J. LAURENS BARNt Keys to the Hawaiian

Marine Gammaridea,

0-30 Meters

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 58

j . Laurens Barnard Keys to the Hawaiian

Marine Gammaridea,

0-30 Meters

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

I971
ABSTRACT

J. Laurens Barnard. Keys to the Hawaiian Marine Gammaridea, 0-30 Meters.

Smithsonian Contributions to Zoology, 58, 135 pages, 1971.—Methods of collecting

Hawaiian Gammaridea, their preservation, sorting by color schemes and head shapes,

dissection, terminology and identification are presented. Keys to taxa illustrated with

drawings are given for 120 species found in depths from 0 to 30 m. Synonymies, distribution, and notes on crucial characters of each species are given. This paper supplements the formal descriptions of the Hawaiian species contained in Smithsonian

Contributions to Zoology, number 34.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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Contents

Page

Introduction 1

Methods of Presentation 2

Collection of Amphipods 3

Dissection 3

Appendages—Positions and Terms 5

Appendages—Setae and Spines 5

Glossary 6

Diagnosis of the Basic Gammaridean 8

Identifications—Objective 10

Identifications—Non-Objective 12

Body Shape 14

Color in Preservative 14

Head and Eyes 16

Kinds of Uropod 3 18

Taxonomic Arrangement 19

Order Amphipoda 19

Key to Hawaiian Families of Gammaridea 20

Ampeliscidae 22

Amphilochidae . . . '. 25

Ampithoidae 29

Anamixidae 40

Aoridae 40

Biancolinidae 52

Cheluridae 54

Colomastigidae 54

Corophiidae 55

Dexaminidae (=Atylidae) 61

Eusiridae (—Calliopiidae) 61

Gammaridae 66

Isaeidae (=Photidae) 88

Ischyroceridae 97

Leucothoidae (cf. Anamixidae) 102

Liljeborgiidae 103

Lysianassidae 105

Ochlesidae 105

Oedicerotidae 105

Phliantidae 109

Phoxocephalidae 109

Pleustidae 109

Podoceridae 113

Sebidae 117

Stenothoidae 120

Superfamily Talitroidea: Families Hyalidae, Hyalellidae 122

Literature Cited 131

Index 133

j . Lamens Barnard Key s to th e Hawaiia n

Marine Gammaridea,

0-30 Meters

Introduction

Gammaridea are the commonest suborder of Amphipoda on the nearshore benthos of Hawaii and one of

the most common groups of medium-size and small

organisms (1-7 mm body length). Caprellidea, a suborder of skeleton shrimps are also prevalent but less

common and not treated here; Hyperiidea, the third

major suborder, occur primarily in the oceanic

bathypelagic realm.

Hawaiian Gammaridea are very small organisms

barely visible to the naked eye and frequently overlooked by the collector. Most Hawaiian species have

bodies less than 5 mm in length, many adults are approximately 1 mm in length, but a few species attain

lengths of 14 mm. They are abundant in algae and

coral heads and on rubble but are rare on sand bottoms.

Mud bottoms, though rare in shallow waters of Hawaii,

would be expected to have high numbers of amphipods,

as they do on continental shores; mud bottoms have

not been explored in Hawaii and this treatment of

Gammaridea concerns primarily those in epifaunal

biotopes.

A potential wealth of ecologic, genetic, and physiological problems occurs in the Amphipoda. Some of

these are suggested in Barnard (1969). Food habits, as

one measure of the niche, are of primary interest.

Adaptation to food ingestion ranges from epiphytic

herbivory to ectoparasitic sucking and includes mud

ingesting, sand processing, lignivory, detritus feeding,

slime lapping, filter feeding, carrion ingestion, and

broad spectra of omnivory. Predation by benthic Am-

/. Laurens Barnard, Department of Marine Invertebrates,

National Museum of Natural History, Smithsonian Institution, 20560.

phipoda is a very minor consideration but worth investigating. Mechanisms in tube building, swimming,

maintenance of nestling positions, saltatory escape, and

copulation are of considerable interest and almost

unstudied anywhere. Predation and parasitism upon

amphipods by other organisms, life histories, and reproductive rates are all known only rudimentarily, even in

those few species thus far studied.

This paper presents a system, based on the known

species, of identifying Hawaiian Gammaridea dwelling

in the range 0-30 m. Before attempting a first identification, however, the observer must have a clear

understanding of the topics covered in this introduction. "Methods of Presentation," for example, contains

an explanation of the format of the keys and the figures, as well as a brief discussion of the monographs

which the student should have at hand for reference;

and this is followed by sections outlining the basic

method of collection and the techniques of dissection.

Problems may arise from oddities in the position of

appendages and difficulties in seeing certain characters

because of the many setae occluding the chitinous surfaces of the appendages; these problems, which the

observer will note immediately after dissection, are

treated under the sections "Positions and Terms" and

"Setae and Spines." The "Glossary" found on page 6

contains terms especially applicable to the Amphipoda,

and a section on "Identification" discusses in general

terms use of the keys. The "Key to Hawaiian Families,"

page 20, is preceded by a headnote warning of problems that may arise during its use. A principal aid to

use of the keys is the "Diagnosis of the Basic Gammaridean" presented on page 8, which the student is advised to memorize.

For advanced students wanting assistance and ideas

in rapidly sorting complex samples of amphipods ac1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cording to a limited number of characters, there is pre*

sented a special section on "Non-Objective" identification which may be skipped if the reader is interested

primarily in identifying one taxon at a time.

The systematic section of this paper is introduced by

a formal diagnosis of the suborder Gammaridea, and

in this section follow in alphabetical order the keys to

families, and under the families, also in alphabetical

order, keys to the genera and species thus far found in

Hawaiian waters.

Methods of Presentation

To minimize text and to avoid long exclusive diagnoses, the comments appended to keys and taxa are

abbreviated to include only major distinctions of the

taxa, and may consequently appear indiscriminate

or inconsistent. The reader may assume, however, that

when a character is mentioned for one taxon but not

for another, that it is in a negative state in the other

related taxa of that group (family or genus) under

discussion.

No words or pictures presented in such a format can

absolutely distinguish species, especially in a fauna not

completely explored; caution must therefore be exercised in using this work. When questions arise, original descriptions and diagnoses should be consulted,

especially in papers by Schellenberg (1938) and

J. L. Barnard (1955, 1970). Stebbing (1906) and

Sars (1895) remain as invaluable references and

should be at hand, particularly for the comprehensive

world taxonomic treatment with keys offered by Stebbing, and for the magnificent illustrations of various

kinds of amphipods in the Norwegian fauna presented

by Sars.

Perusal of Stebbing's work will quickly familiarize

the reader with the kinds of characters utilized in

gammaridean taxonomy and will remove any overconfidence that the present treatment of Hawaiian Gammaridea is exhaustive. Stebbing's work gives most of

the modern names applied to the genera and species

in Sars' work, and in it one may observe the general

appearance of various genera also occurring in the

tropics—for example, Elasmopus, Ericthonius, Corophium, Gammaropsis (=Eurystheus), Lembos, Leucothoe, Ampelisca, Byblis, Ampithoe, Aoroides (like

Sars' Aora), Pontogeneia, Maera, Ceradocus, Eriopsia (tropical members of this genus vary slightly

from those shown), and Laetmatophilus. Examination

of Sars' work is especially helpful in developing familiarity with the gross appearance of such families as

the Lysianassidae, Ampithoidae, Gammaridae, Stenothoidae, Amphilochidae, Leucothoidae, Aoridae, Isaeidae (=Photidae), Ischyroceridae, Corophiidae,

Dexaminidae, and Liljeborgiidae. But Norwegian

talitroids (Hyale and beachhoppers of the genera Orchestia and Talitrus) are not typical of tropical members. The work of Chevreux and Fage (1925) dealing

with the Mediterranean (including warm-temperate

France) is also useful and slightly more oriented

towards genera likely to occur in the tropics than is

Sars' Norwegian work.

An index to world species sparsely referenced and

keyed to the Zoological Record was published by

J. L. Barnard (1958), but even in this short time it

is out of date because of accelerated study in the group.

In the present work, most of the key characters are

illustrated in accompanying figures by outline drawings. Occasionally the drawings are semidiagrammatic

and, for clarity of shape, the setae are removed, being

indicated by small circles at the points of attachment.

To facilitate readability of keys and diagnoses, the

figure citations are minimized, but figures can be found

to illustrate most characters by searching plates concerned with the appropriate family. Capital letters on

the drawings of Figures 5-68 form a code to the species

found in each figure legend, whereas lower-case letters

indicate the appendage shown. The code to this system

is found on page 9. The drawings, unless otherwise

marked, represent males but often they actually represent both sexes, since, in many species no significant

differences occur between the sexes. The female sign

is attached to those appendages in which the female

form exhibits a significant taxonomic difference from

the male.

The key to families (p. 20) is presented in an abbreviated form in which certain families are keyed out

at several points. To most of the families and genera

keys have been written to cover both sexes of the

various taxa, but a few concern only adult males. The

females of these taxa have not yet been adequately differentiated, and this statement may also apply to

juveniles, since these usually resemble adult females.

Species of Amphipoda not easily identified by these

keys should be presumed to be unrecorded from

Hawaii, and for their identification to genus one must

resort to Barnard's (1969) handbook. New species and

newly recorded species remain to be discovered; for

although the species that are abundant and widely
NUMBER 5 8 3

distributed in littoral algae and on ordinary rubble and

coral bottoms probably have been captured, exploration has scarcely begun on the specialized habitats,

especially those open only to the Scuba diver. Several

species not found elsewhere have, for example, been

collected in the molluscan assemblage associated with

the black-coral Antipathes; and another species,

Elasmopus calliactis, is found only in association with

calliactid sea-anemones attached to shells of Dolium

inhabited by the hermit-crab Dardanus. Among the

wealth of very specialized habitats such as these must

occur many more amphipods which, although of very

limited distribution, may be extremely abundant

within their niches. More than a dozen Hawaiian

species are based on a few specimens; their apparent

rarity is in part a measure of the difficulty in sampling

their special habitats with gross collecting methods so

far utilized.

Collection of Amphipods

Amphipoda are most easily collected by washing substrates in a bucket of seawater mixed with 1-2 ounces

of formaldehyde. This poisoned water causes amphipods to swim out of their concealment, die, and fall to

the bottom of the bucket amid other debris. The coarse

fraction of the debris is removed from the bucket by

hand and the remaining contents then strained

through a 100 mesh/inch Tyler screen, the solid contents retained by the screen being preserved in a container with a mixture of 5 percent formaldehyde

(concentrated formaldehyde diluted 9:1) and seawater. In the laboratory the sample is washed in freshwater; the species are then sorted under a stereoscope

and represerved in 70-percent alcohol. Because specimens of a few species remain in their place of

concealment, substrate samples should be preserved

and examined in the laboratory.

Dissection

Under normal circumstances the student identifying

Gammaridea to genera should use the entire procedure of dissection and observation described by

Barnard (1969). The fauna of Hawaii is now sufficiently well known, however, that one may initially be

able to use the swifter methods described below.

The keys require that the material be in adequate

condition, with unbroken uropods and antennae.

Broken specimens, except those missing the distal ends

of antennae,* should be avoided until experience is

gained. The identification requires the initial minimum

dissection of the amphipod and observation of other

characters not initially removed from the body of the

animal; but one must be prepared to continue dissecting off parts of the organism in the event a particular

shape has to be observed. Keys, diagrams, and discussion should be perused thoroughly before undertaking

the dissection and identification.

Amphipoda vary morphologically from specimen to

specimen; if strong sexual dimorphism occurs, there

will be considerable difference in male gnathopod

2, which will be generally larger and more strongly

ornamented than in females and juveniles; male antennae may be longer; uropod 3 more setose; and eyes

larger than in females. A few species have an axial

reversal in gnathopodal dominance so that gnathopod 1

is larger and, in males, more complex than gnathopod

2, the latter assuming the character of a first gnathopod. Ovigerous females carry eggs in the ventral

brood pouch formed by the lamellae attached to the

medial faces of the coxae; on the benthos adult females

are generally more common than are males. Largebodied individuals without brood plates are not always

males; these plates may be lost by gerontic females.

Males may be determined by microscopic examination

of the ventral surface of thoracic segment 7 between

the members of pereopod 5. A small pair of penial

projections, occasionally minutely spinose, occurs on

adult males but they should not be confused with gills

occasionally attached to the coxae of that segment.

Gills break off easily whereas penial processes do not.

So-called intersexual adults may lack any marks of

their sex. Juveniles look generally like females.

The specific identity of females occasionally has to

be assumed from their repeated association with males

in many samples; samples frequently have two or more

species of the same genus, with females and juveniles

very difficult to distinguish from species to species.

For dissecting and mounting amphipods for microscopic examination, the procedures detailed below

have been found useful.

From a sample of Amphipoda preserved in alcohol,

or transferred to freshwater, should be isolated a few

specimens grossly similar to one another. One is

•Specimens bearing at least one member of pereopods 3-5

(6 legs) and one member of pereopods 1-2 (4 legs) can also

be utilized.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

chosen as that to be dissected and the others saved

for later verification and as source material for opposite sexes and juveniles. The subject amphipod is

placed in a Syracuse dish and covered deeply with

alcohol.

SLIDE 1.—A small drop of glycerine is placed on

a glass slide. The left uropod 3 of the specimen (see

Figures 1, 2, and 4) is detached by means of a

jewelers forceps plus dissecting needle (insect pin

mounted on a wooden stick) and placed in the drop

of glycerine. The drop will disperse momentarily in

mixing with alcohol but will coalesce quickly if a

light breath of air is applied. The uropod on the

slide is observed with the stereoscope and arranged

so that its dorsal side is up. Observation of the uropod before dissection almost invariably demonstrates sufficient differences in the two rami and

shape of peduncle so that the dorsal side can be distinguished after dissection. A glass cover slip is applied gently and manipulated with the forceps so

as to prevent the uropod from rolling over while the

glycerine spreads.

SLIDE 2.—The carcass of the amphipod with its

left side up is placed in a depression slide filled with

glycerine and a cover slip is applied. Most Hawaiian

amphipods fit snugly into a standard depression slide

but a few are so large that supports for the cover

slip are required so as to elevate it and prevent

crushing of the body. Paper clips, staples, sand grains

and broken bits of amphipod appendages are useful as supports of varying thickness.

Glycerine may shrink parts for a few hours but they

usually return to normal condition. Many specialists

prepare their material by slowly adding glycerine to

alcohol so as to adjust the specimens to this medium,

and later may dissect the animals in glycerine or warm

glycerine jelly. Numerous other media are also

satisfactory.

If the urosome (posterior half of abdomen) of the

amphipod is strongly flattened and if uropod 3 is not

clearly apparent, the full urosome should be removed

to slide 1 and mounted dorsal side up. If uropod 3 or

its empty socket is still invisible one may have to turn

the urosome over in order to find a vestigial uropod 3

appearing as a pair of flaps on the ventral side of the

anus and telson.

Uropod 3 on slide 1 is examined with a compound

microscope and the observations utilized to commence

the basic key (p. 20).

Many of the other observations necessary in later

parts of the keys may be made from slide 2 of the

body, e.g., coxa 1, antennal length, extent of dorsal

depression of the body, gnathopods 1-2 if not strongly

twisted, head, urosomite 1, and pereopods. Coxa 1 may

be obscured by the thickness of the body interrupting

transmitted light, and a slide must be made of it after

dissection. If examination of mouthparts, telson, and

accessory flagellum (because of its diminutiveness) are

required in the keys, further dissection and slide

preparation are necessary. The following additional

glycerine slides are made by dissecting parts from the

carcass of slide 2:

SLIDE 3.—Thoracic legs: all thoracic (pereonal)

legs from the right side are removed by using forceps

to pull off the legs and their coxae, and are placed

on the slide so as to appear to be from the left side;

while disengaging these legs one must hold the

amphipod in the depression slide (or return it to

alcohol in a Syracuse dish) with a forceps or pin in

the hand opposite to that grasping the leg; a pin is

preferable in holding the amphipod on the segment

just above the coxa-leg being removed, so as to prevent breaking the body.

SLIDE 4.—Maxillipeds: the posterior mouthpart,

composed of 2 halves basally joined, are easily

snapped off with forceps and mounted so that the

inner and outer plates (lamellae) face upwards; the

curved base may be removed so that the cover-slip

fits tightly. Use of the term in the singular (maxilliped) indicates half of the pair.

SLIDE 5.—Maxillae 1-2: these 2 pairs of setose

mouthparts lie below the maxillipeds, toward the

mandibles, and cover the lower lip; dissection of

maxilla 1 requires removal of the inner plates that

often are attached more deeply than the remainder

of the appendage.

SLIDE 6.—Lower lip: this is often and easily

destroyed by tearing while removing maxilla 1, and

is difficult to separate because of scleritic connections

to mandibles or upper lip; rarely required for purposes of keys herein, it is mounted in glycerine

scattered with sand grains to elevate the cover slip.

SLIDE 7.—Mandibles: these are attached by

powerful muscles and must be firmly grasped at

their muscular bases in order to pull them loose.

Knowledge of the palp is essential and observation

of palps before dissection will ensure against their
NUMBER 5 8

unobserved accidental loss, for the mandible can

often be pulled away, leaving its palp. Mandibles

often are brittle and easily shattered: they are placed

on the slide with the molars, if any, facing upward;

otherwise a left mandible should be seated so that

the toothed incisor points "north" while the palp

points "west"; a right mandible has the palp pointing "east" if the incisor points "north." Sand grains,

are usually necessary to support the cover slip.

SLIDE 8.—Right antennae 1 and 2: these are

mounted so as to appear left sided, with medial

surfaces facing upward. The accessory flagellum of

antenna 1, often tiny, occurs on the apicomedial end

of peduncular article 3.

SLIDE 9.—Telson: this is the terminal flap on

pleonite 6, often bilobed. One must be sure that

both lobes are removed, for a single lobe mounted

on a slide will appear as an unlobed telson and lead

to erroneous identification.

SLIDE 10.—Gnathopods 1 or 2: if these are thick,

they should be removed to a separate slide with

support for the cover slip.

The carcass is returned to slide 2 and a cover slip

replaced; right uropods 1-3, pleopods, upper lip and

epistome and all left sided appendages except uropod 3

remain on the animal; if the projection of the epistome

must be observed the carcass can be flipped over with

its right side up, as the removal of antennae 1-2 on

that side permit its view. Pleopods are not utilized in

the keys but they can be removed to a storage vial

(Vie

 o r

 Vs dram) so that epimera 1-3 are more clearly

visible.

As the right thoracic legs and their coxae are removed from the body they should be observed carefully

so as to keep them in numerical sequence; usually the

gnathopods are prehensile and gnathopod 2 (coxa 2)

is larger than gnathopod 1 (coxa 1); pereopods 1-2

usually are oriented so as to point or function towards

the anterior, and coxa 4 (of pereopod 2) is usually

larger than coxa 3; pereopods 3-5 (coxae 5-7) usually

have coxae smaller than l-A (except in some tubicolous species) with coxa 5 largest and bilobed and coxa

7 smallest and simple, coxa 6 being intermediate in

size and weakly bilobed; pereopods 3-5 usually increase

in size successively but if pereopod 5 is shorter than

the others it is usually of distinct morphology. The

observer will note that the legs are not numbered in

sequence with their coxae, as the first 2 pairs are called

gnathopods, even though they are equivalent ontogenetically to pereopods. Gnathopod 2 and pereopods

1—5 or 1-4 have a gill attached to the coxa, and females have a second kind of lamella—the setose brood

plate attached one each to coxae 2, 3, 4, and 5. Brood

plates may be removed and preserved in the small vial

with the pleopods before mounting the legs.

Slides may be labeled with grease pencil and stored

flat in boxes to prevent accumulation of dust; storage

for more than 2 weeks is inadvisable, as parts become

very soft and commence dispersion in glycerine. Slides

utilizing permanent mounting media may be made

for long-term storage of holotypes or special material,

but the time needed makes it inadvisable for a beginning student, who should be involved with dissecting

and observation of as many specimens as possible. Many

taxonomists preserve their dissected organisms in small

vials in alcohol. Parts erroneously mounted upside

down are difficult to right in permanent media and it

is all too easy to make such mistakes with tiny appendages.

Appendages—Positions and Terms

Gnathopods of amphipods are bent like bird-legs, so

that the free ends point forward; other legs or their

dactyls also are normally oriented in other than vertical axes. In describing their various parts taxonomists

visualize the legs as being straightened out into a

vertical axis. Hence the anterodistal corner of article 6

on gnathopod 2 of a vertically oriented appendage appears anterodorsal when the leg is normally positioned

by the animal. The true posterior edge of a flexed dactyl on pereopod 5 may appear on the animal to have

its face pointing anteriorly or ventrally.

Antennae are described as if they have been fully

stretched horizontally to the front.

Uropods are inconsistently described as also occurring in a horizontal plane though they are derivatives

of vertically oriented appendages.

The Glossary (p. 6) contains terms used to describe

these appendages and positions.

Appendages—Setae and Spines

Gammarideans are especially setose and spinose crustaceans. Setae and spines are a headache to the descriptive taxonomist, who must spend a great deal of time

illustrating them; but they often have value in classi-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fication as well as identification, though they occasionally hide other good characters. Except in the taxonomy of freshwater amphipods, the minute structure

of setae and spines has rarely been used systematically

but it is now being employed more frequently by marine taxonomists. Setae and spines are fundamentally

similar, as both have a nerve canal; but spines are

shorter, stouter, and stiffer than setae. Some authors

correctly use the term setal spines for spines but herein

the simpler term spine is used. Many spines (and setae)

have apical or subapical branches possibly acting as

triggers to the neuron. Some amphipod setae are as

complex as those in polychaetes, being composed of

two or more articulate parts. Setae and spines are

basally articulate but stout spines often become immovably fused to their parent appendage. A good rule

of thumb is to call spines those setae that will not bend

in the middle without breaking.

In the drawings accompanying this paper most of

the plumose setae and many of the triggered spines are

drawn as simple lines, because the plumes and triggers

are too small or thin to be represented and the added

drafting would consume double the time in making all

the other figures.

Many gnathopods have a small stiff spine on the

palm at the base of the dactyl that in some cases may

act to spring open the dactyl or in others to lock it,

although the latter is probably rare, as the muscles of

the gnathopod appear to be capable only of closing the

dactyl. On pereopods the distal end of article 6 often

has special spines called "locking spines" and these are

often found also defining the proximal end of the palm

on the gnathopods. Their purpose is probably to prevent lateral or medial movement of dactyls rather than

to lock them into a flexed position; or they may function in ecdysis or prehension. The minute structure

of these spines is of special value in recognizing species

in the genera Hyale and Ampithoe, as the spines may

be large or small, striate or unstriate, or grossly striate

spirally, with the appearance of a screw.

The extent of setation in the gnathopods of Lembos,

especially in terminal males, is valuable for identification; brushes of setae on antennae of certain species

of Hyale are also important.

The general and specific setosity of various species

has, as yet, scarcely been conveyed by words or statistics, but the taxonomist often relies on gross setal differences as specific characters.

