancoUna is entirely distinctive. Assignment to Corophioidea is based on similarities of cephalic

appearance to corophiins, the heavily setose peduncle of uropod 1 (rare in Amphipoda but occasionally present in corophioideans), the fleshy telson

and prehensile pereopods similar to those of various ampithoids and Isaea.

PODOCERIDAE

DIAGNOSIS.—Corophioidean with elongate and

free urosomite 1, usually more than twice as long

as next segment; uropod 3 variable, generally vestigial or absent; pereopods not glandular; abdomen

generally capable of flexion beneath thorax

(unique). See Barnard (1969a) for key and diagnoses.

CHELURIDAE

DIAGNOSIS.—Corophioidean with urosomite 1

elongate but coalesced with other urosomites into

large rigid box; uropod 3 large and flabellate, ramus longer than peduncle, inner ramus, if present,

scalelike, main ramus setose strongly or weakly;

pereopods not glandular; uropod 2 flabellate

(unique). See Barnard (1969a) for key and diagnoses.

Literature Cited

Barnard, J. L.

1958. A Remarkable Xew Genus of Corophiid Amphipod

from Coastal Marine Bottoms of Southern California. Bulletin of the Southern California Academy of Sciences, 57:85-90, plates 26-28.

1962. Benthic Marine Amphipoda of Southern California.

1: Families Aoridac, Photidae, Ischyroceridae,

Corophiidae, Podoceridae. Pacific Naturalist,

3:1-72, 32 figures.

1967. Bathyal and Abyssal Gammaridcan Amphipoda of

Cedros Trench, Baja California. United States National Museum Bulletin, 260:1-205, figures 1-92.

1969a. The Families and Genera of Marine Gammaridean

Amphipoda. United States National Museum Bulletin, 271: 535 pages, 173 figures.

19fi9b. A Biological Survey of Bahia de Los Angeles Gulf

of California, Mexico. IV: Benthic Amphipoda

(Crustacea). Transactions of the San Diego Society of Natural History 15:175-228, 30 figures.

1970. Sublittoral Gammaridea (Amphipoda) of the

Hawaiian Islands. Smithsonian Contributions to

/.oology, 34: 286 pages, 180 figures.

Kudrjaschov, V. A., and S. V. Vassilenko

1966. A New Family Caprogammaridac (Amphipoda,

Gammaridea) Found in the North-west Pacific.

Crustaceana, 10:192-198, 4 figures.

Myers, A. A.

1968. A New Genus and Two New Species of Gammaridean Amphipoda from Central America. Journal

of the Linnean Society of London Zoology, 47:527-

531, 2 figures.

1969. A Revision of the Amphipod Genus Microdeutopus

Costa (Gammaridea: Aoridae) . Bulletin of the

British Museum (Natural History), 17:93-148, figures 1-22, plate 1.

Stebbing. T.R.R.

1910. Crustacea. Part 5: Amphipoda. Scientific Results

Trawling Expedition H.M.C.S. "Thetis." Australian

Museum, Memoir 4. volume 2: 565-568, plates

47-60.

U. S. GOVERNMENT PRINTING OFFICE: IS73 5t6-22l/l7

Publication in Smithsonian Contributions to Zoology

Manuscripts for serial publications are accepted by the Smithsonian Institution Press, subject to substantive review, only through departments of the various Smithsonian museums. NonSmithsonian authors should address inquiries to the appropriate department. If submission is

invited, the following format requirements of the Press will govern the preparation of copy.

Copy must be typewritten, double-spaced, on one side of standard white bond paper, with

\W top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompanied by the anginal artwork. Duplicate copies of all material, including illustrations, should be

retained by the author. There may be several paragraphs to a page, but each page should begin

with a new paragraph. Number consecutively all pages, including title page, abstract, text, literature cited, legends, and tables. The minimum length is 30 pages, including typescript and illustrations.

The title should be complete and clear for easy indexing by abstracting services. Taxonomic

titles will carry a final line indicating the higher categories to which the taxon is referable:

"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify

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.
K

1

•

"iHr

^ ^MBI