Glossary

[The italic references in parentheses are those used on

Figures 5-68 to identify the parts named. A complete

list of these is to be found on page 9.]

accessory flagellum {ac, af). The secondary ramus of

antenna (a) 1, often absent or vestigial, and attached

medially to peduncular article 3.

aesthetasc, aesthete. Sensory setae of antennae, flattened

and nontapering.

article. The segment of an appendage.

calceolus. A small globular or helmet-shaped, articulate

sense organ on the antennae; presumably a modified

aesthetasc; of rare occurrence in Gammaridea and

most often seen in Eusiridae.

carpochelate. Immovable finger of prehensile appendage occurring on carpus (article 5); example:

Leucothoe.

chela. Immovable finger of prehensile appendage.

chelate. Descriptive of the palm of a gnathopod protruding as an immovable finger on which the dactyl

closes. See parachelate, carpochelate, propodochelate, merochelate.

claw, claw-like. Descriptive of a talon or simple, tapering nail. (Not descriptive of chelae as used in decapod terminology.)

compressed. Flattened from side to side.

conjoint. Describing the basal amalgamation of flagellar articles on antennae.

corneal lens. A biconvex cuticular body occurring

directly in or on the chitinous cephalic surface in Arapeliscidae; contrasted with subcuticular ommatidia.

coxa (c), coxal plate. (Terms used synonymously

herein.) Article 1 of a pereonal appendage, expanded into a lateral lamella. (Terms for other articles of the appendages such as basis, ischium, merus

carpus, propodus, and dactyl are frequently but not

universally used in Gammaridea; instead, the articles are simply numbered.)

dactyl. Talon-like terminal article of pereopods (article 7) or maxillipeds (articles 3 or 4).

depressed. Flattened dorsoventrally.

emarginate. Descriptive of the concave posterior end of

an uncleft telson.

entire. Descriptive of an uncleft telson.

epimeron (ep). A lateral pleuron of pleonites 1-3; the

ventrolateral plate-like extension of the body

segment.

epistome. The anterior surface of the head above the

labrum; this area is often extended ventrally to ap-
NUMBER 5 8

pear as a part of the labrum and may be anteriorly

produced as a cusp or lobe.

flagellum. The distal portion of either antenna 1 or 2;

on antenna 1 it commences with article 4, on antenna

2 with article 6; because basal peduncular articles

of antenna 2 are often difficult to resolve, the juncture may be recognized between the elongated final

peduncular article and the shortened first flagellar

article which is followed by similar short articles; on

antenna 1, however, article 1 of the flagellum is occasionally elongate and apparently composed of nonsegregated (thus conjoint) daughter articles.

galeate. Descriptive of the helmet-shaped heads of

various oedicerotids and synopiids.

gnathopod (g). One member of the first two pairs of

free thoracic appendages; these appendages differ in

function and usually in appearance from following

pereopods; often called pereopods.

hand. Article 6 of a gnathopod.

incisor (in). The apical portion of the mandible,

usually formed into a toothed chewing edge or untoothed chopping plate.

joint. The juncture between two articles of an

appendage.

labrum. (See upper lip.)

lacinia mobilis (lm). Articulated accessory plate proximal to the mandibular incisor, often absent or missing on either left or right mandibles, occasionally indistinguishable from a spine of the spine-row.

locking spine. The distal-most spine(s) on article 6 of

pereopods 1-5, near the base of the dactyl.

lower lip (//) (labium). A fleshy complex posterior to

the mandibles, always composed of at least one pair

of lobes (outer), often with a medioproximal pair of

inner lobes; the lateroproximal ends of the outer

lobes are often attenuated as alae and are denoted

as mandibular lobes.

mandible (m). The anterior movable appendage of the

buccal group; usually composed of a body bearing a

distal incisor, a lacinia mobilis, spine row, molar, and

3-articulate palp.

massive. A term applied to the heads of Synopiidae and

Oedicerotidae; head as long as pereonites 1-3 combined and as tall as or taller than long (length not

including rostrum). Heads of Ampeliscidae and

Phoxocephalidae are elongate but not massive; heads

of Acanthonotozomatidae are as tall as long but are

not as long as pereonites 1-3 combined.

maxilla (x) 1. A pair of cephalic appendages posterior

to the lower lip; for taxonomic purposes only three

portions of each member are named: the medial lobe

(plate) usually bearing marginal setae, the lateral

and larger lobe (plate) bearing terminal spines, and,

attached to the outer lobe, a palp usually composed

of two articles but occasionally absent.

maxilla (x) 2. A pair of cephalic appendages posterior

to maxilla 1; for taxonomic purposes each member

recognized as a pair of lobes (plates) medial and

lateral, usually strongly setose.

maxillipeds (xp). The posteriormost pair of "cephalic"

appendages, representing the primitive first thoracic

segment now amalgamated with the head but in

amphipod taxonomy not included in the sequential

numbering of thoracic appendages; for taxonomic

purposes recognized as a pair of basally amalgamated appendages, each member composed of a

proximal (inner) plate, a distal (outer) plate, and

a palp of four articles, rarely reduced to 3 or 2

articles or absent.

merochelate. Immovable finger of prehensile appendage occurring on merus (article 4) ; example:

gnathopod 1 of Aoroides.

metasome. Pleonites 1-3. [Term rarely used.]

molar {mo). A process of the mandible, located on the

midmedial margin; when completely developed it is

a large, massive, subcylindrical body with a surface

of ridges and teeth used for grinding (triturative).

ommatidium (singular), ommatidia (plural), ommatidial (adjective). Terms applying to the parts of the

subintegumentary compound eye, not to be confused with the comeal lenses of the integument of

Ampeliscidae.

palm. A posterior surface or margin of article 6 of a

gnathopod or pereopod on which article 7 (dactyl)

closes for the purpose of prehension; usually recognizable because of expansion of article 6 or by occurrence of special spines or ornamentation and

usually with a proximal defining limit marked by a

change in marginal slope or occurrence of special

spines.

palp {pal). Terminal articles of a buccal appendage,

in Amphipoda occurring only on mandibles, first

maxillae, and maxillipeds as the stenopodous terminal articles distal to the expanded outer plates or

main body.

parachelate. A weakly chelate condition like that

shown herein for gnathopods 1-2 of Biancolina.

peduncle. The basal articles of a fundamentally biramous appendage; in Amphipoda applied to an-
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tennae, pleopods, and uropods; antenna 1 with three

peduncular articles, antenna 2 with five peduncular

articles (but appendage not biramous); pleopods

with one definitive peduncular article but remnants

of others occurring proximally; uropods each with

one peduncular article.

pereon. The complex of seven free thoracic segments

bearing gnathopods and pereopods, not including

the maxillipeds.

pereonite. A segment of the pereon.

pereopod (p). A walking, grasping, standing, or feeding appendage attached to a pereonite; normally

composed of seven articles, including coxa; in Araphipoda the first two pairs are often termed gnathopods and only the last five pairs of thoracic legs are

called pereopods.

plate. A flattened lobe on an article of a maxilla or

maxilliped.

pleopod (n). A biramous swimming appendage on

pleonites 1-3, one pair for each pleonite.

pleon (pi). The abdomen (of six free segments in

Gammaridea, rarely with some segments coalesced).

See metasome and urosome.

pleonite. A segment of the pleon.

prehensile. Adapted for seizing or grasping; applicable

to but rarely used for gammaridean gnathopods;

especially useful in denoting pereopods of several

species of Ampithoe which either are subchelate or

chelate, or have distinct, spinose palms or nonskid

surfaces indicating their use in grasping.

propodochelate. Synonymous with chelate.

propodus. The sixth article of a thoracic appendage

(especially used to denote the palmar article of a

gnathopod).

scale, scale-like. Terms applied to the accessory flagellum (ac) when forming a small lamella immovably

fused to article 3 of antenna (a) 1; and to the inner

ramus of uropod (u) 3 when strongly reduced and

plate-like.

simple. Used in amphipod taxonomy to denote the absence of spines or setae on appendages; or the occurrence of but a single article in the ramus of a

uropod; or especially to the absence of a palm on a

gnathopod (g) or pereopod (p). Distinction between subchelate and simple is often weak.

subchelate. Article 6 of a gnathopod or pereopod having a distal palm against which article 7 closes; a

prehensile condition in which the palm is not produced to form a finger; intermediate in condition

between chelate and simple. Complexly subchelate

or complexly chelate are terms referring to the formation of a false chela by protrusion of teeth, cusps,

or lobes from articles other than the sixth and upon

which article 7 impinges to form a prehensile condition ; occurring especially in Aoridae, Corophiidae,

and Leucothoidae. See "simple."

telson (t). A flap dorsal to the anus attached to

pleonite 6, primitively bilobed but usually in

Amphipoda with bases coalesced and often with

lobes completely coalesced to form a single plate.

terminal. Male or female in final stage of adulthood.

triturative. Descriptive of the rasp-like surface of a

mandibular molar, composed of teeth, ridges, and

cusps.

upper lip (ul) (labrum). A fleshy lobe attached to the

anterior cephalic margin in front of the mandibles;

occasionally the anterior surface of the labrum protrudes as a lobe or cusp; often the cephalic area to

which the labrum is attached is recognizable as an

"epistome" and may also be lobed; or both labrum

and epistome may be indistinguishable and produced together as a single lobe.

uropod (u). One member of the three pairs of terminal pleonal appendages, each formed of a peduncle

and two rami (occasionally rami of uropod 3 reduced or absent, rarely rami of uropods 1-2 absent

or reduced).

urosome (ur). The complex of pleonites 4, 5, 6, carrying uropods (u), and telson (t). Often numbered as

urosomites 1, 2, 3.

urosomite. A segment of the urosome (ur).

Diagnosis of the Basic Marine Gammaridean

FIGURES 1, 2

Accessory flagellum well developed, with 4 or more articles. Primary flagellum of antenna 1 not basally

conjoint (Figure 2) or elongate.

Mouthpart field quadratiform from lateral view (as

contrasted to coniform).

Each mandible with 3-articulate palp, article 3 longer

than article 1; molar present, with grinding surface

composed of ridges and teeth ( = triturative).

Lower lip (labium) with principal (outer) lobes undivided or unnotched and not widely separate, lobes

with sharp or blunt lateroproximal extensions

("mandibular extensions"), without inner lobes.

Each maxilla 1 bearing inner lobe, outer spinose lobe

and strong, straight palp of 2 articles.
NUMBER 5 8

Pereon

Head

Epimeron

Pleopod

Telson

Gnathopods

Accessory Flagellum

Uropods

Pereon Pleon 1-3 Urosome Pleopod Coxae

FIGURE 1.—The basic gammaridean.

The following letter symbols are used to identify parts

on Figures 5-68. For additional details of definition,

also Figure 1.

a antenna

ac,qf accessory flagellum

b prebuccal mass from

lateral view

c coxa

d dorsal

tp epimeron

g gnathopod

h head

in rnandibular incisor

ip inner plate

ir inner ramus

j juvenile

/ lateral

// lower lip

Im lacinia mobilis

m mandible

mo rnandibular molar

mp mandibular palp

ml mouthparts

n pleopod

op outer plate

or outer ramus

p pereopod

pal palp

pi pleon

of the amphipods illustrated

see Glossary (page 6) and

r medial

t telson

tl telson, lateral

u uropod

ul upper lip

ur urosome

v ventral teeth of pereon

w side view

x maxilla

xp maxilliped

z gill
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Each maxilla 2 with 2 well-developed, setose lobes.

Each side of maxillipeds with large inner (proximal)

lobe and outer (distal) lobe, and 4-articulate palp,

article 4 claw-shaped (unguiform).

Gnathopods well developed and subchelate ("powerful" ), non-lysianassid in form. (See lysianassid in

Figure 52.)

Gnathopod 2 larger than 1 in male, gnathopod 1 never

larger than 2 in female, article 3 of gnathopod 2

short.

Article 4 of pereopods 1-2 not extensively elongate.

Pereopods 3-5 of congruent structure and successively

slightly longer.

Three pairs of large and subequal uropods present, all

biramous, rami subequal in length, lanceolate

(Figure 4E) ; peduncle of uropod 3 not elongate.

Telson deeply cleft, of medium length.

Coxae forming elongate, rectangular plates with

quadrate or rounded distal edges, coxae 1-4 of uniform shape or slightly increasing in size consecutively, coxa 4 excavate posterodorsally, coxae not

splayed.

All body segments free. Metasome (pleonites 1-3)

only as long as last 5 pereonites combined; head

subcuboidal, not massive, rostrum small. Body laterally compressed.

Eyes if present ommatidial, not corneal and composed

of one eye on each side of head.

Figure 2 shows ordinary kinds of dissected parts and

illustrates variations in telsons, the fleshy telson, the

appearance of quadratiform and conical mouthpart

bundles, and a conjoint primary flagellum of antenna

1. Other variations are developed in the figures cited

in the Keys.

Identifications—Objective

The concept of a basic marine gammaridean is useful to the observer using the key to families (p. 20).

The diagnosis of the basic gammaridean on

page 8 contains the framework of an ordinary gammaridean as illustrated in Figures 1 and 2 (upper). In

a specimen being identified any morphological departure from this plan signals an extraordinary character to be noted. This may hasten the process of operating the various keys, because the observer has at

hand a special memory device. For example, the specimen may fit the model of the basic gammaridean in all

characters except for the presence of a non-triturative

mandibular molar, a condition which should lead one

fairly rapidly through the key to the Liljeborgiidae. Or

the specimen may fit the basic gammaridean except

for having spinose antennae and a short, morphologically distinctive pereopod 5, a condition which should

lead to the Phoxocephalidae.

Because of the abbreviated form of the keys their

various alternatives should be traced to conclusion,

especially where one has difficulty in making a value

judgment on a character. For example, in couplet 11

of the first section in the key to families, one may often

choose either pathway and trace each to its conclusion.

Suppose that the organism truly belongs to the Aoridae

of couplet 12 but one is tracing it through Section D

(leading from couplet 11) : various incongruities will

crop up as one terminates the key, perhaps in the Gammaridae, and if the organism is then traced through

the key to the Gammaridae (p. 68), it will become

clear that no genus or species of Gammaridae has a

third uropod precisely like that found in various aorids.

One must therefore return to couplet 12. The use of

figure 4 in classifying kinds of uropod 3 can be of

assistance, as can the classification of heads and eyes

in figure 3, although the observations to be made from

the head are of a higher order of difficulty than those

from uropod 3.

U SE OF THE KEYS AND MAJOR PITFALLS.—No Hawaiian gammaridean so far discovered completely

lacks a uropod 3 but several taxa (e.g., Podocerus,

Corophium, Palinnotus) have such small third uropods that superficially these appear to be absent. The

podocerid and phliantid uropod 3 is a tiny flap below

FIGURE 2.—Upper half, The basic gammaridean; lower half, variations in pertinent structures of

Gammaridea not otherwise shown in subsequent figures:

al conjoint main flagellar

base (stippled) rarely

occurring

ac variations in accessory

flagellum

head of a species with //-/

conically arranged

mouthparts

lower lip of species tl—2

with outer lobes

incised

lateral view of normal

telson with keel

stippled

lateral view of fleshy

telson
NUMBER 5 8 11

ac ac

• CD.
12

the telson and has to be observed by mounting the

urosome ventral side up on a depression slide and focusing with high power on a compound microscope.

Many taxa of Hawaiian amphipods frequently shed

uropod 3 on preservation (examples are Gitanopsis,

Melita, Ceradocus, Maera, Leucothoe, Kanaloa; these

genera have exceptionally elongated uropod 3. The

socket or imagination on pleonito 6 remain as marks

of the presence of the uropod. Until some familiarity is

gained with other characters one prefers to avoid

specimens lacking uropod 3 but, as a last resort, one

may examine brief remarks on each family herein to

identify the material by elimination.

The keys cannot be used to elaborate diagnoses of

taxa because several are cited twice, in some cases in

erroneous positions so as to handle possible observational errors. For example, Oedicerotitdae in couplet 7

of the basic key are misplaced, as they have an elongate peduncle on uropod 3 and belong legitimately

where placed in couplet 6 of Key Section B.

Identifications—Non-Objective

FIGURES 3, 4

The term non-objective is used here in place of "subjective" because the methods discussed in this section,

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

when fully elucidated, will probably be perfectly

objective.

Once a specialist learns the main species of a fauna

such as that in Hawaii, he is generally familiar with

the overall appearance of those species or, by trial and

error, has memorized various clues to their identity.

At this point, if he is processing large quantities of

material for ecological studies, he does not necessarily

pursue a formal identification with each specimen but

may separate them into groups by means of nonobjective criteria. Shape of body and head, eye color,

and structure of uropod 3 are often characters of first

recall. They can also be used by the nonspecialist to

sort materials into groups before pursuing the formal

identifications. Characters of antennae, pereopods and

gnathopods can also be useful; but antennae and

pereopods often become separated from preserved

specimens, and gnathopods often widely differ between

the sexes and according to the maturity of the amphipod. Uropod 3 also often falls off specimens, and such

specimens should generally be avoided. If uropod 3 is

absent one may generally conclude that it was very

large or elongate, conditions which are clues to the

identity of the amphipod.

The following four sections present several schemes

for non-objective identifications based on general body

FIGURE 3.—Heads and eyes of Hawaiian Gammaridea. Heads of Biancolina, Ochlesis and Palinnotus are omitted, since for them body shape is more important than heads as a recognition

character. The left hand side of the figure is mostly composed of black eyes (solid ink), the right

red-clear eyes (outlines and clear ommatidia only) whereas in the middle various stippled eyes

indicate that either dark or red-clear eyes occur in different species or genera with the head

designated. Identifications are as follows:

A

AN

AO

AS

C

CEM

D

E

ER

ET

F

GAN

H

3

L

Ampeltsca

Arumixis

Aoroides

AmphilochusGitanopis-Gitana

Colomastix

Ceradocus-ElasmopusMaera (rarely)—

Melita

Parapleustes

Eusiroides

Eriopisa-Eriopisella

Gammaropsis atlantica

Leucothoidae

Gammaropsis-AloiloiNeomicrodeiUopus

Hyalidae and

Parhyalella

Jassa and Ischyrocerus

Liljeborgia

LM

M

MO

N

0

P

PS

PT

R

RI

S

St

T

U

UX

V

VN

Lembos and Maera

Ampithoidae

Mokuoloe

(Amphilochidae)

Nuuanu

Kanaloa

(Oedicerotidae)

Podoceridae

Photis

Pontogeneia

Corophium

Ericthonius

Seba (glands shown)

Stenothoe

Atylus

Wailele

Paraphoxus

Ventqjassa and

Parajassa

Chevalia

X

r

0

V

w

y

yy

z

Chelura (eyes amber

in alcohol)

Lysianassa

Eyes:

absent

medium brown or

caramel

dark, black, brown,

deep purple

red, orange, yellow,

clear

permanently yellow

medium to pale

purple
NUMBER 5 8 13

.v

383-197 O - 71 - 2
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

shape, overall color patterns, the specific pattern of

head shape combined with eye shape and color, and

the morphology of uropod 3. The last method is least

subjective but it is generally useful to classify only to

generic level. By more extensive study, however, one

may be able to develop it to greater precision. The

observation of heads is more precise to specific level

than the other methods, but its use requires considerable experience. The overall body shapes are useful

only in a few genera. The general color patterns of

bodies in various preservatives and in life hold great

promise in making rapid identifications, but far more

study is required before this method is feasible. These

sections are presented mainly as a stimulus to biologists

residing in Hawaii, in the hope that they can perfect

these methods of identification.

BODY SHAPE.—Because several species of Gammaridea have characteristic body appearances, they will

be recognized long before a look at uropod 3 or the

head is necessary. For example, the following taxa are

best recognized by their general appearance (see

figures herein and in Sars, 1895) : Ochlesidae, Podoceridae, Colomastigidae, Corophium, Cheluridae, and

Biancolinidae. The hard, clear white, glossy bodies of

Lysianassidae and Sebidae are likewise characteristic.

COLOR IN PRESERVATIVE.—Amphipods preserved in

formaldehyde-seawater, though changed in color from

life, usually retain a specific color pattern for a few

days; and once this color pattern has been matched

to certain species, they may be recognized and sorted

into groups with rapidity and fair precision. Since

they are most easily and inexpensively preserved

initially in a formaldehyde-seawater solution and since

it is desirable to examine the carcasses in freshwater

within 1-2 days after collection, washing the amphipods in freshwater is convenient and it removes the

offensive preservative as well; they should, of course,

be represented in alcohol. Species of some genera like

Amphilochus have several color phenotypes in a 2-day

formaldehyde solution, whereas others turn translucent

or white but often retain a fairly stable color in the

eyes. The frosted appearance of amphipod eyes is lost

after death. The colors of orange, red, buff, yellow,

green, pink and gray fade after death, and quite

rapidly in formaldehyde, but black and brown in the

eyes are usually retained both in formaldehyde and in

alcohol, though they may be seen under high power

magnification to be brownish or purple. Hence the

distinction between black (brown, purple) and nonblack can be useful in identifying species, and this

works well also for specimens preserved in alcohol,

since eyes that have become translucent in alcohol

generally would have become red or yellow in formaldehyde. This distinction between black and non-black

can be used for example, to distinguish juveniles of

Elasmopus calliactis, which lack the dark ocular pigment, from juveniles of similar species, which have it.

The retention of purplish brown pigments on the

body is very useful in distinguishing species of Lembos

(p. 50) and may become useful in other genera when

they can be studied extensively. The patterns are best

seen under low-power stereoscopic microscopy. Dark

pigments such as black, purple, or brown may actually

be covered in life with some other color, like red. Red

FIGURE 4.—Kinds of uropod 3 in Hawaiian Gammarida; all are drawn as left-sided so that the

outer ramus of each occurs towards the top of this illustration. Letters in capitals refer to familial

groups (sometimes used to identify genus when family is monotypic); letters in lower case refer

to infra-familial groups, mainly genera.

A Arapeliscidac

B Biancolinidae

C Colomastigidae

D Pleustidae

E Eusiridae

F Leucothoidae

G Gammaridae

H Hyalidae-Hyalellidae

/ Ischyroceridae

J Oedicerotidae

L Liljeborgiidae

M Ampithoidae

N Anamixdae

O Ochlesidae

P Podoceridae

Q, Stenothoidae

R Corophiidae

S Sebidae

T Atylidae-Dexaminidae

U Isaeidae-Aoridae

V Phliantidae

W Phoxocephalidae

(Paraphoxus)

X Cheluridae

T Lysianassidae

(Lysianassa)

Z Amphilochidae

e Elasmopus

h Hyale and Parhyalella

i Ischyrocerus

j Jassa

I Chevalia

m Melita

n Nuuanu

o Corophium

p Photis

q Gammaropsis

r Eriopsa

s aorids

t Neomicrodeutopus

v Ventojassa

w Ericthonius

y Parhyale

Z Maera
NUMBER 5 8 15

Hy
16

pigment dissolves easily in preservatives under most

circumstances, either turning yellow or revealing a yellow pigment mixed with it, and this yellow usually

disappears some days later to leave the cuticle or eye

translucent. Dissolution of the pigment may occur

differentially in various specimens, possibly indicating

different phenotypes recognizable in life as red or

white-eyed. This is especially noticeable in Ericthonius

brasiliensis; a sample of this species may contain specimens with red, yellow, and clear eyes, but all of these,

after a week of preservation, will bear clear eyes. Eyes

deteriorate in any preservative, so that museum specimens often have amber eyes or the eyes undergo chemical reorganization and resemble egg yolks.

A well-developed scheme of identification based on

color cannot be presented here, as I was unable to

amass sufficient information on all of the Hawaiian

species, but the eyes alone have been examined for

color in all species. Most of these observations were

made within two days of collection on specimens preserved in formaldehyde; this operation became a

standard procedure and is so noted in the following

sections and in the systematic section of this paper.

HEAD AND EYES (In this section, italic references

in parentheses are to parts of Figure 3).—Cephalic

outline and attendant eye shape are highly characteristic of Phoxocephalidae and Ampeliscidae, even

though their general body appearance is not. Phoxocephalids have visor-shaped rostra (UX) and Ampeliscidae have subcuboidal heads with 2 clear jewel-like

corneal lenses on each side of the head near the anterior margin. Phoxocephalids have dark eyes, and

ampeliscids have red pigmentary masses behind the

lenses and in the center of the head; the red masses

may bleach in alcohol (some genera like Byblis have

brown or black pigmentary masses, but such genera

have not yet been found in shallow waters of Hawaii).

The galeate head of the oedicerotid Kanaloa is also

distinctive (O). The eyes of this genus are deep yellow in either alcohol or formaldehyde, but other oedicerotids yet uncollected in Hawaii are known to have

a wide variety of eye colors, and the tironid Synopia

may also have yellow eyes like Kanaloa.

Heads of Podoceridae (P) have a columnar anterior projection; the eyes are reddish, with cores often

retaining muddy pink-purple pigment in alcohol; and

the species Laetmatophilus hala has dorsal cephalic

protuberances, a highly distinctive feature.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The remaining kinds of gammaridean heads in Figure 3 are less distinctive than those already discussed

and are best classified in groups (left and right in the

figure) as to eye color. Genera with eyes drawn in

solid black are black-brown-purple (="dark") in

formaldehyde after 2 days preservation and they generally remain a dark color after transferral to alcohol.

This color retention is of medium extent in a few

species of Elasmopus, the eyes of which may turn

medium purple in alcohol; whereas in two species of

Amphilochidae, one of Colomastigidae, and three of

Ischyroceridae the eyes may turn muddy magenta or

rusty in either formaldehyde or alcohol. Eyes with

stipples or bearing the code "wy" represent a group in

which eyes may be either dark, red, partly pigmented,

or clear: for example, the head shape in LM is common

to Lembos and Mcera, in most species of which the

eyes are dark except for a few of Maera, which have

red or clear eyes. Eyes drawn in outline represent

heads (and species) in which eyes are red, yellow, or

clear in any preservative.

The distance of observation often makes a difference

in the color observed (one species of Maera has bloodred eyes that appear black from a distance), and purple eyes often appear more strongly pink upon close

observation.

Within a few genera, the species of which have heads

of similar shape, and frequently with ordinary specific

characters obscured in females or juveniles, the species

may often best be distinguished by eye color; examples

are the genera Elasmopus, Maera, Colomastix.

Amphilochus.

Recognition of cephalic outline is difficult as a quick

method of identification unless a great study effort is

undertaken. Here it should be noted that the illustraions of Figure 3 are designedly fragmentary because

they are the views one must become acquainted with.

Many pitfalls exist in their use but the cephalic shape

can be very useful if a wide variety of materials is

available and the student must identify and count numerous specimens for ecological analyses. In addition

to the distinctive head shapes discussed above,

three main points of focus occur in the head: (1) the

shape and size of the rostrum; (2) the protrusions

or indentations at the anteroventral corner of the head,

partially obscured by coxa 1; and (3) the base of

antenna 2, often complex, with obscure sutures and

often with a glandcone on article 2 resembling a tooth

on the head.
NUMBE

R 5 8

17

Identification by means of head characters is ex-

pedited by the ability to manipulate tools and brushes

dexterously, so that the amphipod can be handled with

rapid and gentle movements, and by memorizing

shapes sufficiently to recognize them from any view.

If the amphipod is held with one tool and the pair of

first antennae depressed with another tool, the rostrum

can usually be seen easily. Then coxa 1 can be turned

aside, dirt brushed away, and the head tilted so that

reflected light shines on the sutures at the junction be-

tween the head and antenna 2. With a little practice,

one may determine whether the head is deeply re-

cessed and one may distinguish between a tooth

(glandcone) on article 2 of antenna 2 and a tooth or

excavation on the head proper. For such examination,

fresh specimens with pigmentary stain are superior to

bleached museum specimens, unless special stains are

applied.

The lists below classify the species of Hawaiian

Gammaridea according to eye color, the genera within

each list being arranged alphabetically. The conditions

noted at the head of each list assume observations of a

mounted head, with left coxa 1 removed, under a

compound microscope.

Eyes are absent in Eriopisa laakona and Seba ekepuu

(glandular tissue is shown in outline in figure 3).

List 1. Eyes black or brown in formaldehyde

(2-day) solution, retaining black-brown, deep purple

in alcohol, or eyes with residual black after veneer of

garnet dissolved in alcohol (marked with +); species

retaining opaque blotches of light purple, muddy

brown, and rusty pigment in alcohol are marked with

(x); coffee-brown is indicated by (B), caramel

orange-brown by (C):

Amphilochus menehune x

Ampithoe kaneohe (tiny black spot in alcohol)

Aoroides nahili

Geradocus hawaiensis

Chevalia aviculae (tiny black spot in alcohol) +

Colomastix lunalilo x

Corophium acherusicum

Corophium baconi

Corophium insidiosum

Elasmopus diplonyx x

Elasmopus ecuadorensis hawaiensis

Elasmopus hooheno

Elasmopus molokai

Elasmopus piikoi

Elasmopus pocillimanus

Elasmopus rapax

Elasmopus spinidactylus

Eriopisella sechellensis

Eusiroides diplonyx

Gammaropsis af ra B—C

Gammaropsis alamoana

Gammaropsis haleiwa

Gammaropsis kaumaka

Gammaropsis pali +

Gammaropsis pokipoki

Gitanopsis pele

Hyale affinis

Hyale ayeli

Hyale grandicornis bishopae

Hyale honoluluensis +

Hyale iole

Hyale laie

Hyale sp.

Ischyrocerus kapu x

Ischyrocerus oahu x

Jassa lilipuna x

Konatopus paao C

Lembos aequimanus

Lembos intermedius

Lembos kamanu x

Lembos leapakahi B

Lembos macromanus B

Lembos pualani

Lembos waipio B

Lembos sp.

Liljeborgia heeia

Liljeborgia laniloa

Lysianassa ewa B

Maera insignis

Maera pacifica

Maera quadrimana

Maera sp. A (a few)

Maera sp. B

Melita pahuwai

Neomicrodeutopus makena

Palinnotus alaniphalias

Parajassa angularis

Paraphoxus centralis

Parapleustes derzhavini

Parhyale hawaiensis

Parhyalella pietschmanni +

Photis kapapa

Pontogeneia pacifica

Ventojassa ventosa

Wailele manene
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

List 2. Eyes red (R, known red), yellow or clear

in formaldehyde, turning yellow or clear in alcohol

(CA, known only), except (+) eyes turning black or

purple in alcohol after garnet veneer dissolved away;

(C) indicates caramel-orange; (x), in alcohol portions of eyes with opaque muddy purple, rusty or pink

stain (eyes otherwise in this category are translucent,

never opaque in alcohol, except Kanaloa with permanently yellow pigment); (P), translucent purple

in alcohol, unknown in formaldehyde; (L), head with

4 clear corneal lenses and 5 large red internal pigment

masses; (O), orange in alcohol:

Aloiloi nenue +

Ampelisca schellenbergi L

Amphilochus kailua CA

Amphilochus likelike R

Amphilochus menehune x

Ampithoe akuolaka P

Ampithoe kaneohe +, P

Ampithoe orientalis CA

Ampithoe poipu CA

Ampithoe ramondi

Ampithoe waialua P

Ampithoe spp. CA

Anamixis stebbingi R

Aoroides columbiae C, O

Atylus nani R

Biancolina mauihina R

Chevalia aviculae R

Colomastix kapiolani R

Colomastix pusilla R

Colomastix sp. R

Cymadusa filosa CA

Cymadusa hawaiensis CA

Cymadusa oceanica CA

Elasmopus calliactis R

Elasmopus pectenicrus CA

Ericthonius brasiliensis R

Eriopisa hamakua O

Gammaropsis atlantica R

Gammaropsis pali +

Gitana liliuokalaniae R

Hyale honoluluensis +

Hyale waimea R

Ischyrocerus sp. C O

Jassa lilipuna x

Kanaloa manoa Y

Konatopus paao C

Laetmatophilus hala R

Lembos kamanu P

Leucothoe hyhelia R

Leucothoe lihue R

Leucothoe tridens R

Leucothoides pottsi R

Maera kaiulani O

Maera serrata P

Maera sp.A R

Melita appendiculata CA

Mokuoloe ninole R

Nuuanu amikai O

Ochlesis alii O

Paragrubia vorax CA, P

Parapleustes honomu R

Parhyalella pietschmanni +

Photis aina CA

Photis hawaiensis R

Podocerus brasiliensis R

Podocerus hanapepe R

Podocerus talegus R

Stenothoe haleloke R

Stenothoe valida R

Stenothoe sp.A R

Stenothoe sp.B R

Tropichelura insulae (eyes amber in alcohol)

KINDS OF UROPOD 3 (In this section italic references in parentheses are to parts of Figure 4).—The

morphology of uropod 3 can be helpful in rapidly

identifying a specimen to the familial or generic level.

The basic uropod 3 (E) has a short peduncle and long

lanceolate rami characteristic of Eusiridae (and nonHawaiian Gammaridae). The Dexaminidae (=Atylidae) (L and T, dotted lines) have the basic uropod 3

whereas the Ampeliscidae (A) and Liljeborgiidae (L,

solid lines), have the outer ramus slightly shortened.

The Pleustidae (D) have the outer ramus strongly

shortened and the Colomastigidae (C) and Ochlesidae

(O) have the outer shortened and the peduncle slightly

elongate. The Phoxocephalidae (W) have a 2-articulate outer ramus with the inner ramus slightly shortened but the rami remain thin, whereas in Gammaridae

such as Maera and Elasmopus (Gez) the rami are

broad and blunt and have multiple spine groups and

the outer ramus may or may not have a second article.

In Melita, Eriopisa-Eriopisella, and Nuuanu (Gmnr),

the inner ramus becomes very short and the outer

ramus very elongate. These long uropods (that of

FJNZ, plus a long version of Gez) may often be lost in

preserved amphipods.
NUMBER 5 8 19

The isaeid (and aorid) uropod 3 (U) is difficult to

classify, in that the peduncle is slightly to moderately

elongate, but the rami equal or exceed the peduncle in

length; they are usually thicker and more cylindrioid

or cubiform than in the other taxa discussed above, and

the inner ramus in Photis is shortened and absent in

Neomicrodeutopus.

Biancolina has a uropod 3 (B) similar to that of the

isaeids but it is very small and has short, weak spinesetae.

The Oedicerotidae, Amphilochidae, Leucothoidae

and Anamixidae are characterized by a very elongate

uropod 3 with both peduncle and rami elongate; the

outer ramus occasionally is shortened, especially in the

Amphilochidae (FJNZ). A shorter and stouter version

of this uropod is seen in Lysianassa (Y).

An elongate peduncle and short rami characterize

the Ischyroceridae (/), and to some extent the Ampithoidae (M) but in Jassa the peduncle may not be so

conspicuously elongate as in other ischyrocerids (see

dotted line in /). Ischyroceridae lack the long setae on

the inner ramus of uropod 3 found in the Ampithoidae,

and the latter have one or two large hooks on the outer

ramus, these hooks being larger than in Jassa. Ischyrocerids generally have small denticles, cusps, or an

apical uncinus on the outer ramus (see variations in

Iv, Ij, Ii). Ventojassa has 1-3 stiff setae on the outer

ramus but it is an exceptional ischyrocerid (Iv). The

outer ramus of ischyrocerids generally has to be viewed

with an oil-immersion lens, though uropod 3 of Jassa

is clear under high power.

The remaining kinds of uropod 3 shown in Figure 4

all lack an inner ramus or have no ramus at all. Neomicrodeutopus in the Isaeidae (Aoridae) lacks this ramus

and so does Ericthonius in the Corophiidae (Rw).

Ericthonius has the long peduncle characteristic of

Ischyroceridae and the only ramus present appears also

to be ischyrocerid in structure. The other Hawaiian

corophiid, Corophium (Ro), has a small flabellate

uropod 3 with a short peduncle and slightly longer

ramus; it requires some patience to distinguish the

appendage from the nearby telson and uropod 3.

Palinnotus (V) has a thin replica of the corophiid

uropod 3 and Tropichelura (X) has an enormously

long replica of uropod 3 seen in Corophium.

Podocerus (P) has a reduced uropod 3, lacking a

ramus and not visible from the side, that lies underneath a large fleshy telson, as shown in the figure.

The talitroids Hyale and Parhyalella (Hh) and

Parhyale [Hy) have an easily visible uropod 3, but the

ramus and peduncle are small and subequal in length

and have short, stout spines in contrast to Neomicrodeutopus and Palinnotus. Parhyale has a scale-like

inner ramus.

The final morph is the 3-piece uropod 3 composed

of a peduncle and a 2-articulate ramus found in the

Stenothoidae (Q) and in Seba (S). The latter has a

broadly flattened uropod 3 with shortened article 2

of the ramus in contrast to that of the Stenothoidae.

Taxonomic Arrangement

Because of the convenience to identification, portions

of the old-fashioned system of classification of gammarideans are retained herein. The families Aoridae,

Corophiidae and Isaeidae ( = Photidae) are maintained distinctly, as Corophiidae have only two easily

recognizable genera in Hawaii and because the

enlargement of gnathopod 1 in Aoridae is a convenient

way to separate the many species of aorids and isaeids

in Hawaiian waters.

Referral of corophiids to the isaeid complex is a

semantic error, as the term corophiid has priority. The

ultimate referral of photids, aorids, isaeids, ischyrocerids, ampithoids, podocerids, chelurids and other

groups to a broadly conceived superfamily of corophiids should occur soon and the preferable term

corophiidean applied to the complex. Not only is the

Corophiidae the oldest rooted name in the group, it

has also been most widely used in textbooks and thus

is familiar to the general zoologist. The transition from

photid to isaeid to corophiidean that has occupied

the past two decades is an unfortunate contemporary

unheaval necessitated by the correlation of names to

familial concepts as they are slowly but carefully

studied and amalgamated. If Corophium, lending its

name to the oldest familial root, were not one of the

most atypical of the isaeid-photid-corophiid members,

the reestablishment of an inclusive name for the whole

group would, undoubtedly, have occurred sooner.

Order AMPHIPODA

Suborder GAMMARIDEA

DIAGNOSIS (restricted to Hawaiian marine species,

0-30m).—Pleon (abdomen) distinctly divided into

anterior and posterior sections of 3 segments (or their

vestiges) each, first section (metasome) bearing 3 pairs
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of flexible pleopods, second section (urosome) bearing

2-3 pairs of stiff uropods; pleopods usually partially

hidden by lateral pleurae (epimera), terminal segment

of urosome with a flap (telson); carapace absent,

thorax (mesosome) composed of 7 freely visible segments, legs attached to segments composed of 2 pairs

of prehensile gnathopods (segments 1-2), 2 pairs of

forward acting pereopods (segments 3-4) and 3 pairs

of lateroposteriorly acting and/or laterally flattened

pereopods (segments 5-7); coxae conspicuous; head

with 2 pairs of antennae but no movable scales and

only antenna 1 occasionally weakly biramous; eyes

sessile and enclosed within head or rarely formed of

corneal lenses on surface.

REMARKS.—Only tanaids and isopods might be

confused with Gammaridea but neither of the first two

orders has precisely 3 pairs of pleopods nor more than

one pair of uropods. The lateral plate-like coxae (the

first articles of the thoracic appendages) are enlarged

in Hawaiian marine gammarideans to an extent not

seen in other local crustaceans. Unlike various shrimps

and mysids, amphipods lack a carapace.

Key to Hawaiian Families of Gammaridea

Figure Numbers Referenced in Italics

Notes:

COUPLET 1.—The inner ramus of Parhyale cannot be seen if uropod 3 is mounted upside

down; if the uropod appears uniramous the cover slip should be moved so as to roll the uropod

for examination on all surfaces; occasionally one ramus will break off but the socket remains.

COUPLET 2.—A failure to clip off all of the appendage can result in an abnormally

shortened uropod 3; the appendage should also be checked on slide 2, before dissection.

COUPLET 4.—Coxa 1 is counted as article 1.

COUPLET 5.—Lenses usually are difficult to see except with finely focused compound

microscope on slide 2, medium power.

COUPLET 7.—Accessory flagellum is occasionally broken, or antenna 1 is frequently broken

between articles 1 and 2; the accessory flagellum is on the mediodistal margin of article 3;

antenna 1 on slide 8 should be rotated to see all surfaces with high power.

COUPLET 8.—Extremely rare occurrence; maxillipedal palps universally present in Gammaridea except for this genus and a few deep-sea taxa.

COUPLET 9.—Both sexes may be required.

SECTION A, COUPLET 3.—The palp is often broken off during dissection; slide 2 should be

checked before dissecting mouthparts; mandibular palp projects forward between the second

antennae and the base of the palp may be seen attached to the mandible just below the anteroventral corner of the head.

SECTION B, COUPLET 4.—Oil immersion objective may be required to see these tiny

ornaments.

1. Uropod 3 with one ramus or without rami (4, lower) KEY SECTION A

Uropod 3 with 2 rami, inner occasionally very small (4, upper) 2

2. Peduncle of uropod 3 of normal dimensions, short (4E£^A) 3

Peduncle of uropod 3 elongate, either actually, or relative to rami (4FJNZJMJ,Y).

KEY SECTION B

3. Inner ramus of third uropod less than half as long as outer ramus (4Hy,Up,Gmnr).

KEY SECTION C

Inner ramus of third uropod 75 percent or more as long as outer ramus (occasionally outer

ramus shorter than inner) (4Ge) 4

4. Article 3 of gnathopod 2 elongate and gnathopod 2 like Figure 52 g2 + body arrow.

Lysianassidae (p. 105)

Article 3 of gnathopod 2 not elongate or gnathopod 2 not like Figure 52g2 + body arrow 5

5. Head elongate and bearing small external cuticular lenses 4 in number (3A,5Ah) (coxae

long and urosomites 2-3 coalesced) Ampeliscidae (p. 22)

If elongate, head massive or very flat, lacking special lenses, eyes otherwise ommatidial,

internal, and multifaceted 6

6. Head massive, galeate, eyes large and essentially coalesced into one mass (seen from dorsal

view) (eyes not black in alcohol) (3O) 7

Head not massive, not galeate, eyes if present separate 8
NUMBER 5 8 21

7. Accessory flagellum 3-f- articulate, pereopod 5 if elongate much less than 1.5 times as long

as pereopod 4, pereopods 3-4 not strongly shortened Synopiidac*

Accessory flagellum absent or vestigial, pereopod 5 remarkably elongate, at least 1.5 times

as long as shortened pereopods 3-4 (54, body) Oedicerotidae (p. 105)

8. Maxillipeds lacking palp (53xp), mouthparts arranged in conical bundle, body with form

and coxae of Figure 53 Ochlesidac (p. 105)

Maxilliped with palp (2xp), body and coxae not like Figure 53 9

9. Body cylindrical or subcylindrical (22, 24), mandible lacking palp 10

Body compressed laterally or not distinctly subcylindrical, mandible with palp 11

10. Urosomites 2-3 coalesced, uropod 3 of medium length, at least one ramus elongate,

antennae 1-2 stout, short, stiff, projecting anteriorly, head short and thick (24).

Colomastigidae (p. 54)

Urosomites 2-3 free, uropod 3 short, rami and peduncle equal to each other, peduncle

flat, with gap between attached rami, antennae 1-2 thin, elongate, turned posteriorly,

head flat like that of insect or spherical and bearing constricted neck (22).

Biancolinidae (p. 52)

11. Telson thick, short, entire, fishy (2tl2) 12

Telson cleft or entire, short or long, often with ventral keel, but thin and not fleshy (2tll).

KEY SECTION D

12. Gnathopod 1 showing sexual dimorphism and longer or stouter than gnathopod 2 (16C,W).

Aoridae (p. 40)

Gnathopod 1 not showing sexual dimorphism and smaller than gnathopod 2, gnathopodal

dimorphism, if present, confined to gnathopod 2 (7) Isaeidae ( = Photidae) (p. 88)

SECTION A

1. Both pairs of gnathopods chelate Sebidae (p. 117)

Gnathopods usually subchelate or simple, both pairs never chelate together 2

2. Gnathopod 2 typical of Figure 52g2 + body arrow Lysianassidac (p. 105)

Gnathopod 2 not fitting Figure 52 3

3. Coxa 1 small and covered by coxa 2 (and 3) (62H) Stenothoidae (p. 120)

Coxa 1 visible and not less than half surface area of coxa 2 (1) 4

4. Body flat dorsoventrally, or cylindrical, often broadened and/or rugose (23, 55, 58) 6

Body compressed laterally (normal gammaridean) (/) 5

5. Mandible lacking palp Talitroidea (p. 122)

Mandible with palp Isaeidae(=Photidae) (in part, p. 88)

6. Uropod 3 very large, almost as long as urosome, uropod 2 of abnormal, elephantine structure (23) Cheluridae (p. 54)

Uropod 3 very small or seemingly absent, uropod 2 normally slender 7

7. Mandible lacking palp, molar vestigial or absent 8

Mandible with palp and triturative molar 9

8. Body cylindrical, coxae very small and not splayed Eophliantidac*

Body very broad, rugose, armored, depressed, coxae large and splayed (55).

Phliantidac (p. 109)

9. Urosomite 1 elongate (58L,T) Podoceridae (p. 113)

Urosomite 1 not elongate (/) Corophiidae (p. 55)

SECTION B

1. Gnathopod 2 typical of Figure 52g2 + body arrow Lysianassidae (p. 105)

Gnathopod 2 not fitting Figure 52 2

2. Gnathopod 1 either absent or with the form of Figure 14Lgl and Agl 3

Gnathopod 1 present and not like Figure 14 4

3. Mandibles and maxillae absent or extremely vestigial and replaced by large, easily visible

ventral keel (15Smt) Anamixidac (p. 40)

Mandibles and maxillae present in ordinary condition Lcucothoidac (p. 102)

*not yet found in Hawaii.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4. Uropod 3 with special form of Figure 4M, outer ramus with 2 large hooks (rarely one in

non-Hawaiian genus) Ampithoidae (p. 29)

Uropod 3 not like Figure 4M 5

5. Rami of uropod 3 elongate (4FJNZ), outer not uncinate 6

Kami of uropod 3 shorter than peduncle, outer minutely uncinate and subapically ornamented

with tiny denticles (4Iijv) Ischyroceridae (p. 97)

6. Coxa 1 small and partially hidden by coxa 2, pereopod 5 not grossly longer than pereopod

4 (6P) Amphilochidae (p. 25)

Coxa 1 large and visible, pereopod 5 much longer than pereopod 4 (54, body).

Oedicerotidae (p. 105)

SECTION C

1. Inner ramus of uropod 3 scalelike and partially appressed to outer ramus (4Hy), mandible

lacking palp Talitroidea (in part, p. 122)

Inner ramus of uropod 3 easily visible and distinct from outer ramus (4E, Up), mandible

with palp 2

2. Head scarcely rostrate, telson uncleft (fleshy), pereopod 5 resembHng 4.

Isaeidae ( = Photidae) (p. 88)

Head with visor rostrum (3UX), telson cleft, pereopods 4 and 5 grossly different (5Pp4 and

Pp5) Phoxocephalidae (p. 109)

Head poorly rostrate, telson cleft, pereopod 5 resembling 4 (/) Gammaridae (p. 66)

SECTION D

1. Urosomites 2—3 or 1—3 coalesced and mandibular palp absent.

Dexaminidac (=Atylidae) (p. 61)

Urosomites 2-3 free and mandibular palp present 2

2. Mandibular molar not triturative (51 Hm), but often bearing articulate spines 3

Mandibular molar triturative (bearing ridges and cusps) (2mo) 5

3. Pereopods 3-5 and peduncle of antenna 2 strongly spinose and setose, pereopod 5 shorter

of strongly different shape from pereopod 4 (5) Phoxocephalidae (p. 109)

Pereopods 3-5 and peduncle of antenna 2 weakly setose, pereopod 5 like pereopod 4 4

4. Telson cleft, accessory flagellum very long and stout (51H) Liljeborgiidae (p. 103)

Telson entire, accessory flagellum vestigial or absent, not visible grossly (high-power microscope necessary) Pleustidae (p. 109)

5. Inner ramus of uropod 3 much shorter than outer (less than one third).

Gammaridae (in part, p. 66)

Rami of uropod 3 subequal to each other in length or outer slightly shorter than inner 6

6. Accessory flagellum with 2 or more articles Gammaridae (in part, p. 66)

Accessory flagullum 1-articulate or absent 7

7. Telson cleft Eusiridae (p. 61)

Telson entire Pleustidae (p. 109)

Family AMPELISCIDAE

Genus AmpeUsca Kr0yer

Ampelisca schellenbergi Shoemaker

FIOURE 5A

Ampelisca schellenbergi. Shoemaker, 1933b: 3-5, fig. 2; J. L.

Barnard, 1954a: 14-16, pis. 7, 8; J. L. Barnard, 1967:8-

10, figs. 1 a-m.

Recognized by the elongate, tall head, coalesced urosomites 2-3, the presence of 4 small, shiny, surficial corneal lenses on the head (2 on each anterolateral side),

head with bright red pigment in 5 large spots (before

placing in alcohol), the characteristic pereopods 4 and

5, 5 being shorter and of different morphology from

4 (5 Ap4 and Ap5). The family differs from phoxocephalids among many characters in the absence of an

accessory flagellum, lack of rostrum, relatively short

pereopod 4 and coalesced urosomites 2-3.

Ampelisca vaguely resembles Chevalia in the Isaeidae but the latter genus has urosomites 1-2 coalesced

and 3 is free, while the eyes are ommatidial, internal,

and multifaceted.

Ampeliscids build soft domiciliary tubes on mud

bottoms but this Pan-American species is also associated with surf grasses on the continent.

Pan-American; off Honolulu, 93-229 m.
NUMBER 5 8 23

FIGURE 5.—Ampcliscidae and Phoxocephalidae: A, Ampelisca schellenbergi Shoemaker; p, Paraphoxus centralis (Schellenberg).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 5 8 25

Family AMPHILOCHIDAE

FIGURES 6-8

This family is characterized by the small coxa 1 (and

occasionally small coxa 2), elongate biramous uropod

3 with long peduncle, large coxa 4, conspicuous rostrum, and shortened uropod 2. It differs from the superficially similar Ischyroceridae by the presence of a

conspicuous rostrum, the grossly excavate coxa 4, and

thin telson. Stenothoidae differ from amphilochids in

the uniramous uropod 3. Hawaiian anamixids and

those few leucothoids with small coxa 1 differ from

amphilochids in the special configuration of gnathopod

1 with its exaggerated carpochelation.

Amphilochids are very small (0.7 to 2.8 mm long)

and are difficult to dissect, but presumably we now

have in hand the most common Oahuan species and

they can be identified by small scale characters, usually

without dissection. But it is advisable to confirm initial

identifications by dissecting mouthparts and checking

them for generic validity until one becomes routinely

familiar with the general appearance of amphilochids.

A mount of the whole animal on slide 2 is examined

with a compound microscope to observe gnathopods

1-2, pereopods and uropods; gnathopods should be

extended before mounting and dirt removed by camelshair brush or syringing with alcohol, and gnathopodal

dactyls slightly extended away from the palmar surfaces so as to observe dactylar pectinations—these are

difficult to see on dirty specimens with dactyls fully

closed. The characters may be checked in the keys but

a short diagnosis of each species follows so as to balance

all characters.

Hawaiian amphilochuses appear to be strongly associated with scleractinian corals of the genus Pocillopora. Many of the amphipods have the same coffee or

bronze color of the coral; possibly they feed on the

mucus exuded by the coelenterate.

Schellenberg's (1938) identification of Amphilochus

marionis Stebbing, from Oahu, has not been confirmed

in the present study. I am at a loss to recognize that

species among hundreds of amphilochids collected recently. The surprising diversity of the family and the

presumed high endemicity of the species indicate the

improbability of a subantarctic species occurring in

Hawaii.

Key to Genera of Amphilochidae

Figure Numbers Referenced in Italics

1. Coxae 3 and 4 contiguous and not overlapping, very large and covering reduced coxae 1

and 2 (8, body) (Cyproideinae) Afokuoloe J. L. Barnard

Coxa 2 large, visible, coxae 2-4 overlapping each other, coxa 2 covering reduced coxa 1

(6P) (Amphilochinae) 2

2. Gnathopods extremely weak, essentially simple (6Lgl), palp of maxilla 1 uniarticulate.

Gitana Boeck

Gnathopods of medium to large size, palms well developed and transverse (6Kgl), palp of

maxilla 1 biarticulate 3

3. Mandibular molar weakly triturative {7Mm) and elongate-conical Amphilochus Bate

Mandibular molar heavily triturative and cushion-shaped (7Pm) Gitanopsis Sars

FIGURE 6.—Amphilochidae: i, Amphilochus likelike J. L.

Barnard; K, A. kailua J. L. Barnard L, Gitana liliuokalaniae

J. L. Barnard; M, Amphilochus menehune J. L. Barnard; P,

Gitanopsis pele J. L. Barnard.

Genus Gitana Boeck

Gitana liliuokalaniae J. L. Barnard

Gitana liliuokalaniae J. L. Barnard, 1970:37-39, figs. 7, 8.

This species differs from other Hawaiian amphilochids

in the extremely weak and characteristically shaped

gnathopods {6 Lgl and Lg2). Only one specimen has

so far been collected. Eyes red in formaldehyde.

Kaneohe Bay, 3-4 m.
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Amphilochidae: K, Amphilochus kailua J. L. Barnard; L, Gitana likiuokalaniae J. L.

Barnard; M, Amphilochus menehune J. L. Barnard; p, Gitanopsis pele J. L. Barnard.
NUMBER 5 8 27

xp

FIGURE 8.—Amphilochidae: Mokuoloe ninole J. L. Barnard.
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Hawaiian Species of Genera Amphilochus Bate and Gitanopsis Sars

Figure Numbers Referenced in Italics

1. Gnathopods 1 and 2 small to medium (6P), hands smaller than coxa 2, mandibular molar

flat, cushion-shaped, extremely triturative (7Pm) G. pele

Gnathopods 1 and 2 large, hands larger than or subequal to coxa 2, mandibular molar long,

subconical, weakly triturative (7Mm) 2

2. Article 4 of gnathopod 2 with 1-3 stout posterior spines, anterodistal end of hand (article 6)

sharp, dactyl of gnathopods 1-2 with minute serrations, scales or pectinae on inner margin

besides large tooth (6Mg2) A. menehune

Article 4 of gnathopod 2 without stout spines, anterodistal end of hand softly rounded, dactyls

of gnathopods 1-2 lacking serrations, scales or pectinae except for large tooth (6Kgl).... 3

3. Dactyls of pereopods 1-5 with large mamilliform hump (7Kp2), posterodistal end of article 2

on gnathopod 2 lacking stout spines (6Kg2) A. kailua

Dactyls of pereopods 1-5 simple, posterodistal end of article 2 on gnathopod 2 bearing stout

spine (6Igl) A. likelike

Amphilochus kailua J. L. Barnard

Amphilochus kailua J. L. Barnard, 1970:30-33, figs. 2, 3.

Gnathopods 1-2 large, hands equal to or larger than

coxa 2, article 4 of gnathopod 2 lacking large stout

posterior spines (but possibly with setae), anterodistal

end of hand of gnathopod 2 softly rounded, dactyls

of gnathopods lacking serrations except for main tooth;

dactyls of pereopods 1-5 with relatively large mamilliform hump; posterodistal end of article 2 of gnathopod 2 with slender or no seta. Dissection: mandibular

molar of medium size, subconical and weakly triturative; outer plate of maxilliped with convex, unpectinate medial margin and several mediomarginal setae.

South shore of Oahu, intertidal algae, to 5 m on

sand bottom.

Amphilochus likelike J. L. Barnard

Amphilochus likelike J. L. Barnard, 1970:33, fig. 4.

Gnathopods 1-2 large, hands equal to or larger than

coxa 2, article 4 of gnathopod 2 lacking stout posterior

spines (but possibly with setae), anterodistal end of

hand on gnathopod 2 softly rounded; dactyls of gnathopods lacking serrations except for main tooth; dactyls

of pereopods 1-5 simple; posterodistal end of article

2 on gnathopod 1 with a stout spine. Dissection: mandibular molar of medium size, subconical and weakly

triturative; outer plate of maxilliped with slightly

excavate and pectinate distomedial margin, distolateral

end with 1 large sabre-spine.

Oahu, 2-30 m, especially on Pocillopora.

Amphilochus menehune J. L. Barnard

Amphilochus menehune J. L. Barnard, 1970:33-37, figs. 5, 6.

Gnathopods 1-2 large, hands equal to or larger than

coxa 2, article 4 of gnathopod 2 with 1-3 stout, posterior spines, anterodistal end of hand of gnathopod

2 sharply extended (but minutely); dactyls of gnathopods pectinate or serrate on inner edge apart from

main tooth; dactyls of pereopods 1—5 simple; posterodistal end of article 2 on gnathopod 2 with slender or

no seta. Dissection: mandibular molar of medium size,

subconical and weakly triturative; outer plate of maxilliped with slightly excavate and pectinate distomedial

margin, distolateral end with 1 large sabre-spine. Eyes

brown.

Oahu, 0-33 m, widespread and abundant in numerous biotopes.

Gitanopsis pele J. L. Barnard

Gitanopsis pele J. L. Barnard, 1970:40, fig. 9.

Gnathopods 1-2 small, hands smaller than coxa 2,

article 4 of gnathopod 2 lacking posterior spines (possibly with setae), anterodistal end of hand of gnathopod 2 softly rounded; dactyls of gnathpods pectinate
NUMBER 5 8 29

or serrate on inner edge apart from main tooth; dactyls

of pereopods 1-5 simple; posterodistal end of article

2 on gnathopod 2 with slender or no seta. Dissection:

mandibular molar very large, flat, cushion-shaped, extremely triturative; outer plate of maxilliped with

slightly excavate and pectinate distomedial margin, distolateral end with 1 large sabre-spine. Eyes brown,

unlike other red-eyed amphilochids, except A. menehune.

Oahu, Kaneohe Bay, 2 m, on Pocillopora reef.

Genus Mokuoloe J. L. Barnard

Mokuoloe ninole J. L. Barnard

Mokuoloe ninole J. L. Barnard, 1970:40-44, figs. 10, 11.

Mokuoloe is recognized by the grossly enlarged coxa 4,

remarkable for amphilochids; coxa 4 is almost shieldlike, abuts coxa 3 and the latter covers the reduced

coxae 1 and 2. These are characters of Cyproideinae,

one subfamily of amphilochids and Mokuoloe is the

only cyproidein so far found in Hawaii, though others

probably will be discovered. A short diagnosis of the

genus is given so that other cyproideins may be

recognized.

Antenna 2 with only 2 articles in flagellum; rostrum

short; mandible with palp and triturative molar; maxilla 1 with 1-articulate palp armed with strap-shaped

setae; maxilliped with thin palp, article 3 bearing a

thumb that forms a claw with dactyl, outer plate large,

not excavate; gnathopods 1-2 subchelate, of medium

size, palms oblique and deeply serrate in fan-shaped

fashion, sharp dactyl of gnathopod 1 much longer than

palm, lobe on article 5 weak, dactyl of gnathopod 2

fitting palm, hand with sharp anterodistal extension,

lobe on article 5 long, thick, and guarding article 6;

article 2 of pereopods 3-4 thin, of pereopod 5 expanded; outer ramus of uropod 3 shorter than inner

ramus; pleonites 4 and 6 weakly keeled dorsally on

both sides but pleonite 6 not vaulted over telson; telson

long, broad, triangular and weakly keeled dorsally on

both margins, thus boat-shaped. Cyproidins resemble

Stenothoidae but differ in the biramous uropod 3.

South Oahu, 30 m, on bottom dominated by coralline "footballs."

Family AMPITHOIDAE

FIGURES 9-13

Ampithoids belong with the isaeid group of families

bearing fleshy telson and glandular pereopods 1-2;

coxa 4 is scarcely or not excavate posteriorly; uropod 3

outer ramus has 2 hooks (one hook only in a nonHawaiian genus); the rami are much shorter than the

peduncle, the latter being subcylindrical and the rami

being slightly flattened and subovate (12Wu3). Ampithoids may be confused with ischyrocerids but the latter family has thinner, pointed-rami on uropod 3 and

with one exception their members have only one hook

or simply a curved (uncinate) distal end with minute

denticles lining the subapical margin of the outer

ramus.

Specimens without antenna 1 are difficult, if not

impossible to identify generically unless they are members of Paragrubia. Juveniles and often females of

Ampithoe and Cymadusa are almost impossible to

identify specifically, as their morphological distinctions

have not been clarified in Hawaiian species. Considerable ecotypic, seasonal and "founder effect" morphological variation seems to occur in the species of this

family. Fully mature males are rare.

Probably the largest species of amphipods in the

Hawaiian chain belong to this family. They build

tubes of parchment but are often associated with algae

presumably to which the tubes are attached, possibly

near or among the rhizomes. Members of this family in

California have been observed to roll up edges of kelp

blades and cement them into tubes; this may also occur in Hawaiian algae and observations should be

noted in the literature as this phenomenon has never

been adequately described.

Apparently all species in Hawaii have red eyes or no

black-brown; A. kaneohe however retains a dark spot

in the eye after bleaching in alcohol.

The peduncular tooth frequently mentioned on uropods 1-2 lies distalwards and between the rami so that

the uropod must be viewed ventrally in order to determine the length of the tooth {13Oul).

M3-19T O - 71 - 3
FIGURE 9.—Ampithoidae: A, Ampithoe akuolaka J. L. Barnard; c, Cymadusa oceanic a J. L.

Barnard; R, C. hawaiensis (Schellenberg); K, Ampithoe kaneohe J. L. Barnard; o, A. orienialis

Dana; p, A. (Pleonexes) poipu J. L. Barnard; R, A. ramondi Audouin; s, Ampithoe sp.; v,

Paragrubia vorax Chevreux; w, A. waialua J. L. Barnard.
FIGURE 10.—Ampithoidae: A, Ampithoe akuolaka J. L. Barnard; c, Cymadusa oceanic a J. L.

Barnard; F, C. filosa Savigny; H, C. hawaiensis (Schellenberg); K, A. kaneohe J. L. Barnard;

o, A. orientate Dana; r, A. (Pleonexes) poipu J. L. Barnard; R, A. ramondi Audouin; s, Ampithoe

sp.; v, Paragrubia vorax Chevrcux; w, A. waialua J. L. Barnard.
FIGURE 11.—Ampithoidae, male gnathopod 2: A, Ampithoe akuolaka J. L. Barnard; c, Cymadusa

oceanica J. L. Barnard; F, C. filosa Savigny; H, C. hawaiensis (Schellenberg); K, A. kaneohe

J. L. Barnard; o, A. orientalis Dana; p, A. (Pleonexes) poipu J. L. Barnard; R, A. ramondi

Audouin; w, A. waialuaj. L. Barnard.

FIGURE 12.—Ampithoidae: A, Ampithoe akuolaka J. L. Barnard; F, Cymadusa filosa Savigny;

H, C. hawaiensis (Schellenberg); K, Ampithoe kaneohe J. L. Barnard; P, A. (Pleonexes) poipu

J. L. Barnard; R, A. ramondi Audouin; s, Ampithoe sp.; v, Paragrubia vorax Chevreux; w,

Ampithoe waialua J. L. Barnard.
NUMBER 5 8

Spk
3 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Genera of Ampithoidae

Figure Numbers Referenced in Italics

1. Gnathopod 1 larger than gnathopod 2 in both sexes Paragrubia Chevreux

Gnathopod 1 smaller than or equal to gnathopod 2 in both sexes 2

2. Antenna 1 with accessory flagellum Cymadusa Savigny

Antenna 1 without accessory flagellum 3

3. Telson with very large dorsoposterior hooks (12Pt) Ampithoe (Pleonexes) Bate

Telson with small dorsoposterior hooks, slits or nobs (12At) Ampithoe Leach

Genus Ampithoe Leach The peduncular tooth is present in A. orientalis and

__. . .__ .. , , . _ „ , Ampithoe sp., both otherwise easy to distinguish from

Diagnosis (Hawaii only) : Accessory flagellum absent, ,

r

 „ .. . , . , . ., .. . ., ,

^ , , ,

 7 ; . :; , , other Hawaiian ampithoids in the very thin gnathopod

ventral peduncular tooth of uropod 1 absent, or very , . , „ , , . , , .

, \ , , , , .

 r

 . .' ,

 7 1 with small transverse or parachelate palms, and in

short and blunt or long and sharp, gnathopod I larger . . , , * . , .. . ...

, , , « • , i o i i A- kaneohe m which the tooth is short and blunt unlike

than gnathopod 1 in male, uropod 3 normal, telson .

... „ , -i i i that in Cymadusa.

with small dorsoposterior hooks or nobs.

 J

Uropod 1 of Hawaiian ampithoes often lacks a Probably

 severa l mor e

 Hawaiian ampithoes remain

peduncular tooth; that character makes the separation

 t o b e

 discovered and others might have a peduncular

of most members of the genus from other Hawaiian

 toot h o n

 uropod 1. Fortunately most specimens retain

ampithoids relatively easy in juveniles and females that at least one member of pereopods 3-5, thus permitting

may have missing antennae, or gnathopods that are recognition of A. kaneohe and A. poipu especially but

difficult to distinguish from each other in size. the former species also has significantly small eyes.

Key to Hawaiian Species of Genus Ampithoe Leach

Figure Numbers Referenced in Italics

1. Gnathopod 1 of both sexes with narrow, rectangular article 6 bearing distinctly transverse

or parachelate palm hidden in dense setal brush (10S1) 2

Gnathopod 1 of both sexes with slightly narrowed to broadened article 6 bearing distinctly

oblique, occasionally weak palm not hidden by setal brush (10P1) 3

2. Gnathopods 1-2 of female and gnathopod 1 of male, parachelate (10S1) Amphithoe sp.

Gnathopods 1-2 of female and gnathopod 1 of male with perfectly transverse palm (10 Ol).

A. orientalis

3. Pereopods 3-5 strongly subprehensile, sixth articles with special palmar spine sabre-shaped

and guarding palm (12Kp4) 4

Pereopods 3-5 weakly or not subprehensile, lacking special sabre-spine guarding a palm

(12Sp4) 6

4. Telson with 2 strong reverted dorsal hooks easily seen from lateral view (12Pt).

A. (Pleonexes) poipu

Telson with weak dorsal nob scarcely shaped as hook (12Rt) 5

5. Eyes very small, garnet red in formaldehyde, brown to black to faint purple core in alcohol,

pereopods 3-5 with striate sabre spine strongly guarding distinct palm (12Kp4).

A. kaneohe

Eyes large, presumably red in life, clear in alcohol in adult or with faint purple core in

juvenile, pereopods 3-5 with pair of sub-sabre striate distal locking spines weakly guarding

obsolescent palm (12Rp3) A. ramondi (in part)

6. Pleonal epimera with slight notch at posteroventral corner, bearing weak lateral ridges or

indented line (13Aep), apical lobules of lower lip very short and tumid (9 All), pereopods

3-5 with pair of straight, unstriate distal locking spines lacking any proximal partners

(12Ap5) A.akuolaka

Pleonal epimera smoothly rounded-quadrate posteroventrally (13Kep), lacking lateral ridges,

apicolateral lobule of lower lip thin and projecting farther than medial lobule (9KU),

pereopods 3-5 with pair of sub-sabre striate locking spines with at least one proximal

partner (12Pp3) 7
NUMBER 5 8 35

Pereopods 3-5 with at least 2 spines in tandem proximal to locking pair (12Rp3), article 2

of pereopod 3 with weak anterior spines or notches if any, palm of gnathopod 1 weak

(10R1) A. ramondi (in part)

Pereopods 3-5 with only 1 spine in tandem proximal to locking pair, article 2 of pereopod 3

with 1-2 large anterior notches bearing very stout spine (9Wp3), palm of gnathopod 1

strong {10W1) A. waialua

Ampithoe akuolaka J. L. Barnard

Ampithoe akuolaka J. L. Barnard, 1970:44, figs. 12, 13.

Male gnathopod 1 with articles 5 and 6 equal to each

other in length, article 6 very thin and slightly tapering

distally, palm oblique and weak, defining spine on

posterior margin of hand, article 5 with sharp posterodistal extension; female gnathopod 1 with article 5

lobe trapezoidal, unproduced; male gnathopod 2 with

strong anterodistal lobe on article 2, hand slightly

elongate, not strongly enlarged, palmar region bearing

deep slit separating off long posterior tooth, palmar

tangent oblique but no distinct palmar margin present,

dactyl strongly overlapping palm; female gnathopod

2 with oblique palm distinctly marked by spine, article 5

lobe triangular; pereopods 3-5 simple, bearing pair

of straight, unstriate distal locking spines, with no

closely adjacent proximal partners, main locking spine

on pereopod 3 long, on pereopods 4-5 short; pleonal

epimera with rounded posteroventral cusp, notch and

setule, lateral faces with ridge; eyes of medium size,

pale purple in alcohol; uropod 1 lacking peduncular

process between rami; lower lip with especially short

and tumid apical lobules.

Oahu, intertidal, on Sargassum, Ulva, and rocks.

Ampithoe kaneohe J. L. Barnard

Ampithoe kaneohe J. L. Barnard, 1970:44-^7, figs. 14, 15,

16, 24/.

Male gnathopod 1 with article 6 longer than 5 and

ovatorectangular, well developed oblique palm defined

by spine, article 5 with weak, broad, slightly coniform

posterior lobe, dactyl fitting palm; female gnathopod

1 with lobe on article 5 truncate; male gnathopod 2

with large anterodistal lobe on article 2, hand very

elongate, palm and posterior margin of hand congruent, slightly setose, dactyl about half as long as

hand, medioterminal face of hand with long setal

brush; female gnathopod 2 like gnathopod 1 but lobe

of article 5 sharp; pereopods 4-5 strongly subprehensile, bearing palm guarded by prostrate striate sabrespine and 3 other erect striate spines; pleonal epimera

1-3 rounded posteroventrally; eyes very small, clear in

alcohol or bearing slight dark core near posterior edge;

uropod 1 with short truncate peduncular process between rami.

Oahu, intertidal and 30 m.

Ampithoe orientalis Dana

Ampithoe orientalis Dana, 1853:937-939, pi. 64, fig. 2.

Ampithoe orientalis J. L. Barnard, 1955:26-28, fig. 14; 1970:

47, fig. 17a-t.

Male and female gnathopod 1 with articles 5 and 6

subequal to each other in length, both thin, article 5

rectangular, bearing short transverse palm hidden by

setae, article 5 unlobate; male gnathopod 2 with weak

anterodistal lobe on article 2, hand elongate, palm

and posterior margin of hand congruent, distally

sinuous, weakly setose, dactyl as long as hand, in young

male dactyl much shorter and palm with weak distal

excavation; female gnathopod 2 like gnathopod 1 but

much stouter, article 5 slightly shorter and lobate and

article 6 slightly longer; pereopods 3-5 simple, distal

pair of locking spines slightly striate and weakly sabreshaped, with strong spine in tandem proximally; pleonal epimera rounded posteroventrally and bearing

lateral ridges; eyes large, clear in alcohol; ommatidia

large; uropod 1 with long sharp peduncular process between rami; lower lip with apicolateral lobules distinctly longer than medial but neither large.

Close to Ampithoe sp. but female (and male) antenna 2 thin, with flagellum about as long as articles

4-5 of peduncle and with other characters mentioned under Ampithoe sp.

Philippines; Oahu, intertidal, on Sargassum, Ulva,

and rocks.
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Aep

FIGURE 13.—Ampithoidac: A, Ampithoe akuolaka J. L. Barnard; H, Cymadusa hawaiensis

(Schellenberg); K, Ampithoe kaneohe J. L. Barnard; o, A. orientate Dana; p, A. poipu J. L.

Barnard; w, A. waialua J. L. Barnard.

Ampithoe (Pleonexes) poipu J. L. Barnard

Ampithoe (Pleonexes) poipu J. L. Barnard, 1970:57-61,

figs. 22,23, 24<z-Jt.

Male and female gnathopod 1 with article 6 much

longer than 5 and rectangular, poorly developed

oblique palm defined by large spine actually on posterior margin of hand, article 5 with weak, narrow,

slightly twisted posterior lobe, dactyl fitting palm;

male gnathopod 2 not strongly enlarged and simply

a stouter version of gnathopod 1; female gnathopod

2 like gnathopod 1; pereopods 4—5 strongly subprehensile, bearing palm slightly weaker than in

A. kaneohe, with similar striate sabre-spine not as

strongly guarding palm as in A. kaneohe and with 3

other erect striate, straight spines; pleonal epimera 1-3

rounded posteroventrally; eyes of medium size, clear

in alcohol; uropod 1 with short truncate peduncular

process between rami; telson with 2 dorsal nobs enlarged into strong reverted hooks from lateral view not

present in other Hawaiian ampithoids, hooks appearing as large posterior papillae from dorsal view.

The large dorsal hooks of the telson cast this species

into the genus Pleonexes Bate but I have reduced that

genus to subgeneric status under Ampithoe and consider that it signals only a grade of evolution and not

a clade.

Kauai, Poipu Beach.

Ampithoe ramondi Audouin

Ampithoe ramondi Audouin.—J. L. Barnard, 1955:28-29

(with references); 1970:47, figs. 18,19.

Male and female gnathopod 1 with article 6 longer

than 5, article 6 narrowly ovate and slightly tapering,

palm very weak and defining spine on posterior margin

of hand, article 5 with broad posterior margin, scarcely

lobate; male gnathopod 2 with strong anterodistal

lobe on article 2, hand slightly elongate and expanded,

in terminal stage anterodistal corner strongly inflated

and expanded, setose, palm deeply sigmoid and defined

by strong tooth or bearing deep slit guarded by very

stout, short thumb; female gnathopod 2 with slightly

s-shaped palm and narrow lobe on article 5; pereopods

4—5 slightly subprehensile, with weak palm bearing 2

striate sabre-spines not guarding palm and bearing

2 or more proximal neighboring spines in tandem;

pleonal epimera rounded posteroventrally; eyes large,

clear in alcohol, red in formaldehyde; uropod 1 lacking

peduncular process between rami.
NUMBER 5 8 37

Pereopods 3-5 with different spinal arrangement, in

juveniles with only 1 of the distal locking spines appearing distinctly sabre-shaped, the other much smaller

and straight.

Circumtropical.

Ampithoe waialua J. L. Barnard

Ampithoe waialua J. L. Barnard, 1970:53-54, figs. 20, 21.

Male gnathopod 1 with article 6 longer than 5 and

rectangular, with moderately well developed, oblique

and excavate palm, article 5 with small lobe distally

sharp and slightly extended, dactyl overlapping palm;

female gnathopod 1 with thinner hand, even palm,

article 5 lobe smaller; male gnathopod 2 small, article

2 with strong anterodistal lobe, hand small, slightly

expanded, palm very oblique and sigmoid, defined by

strong cusp, dactyl not fitting palm; female gnathopod

2 with hand stouter than on gnathopod 1, palm slightly

s-shaped, lobe on article 5 subtriangular; pereopods

3-5 simple, distal pair of locking spines straight, striate,

erect, bearing adjacent proximal spine; pleonal

epimera rounded posteroventrally; eyes large, clear

in alcohol; uropod 1 with obsolescent, leaf-like

peduncular tooth between rami.

Females of this species and A. ramondi are grossly

indistinguishable.

Oahu, intertidal to 30 m.

Ampithoe sp., J. L. Barnard

Ampithoe sp., J. L. Barnard, 1970:54-57, fig. 17 j-POnly female known: gnathopods 1 and 2 with article

6 longer than 5, more so in gnathopod 2 than in

gnathopod 1, article 5 unlobate, article 6 very slender,

palms parachelate and partially hidden by setae;

pereopods 3-5 simple, distal pair of locking spines

slightly striate, slightly sabre-shaped, adjacent to large

proximal spine in tandem; pleonal epimera rounded

posteroventrally and bearing lateral ridges; eyes of

medium size, ommatidia small, with pale purple core

in alcohol; uropod 1 with long sharp peduncular process between rami; lower lip with weak apical lobules

extending almost equally.

Differs from A. orientalis not only in gnathopods

but female antenna 2 very stout and short, flagellum

scarcely longer than article 5; articles 2 and 4 of

pereopods 1-2 especially tumid, though article 2 of

A. orientalis very stout and distally subquadrate; differs

also from other Hawaiian ampithoes in especially enlarged uropod 3.

Oahu, intertidal on Sargassum, Ulva, and rocks.

Genus Cymadusa Savigny

Diagnosis: Accessory flagellum present, ventral

peduncular tooth of uropod 1 long and sharp, gnathopod 2 larger than gnathopod 1, uropod 3 normal;

telson without large hooks.

The two species of this genus endemic to Hawaii are

not yet well defined in all stages as they have been

rarely collected and materials are sparse. By coincidence, they and Paragrubia vorax plus the cosmopolitan Cymadusa filosa have a long peduncular cusp

between the rami of uropod 1 and thus are distinguished from all but one known species of Ampithoe

in Hawaii. But in other regions Ampithoe often has the

same tooth. Members of Cymadusa have a small 1-4

articulate accessory flagellum on antenna 1 and under

preservation many cymadusas retain that antenna, so

that they are easy to distinguish from Ampithoe, which

lacks an accessory flagellum. No member of Hawaiian

Cymadusa has the large rounded anterodistal lobe on

article 2 of the gnathopods seen in Paragrubia. AH

cymadusas have an elongate article 5 on male gnathopod 1 rarely occurring in Hawaiian ampithoes and

the posterior margin of that article is very flat and

often slightly produced or slightly angular at the posterodistal end. Article 4 on gnathopod 1 is triangular

and most of the triangular lobate portion is separated

from, but matches contiguously, the oblique proximal

margin of article 5.

Key to Hawaiian Males of Genus Cymadusa Savigny

Figure Numbers Referenced in Italics

1. Pleonal epimera 1-3 with very small tooth and notch on posteroventral corners (observe

from slide 2 on medium-power compound microscope) (13Hep) C. hmwmensu

Pleonal epimera 1-3 rounded-quadrate posteroventrally, occasionally with extremely weak

indication of cusp (13Oep) 2
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2. Palm of gnathopod 2 semilunar (11C) posterior lobe on article 5 as broad as length of

article 4 C* oceontca

Palm of gnathopod 2 not excavate, often with small hump in middle {11F), posterior lobe

on article 5 narrow, much less than half as broad as length of article 4 C. filosa

Cymadusa filosa Savigny

Cymadusa filosa Savigny.—J. L. Barnard, 1955: 29, fig. 15

(with synonymy); 1970: 61.

Article 5 of male gnathopod 1 longer than article 6,

with weakly produced posterodistal corner, article 6

subovate and strongly palmate; article 5 of gnathopod

2 short, posterior lobe thin and tumid, article 6 of

medium enlargement, palm short, often with small

hump in middle, dactyl very stout and fitting palm.

This species has gone under various names throughout the world and apparently is highly variable; the

Hawaiian population is especially noted for the weak

setation on the gnathopods.

Cosmopolitan in the tropics; in Hawaii generally on

seaweeds in intertidal zone.

Cymadusa hawaiensis (Schellenberg)

Grubia hawaiensis Schellenberg, 1938:88-90, fig. 45.

Cymadusa hawaiensis.—J. L. Barnard, 1970:61, figs. 25, 26.

Articles 5 and 6 of male gnathopod 1 equal to each

other in length, article 6 broadly subrectangular, palm

of medium distinctness; article 5 of gnathopod 2 with

truncate posterior lobe of medium breadth, as broad

as length of article 4, article 6 longer than 5, subrectangular and similar to that of gnathopod 1 but

palm slightly longer, oblique, straight, dactyl fitting

palm. Gnathopods 1-2 are very elongate in comparison to those of other species but juveniles have not been

described adequately.

Oahu, intertidal, on seaweeds.

Cymadusa oceanica J. L. Barnard

Cymadusa oceanica J. L. Barnard, 1955:31, fig. 16.

Articles 5-6 of male gnathopod 1 subequal to each

other in length, with strongly angular posterodistal

corner on article 5, article 6 thin and poorly palmate,

article 5 of gnathopod 2 with truncate posterior lobe

of medium breadth but as broad as length of article 4,

article 6 slightly longer than 5, generally small, palm

short and excavate, dactyl overlapping palm.

Waimanalo, Oahu, depth unknown but presumably

intertidal among algae.

Genus Paragrubia Chevreux

DIAGNOSIS.—Accessory flagellum absent, ventral

peduncular tooth of uropod 1 present, gnathopod 1

larger than gnathopod 2 (but very little in female or

juvenile), uropod 3 with special form slightly distinct

from other arnpithoids in combination of characters;

rami narrow and elongate, hook spines of outer ramus

differ strongly from each other in size and proximal

hook not distinctly reverted toward head of animal,

inner ramus with stout distal spines; telson without

large hooks.

Paragrubia vorax Chevreux

Paragrubia vorax Chevreux, 1901:427-431, figs. 50-55.—

Schellenberg, 1938: 90.—J. L. Barnard, 1955:31-34, fig.

17; 1970:61, fig. 32 h-m.

Apparently Paragrubia is monotypic and distributed

throughout the Indo-Pacific tropics. Gnathopod 1 is

larger than gnathopod 2 in both sexes though scarcely

discernible in the female. Uropod 3 in combination

with gnathopod 1 would provide immediate recognition of this as an ampithoid and not an aorid if uropod

3 were precisely typical of ampithoids but it is not. The

2 hooks on the outer ramus are not as strongly reverted

as in other ampithoids and one of those spines is larger

than the other. The rami appear thinner than in other

ampithoids. The resemblance to Isaeidae (= Photidae) and Aoridae is confusing in undissected individuals but once uropod 3 is dissected and observed

from flattened dorsal view the characteristic apical

FIGURE 14.—Anamixidae and Leucothoidae: A, Anamixis stebbingi Walker; H, Leucothoe hyhelia J. L. Barnard; L, L. lihue J. L. Barnard; p, Leucothoides pottsi Shoemaker; T, Leucothoe

tridens Stebbing.
NUMBER 5 8
40

spines are revealed from among various setae and the

cylindrical or planocylindrical character of the peduncle and rami is also more readily seen.

Male gnathopod 1 in terminal stages is very bulky,

articles 5 and 6 very broad, the dactyl almost semilunar

and overlapping a scarcely oblique, slightly concave

palm (10V2). The smaller male gnathopod 2 palm is

also concave.

Eyes clear or pale purple in alcohol.

Presumably throughout Hawaii among shallowwater algae.

Family ANAMIXIDAE

Genus Anamixis Stebbing

Anamixis stebbingi Walker

FIGURES 14A, 15S (Except Sgl)

Anamixis stebbingi Walker, 1904:259-261, pi. 3, fig. 18.—

J. L. Barnard, 1965:488-489, fig. 4; 1970:61-63, fig.

27.

Anamixids have the leucothoid morphology with a

carpochelate gnathopod 1 (14Agl), and large

gnathopod 2 also seeming to be carpochelate but with

an inflated article 6 (hand). Anamixids have a small

coxa 1 like some leucothoids but differ in the loss or

extreme reduction of mandibles and maxillae, a ventral

cephalic keel prominently replacing these mouthparts.

The keel may be seen by focusing ventromedially on

the head (slide 2). Some anamixids (Paranamixis) lose

gnathopod 1 completely. Gnathopod 2 of Hawaii's

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

anamixid is very characteristic (15Sg2) and distinguishes it from any leucothoid. See the latter family

(page 102) for color notes.

Presumably anamixids pierce and suck tissues of

sessile invertebrates, possibly sponges, coelenterates, or

ascidians.

Ceylon to Hawaii. Oahu, intertidal to 30 m,

presumably piercing tissues of sessile invertebrates.

Family AORIDAE

FIGURES 16-21

Aorids belong with the greater isaeid (= photid) complex in having a fleshy telson and glandular pereopods

1-2. Aoridae are probably broadly polyphyletic descendents of various isaeids in which a reversal in

dominance of gnathopods has occurred, gnathopod 1

becoming large and complex in males and simply

enlarged in females.

Like other isaeids, aorids build tubes attached to

algae and hard substrates or they apparently line

existing holes with their parchment-like material spun

from glands in pereopods 1-2.

Species of this family often shed antennae and

pereopods 3-5 on preservation. They may be distinguished from the commonly occuring ampithoid

Paragrubia in the absence of distinct hooks on the

stout outer ramus of uropod 3. The uropod should be

mounted and observed from dorsal view to make this

distinction.

Key to Genera of Aoridae

Figure Numbers Referenced in Italics

1. Antenna 1 lacking accessory flagellum, male gnathopod 1 with article 4 produced strongly

into long free acute tooth (merochelate) (18Cgl) Aoroides Walker

Antenna 1 bearing accessory flagellum, male gnathopod 1 lacking long free tooth on article 4.. 2

2. Article 6 of male gnathopod 1 narrower and/or shorter than article 5, latter with strong or

rudimentary posterodistal tooth (18Kgl, Mgl) 3

Article 6 of male gnathopod 1 as broad as or broader than article 5, article 6 longer than

article 5 and strongly palmate, latter without teeth (18Agl, Pgl) 4

3. Uropod 3 biramous, head deeply recessed ventrally for attachment of antenna 2 (17Kh).

Konatopus J. L. Barnard

Uropod 3 uniramous, head weakly recessed ventrally for attachment of antenna 2 (17Nh).

Neomicrodeutopus(?) Schellenberg

4. Antennae short and extending equally, article 3 of antenna 1 as long as article 1, inner plate

of maxilla 1 with numerous medial setae, peduncle of uropod 2 with vestigial but sharp

tooth between rami (20Qu2) Aloiloi J. L. Barnard

Antenna 1 much longer than antenna 2, article 3 much shorter than article 1, inner plate

of maxilla 1 bearing only 1 long terminal seta, peduncle of uropod 2 with long sharp tooth

between rami (21Bur) Lembos Bate
NUMBER 5 8 41

Females of Aoridae

1. Uropod 3 uniramous Neomicrodeutopus (?) Schellenberg

Uropod 3 biramous 2

2. Inner plate of maxilla 1 with many medial setae 3

Inner plate of maxilla 1 with only terminal seta(e) 4

3. Head deeply recessed for attachment of antenna 2, eyes caramel-brown or reddish brown in

alcohol (!7Kh) Konatopus J. L. Barnard

Head weakly recessed for attachment of antenna 2, eyes black or purplish brown in alcohol

{17Ah) Aloiloi J. L. Barnard

4. Accessory flagellum absent Aoroides Walker

Accessory flagellum present Lembos Bate

Genus Aloiloi J. L. Barnard

Aloiloi nenue J. L. Barnard

Aloiloi nenue J. L. Barnard, 1970:63-68, figs. 28-30.

This monotypic Hawaiian genus has a gross resemblance to various members of Lembos. Gnathopod 1

appears very similar to that of Lembos but closer examination demonstrates many characters in common

with Gammaropsis pali in the Isaeidae. Gnathopod 2

is larger than normal for Hawaiian members of Lembos

and resembles that of various members of Gammaropsis. Antenna 1 is short like antenna 2 and has an

elongate article 3 like that of Gammaropsis and unlike

that of Lembos. The inner plate of maxilla 1 is unlike

other Hawaiian aorids in the presence of medial setae

on the inner plate, and it thus resembles several species

of Gammaropsis. Uropod 2 has only a vestigial tooth

between the rami, unlike various members of Lembos

in Hawaii, and it thus resembles most species of

Gammaropsis.

The weakly excavate palm of gnathopod 1 bearing

a minutely ridged defining corner and the giant palmar

spine of gnathopod 2 are uncharacteristic of Lembos.

Uropod 3 has rami equal to each other in length, not

longer than the peduncle, the outer with barrel-shaped

article 2. Pleonal epimera 1 and 3 bulge posteriorly

and have a small or medium posteroventral cusp and

notch, epimeron 2 with these characters weak.

Article 2 of pereopods 3-5 is narrowly rectangular

like those of Gammaropsis pali. The head is of moderate recessment for the attachment of antenna 2 and

the eyes are irregularly ovate, garnet red in formaldehyde, pale purple in alcohol. Article 3 of antenna 1

and the distal ends of the flagella of both antennae are

bright pink in formaldehyde. The stout mandibular

palp strongly dominates the remainder of the mandible.

Oahu, Kaneohe Bay, 3-4 m. Rare.

Key to Hawaiian Species of Genus Aoroides Walker

Eyes black or deep brownish purple in formaldehyde and alcohol, uropod 1 with, but uropod 2

lacking long ventral cusp between rami A. nahili

Eyes orange or clear in formaldehyde and alcohol, uropod 2 with long acute cusp between rami

on both uropods 1-2 A. Columbia*

Aoroides columbiae Walker

Aoroides columbiae Walker, 1898:285, pi. 16, figs. 7-10.—

Thorsteinson, 1941:83-84, pi. 6, figs. 65-66.—J. L. Barnard, 1954b:24-26, pi. 22; 1970:68-70, figs. 31, 32 a-g.

Aoroides californica Alderman, 1936: 63-66, figs. 33-38.

Male gnathopod 1 has an elongate sharp cusp on article 4 like that of the genus Aora (IBCgl), but antenna

1 lacks all but a vestige of an accessory flagellum. Article 6 is elongate and rectangular and bears a large

dactyl. Females have gnathopods of median stoutness

for amphipods and both sexes differ from species of

Lembos and Aoroides nahili by the orange or pale, not

black, eyes. The orange-eyed Konatopus has a deep

ventral recessment on the head for the attachment of

antenna 2.

Pacific America, warm and cold temperate, to

Hawaii. Oahu, 3-30 m, rarely and questionably

intertidal.

Aoroides nahili J. L. Barnard

Aoroides nahili J. L. Barnard, 1970:70, fig. 33.

Gnathopods 1-2 of the female and gnathopod 2 of the

male are conspicuously thinner in this species than in
Sgl

FIGURE 15.—Anamixidae and Lcucothoidae: H, Leucothoe hyhelia J. L. Barnard; L, L. lihue

J. L. Barnard; p (and Sgl), Leucoth.oid.es pottsi Shoemaker; s (except Sgl), Anamixis stebbingi

Walker; T, Leucothoe tridens Stebbing.

Hg2
FIGURE 16.—Aoridae (Isaeidae): A, Aloiloi nenue J. L. Barnard; c, Aoroides columbine Walker;

l, Lembos intermedius Schellenberg; K, L. kamanu J. L. Barnard; L, L. leapakahi J. L. Barnard;

M, L. macromanus (Shoemaker); N, Neomierodeutopus (?) makena J. L. Barnard, w, Lembos

waipio J. L. Barnard.
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 17.—Aoridac (Isaeidae and Corophiidae): A, Aloiloi nenue J. L. Barnard; B, Aoroides

nahili J. L. Barnard; c, Aoroides columbiae Walker; E, Ericthonius brasiliensis Dana; K,

Konatopus paao J. L. Barnard; L, Letnbos leapakahi J. L. Barnard; M, Lembos kamanu J. L.

Barnard; N, Neomicrodeutopus (? ) makena J.L. Barnard; o, Corophium acherusicum Costa

(See also Figure 26); p, Lembos pualani J. L. Barnard; Q, L. aequimanus Schellenberg.
FIGURE 18.—Aoridae (Isaeidae), gnathopod 1: A, Aloiloi nenue J. L. Barnard; c, Aoroides

columbine Walker; D, Neomicrodeutopus makena J. L. Barnard; i, Lembos intermedius Schellenberg; K, Konotopus paao J. L. Barnard; L, Lembos leapakahi J. L. Barnard; H, L. macromanus

(Shoemaker); N, Aoroides nahili J. L. Barnard; p, L. pualani J. L. Barnard; Q, L. aequimanus

Schellenberg; s, Lembos sp.; w, L. waipio J. L. Barnard.

383-X97 O - 71 - 4
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Aoridae (Isaeidae) gnathopod 2: A, Aloiloi nenue J. L. Barnard (also mandible);

c, Aoroides columbine Walker; K, Konatopus paao J. L. Barnard; L, Lembos leapakahi J. L.

Barnard; M, L. macromanus (Shoemaker); N, Neomicrodeutopus (?) makena J. L. Barnard

(also mandible); P, Lembos pualani J. L. Barnard; z, Aoroides nahili J. L. Barnard.
FIGURE 20.—Aoridae (Isaeidae): A, Aloiloi nenue J. L. Barnard; B, Lembos kamanu J. L.

Barnard; c, Aoroides columbiae Walker; i, Lemos intermedius Schellenberg; K, Konatopus paao

J. L. Barnard; L, Lembos leapakahi J. L. Barnard; M, L. macromanus (Shoemaker); N, Neomicrodeutopus (?) makena J. L. Barnard; Q, Lembos aequimanus Schellenberg; z, Aoroides

nahilij. L. Barnard.
FIGURE 21.—Aoridae (Isaeidae): A, Aloiloi nenue J. L. Barnard; B, Lembos kamanu J. L.

Barnard; c, Aoroides columbiae Walker; i, Lembos intermedium Schellenberg; K, Konatopus paao

J. L. Barnard; L, Lembos leapakahi J. L. Barnard; N, Neomicrodeutopus (?) makena J. L.

Barnard; p, Lembos pualani J. L. Barnard; w, L. waipio J. L. Barnard.
NUMBER 5 8 49

A. columbiae but both species have to be observed together to note the differences until experience is

gained; otherwise the small gnathopods have to be dissected, mounted flat, and compared with Figures

19Cg2 and Zg2. Male gnathopod 1 differs slightly

from that on A. columbiae in the turgid proximoanterior margin of article 5. The black eyes are combined

with brownish purple pigment occurring in most specimens on thoracic segment 5.

Oahu, 0-4 m, generally phycophilous.

Genus Konatopus J. L. Barnard

Konatopus paao J. L. Barnard

Konatopus paao J. L. Barnard, 1970:70-72, figs. 34, 35.

Male gnathopod 1 is like that of Microdeutopus Costa

(see Sars, 1895, plate 192) but the distal tooth of article 5 is rudimentary and article 6, though shorter and

more slender than article 5, has a minute palm. The

head has a deep recessment for the attachment of antenna 2 and the lateral cephalic lobes are much longer

than in other Hawaiian aorids. Article 3 of antenna 1

peduncle is as long as article 1 and the inner plate of

maxilla 1 has medial setae; it thus resembles Aloiloi

alone among other Hawaiian aorids in these characters. The rami of uropod 3 are only slightly longer than

the peduncle and the outer has a barrel-shaped article

2; uropod 2 lacks a peduncular process between the

rami. Pleonal epimera 1-3 have the bulging posterior

margins and small posteroventral tooth and notch.

Article 2 of pereopods 3-5 is slender. Female gnathopods 1-2 are scarcely different in size and females thus

might be classified as Isaeidae.

The ochraceous, caramel or reddish brown eyes are

unique in the Aoridae except for the weak orange eyes

of Aoroides columbiae. Other aorids have black or

purple-brown in the eyes, even in alcohol. Eyes should

be examined under medium-power compound microscopy as they may appear black from a distance under

low-power stereoscopy.

Male pereonite 1 and coxa 1 become extremely

elongate anteroposteriorly in terminal stages.

Oahu. intertidal to 30 m.

Genus Lembos Bate

Members of Lembos are numerous in Hawaii and

often difficult to distinguish from each other in female

and juvenile stages. A sample with 3 or 4 species may

seem impossible to partition into taxa but if one commences looking for L. leapakahi, the most easy to distinguish by its pure white color in alcohol and large

smooth gnathopods, next L. intermedius by the color

pattern of head, pereonite 1 and pereonite 6, followed

by L. waipio with distinct pale brown color on the

dorsal sides of pereonites 2-6, one is then faced only

with the problem of distinguishing L. macromanus

from L. pualani by observing the stairstepping of the

urosomal segments dorsally and the dorsally short

urosomite 2 in L. macromanus. L. pualani has dull

eyes with distinct ommatidia separated from each other

by clear space. L. kamanu and other taxa are distinguished from L. pualani by small differences and are

known only in the female condition as yet.

Males are very easy to distinguish in their terminal

stages according to teeth on the belly of the thorax.

Sternal teeth must be observed from direct lateral view

after coxae 1-5 have been dissected off both sides of

the carcass (see Figure 16Lv, etc.). Article 2 on the

outer ramus of uropod 3 is missing in one species only,

but occasionally an aberrant specimen of other species

may lack this article on a regenerating ramus. Collections older than 2 months and preserved in alcohol are

not reliable for color differentiation.

All terminal males except those of L. leapakahi have

a deep notch on the palm of gnathopod 1. Lembos

macromanus males have especially dense setae projecting anteriorly from article 5 of gnathopod 2 and L.

pualani has a very dense brush of setae projecting distomedially from article 6 of gnathopod 1. Coxa 1 is

highly modified in L. pualani.

Female gnathopod 1 is larger than 2 but relatively

plain, with a simple palm defined by a spine; in L.

leapakahi it resembles that of the male and is thus easy

to distinguish but only in L. macromanus of the other

Hawaiian species is there a slightly increased tumidity

of articles 5-6. Female gnathopod 2 does not always

match that of the male precisely in setation or lobation

on article 2.

Numerous other characters occur on members of

Lembos but they have not been studied adequately in

relation to various instars of the species. For example,

the following offer promise as characters of qualitative

value to distinguish all stages of each species instead of

the present necessity to rely on the artifact of color

preservation in alcohol: (1) The extent of distal setation on the outer ramus of uropod 3; (2) the number
5 0 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of setae and condition of ornaments on the telson; precise structure of shark-tooth flake on left mandibu-

(3) mixture of spines and setae on article 2 of pereo- lar molar.

pods 3-5; (4) relative length to antenna 1 and stout- Eyes of this genus are brown, from bright coffee to

ness of antenna 2 peduncle; (5) ocular structure; (6) dull deep brown, except for L. kamanu.

Key to Hawaiian Species of Genus Lembos Bate

Figure Numbers Referenced in Italics

1. Gnathopod 1 of both sexes with large subcircular article 6, palm convexly transverse, lacking

deep notch, article 6 not significantly longer than broad article 5 (18Lgl), pleonal epimera

2-3 rounded posteroventrally, gnathopod 2 especially stout in both sexes, body usually

pure white and lacking any pigmentary spots in alcohol L. leapakahi

Gnathopod 1 of male larger and more complex than that of female, article 6 subrectangular

(18Igl), palm transverse or oblique but in male bearing deep notch with denning tooth,

article 6 significantly longer than article 5, all pleonal epimera with small posteroventral

notch and tooth, gnathopod 2 not especially stout, body with small to large amount of

pigment in alcohol 2

2. Uropod 3 lacking article 2 on outer ramus, thoracic sternal teeth on male bifid (16Iv), body

pigment in alcohol forming broad dark stain on posterior half of head, anterior half of

pereonite 1 and posterior half of pereonite 6 L. intermedius

Uropod 3 with small barrel-shaped article 2 on outer ramus, thoracic sternal teeth of male

simple if present (16Mv), body pigment composed either of spots or blotches more widely

scattered than just on head, pereonite 1 and pereonite 6 3

3. Dorsal margin of urosomite 2 not especially short, male gnathopod 1 relatively narrow, article 5

long, article 6 strongly setose medially (18Pgl), male thoracic sternites lacking teeth, each

ocular ommatidium distinctly visible L. pualani

(Proceed also to subkey, below)

Dorsal margin of urosomite 2 especially short and full dorsal margin of urosome conspicuously

stairstepped, male gnathopod 1 stout, article 5 short, article 6 poorly setose medially

(18Mgl), male thoracic sternites 2, 3 or 4 with teeth, ommatidia not clearly distinct 4

4. Body background buff in alcohol, covered with dark spots, eyes bright coffee color, male sternal

teeth on thoracic segments 2-3 only (16Mv) L. macromanus

Body background pure white in alcohol, with pale brown dorsal stripes or blotches on

pereonites 2-6 or 3-6, eyes medium-dull brown, male sternal teeth on thoracic segments

2-4 (16Wv) L. waipio

Some Females of Lembos from Couplet 3

1. Gnathopod 1 normal for female of Lembos, palm evenly convex L. pualani

Gnathopod 1 with slightly scalloped palm (uniquely also with shark-tooth on left mandibular

molar, and strongly depressed urosomite 3 (20Bgl) L. kamanu

Gnathopod 1 with sinuous or excavate palm 2

2. Lateral lobe of head narrow, rami of uropod 3 elongate, peduncular tooth of uropod 2 short.

L. aequimanus

Lateral lobe of head normally broad, rami of uropod 3 short, peduncular tooth of uropod 2 long.

Lembos sp.

Lembos ? aequimanus Schellenberg a narrow gnathopod 1 with excavate palm bearing a

?Lembos {Bemlos) aequimanus Schellenberg, 1938:76-77, weak distal protrusion; the urosome is normal. Rami of

fig. 39. ' uropod 3 are elongate. Lateral cephalic lobe narrow

Not Lembos {Bemlos) aequimanus.—}.!,. Barnard, 1955:34. and mammilliform, anteroventral corner of head

Lembos ?aequimanus.—]. L. Barnard, 1970:72-76, fig. 36 quadrate.

a—e.

Peduncular tooth of uropod 1 shorter than in other

Only the female stage of this species is known; it re- species of Lembos and tooth of uropod 2 uniquely

sembles the female of L. pualani in many ways but has obsolescent.
NUMBER 5 8 51

Body ground color slightly brownish, with scattered

darker spots.

Gilbert; Fiji; Oahu, intertidal, Sargassum.

Lembos intermedius Schellenberg

Lembos intermedius Schellenberg, 1938:77-78, fig. 40.—

J. L. Barnard, 1955:34; 1970:76-77, fig. 37.

Male coxa 1 with quadrate anteroventral corner; article 2 of gnathopod 2 with strong, thin mammilliform

anterodistal lobe, article 5 less than half as long as

article 6, latter expanded, anteriorly bulging, moderately setose medially, palm oblique, defining process

thus short, dactyl scarcely overlapping palm; article 2

of gnathopod 2 with large lobe similar to that on gnathopod 1, palm nearly transverse, setation dense medically; thoracic sternites 2-5 each with low broad bifid

tooth, sternite 1 with obtuse simple tooth, 6 with tiny

cusp; uropod 3 lacking article 2 on outer ramus; female gnathopod 1 normal, article 6 longer than 5, article 2 of gnathopod 2 with anterodistal corner sharply

quadrate.

Posterior portion of head, anterior half of pereonite

1, and posterior half of pereonite 6 with dark brownishpurple stain.

Oahu, 0-4 m.

Lembos kamanu J. L. Barnard

Lembos kamanu J. L. Barnard, 1970:77-79, fig. 38.

Only the female stage of this species is known; it resembles the female of L. pualani but has a weakly scalloped palm on gnathopod 1, has a strongly depressed

urosomite 3 that drops strongly from the dorsal line of

urosomite 2; but the latter is elongate like that of L.

pualani. Uropod 3 has a short peduncle with very long

rami in comparison to those of several species of

Lembos but not extraordinarily different from that

of L. pualani; the anteroventral cephalic tooth

is thick. The main spines on the inner plate of

the maxilliped are extremely broad and flabellate;

they are difficult to see because of their transparency

and concealment by various other setae.

Body very white in alcohol, bearing scattered pigment spots, eyes pale purple in alcohol, central cores

surrounded by one layer of large ommatidia.

Oahu, intertidal.

Lembos leapakahi J. L. Barnard

Lembos leapakahi J. L. Barnard, 1970:79-81, figs. 39, 40.

This beautiful amphipod has a large smooth gnathopod 1 in both sexes and juveniles; article 6 appears

almost circular, and article 5 is nearly as long as 6.

Gnathopod 2 is slightly larger than in any other Hawaiian member of Lembos. Males have simple teeth on

pereonal sternites 2-5.

Body pure white in alcohol, eyes ochraceous brown

and bright.

Oahu, 0-30 m.

Lembos macromanus (Shoemaker)

Bemlos macromanus Shoemaker, 1925:36-41, figs. 10-13.

Lembos macromanus.—Schellenberg, 1938:79.—J. L. Barnard, 1962:9, fig. 3; 1970:82, figs. 41, 42 a-m, o.

Male coxa 1 quadrate, with subrounded anteroventral

corner; article 2 of gnathopod 1 with sharp downward

pointing submammilliform lobe, article 5 about half or

much less as long as article 6, latter very broad in terminal male, rectangular, weakly setose medially, palm

transverse, main process slightly rounded apically and

marked basally with suture in terminal stage, dactyl

strongly overlapping palm; article 2 of gnathopod 2

with quadrate or weakly produced anterodistal corner,

palm nearly transverse, setation dense medially and

on anterior margin of article 5; thoracic sternites with

weak tooth on segments 2-3; female gnathopod 1

stouter than normal, article 5 shorter than 6, gnathopod 2 lacking anterodistal lobe on article 2 and article 5

poorly setose in contrast to male. Pereopods 4-5 with

especially dense plumose setae on the posterior margin

of article 2.

Body buff in alcohol, with numerous dark spots and

blotches, eye brightly coffee brown throughout, ommatidia indistinct.

Pacific America; Philippines; Oahu, 0-30 m.

Lembos pualani J. L. Barnard

Lembos (Bemlos) aequimanus.—J. L. Barnard, 1955:34 (not

Schellenberg).

Lembos pualani J. L. Barnard, 1970:82-85, figs. 42 n, 43.

Male coxa 1 quadrate, anterior and ventral margins

deeply concave, anteroventral corner sharply attenuate ; article 2 of gnathopod 1 with medium-size mammilliform anterodistal lobe, article 5 about 70 percent

as long as article 6, latter rectangular, strongly setose
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

medially, dactyl strongly overlapping palm; article 2

of gnathopod 2 with long, thin, mammilliform lobe,

articles 5 and 6 equal to each other in length, palm

nearly transverse, setation dense medially; thoracic

sternites lacking teeth; female gnathopod 1 normal,

article 6 longer than 5, gnathopod 2 lacking anterodistal lobe on article 2.

Body very white in alcohol, bearing scattered pigment spots, eyes dull purplish brown and each

ommatidium distinct from another.

Oahu, 0-4 m.

Lembos waipio J. L. Barnard

Lembos waipio J. L. Barnard, 1970:85-87, figs. 44, 45.

Male coxa 1 with rounded anteroventral corner; article 2 of gnathopod 1 with anterodistal process scarcely

evident, article 5 half as long as 6, latter slightly expanded, subrectangular, tapering distally, poorly setose, defining process with stout posterior spine, dactyl

strongly overlapping palm; article 2 of gnathopod 2

with weak anterodistal cusp, article 5 longer than 6,

palm oblique, setation on gnathopod 2 of moderate

extent; thoracic sternites 2-4 with one spine tooth each;

female gnathopod 1 normal, article 5 subequal to 6

in length, gnathopod 2 lacking anterodistal lobe on

article 2.

Body very white in alcohol, with gross pale purplebrown dorsal stripes or blotches on pereonites 2-6 or

3-6; eyes intermediate between dull and bright, ommatidia indistinct.

Oahu, 0-30 m.

Lembos sp., J. L. Barnard

Lembos sp., J. L. Barnard, 1970:87-90, fig. 36/-*.

Only the female stage of this species is known; it resembles the female of L. pualani in many ways but the

normal-sized gnathopod 1 has a slightly sinuous palm

with a posterior excavation marking a weak defining

cusp; the urosome and lateral cephalic lobe are normal. Peduncular teeth of uropods 1-2 normally long.

Body ground color white with scattered darker spots.

Oahu, intertidal.

Genus Neomicrodeutopus (?) Schellenberg

Neomicrodeutopus (?) makena J. L. Barnard

Neomicrodeutopus (?) makena J. L. Barnard, 1970:90-92,

figs. 19/, 46,47.

The uniramous uropod 3 is diagnostic and male

gnathopod 1 has the appearance of that on various

boreal species of Microdeutopus, a stout but long article

5 bearing one long and one short posterodistal tooth

guarding the stout, almost simple article 6 with medium sized dactyl; gnathopod 2 is small, slender, palm

nearly transverse; pleonal epimera 1—3 are fairly unusual in their posteriorly truncate appearance, each

with small posteroventral bevel-notch bearing a stout

seta; ventral margins of those epimera naked.

Female gnathopod 1 generally like that of female

Lembos but distinguishable from Hawaiian species of

that genus in the combination of uropod 3, pleonal

epimera and non-extended anteroventral corner of

head.

Eyes maintaining dark pigment core in irregularly

circular ommatidial group.

Oahu, Makapuu Point, on Zoanthus.

Family BIANCOLINIDAE

Genus Biancolina Delia Valle

Biancolina mauihina J. L. Barnard

FIGURE 22

Biancolina mauihina J. L. Barnard, 1970:103, figs. 57, 58.

Body subcylindrical, urosome depressed, coxae short

but family differing from Corophiidae in the fully biramous, though small uropod 3 and from the Colomastigidae in the presence of long flexible antennae

curling backwards from a flattened head resembling

that of some isopods and insects, uropod 3 short, rami

stubby and bearing one or 2 apical spines and ventrolateral margins of uropods 1-2 bearing long plumose

setae like those of Parajassa angularis; differing from

latter in shorter uropod 3 with short peduncle, flexible

antennae, small almost parachelate gnathopods and

prehensile pereopods 3-5. The telson of Biancolina is

very short and in this species minutely notched. Uropods 1 and 2 are diverse: in some specimens the outer

rami are simple, stubby, nearly half as long as inner

rami; whereas in other specimens the outer ramus of

uropod 2 has a long stout spine reaching to the apex

of the inner ramus, and on uropod 1 the outer ramus

is a grotesquely armed piece nearly as long as the inner

ramus. These differences have not been resolved as to
NUMBER 5 8 53

FIGURE 22.—Biancolinidae: Biancolina mauihina J. L. Barnard.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

whether they denote different species or different sexes

of the same species. Eyes red in formaldehyde.

Probably bores into stems of algae. Oahu, Kawela

Bay, intertidal.

Family CHELURIDAE

Genus Tropichelura J. L. Barnard

Tropichelura insulae (Caiman)

FIGURE 23

Chelura insulae Caiman, 1910:182, pi. 5, figs. 1-6.—J. L.

Barnard, 1955:39.

Tropichelura insulae.—J. L. Barnard, 1959: 6, figs. 2, 4C,

D, E.

Uropod 2 with its broadly lamellar, setose peduncle

and minute rami is an unmistakable character of this

species. Uropod 3 is very elongate and the inner ramus

is absent. The urosome is elongate and the segments

coalesced together to form a shallow box. Gnathopod

1 is enormously enlarged, like gnathopod 2 of a Maera,

whereas gnathopod 2 is very small.

Eyes dull ochraceous to amber in alcohol.

Bores or rasps wood, occurring mainly in tunnels already created by limnoriid isopods; Honolulu Harbor,

pilings and floating docks.

Circumtropical.

Family COLOMASTIGIDAE

Genus Colomastix Grube

FIGURES 24, 25

Colomastigids resemble some corophiids in the cylindrical body plan and very short coxae; but urosomites

2-3 are distinctly coalesced, whereas, in corophiids

they are either free or coalesced with urosomite 1; and

colomastigids have a distinctly biramous uropod 3 and

large telson. The mandible lacks palp and incisor and

bears large terminal spines curved medially (24 Lin).

The maxillipeds are thin like those of stenothoids and

leucothoids and have either free or coalesced inner

lobes, broad but short outer lobes and a thin palp;

the basal articles may form a large shell-like hood (seen

FIGURE 23.—Cheluridae: Tropichelura insulae (Caiman).
NUMBER 5 8 55

from lateral view before dissection). Gnathopod 1 is

very thin and usually bears a distal brush of setae with

dactyl concealed or unidentifiable, but the male of

C. pusilla has a shortened gnathopod 1 with stout,

short dactyl. Male gnathopod 2 becomes enlarged and

distinctly subchelate, often grotesquely. The antennal

flagella are short.

Colomastigids in Hawaii are very small and the

obscure mouthparts are very difficult to extirpate.

They can be checked in situ by rolling the head ventral

side up so that the spiny mandibles, protruding

epistome, and midcephalic keel can be seen, along with

vague outlines of the maxillae; the maxillipeds should

be removed first.

Key to Hawaiian Species of Genus Colomastix Grube

Figure Numbers Referenced in Italics

1. Outer ramus of third uropod half or less as long as inner ramus (25Ltu3) C. lunalilo

Outer ramus of third uropod 75-100 percent as long as inner ramus 2

2. Uropod 1 extending the same distance as uropod 2 (coxae 1-4 broadly rounded anteriorly,

outer rami of uropods 1-3 varying from 66 to 75 percent as long as inner rami, inner plates

of maxillipeds freely split) C. kapiolani

Uropod 1 short, extending only as far as middle of rami of uropod 2 3

3. Each pair of rami on uropods 1-3 equal in length (coxae 1-4 narrowed and attenuate but

not pointed anteriorly, inner plates of maxilliped coalesced) C. pusilla

Outer ramus of uropod 3 slightly shorter than inner ramus (other characters unknown).

Colomastix sp.

Colomastix kapiolani J. L. Barnard

Colomastix kapiolani J. L. Barnard, 1970:96, fig. 50.

Terminal male gnathopod 2 has a characteristic shape

(25Kg2).

Oahu, 2-30 m, especially on Pocillopora bottoms.

Colomastix lunalilo J. L. Barnard

Colomastix lunalilo J. L. Barnard, 1970:96-100, figs. 51, 52.

The eyes are brown in formaldehyde and remain dark

in alcohol; all other species of the genus so far collected in Hawaii have red eyes in formaldehyde that

become rusty-pink or clear in alcohol. The anterior

coxae are not attenuate anteriorly but have small

nipple-like points; the inner plates of the maxilliped

are coalesced like those of C. pusilla.

Oahu, Kaneohe Bay, 3-4 m, on bottom of corals,

algae, and rubble.

Colomastix pusilla Grube

Colomastix pusilla Grube.—J. L. Barnard, 1955:39-42, fig.

20 (with references).

Male gnathopod 1 is shortened and has a short stout

dactyl unlike females of C. pusilla and both sexes of the

other species of the genus that all have a distal setal

brush on gnathopod 1.

Possibly cosmopolitan in tropical and temperate

seas. Oahu, Kaneohe Bay, depth and biotopes

unknown.

Colomastix sp., J. L. Barnard

Colomastix sp., J. L. Barnard, 1970:100.

A single juvenile of this species has been collected; it

may represent a stage of C. pusilla but gnathopod 1

has only a single long hooked seta (Fdactyl) and is

possibly a stage in the transformation between the

juvenile-female state to that of the male; however, the

outer ramus of uropod 3 is slightly shortened, unlike

C. pusilla. Coxae and maxillipeds have not been

studied.

S. Oahu, 30 m, bottom dominated by coralline

"footballs."

Family COROPHIIDAE

FIGURES 1 7E, O, 26

Corophiids belong to the isaeid-complex, with fleshy

telson and grandular pereopods 1-2 but corophiids

have a strongly flattened urosome; they differ from

podocerids in the relatively short urosomite 1; uropod

3 is uniramous in Hawaiian corophiids. Keys to

Isaeidae (page 88), Ischyroceridae (page 97), and

Aoridae (page 40) should also be examined for species

superficially similar to Corophiidae.
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Kxp

FIGURE 24.—Colomastigidae: K, Colomastix kapiolani J. L. Barnard; L, C. lunalilo J. L. Barnard;

p, C. pusilla Grube.

FIGURE 25.—Colomastigidae: K, Colomastix kapiolani J. L. Barnard; L, C. lunalilo J. L. Barnard;

p, C. pusilla Grube.
NUMBER 5 8 57

Lgl Lg2
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Corophiidae: A, Corophium acherusicum Costa (see also Figure 17); B, C. Pbaconi

Shoemaker; i, C. insidiosum Crawford.
NUMBER 5 8 59

Key to Genera of Corophiidae

Figure Numbers Referenced in Italics

Article 3 of antenna 1 shorter than article 1, antenna 2 very stout in male (261) and article 4

with tooth in male or strong spines in female; mandibular palp 2-articulate (26Bmp), gnathopods small but gnathopod 2 grossly setose; uropod 3 flat (26Bur), eyes dark.

Corophium Latreille

Article 3 of antenna 1 longer than article 1, antenna 2 slender in male (17 E), article 4 lacking

tooth or strong spines (but setose); mandibular palp 3-articulate; gnathopod 1 small, gnathopod

2 in male very large, carpochelate (17E); uropod 3 subcylindrical and with special form of

Figure 4Rw, eyes red or clear Ericthoniw Milne-Edwards

Key to Hawaiian Species of Genus Corophium Latreille

Figure Numbers Referenced in Italics

1. Urosome boat- or saucer-shaped, segments all coalesced, forming a lateral upturned rim with

a slight notch on each side, uropods 1 and 2 attached ventrally (26Bur) C. ?baconi

Urosomites separate (or if coalesced not forming a saucer), marked with lateral stairsteps or

notches for insertion of uropods 1-2 (26Iur) 2

2. Antenna 2 with large distal tooth on article 4 (26Ia2, left) . . . males 3

Antenna 2 lacking tooth, article 4 bearing large spines ventrally (26Ba2) . . . females 4

Antenna 2 with both a tooth and spines on article 4 (26Ia2, right).

aberrant kinds of C. insidiosum

3. Rostrum large from dorsal view (26Bhd) C. insidiosum

Rostrum obsolete, head deeply invaginated from dorsal view (26Ahd) C. acherusicum

4. Spines of article 4 on antenna 2 occurring in 2 pairs and a single terminal C. insidiosum

Spines of article 2 on antenna 2 occurring in 3 pairs and a single terminal C. acherusicum

Corophium acherusicum Costa

Corophium acherusicum Costa.—Crawford, 1937:617-620,

fig. 2.—Shoemaker, 1947:53, figs. 2, 3; 1949:76.—J. L.

Barnard, 1955:37.

Microscopic examination of the large tooth on male

antenna 2 demonstrates that it is really two teeth, the

second very small; the same is true of C. insidiosum

males. The dactyl of gnathopod 2 has two accessory

teeth. The female head is normally rostrate like males

of C. baconi. Eyes black-brown like other species of

Corophium herein.

This species, like C. insidiosum, is cosmopolitan in

harbors; it builds tubes on pilings, boats and docks.

Oahu, Waikiki, on bottom of small boat.

Corophium ? baconi Shoemaker

?Corophium baconi Shoemaker, 1934:356-359, fig. I;

PShoemaker, 1949:82, figs. 5, g, h.—J. L. Barnard,

1970:100-101, fig. 53.

Male and female rostra are normal. Male antenna 2

is generally like that of C. acherusicum and normal

C. insidiosum. The female lacks spines on article 5

of antenna 2 unlike females of C. acherusicum and C.

insidiosum. The urosome (26 Bur) is characteristic;

this is the only species from the Pacific Ocean as yet

known to have this kind of urosome. The dactyl of

gnathopod 2 has only 1 accessory tooth.

Eastern and Central Pacific in the quadrangle from

Bering Sea to Hawaii to California to Peru. Generally

a species of the open seas, in contrast to its Hawaiian

congeners. Oahu, Kaneohe Bay, Hawaii Marine

Laboratory dock area.

Corophium insidiosum Crawford

Corophium insidiosum Crawford, 1937: 615, fig. 2 a-g.—

Shoemaker, 1947: 53, figs. 6, 7.—J. L. Barnard, 1970: 101-

102, fig. 54.

See notes on C. acherusicum. Dactyl of gnathopod 2

with 2-3 accessory teeth. The population at Hilo, only

record so far of the species from Hawaii, is highly

aberrant. Only a few individuals are normal; most

males and females display one or more characters of

the opposite sex. Males may have a few stout spines

on the ventral margins of antenna 2 and females may

have a distal tooth on article 4 like males. This may

represent the loss of phenotypic stability as the result
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 27.—Dexaminidae (=Atylidae): A, Atylus (Kamehatylus) nani J. L. Barnard; w,

Paradexamine (Wailele) maunaioa J. L. Barnard.
NUMBER 58

of a "founder effect." The species is presumed to have

been introduced, like C. acherusicum, by shipping in

the last 2 centuries.

Hawaii, Hilo.

Genus Ericthonius Milne-Edwards

Ericthonius brasiliensis (Dana)

Ericthonius brasiliensis.—Stebbing, 1906:671-672.—Schellenberg, 1938:90.—J.L. Barnard, 1955:37-38.

Erichthonius abditus.—Sars, 1895:602-603, pi. 215.

The ocular (cephalic) lobes are long, the eyes red in

life, turning yellow or clear in spirit; the telson is very

short and broad and covered with tiny hooks; uropod

3 is unmistakable.

Cosmopolitan in tropical and temperate seas, forming great masses of silty tubes attached to piles and

docks in harbors of Oahu but also occurring sparsely

in the open sea to depths of at least 30 m.

61

Family DEXAMINIDAE ( = Atylidae)

FIGURES 27, 28

The Atylidae were recently synonymized with the

Dexaminidae (J. L. Barnard, 1970, p. 26, n. 10). The

family is recognized by the coalesced urosomites 2-3,

the cleft telson, long antennae with vestigial or obsolete

accessory flagellum, and small but subchelate gnathopods. Dexaminids are distinguished from Ampeliscidae

in their compound ommatidial eyes; from Sebidae in

the biramous uropod 3; and from Colomastigidae by

the lateral body compression, the presence of a subchelate gnathopod 1, and the occurrence of mandibular incisors and elongate coxae.

The only atylin of Hawaii is Atylus {Kamehatylus)

nani. It lacks a mandibular palp and thus differs from

the majority of Atylus, but it is so clearly derived from

a line of atylins especially represented by a Japanese

species that it must remain within the genus Atylus.

Key to Genera of Dexaminidae

All urososmites coalesced, palp of maxilliped extending well beyond outer plate, inner plate

of maxilliped well developed, palp of maxilla 1 reaching end of outer plate and 2-articulate,

lower lip lacking inner lobes, pleonites with only 1, or less than 1, dorsal tooth each.

Atylus (Kamehatylus)

Only urosomites 2-3 coalesced, palp of maxilliped not extending beyond outer plate, inner plates

absent, palp of maxilla 1 very short and 1-articulate, lower lip with inner lobes, pleonites 3

and 4 with dorsolateral tooth besides dorsal tooth Paradexamine (Wailele)

Genus Atylus Leach

Atylus (Kamehatylus) nani J. L. Barnard

Atylus (Kamehatylus) nani J. L. Barnard, 1970: 93-96,

figs. 48, 49.

This species differs from all other atyluses in the coalescence of all urosomites and from most atyluses in the

absence of a mandibular palp and the presence of 2

articulate spines on each side of the urosome, like

various other genera of dexaminids.

Body with overall ochraceous-orange color of Sargassum sp., but composed of mottled orange, ochre,

rust, yellow, and buff; eyes red in formaldehyde,

bleaching clear in alcohol. Probably free-living in Sargassum sp.

Oahu, Kawela, intertidal.

Genus Paradexamine Stebbing

Paradexamine (Wailele) maunaloa J. L. Barnard

Paradexamine (Wailele) maunaloa J. L. Barnard, 1970:102-

103, figs. 55, 56.

Body white, eyes black in formaldehyde and alcohol.

Possibly inhabits ascidians.

Oahu, near Kualoa Point, intertidal, on rock covered with ascidians.

Family EUSIRIDAE ( = Calliopiidae)

FIGURES 29, 30

These are basic gammarideans like the family Gammaridae and differ only in degree from various gammarids. The accessory flagellum is either absent or

is composed only of 1 article in Hawaiian eusirids. Both

383-197 O - 71 - 5
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 28.—Dexaminidae (=Atylidae): Paradexamine (Wailele) maunaloa J. L. Barnard

and Atylus (Kamehatylus) nanij. L. Barnard.

species from the Hawiian Islands have large dark

eyes, long thin antennae with short peduncles, article 2

of the peduncle on antenna 1 is slightly shorter than

article 1 and article 3 is very short, uropod 3 has long

triangular, sharply pointed rami bearing long plumose

setae and short spines, and the outer rami of uropods

1-2 are shorter than the inner.

Eusirids differ from liljeborgiids in the vestigial accessory flagella, the weakly toothed or smooth dactyls

of the gnathopods, and for both Hawaiian genera of

eusirids, strongly triturative mandibular molars.

Eusirids resemble pleustids but differ in the cleft

telson and normal lower lip.
NUMBER 5 8 63

Exp

FIGURE 29.—Eusiridae: E, Eusiroides monoculoides (Haswell); p, Pontogeneia pacifica

Schellenberg.
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 30.—Eusiridae: E, Eusiroides monoculoides (Haswell); P, Pontogeneia pacifica

Schellenberg.

FIGURE 31.—Gammaridae: A, Maera sp. A; B, Maera sp. B; c, Ceradocus hawaiensis J. L. Barnard; i, Maera insignis (Chevreux); K, M. kaiulani J. L. Barnard; p, M. pacifica Schellenberg;

Q, M. quadrimana (Dana); s, M. serrata Schellenberg.
506
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Hawaiian Species of Family Eusiridae

Figure Numbers Referenced in Italics

Accessory flagellum absent, inner plate of maxilla 1 with 4 large plumose setae (possibly 3-5)

in adult, gnathopods small, palms weakly setose (29Pg2), uropod 2 much shorter than uropod 1,

inner plate of maxilla 2 narrower than outer, strongly and regularly setose (30Px2).

Pontogeneia pacifica

Accessory flagellum present, small, (30Eaf), inner plate of maxilla 1 with 2 terminal setae,

gnathopods large, palms bearing line of very stout spines (29Egl£), uropod 2 reaching end

of uropod 1, inner plate of maxilla 2 large, flabellate, irregularly setulose (30Ex2).

Eusiroides diplonyx

Genus Eusiroides Stebbing

Eusiroides diplonyx Walker

Eusiroides diplonyx Walker, 1909:333-334, pi. 43, fig. 4.—

Pirlot, 1936:302-303, figs. 126-128.—Schellenberg, 1938:

35.—J. L. Barnard, 1970:109-110, figs. 59-61.

With large black eyes and large gnathopods 1-2 this

species resembles Liljeborgia, but the accessory flagellum is minute and 1-articulate and the palms of the

gnathopods bear a line of extremely stout spines in

contrast to the cycles of thin spines in Liljeborgia.

Maxilla 2 of Eusiroides has an extremely enlarged,

flabellate inner plate with short setae organized into a

facially dispersed group rather than in normal rows or

simply marginal.

Pleonal epimeron 3 with 3-5, occasionally 1-2,

rarely obsolescent, strong or weak, small upturned

teeth at posteroventral corner. Male and female

similar.

Pereopods 1-2 have a very large, sabre-shaped posterodistal locking spine on article 6.

Oahu, intertidal to 30 m.

Genus Pontogeneia Boeck

Pontogeneia pacifica Schellenberg

Pontogeneia pacifica Schellenberg, 1938:35-37, fig. 17.—

J. L. Barnard, 1955:5; 1970:110-111, figs. 62-64.

The eyes of this species also are large and dark in

formaldehyde but fade to pale purple in alcohol. The

head has a long, thin but blunt rostrum projecting between the antennae; its shape is variable, often projecting straight, or curved downwards. The accessory

flagellum is absent; but article 3 of the peduncle, especially in the male, has a minute quadrate extension on

the medial side. Peduncles of the male antennae are

strongly notched and bear on peduncles and flagella

numerous anthurium-shaped, transparent calceoli.

Gnathopods of both sexes are similar to each other,

small, article 6 rectangular, palm oblique but shorter

than posterior margin of hand, defined by 2 pairs of

medial and lateral spines, dactyl fitting weakly setose

palm, dactyls weakly toothed (in contrast to strongly

toothed in Liljeborgia). article 5 of gnathopod 1 with

weak triangular lobe, of gnathopod 2 with strongly

triangular lobe extended distally. Maxilla 2 normally

slender and with large setae on medial margin of inner

plate.

Pleonal epimeron 1 with small posteroventral notch,

2 and 3 with bulging posterior margin, slightly sinuous

posteroventral margin and sharply quadrate posteroventral corners, lacking serrations.

Hawaiian Islands, shallow water.

Family GAMMARIDAE

FIGURES 31-41

Hawaiian taxa of this family conform to the basic gammaridean plan, with normal mouthparts and cleft

telson, 2+-articulate accessory flagellum (one species

has 1-articulate condition in juveniles), uropod 3 with

short peduncle and other parts generally normal, but

often with various modifications of uropod 3 such as

elongation of the outer ramus, the frequent occurrence

of a long article 2 on that ramus and occasionally with

the inner ramus reduced to a leaf-like scale.

Coxa 4 is posteriorly excavate in many Gammaridae

but often it is very weakly excavate or totally

unexcavate.

Many Hawaiian Gammaridae have a strong slit or

notch on the head below the lateral ocular lobe but

the alternates are numerous.

Gammaridae have medium-length to short cleft telsons with one or more spines on each lobe, often
NUMBER 5 8 67

FIGURE 32.—Gammaridae: A, Melita appendiculata (Say); c, Elasmopus calliactis Edmondson;

D, E. Pdiplonyx Schellenberg; i, E. piikoi J. L. Barnard; if, E. molokai J. L. Barnard; N, Nuuanu

amikai J. L. Barnard; p, E. pocillimanus (Bate); R, E. rapax Costa.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

multispinose. Antenna 1 is usually much longer than

antenna 2 but both are stiff and well constructed, with

elongate peduncle on antenna 1, and they are rarely

lost during preservation. The accessory flagellum is

occasionally difficult to see because it is only 2-articulate and article 2 is very minute. Gnathopod 2 is

usually large in males but it is weak in the eriopisaeriopisella group.

The most common Hawaiian gammarids belong to

the genus Elasmopus, characterized by short uropod 3

with broad, truncate, spinose rami, and a large mandibular palp projecting forward with article 3 subfalcate and strongly setose.

Uropod 3 is frequently lost during preservation in

Melita, Eriopisa, Eriopisella, Nuuanu, Ceradocus and

in those species of Maera with elongate rami. Only in

Elasmopus and those species of Maera and Ceradocus

with short rami is uropod 3 consistently present. In

Hawaiian waters Eriopisa, Eriopisella and most members of Maera may be recognized by the absence of an

anteroventral cephalic slit, and the two former genera

are differentiated from each other by the fully setose

inner plates of Eriopisa. The genera with cephalic slit

are Ceradocus and Nuuanu, with medially setose inner

plates of maxillae 1-2, and Melita, and (rarely)

Maera with only terminal setae on those plates.

Nuuanu has characteristic pereopods and weak eyes in

comparison with Ceradocus. Females of Melita have a

hooked coxa 6 and females of Maera do not. Males of

the two genera all have individualistic gnathopods, but

juveniles of the two genera may be difficult to separate

until various minute characters of adults are traced

down through juvenile stages.

The comb row of spines on the mediodistal end of

uropod 2 in Eriopisa and Eriopisella can be seen only

by dissection or strong twisting of the attached appendage ; the comb rows distinguish those genera from

species of Maera with long uropod 3, often lost.

Keys to Genera of Gammaridae

Figure Numbers Referenced in Italics

1. Inner ramus of uropod 3 short and scale-like (37Mu3), outer ramus elongate 2

Rami of uropod 3 subequal to each other in length, rarely in juvenile inner ramus half as

long as outer 5

2. Inner plates of maxillae 1-2 with medial edges lacking setae 3

Inner plates of maxillae 1-2 with medial edges strongly setose their full lengths 4

3. Head rounded anteroventrally and lacking notch or slit, uropod 2 with comb-row of spines

on mediodistal end of peduncle (36Hu2), female coxa 6 normal Eriopisella Chevreux

Head with deep anteroventral notch {31 Ah), uropod 2 lacking comb-row, female coxa 6 with

large anterior hooked process (39Mc6) Melita Leach

4. Uropod 3 greatly exceeding uropods 1-2, head lacking anteroventral notch. Eriopisa Stebbing

Uropod 3 scarcely exceeding uropods 1-2, head with deep anteroventral notch.

Nuuanu J. L. Barnard

5. Palp article 3 of mandible falcate and strongly setose on one linear margin (35Dmp).

Elasmopus Costa

Palp article 3 rectolinear and with setae mostly terminal 6

6. Inner plates of maxillae 1-2 strongly setose their full length on medial edges. Ceradocus Costa

Inner plates of maxillae 1-2 with only terminal setae Maera Leach

Alternative Key if Uropod 3 Missing

1. Mandibular palp article 3 weakly to strongly falcate and densely setose along concave margin

(35Cmp) 2

Mandibular palp article 3 rectolinear, setae mostly terminal 4

2. Inner plates of maxillae 1-2 with setae mainly terminal Elasmopus Costa

Inner plates of maxillae 1-2 with medial edges setose their full length 3

FIGURE 33.—Gammaridae, Elasmopus; male gnathopod 2 and epimera: c, E. calliactis Edmondson; D, E. diplonyx Schellenberg; E, E. ecuadorensis hawaiensis Schellenberg; H, E. hooheno

J. L. Barnard; i, E. piikoi J. L. Barnard; M, E. molokai J. L. Barnard; P, E. pocillimanus (Bate);

R, E. rapax Costa; s, E. spinidactylus Chevreux; x, E. pectenicrus (Bate).

70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3. Head with broad, triangular lateral lobes and deep anteroventral notch (3N), uropod 3 not

strongly exceeding uropods 1-2 Nuuanu J. L. Barnard

Head with truncate anterolateral margin and weak anteroventral excavation, no notch (37Lh),

uropod 3 strongly exceeding uropods 1-2 Eriopisa Stebbing

4. Inner plates of maxillae 1-2 densely setose along full medial edges Ceradocus Costa

Setae on inner plates of maxillae 1-2 mostly terminal 5

5. Rami of uropod 3 equal to each other in length, either short or elongate Maera Leach

Inner ramus of uropod 3 very short and scale-like, outer ramus elongate (37Lu3) 6

6. Head broadly truncate or weakly convex on anterolateral margin, uropod 2 with comb-row

of spines on mediodistal end of peduncle (36Hu2) Eriopisella Stebbing

Head with deep anteroventral notch or slit below prominent lateral lobe (37Mh), uropod 2

lacking peduncular comb-row Melita Leach

Genus Ceradocus Costa

Ceradocus hawaiensis J. L. Barnard

Ceradocus hawaiensis J. L. Barnard, 1955:5-8, figs. 2, 3;

1970:115, fig. 65.

Maera insignis, Melita appendiculata, and Ceradocus

hawaiensis all have a pair of triangular dorsal teeth on

urosomite 1 and thus are distinguishable at first glance

from all other Hawaiian gammarideans. Maera insignis

has no other dorsal serrations on the pleon and C.

hawaiensis and Melita appendiculata have other serrations on the pleon while C. hawaiensis may further

be distinguished from M. appendiculata in the presence of serrations on pleonal epimeron 3, the presence

of an anteroventral cephalic notch and a single anteroventral cephalic tooth in contrast to the absence of a

notch and a pair of teeth on M. appendiculata. The

pereopodal dactyls of C. hawaiensis have 1-4 tiny

accessory cusps, whereas dactyls of M. appendiculata

have 1 large accessory tooth. Both males and females

of C. hawaiensis have one large gnathopod 2 and one

small (mixed between right and left), similar between

the sexes, with a transverse deeply toothed palm;

whereas male M. appendiculata has one enlarged

gnathopod 2 (always the left) with extremely attenuate palmar tooth on which the dactyl closes, resembling a shrimp claw. Females of M. appendiculata

have very small gnathopod 2 on both sides, and both

males and females of C. hawaiensis have one of their

second gnathopods small, like those of right side of

male and both sides of female M. appendiculata. The

rami of uropod 3 are of medium and equal extent in

C. hawaiensis, whereas they are highly unequal and

the outer is very long in M. appendiculata; but in

both species they are frequently lost in preservative.

Coxa 1 of C. hawaiensis is sharply attenuate

anterodistally.

The inner plate of maxilla 1 on Ceradocus hawaiensis is lined its full medial margin with stout setae but

the inner plate on M. appendiculata has 2 stout terminal setae and several very thin medial setules.

Eyes black.

Oahu, 0-30 m.

Genus Elasmopus Costa

Elasmopus is the most diverse genus of Gammaridea

in the Hawaiian Islands; it is represented by 8 or 9

species occurring in a variety of habitats from intertidal

zones to depths as great as 30 m; the species are mainly

associated with algae. The short uropod 3 with subequal, flat rami, falcate article 3 of the mandibular

palp, and terminally (but not medially) setose inner

plates of the maxillae, distinguish elasmopuses from

other members of Gammaridae. All have a deep lateral

cephalic notch, but other familial taxa also bear that

character.

Comments on each species are abbreviated as positive statements, the combination of characters being

unique for each species. Eyes are therefore mentioned

only for E. calliactis and E. pectenicrus, since they

are the only species lacking black pigment (brownpurple when faded) ; but locking spines of pereopods

1-2 are mentioned for each species owing to their

diagnostic variability. Sharply serrate epimeron 3 combined with black eyes characterize E. rapax, but E.

pectenicrus females also have those sharp serrations;

the clear eyes (red in life) combined with castelloserrations on one or more article 2 of pereopods 3-5

are better diagnostic features for recognition of E.

pectenicrus than would be a partially negative

statement on epimeron 3.
NUMBER 5 8 71

Keys to Hawaiian Species of Genus Elasmopus

Figure Numbers Referenced in Italics

Terminal Males

1. Palm of gnathopod 2 with marginal cusp or tooth easily visible and defining proximal end of

palm {33Cg2, Rg2) 2

Palm of gnathopod 2 lacking marginal defining tooth, occasionally with locking tooth on

medial face of hand (33Dg2, Xg2) 5

2. Palm of gnathopod 2 with only 1 distal protuberance besides defining tooth, palm otherwise

flat (33Mg2) E. molokai (eyes dark)

Palm of gnathopod 2 with protrusion or tooth in middle besides distal protrusion and proximal

defining tooth (33Hg2) 3

3. Eyes with dark pigment 4

Eyes red in life, clear in alcohol 9

4. Telson squamiform (35Et), main locking spine of pereopods 1-2 a striate sabre (34Hpl).

E. hooheno

Telson normally long (3511), main locking spine of pereopods 1-2 straight, unstriate, apically

chisel-shaped (34Ip2) E. piikoi

5. Hand of gnathopod 2 with large cuplike medial hollow (33Pg2), article 6 of pereopods 1-2

with 2 sabre-shaped striate locking spines (34Pp4) E. pocillimanxu

Hand of gnathopod 2 lacking medial hollow, article 6 of pereopods 1-2 with either simple

locking spines or with only one member sabre-shaped and striate (34Ep2) 6

6. Dactyls of pereopods 1-5 with 2-5 large castelloserrations or villi proximal to main constriction (34Spl) E. spinidactylus

Dactyls of pereopods 1-5 proximally smooth 7

7. Dactyls of pereopods 1-2 with elephantine spine at constriction (plus setules) (34Dp2).

E. diplonyx

Dactyls of pereopods 1-2 with normal seta-spine at constriction (plus setules) (341 p2) 8

8. Palm of gnathopod 2 with 3 distal processes, one long lateral, one small slightly proximomedial

and truncate, one distomedial shallow and spinose, plus mediofacial locking tooth for end

of dactyl, palm poorly setose (33Rg2) E. rapax

Palm of gnathopod 2 with only a small or absent distal process, palm heavily setose, with or

without weak mediofacial locking cusp (33Eg2) E. ecuadorensis hawaiensu

9. Pereopod 4 with posteroventral excavation on article 2 lined with deep crenuloserrations or

castellae (fluted) (35Xp4) E. pectenicrus

Pereopod 4 normally ovatorectangular, unexcavate and with small normal serrations (like

35Hp5) E. calliactis

Females and Juveniles

(Applicable Also to Most Males)

1. Article 6 of pereopods 1-2 with 2 sabre-shaped, striate locking spines on posterodistal end at

base of dactyl (34Pp4) E. pocillimanus

Pair of locking spines composed of 1 sabre-shaped and 1 simple spine (34Spl) 2

Pair of locking spines composed of either 2 simple spines or with one member of the pair

slightly chisel-shaped apically (34Ep5) 4

2. Dactyls of pereopods 1-5 with long castellae or villi proximal to normal constriction (34Spl).

E. spinidactylus

Dactyls lacking accessory villi 3

3. Smaller locking spine of pereopods 1-2 of corkscrew kind (34Hpl), larger locking spine of

pereopods 3-5 striate and sabre-shaped E. hooheno

Smaller locking spine of pereopods 1-2 simple (34Ep2), larger locking spine of pereopods

3-5 straight and simple E. ecuadorensis hawaiensis

4. Ornament of constriction on dactyls of pereopods 1-5 composed of an elephantine spine in

addition to setules (34Dp2) E. diplonyx

Ornament of constriction on dactyls of pereopods 1—5 composed of a stout to slender seta in

addition to setules (34Hpl) 5
FIGURE 34.—Gammaridae, Elasmopus: D, E. diplonyx Schellenberg; E, E. ecuadorensis hawaiensis

Schellenberg; H, E. hooheno J. L. Barnard; i, E. piikoi J. L. Barnard; M, E. molokai J. L. Barnard;

p, E. pocillimanus (Bate); R, E. rapax Costa; s, E. spinidactylus Chevreux; x, E. pectenicrus

(Bate).

FIGURE 35.—Gammaridae, Elasmopus: c, E. calliactis Edmondson; n, E. diplonyx Schellenberg;

E, E. ecuadorensis hawaiensis Schellenberg; H, E. hooheno J. L. Barnard; i, E. piikoi J. LBarnard; M, E. molokai J. L. Barnard; p, E. pocillimanus (Bate); R, E. rapax Costa; s, E.

spinidactylus Chevreux; x, E. pectenicrus (Bate).
NUMBE

R 5 8

73
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

5. Eyes clear in alcohol or red in life and formaldehyde 6

Eyes retaining brown-black-purplish pigment in alcohol 7

6. Article 6 of gnathopod 1 subovate, palm very oblique, outer ramus of uropod 3 with small

article 2 seen in ventral view, pereopods 4-5 with normally small posterior serrations on

article 2, dactyl of gnathopod 2 with few setules only, not wavy or castellate....£. calliactis

Article 6 of gnathopod 1 rectangular, palm nearly transverse, outer ramus of uropod 3 without

article 2, pereopod 4 in female, pereopods 4-5 in juvenile with deep castellations on article 2,

dactyl of gnathopod 2 castelloserrate or wavy on inner margin E. pectenicrus

7. Telson short, scarcely as long as broad (35Mt) E. molokai

Telson long, longer than broad (35Rt) 8

8. Pleonal epimeron 3 with 2+ sharp serrations including posteroventral corner (33Pep), telson

with subapical lateral spines only, with smoothly projecting apical margins (35Rt).

E. rapax

Pleonal epimeron 3 with only one sharp tooth at posteroventral corner, apices of telson

truncate and spinose (35It) E. piikoi

Elastnopus calliactis Edmondson

Elasmopus calliactis Edmondson, 1951:189-191, fig. 3.—

J. L. Barnard, 1970:115, fig. 66.

Elasmopus rapax J. L. Barnard, 1955:11 (only those specimens from Honolulu Aquarium, not Costa).

This species and E. pectenicrus are the only members

of the genus in Hawaii lacking black-brown-purple pigment in the eyes. In life the eyes are red and the body

banded in red but the pigment bleaches in alcohol. Telson long, each lobe with apical excavation hidden by

rank of 5-6 subterminal spines.

Terminal male palm of gnathoped 2 defined by

large cusp and bearing middle tooth and adze-shaped

distal process, dactyl simple.

Elasmopus pectenicrus differs in having deep castello-serrations on article 2 of either or both pereopods

4—5 in both sexes or juveniles.

Oahu, in Honolulu Aquarium and in depths presumably to 30 m, in association with Dardanus hermit

crab bearing Calliactis sea-anemones on its Dolium

shell.

Elasmopus ?diplonyx Schellenberg

?Elasmopus diplonyx Schellenberg, 1938:54, fig. 26.

Elasmopus Pdiplonyx J. L. Barnard, 1970:115-119, figs. 67,

68.

Characterized in all stages by the very stout, sharp

(juvenile) or blunt (adult) spine occupying the constriction of the dactyls on pereopods 1-2. Telson

medium in length, apices excavate and sinuous or

raggedly serrate, each subapex with 2 spines.

Terminal male palm of gnathopod 2 scarcely defined

by acclivity and spines, medial face with locking ridge

and 2 spines, dactyl slightly overriding palmar margin,

palm with small distal protuberance, strongly setose and

lacking middle tooth.

Posterior margin of article 2 on pereopods 4-5

deeply crenuloserrate but distinguished from E. pectenicrus in the dark eyes and dactylar tooth-spine.

Micronesia to Hawaii. Oahu, off Moku Manu

Island, 33 m, on base of black coral in mollusk

association.

Elasmopus ecuadorensis hawaiensis Schellenberg

Elasmopus ecuadorensis hawaiensis Schellenberg, 1938:54—

55, fig. 27.—J. L. Barnard, 1970:119-121, fig. 69.

Article 6 of pereopods 1-2 with one striate sabre-spine

and one simple spine in the locking pair, dactyl normal.

Telson very short, apices slightly excavate, with 2

apicolateral spines on each lobe.

Terminal male gnathopod 2 with undefined s-shaped

heavily setose palm with small distal process, small

ridge on medial palmar face, and small medial locking

spine on ridge at apex of dactyl, latter simple, curved.

Differs from E. hooheno in that the secondary locking spine of pereopod 1 is simple, not ornamented as a

corkscrew and both spines of the locking pair on

pereopods 3-5 are simple. Male gnathopod 2 looks like

that of E. pectenicrus but article 2 of pereopod 4 is unmodified and the locking spines of pereopods 1-2 are

different.

Oahu, KaneoheBay; Hawaii, Hilo; Molokai.

Elasmopus hooheno J. L. Barnard

Elasmopus hooheno J. L. Barnard, 1970:121, fig. 70.

Article 6 of pereopods 1-2 with 1 striate sabre-spine

and 1 uncurved corkscrew spine in the locking pair,
NUMBER 5 8 75

dactyl normal. Telson very short, apices slightly excavate, with 2 apicolateral spines on each lobe.

Terminal male gnathopod 2 palm defined by large

cusp, bearing truncate tooth in middle and slightly

scalloped, spinose distal protrusion, dactyl simple.

Close to E. molokai but main locking spine of pereopods 1-2 sabre-shaped and male gnathopod 2 with

middle palmar tooth.

Oahu, Kawela Bay, intertidal.

Elasmopus molokai J. L. Barnard

Elasmopus molokai J. L. Barnard, 1970:121-125, figs. 71, 72.

Article 6 of pereopods 1-2 with 1 normal short spine

and one long apically chisel-shaped spine in the locking

pair, dactyl normal. Telson very short, apices sinuous,

with 2-4 apical-subapical spines each.

Terminal male gnathopod 2 palm defined by large

cusp, lacking midprocess, bearing subfalcate distal

protrusion, dactyl simple.

Close to E. piikoi, the latter with long telson and

close to E. ecuadorensis hawaiensis, the latter with

male gnathopod 2 of similar, heavily setose, uncuspidate morphology as E. pectenicrus. The telson of

E. rapax (35 Rt) is distinctive of that species.

Oahu, widely distributed from intertidal zones to

depths of 30 m.

Elasmopus pectenicrus (Bate)

Moera pectenicrus Bate, 1862:192, pi. 34, fig. 8.

Elasmopus serrula Walker, 1904:277-278, pi. 8, fig. 37.

Elasmopus pectenicrus.—J. L. Barnard, 1955:8-10, fig. 4

(with references); 1970:125, figs. 73, 74.

Without black pigment in the eyes, thus like E. calliactis. Locking spines of pereopods almost straight and

otherwise normal. Inner ramus of uropod 3 only twothirds as long as outer ramus, latter without article 2.

Telson of medium length, apices truncate, with 4-6

spines each.

Terminal male gnathopod 2 with undefined,

s-shaped, heavily setose palm with small process on

transverse distal end, small ridge on medial palmar

face, and small medial locking cusp near apical extent

of overriding dactyl, latter simple, curved.

Male with article 2 of pereopod 4 excavate posteroventrally and deeply castelloserrate, female with

serrations slightly weaker but not excavate, juvenile

without excavation but with serrations on both

pereopods 4 and 5, unlike adult.

Tropicopolitan. Oahu, Pearl Harbor, on buoys and

docks. Possibly introduced by shipping.

Elasmopus piikoi J. L. Barnard

Elasmopus piikoi J. L. Barnard, 1970:127-128, figs. 75-76.

Article 6 of pereopods 1-2 with 1 normal and 1 apically

chisel-shaped spine in the locking pair, dactyl normal.

Telson of normal length, lobes apically excavate, with

3 terminal spines each.

Terminal male gnathopod 2 palm defined by small

cusp, middle of palm with large truncato-coniform

process, distal end with truncate or slightly bilobate

process; inner margin of dactyl bisinuate, with large

process.

Elasmopus molokai has a short telson and all stages

of E. rapax have 2+ sharp posteroventral serrations

on pleonal epimeron 3, whereas E. piikoi either has 1

serration forming the corner in juveniles and females

or lacks serrations in males. The telson of E. rapax

has spines only on the subapical lateral notch and has

long smooth medial lobes, whereas E. piikoi has spines

terminally.

Oahu, intertidal to 18 m.

Elasmopus pocillimanus (Bate)

Moera pocillimanus Bate, 1862:191, pi. 34, fig. 7.

Elasmopus pocillimanus.—Stebbing, 1906:443-444.—Chevreux and Fage, 1925:246, fig. 257.—Schellenberg, 1938:

56, fig. 28.—J. L. Barnard, 1970:128-131, figs. 77, 78.

Distinguished in all stages by the presence of 2 large,

striate sabre-shaped locking spines on article 6 of

pereopods 1-2.

Telson long, lobes apically extended and smooth,

each lobe with 1 midlateral spine. Eyes densely pigmented with black-brown-purple, more so than in

other species in Hawaii.

Terminal male palm of gnathopod 2 defined by

weak cusp, with slight distal protrusion but medial

surface of hand bearing deep hollow defined proximally by weak spine, dactyl simple, stout, overlapping

palmar face to close within recess.

Cosmopolitan in warm seas. Oahu, (0)-30 m, coral

and coralline bottoms.
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Elasmopus rapax Costa

Elasmopus rapax Costa.—J. L. Barnard, 1955:10-12, fig. 5

(in part, not figs. 5b, g), (with references); 1970:131-

134,685.79,80.

Article 6 of pereopods 1-5 with grossly simple pair of

locking spines, longest spine minutely gnarled, twisted

and pointed apically (seen under oil-immersion lens

only). Telson of normal length, with broad smooth

apices and 2-3 spines on subapical lateral notch,

apices pointed in juveniles and females. Pleonal epimeron 3 with 2-6 sharp serrations including that of

posteroventral corner. Uropod 3 lacking article 2 on

outer ramus.

Terminal male palm of gnathopod 2 defined by

strong tooth on medial face (not seen from lateral

view), middle of palm with tooth slightly medioproximal to long lateral tooth, distal end with shallow spinose process; palm and medial face thus with 4 teeth

in all.

Cosmopolitan in warm and cool seas. Oahu, shallow

water, primarily harbors, possibly introduced by

shipping.

Elasmopus spinidactylus Chevreux

Elasmopus spinidactylus Chevreux, 1908:486-489, figs. 9-

10.—Walker, 1909:336-37.—Schellenberg, 1938:55.—

Shoemaker, 1942:13.—J. L. Barnard, 1965:504; 1970:

136-138, figs. 81,82.

Dactyls of pereopods 1-5 with several villiform or

castelliform teeth on margin proximal to main constriction, condition unique in Hawaii to this species.

Article 6 of pereopods 1-2 with 1 striate sabre and 1

small normal locking spine. Telson short, apices broadly

truncate, each armed with 3 spines.

Terminal male palm of gnathopod 2 unknown in

Hawaii but in southern Polynesia it has a large and

broad fully dominating, falconiform protrusion.

Middle-size males and females have the palm and

posterior margin of article 6 on gnathopod 2 confluent,

heavily setose, distally with a low, broad, spinose protrusion. Pereopods 3-5 have rectangular article 2 with

deep posteroventral notch.

Almost circumtropical. Oahu and Molokai, intertidal.

Key to Hawaiian Species of Genus Eriopisa Stebbing

Figure Numbers Referenced in Italics

Pleonal epimeron 3 with large sharp posteroventral tooth and slightly bulging posterior margin

(36Lur) E. laakona

Pleonal epimeron 3 grossly extended posteriorly as wide obtuse rounded lobe bearing small sharp

cusp on posterodorsal corner (36Hep) E. hamakua

Eriopisa (?) hamakua J. L. Barnard

Eriopisa (?)hamakua J. L. Barnard, 1970:138-140, figs.

83, 84.

Uropod 3 of this species is not known but is presumed

to fit the eriopisa model. The unnotched head lobes

and medially setose inner plates of the maxillae suggest

the generic classification. Gnathopod 2 of the male is

relatively large for the genus, article 5 is short, almost

subcircular, article 6 is very slender and elongate,

densely setose on the posterior margin, the palm defined by a small bulge with 2 spines and the distal end

of the palm with 3 medial spines among the many

setae, palm with weak medial excavation. Gnathopod

1 much more slender, setose and elongate than in E.

laakona.

Eyes very weak and orange in formaldehyde.

S. Oahu, 3-4 m, sandbottom and encrustations.

Eriopisa laakona J. L. Barnard

Eriopisa laakona J. L. Barnard, 1970:140-143, figs. 85, 86.

Both articles 5 and 6 of male gnathopod 2 are elongate, the palm very oblique, weakly setose and defined

by 2 spines, the dactyl slightly overlapping the palm.

Eyes absent.

Oahu, intertidal to 4 m.

Genus Eriopisella Chevreux

Eriopisella sechellensis upolu J. L. Barnard

Eriopisella sechellensis upolu J. L. Barnard, 1970:143-147,

figs. 87, 88.

With its brownish-purple eyes in alcohol and melitalike uropod 3 this species of Eriopisella, unlike mem-
NUMBER 5 8 77

bers of clear-eyed or blind Nuuanu and Eriopisa,

resembles Melita pahuwai closely but differs from

various melitas either in the absence of an anteroventral cephalic notch or in the presence of a comb-row

of closely set spines on the mediodistal end of the peduncle on uropod 2. This appendage must be viewed

from dorsal aspect in order to see the row of spines.

The sexes are similar to each other; they have a small

gnathopod 2 scarcely larger than gnathopod 1 and

similarly shaped, except that article 6 of gnathopod 2

is somewhat protuberant along the posterior margin

of the subrectangular article 6.

Oahu, Kaneohe Bay, 3-4 m, bottom of mixed corals,

corallines and algae.

Genus Alaera Leach

Two groups of Maera occur in Hawaii: those species

with transverse palms of gnathopod 2 in both sexes

plus bifid pereopodal dactyls, and those species with

oblique palms on gnathopod 2 and non-bifid pereopodal dactyls. Maera insignis, a dominant Hawaiian

intertidal gammaridean, is easily recognized by the

pair of sharp dorsal carinae on pleonite 4. Only 2 species of Hawaiian Maera have a lateral cephalic notch

like various species of Elasmopus, and all species of the

latter genus have a conspicuously falcate article 3 of

the mandibular palp, easily seen without dissection.

Uropod 3 of Elasmopus is consistently short but is variable in Maera. Specimens of Elasmopus retain uropod

3 on preservation but specimens of Maera with long

rami may lose that appendage as do specimens of

Ceradocus, Melita, Eriopisella and Eriopisa. Ceradocus

is distinguished from Maera in the densely setose

medial edges of the inner plates of maxillae 1-2;

members of Eriopisella have a comb row of spines on

the mediodistal end of the peduncle on uropod 2; one

species of Melita, in contrast to all but two members

of Maera, has a cephalic notch and the other species

is grossly toothed dorsally on the pleon; and members

of Eriopisa have weakly subfalcate and strongly setose

article 3 on the mandibular palp.

Key to Hawaiian Species of Maera Leach

Figure Numbers Referenced in Italics

1. Pleonite 4 with large triangular dorsal carinae (311) M. insignis

Pleonite 4 dorsally smooth 2

2. Palm of gnathopod 2 oblique 3

Palm of gnathopod 2 transverse 4

3. Coxa 1 sharply hooked forward, head with anteroventral notch.

Maera sp. A (cf. M. hamigera)

Coxa 1 evenly trapezoidal, head lacking anteroventral notch but armed with anteroventral

cusp Maera sp. B

4. Pleonite 3 with 1 or many posterior serrations besides posteroventral tooth M. serrata

Pleonite 3 bearing only posteroventral tooth 5

5. Article 2 of pereopods 3-4 thin and subrectangular (31Qp3), palm of male gnathopod 2 with

3 truncate processes and 2 sinuses (besides sinus adjacent to defining tooth) (38Qg2).

M. quadrimana

Article 2 of pereopods 3-4 moderately broad and subovate (31Pp4), male gnathopod 2 with

2 truncate processes and 1 sinus (38Pg2) 6

6. Eyes dark, purple-brown in alcohol, telsonic lobes weakly excavate-bifid, primarily with medial

cusp on either lobe (4OPt) M. pacifica

Eyes orange or clear in alcohol, telsonic lobes deeply bifid (40Kt) M. kaiulani

Maera insignis (Chevreux)

Elasmopus insignis Chevreux, 1901:406-412, figs. 24-31.

Maera insignis.—Schellenberg, 1938:50-52, fig. 24.—J. L.

Barnard, 1955:12-13; 1970:147 fig. 89.

See remarks under Ceradocus hawaiensis (page 70).

This species differs from all other Hawaiian members of Maera in the pair of sharply triangular dorsal

processes on pleonite 4. The lateral cephalic slit gapes

and the telsonic lobes are stiffly fixed in a slightly spiral

form so as to appear from dorsal view as sharp triangles

with oblique apical margins lined with long spines.

Male pereopods 4-5 are especially broadened and

the male gnathopod 2 palm essentially occupies the

full posterior margin of the hand although the dactyl

383-197 O - 71 - 6
FIGURE 36.—Gammaridae: H, Eriopisa (?)hamakua J. L. Barnard: L, E. laakona J. L. Barnard;

H, Melita pahuwai J. L. Barnard; N, Nuuanu amikai J. L. Barnard; s, Eriopisella sechellensis

upotu J. L. Barnard.
Su3

FIGURE 37.—Gammaridae: A, Melita appendiculata (Say); H, Eriopisa hamakua; L, E. laakona;

M, Melita pahuwai J. L. Barnard; N, Nuuanu amikai J. L. Barnard; s, Eriopisella sechellensis

upolu J. L. Barnard.
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

C92

FIGURE 38.—Gammaridae: A, Maera sp. A; B, Maera sp. B; c, Ceradocus hawaiensis J. L. Barnard; 1, Maera insignis (Chevreux) ,* K, M. kaiulani J. L. Barnard; p, M pacific a Schellenberg;

Q, M. quadrimana (Dana); s, M. serrata Schellenberg.
NUMBER 5 8 81

Ng2

FIGURE 39.—Gammaridae: A, Melita appendiculata (Say); H, Eriopisa hamakua J. L. Barnard;

L, E. laakona J. L. Barnard; M, Melita pahuwai J. L. Barnard; N, Nuuanu amikai J. L. Barnard;

s, Eriopisella sechellensis upolu J. L. Barnard.
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 40.—Gammaridae: A, Macro, sp. A. (cf. M. hamigera); B, Maera sp. B; c, Ceradocus

hawaiensis J. L. Barnard; i, Maera insignis (Chevreux); K, M. kaiulani J. L. Barnard; p, M.

pacifica Schellenberg; Q, M. quadrimana (Dana); s, M. serrata Schellenberg.
NUMBER 5 8 83

Bur

FIGURE 41.—Gammaridae: A, Maera sp. A; B, Maera sp. B; c, Ceradocus hawaiensis J. L. Barnard; i, Maera insignis (Chevreux); K, M. kaiulani J. L. Barnard; p, M. pacifica Schellenberg;

s, M. serrata Schellenberg.
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

reaches only about three-fourths of its distance; the

palm is heavily setose but bears 2 large sharp teeth in

the middle and a smaller bi- or trifid distal protrusion.

The rami of uropod 3 are short and subequal and

the eyes are black.

Indo-Pacific. Oahu, intertidal, common.

Maera kaiulani J. L. Barnard

Maera kaiulani J. L. Barnard, 1970:147-150, figs. 90, 91.

See diagnosis of M. pacifica, below, with following

differences: eyes orange or clear in alcohol; article 4

of gnathopod 2 with strong or weak cusp, palm scalloped distinctly, occasionally in terminal male with

one excavation in middle; article 2 of pereopods 3-5

thinly rectangular only in juveniles; inner ramus of

uropod 3 only half that of outer ramus in juvenile,

rami elongate, and extending strongly beyond uropods

1-2, becoming normally short in adults; telson with

deeply incised lobes with very sharp cusps, 2 large

spines.

Oahu, 4-5 m.

Maera pacifica Schellenberg

Maera pacifica Schellenberg, 1938:42-45, figs. 19-20.—J. L.

Barnard, 1965:511; 1970:150-151, figs. 92, 93.

Eyes bright or brownish purple in alcohol, anteroventral cephalic corner with sharp tooth (rotate head to

see); male and female gnathopod 2 large and similar

to each other but occasionally right and left members

different in size, article 4 without tooth, transverse

palm with strong defining process and very weakly

scalloped palm, male with large excavation in middle,

dactyl often with inner acclivity fitting palmar excavation ; dactyls of pereopods 1-5 complexly bifid; article 2

of pereopods 3-5 ovate, posteroventral lobes broad;

uropod 3 with medium to short rami, thin or of medium

breadth, scarcely exceeding uropods 1-2, rami subequal to each other; telson with truncate apices or

small medial cusp and 3 large spines on each lobe, not

bifid; pleonal epimera 1-3 each with bulging posterior

margin and small to medium-sized sharp posteroventral cusp.

Southern Polynesia; Micronesia; Oahu, 0-30 m.

Maera quadrimana (Dana)

Gammarus quadrimanus Dana, 1853:955-956, pi. 65, fig. 9.

Maera quadrimana.—Schellenberg, 1938:46-48, figs. 21,

22.—J. L. Barnard, 1955:13; 1965, 511-512, fig. 17;

1970:152-155, figs. 94,95.

See diagnosis of M. pacifica, above, with following differences: article 4 of gnathopod 2 slightly extended

distally; male gnathopod 2 with 2 sinuses and 3 truncate processes in middle of palm; dactyl lacking acclivity; article 2 of pereopods 3-4 subrectangular, of 5

subovate, posteroventral lobes narrow and quadrate;

in terminal stages medial cusps on telsonic lobes

obsolescent.

In Hawaii this species and M. pacifica are often difficult to distinguish in female and juvenile stages because pereopods 3-5 often are missing or, if regenerating, article 2 is thin in both species. Study is required

to clarify their distinctions, since variations in rami of

uropod 3 and male gnathopod 2 shape also occur.

Medium-size males of M. quadrimana occasionally1

have one stunted palmar process.

Southern Polynesia; Micronesia; Hawaiian Islands,

0-30 m.

Maera serrata Schellenberg

Maera inaequipes serrata Schellenberg, 1938:41-42, fig. 18.—

J.L.Barnard, 1965:510.

Maera serrata.—]. L. Barnard, 1970:155-156, figs. 96, 97.

See diagnosis of M. pacifica, above, with following differences: eyes red in formaldehyde, pale purple in

alcohol; article 4 of gnathopod 2 with sharp posteroventral extension; male palm of gnathopod 2 with

small excavation in middle, dactyl with small inner

hump fitting excavation; bifidation of dactyls on pereopods weaker than in M. pacifica, M. quadrimana, or

M. kaiulani; article 2 of pereopods 3-5 narrowly subrectangular, with narrow and serrate posteroventral

lobes; rami of uropod 3 equal to each other in length,

thin; narrow telsonic lobes apically bifid, each with

2 pairs of long spines.

Pleonal epimeron 3 deeply serrate posteriorly unlike

other species of Hawaiian Maera. Juveniles have at

least one weak serration and setule besides posteroventral tooth.

Southern Polynesia; Micronesia; Oahu, 0-3 m.

Maera sp. A (cf. M. hamigera), J. L. Barnard

Maera species A J. L. Barnard, 1970:156-159, figs. 98, 99.

A single female is known. Rami of uropod 3 extremely

elongate and palm of gnathopod 2 oblique. Dis-
NUMBER 5 8

tinguished from Maera sp. B in the presence of a lateral

cephalic notch and from juveniles of M. kaiulani in

the dark eyes (blood red to black), cephalic notch,

and oblique palm of gnathopod 2. Dactyls of pereopods

not bifid.

Southern Oahu, 30 m.

85

Maera sp. B, J. L. Barnard

Maera species B J. L. Barnard, 1970:159-161, fig. 100.

A single female is known. Rami of uropod 3 extremely

elongate and palm of gnathopod 2 oblique. Head

without lateral cephalic notch. Eyes black. Dactyls of

pereopods not bifid as in species like M. pacifica.

Oahu, Kawela Bay, intertidal.

Key to Hawaiian Species of Genus Melita Leach

Figure Numbers Referenced in Italics

Pleon with numerous dorsal serrations and cusps (37Mpl), head with bifid corner below lateral

lobe M. appendiculata ( = fresneli)

Pleon bearing only small, almost microscopic dorsolateral cusp on pleonite 4, head with deep

slit below lateral lobe (37Mh) At. pahuwai

Melita appendiculata (Say) ( = fresneli

(Audouin))

Melita appendiculata (Say).—J. L. Barnard, 1970:161, figs.

101, 102.

Melita fresneli (Audouin).—J. L. Barnard, 1955:13-14

(with references).

See remarks under Ceradocus hawaiensis (page 70).

Melita appendiculata is one of the most striking

amphipods in any ocean because of the large decapodous claw of the left male gnathopod 2. In resemblance

to fiddler crabs the right gnathopod 2 is small; both

gnathopods are small in the female.

The conspicuous dorsal serrations and cusps on the

pleon distinguish the species from M. pahuwai but

so many fundamental differences occur between the

two species that they scarely seem to be in the same

genus. Gnathopod 1 of M. appendiculata is of the normally gammarid kind, that of M. pahuwai is complexly ornamented and has transversely bulging palms

and complex dactyls. Melita pahuwai has the normal

cephalic notch whereas M. appendiculata has only a

bifid anteroventral cephalic corner.

Eyes dark orange in alcohol.

Cosmopolitan in warm seas.

Melita pahuwai J. L. Barnard

Melita pahuwai J. L. Barnard, 1970:161-166, figs. 103, 104.

Male gnathopod 2 has a flabellate hand with distally

quadratiform definition, minutely spinose, medial surface with axial channel filled with setae, dactyl overriding palm onto medial surface; male gnathopod 1

with hand distally swollen, palm complexly sinuous and

spinose, dactyl grotesquely bulging basally but apically

unguiform. Outer ramus of uropod 3 with only 1 article

in contrast to M. appendiculata. Both gnathopods of

the male are enlarged.

Pereopodal dactyls with double distal constriction

but no accessory tooth as in M. appendiculata.

Eyes brownish purple in alcohol.

Oahu, Kawela Bay, intertidal.

Genus Nuuanu J. L. Barnard

Nuuanu amikai J. L. Barnard

Nuuanu amikai J. L. Barnard, 1970:166, figs. 105, 106.

Nuuanu is a genus endemic to Hawaii and resembling

Melita but having uropod 3 very short and the outer

ramus composed of 2 articles. In this character Nuuanu also resembles Eriopisa and Eriopisella but differs

from them in the elongate coxae, the tall head with

very widely triangular cheeks (cephalic lobes) bearing

a few scattered clear ommatidia, a weak dorsal tooth

on each of pleonites 1 and 2 and a small dorsolateral

spine on each side of urosomites 2 and 3. Article 2 of

pereopods 3-5 is deeply castello-serrate, like pereopod

4 of Elasmopus pectenicrus and pereopods 4-5 of E.

diplonyx, but the shapes of article 2 (37Np5) are distinctive and the serrations occur on all 3 members of

pereopods 3-5.

Southern Oahu, 18 m, on Pocillopora reef.
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 42.—Isaeidae: Chevalia aviculae Walker.

FIGURE 43.—Isaeidae, Gammaropsis (=Eurystheus Bate): H, Gammaropsis haleiwa J. L. Barnard; K, G. kaumaka J. L. Barnard; L, G. Pali J. L. Barnard; o, G. pokipoki J. L. Barnard; Y, G.

alamoana J. L. Barnard; z, G. atlantica Stebbing.

88

Family ISAEIDAE ( = Photidae)

FIGURES 42-47

The basic member of the isaeid complex, including the

families Ampithoidae, Aoridae, Ischyroceridae, Corophiidae, and, for purposes of this paper, the Podoceridae, has a fleshly telson, very thick, broad, short, and

uncleft, though often dorsally excavate. From a lateral

view the telson projects from behind the lateral wings

of urosomite 3 as a small tent, often armed with at

least 1 thick spine, often with an additional seta, and

occasionally with a conspicuous cornified bead or hook;

in the Corophiidae the telson and urosome may become so flattened that these remarks are no longer true.

Isaeid-complex members, except apparently for podocerids, have glands in articles 2 and 4, and occasionally

3 and 5, of pereopods 1-2. The glands may have a

variety of colors, even purple and orange, but usually

in preserved specimens the glands have been changed

to yellow, ochre, amber, or brown; occasionally they

are colorless. They conceal the muscles normally seen

as axially oriented fibrous bundles. Often a duct

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

may be traced through the appendage to the subapex

or the dactyl where secretions are emitted and spun

into tubes.

Isaeids generally have the appearance of the basic

gammaridean except for the telson, the glandular pereopods and the absence of a posterior excavation on

coxa 4. The unexcavate coxa is also characteristic of

some Gammaridae and it is therefore an unreliable

familial character by itself.

Mouthparts of most isaeid members are grossly monotonous, fulfilling the basic gammaridean concept.

Isaeidae have the fewest modifications of their group,

the outer ramus of uropod 3 lacking spine hooks seen

in Ampithoidae, or the tiny denticles and hooked appearance of Ischyroceridae; and Isaeidae have gnathopod 2 larger than 1 in the male.

Rami of uropod 3 are never flattened or perfectly

lanceolate as they are in many other gammarideans;

generally the rami are thin cylinders or elongate

boxes, and the peduncle is never elongate as it is

ischyroceridae, nor are the rami broadened as in

Ampithoidae.

Key to Genera of Isaeidae

Figure Numbers Referenced in Italics

1. Pleonites 4-5 coalesced (42, body), but pleonite 6 free Chevalia Walker

All pleonites free 2

2. Inner ramus of uropod 3 half or less as long as outer ramus (46Ku3) Photis Krayer

Inner ramus of uropod 3 as long as or longer than outer.

Gammaropsis Liljeborg ( = Eurystheus)

Cheiriphotis, a genus not yet found in Hawaii but articulate. The urosome is tall, so that the coalesence

expected to occur there resembles Photis in having the of the 2 segments is clearly apparent from lateral view,

inner ramus of uropod 3 very short, often absent in Coxae 5-7 are very small and discontiguous and coxae

terminal adults; the peduncle of uropod 3 is platelike 1-4 also are small and barely touch. Antenna 1 is

and very small, and article 3 of antenna 1 is much much longer than 2 and the antennae usually remain

shorter than article 1, whereas in Photis article 3 is as with the animal upon preservation. Chevalia is frelong article 1. quently very abundant in sublittoral samples; but the

organisms are small (3 mm) and may be spotted by

Genus Chevalia Walker their white bodies with red or clear eyes, large gnathor,t ,. . . Xkr „ pod 2 with transverse palm, and long setose antennae

Chevalia aviculae Walker ^ v . ' 6 , , . ,

plus pereopods 3-5 (especially 3-4) and thick pedunFIGURE 42 cle s o n tiie pleopods. The outer rami of uropods 1-2 are

Chevalia aviculae Walker, 1904:288-290, pi. 7, fig. 50, pi. shorter than the inner, the outer bearing conspicuous

8, fig. 50.—J. L. Barnard, 1962:17-20, fig. 5; 1970: terminal spines, but both pairs of rami lack dorsal

' ' ' spines. The rami of uropod 1 have minute setules and

The coalesence of urosomites 1 and 2 is unique to this spines on their lateral margins. The outer ramus of

species of Hawaiian amphipod; urosomite 3 is freely uropod 3 appears shorter than the inner only from
NUMBER 5 8

Yh

FIGURE 44.—Isaeidae, Gammaropsis: H, G. haleiwa J. L. Barnard; K, G. kaumaka J. L. Barnard;

L, G. pali J. L. Barnard; o, G. pokipoki J. L. Barnard; x, G. afra Stebbing; Y, G. alamoana J. L.

Barnard; z, G. atlantica Stebbing.

lateral view. The telson is fleshy, subquadrate, shaped

like a pillow, and strongly setose dorsally. Antenna 1

has a 2-articulate accessory flangellum.

Oahu, 0-30 m, common. Circumtropical and

warm-temperate.

Genus Gammaropsis Liljeborg (= Eurystheus Bate)

FIGURES 43-45

A diverse assemblage of species assigned to this genus

occurs in Hawaii. The several groups may not be

congeneric but the differences among them are minute

and have not been studied carefully in the world fauna

to see their significance. The main group of species

is composed of the atlantica-afra-pali-pokipokikaumaka complex in which uropod 3 has a small article

2 and the inner plate of maxilla 1 has at least 3, often

5 + setae lining the medial margin. Gammaropsis pali

differs from its four other group members in the presence of a fan-shaped transparent flake on the smooth

base of the right mandibular molar, but G. pokipoki

has a rugose surface in the same area. Gammaropsis

haleiwa forms a class by itself: it has 2 articles in the

outer ramus of uropod 3; maxilla 1 has only 1 long
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terminal seta (plus a short one); the right mandibular

molar has a flake; and the maxillipedal palp article 3

bears a distal adze-shaped process, and article 4, unlike

other members of Hawaiian Gammaropsis, is long, almost unguiform, and with a short terminal nail. Other

members have article 4 shorter, stouter, barrel-shaped,

and bearing terminal setal spines as long as the article

itself. Gammaropsis haleiwa and G. kaumaka are

marked by a long ventral peduncular cusp on uropod

2 similar to that seen on uropod 1 of all members of

Gammaropsis. The remaining species, G. alamoana

lacks article 2 on the outer ramus of uropod 3 and

lacks setae on the inner plate of maxilla 1; G. alamoana

has a right molar flake and seta and G. alamoana is

distinguished in its conically acute epistome projecting

forward.

Key to Hawaiian Species of Genus Gammaropsis Liljeborg

Figure Numbers Referenced in Italics

1. Apex of outer ramus on uropod 3 lacking a minute article 2 and bearing 2-5 setae or spines.

(Palm of gnathopod 1 lacking stout defining spines in both sexes, eyes retaining black pigment in alcohol, urosomite 1 with small dorsal pair of protuberances, uropod 2 lacking ventral

peduncular tooth between rami, epistome with long, coniform anterior projection.)

G. alamoana

Apex of outer ramus on uropod 3 with minute cuboid article 2 bearing 2 setae (43Ku3) and

often surrounded or partially hidden by other setae 2

2. Uropod 2 with long sharp peduncular tooth between rami (43Hur, 45Ku2) 3

Uropod 2 lacking long peduncular tooth 4

3. Telson with 2 stout dorsal spines, urosomites smooth, inner plate of maxilla 1 with medial

setae, maxillipedal palp article 3 lacking process, 4 short, with long apical setae.. G. kaumaka

Telson lacking dorsal spines, urosomal margins with weak castellae, inner plate of maxilla 1

with only terminal setae, maxillipedal palp article 3 with strong distal lobe (44HxppaV)

article 4 long, unguiform, with apical nail G. haleiwa

4. Article 5 of gnathopods 1-2 with 2-3 stout anterior spines (43L), article 6 of gnathopod 1

rectangular, palm shorter than posterior margin of article 6, article 2 of pereopods 3-5

slender and rectangular G. pali

Article 5 of gnathopods 1—2 lacking stout anterior spines, article 6 of gnathopod 1 ovate,

palm and posterior margin of article 6 equal to each other in length*, article 2 of pereopods

3—5 ovate or pyriform, especially broad on pereopod 3 (45Yp3) 5

5. Rami of uropod 3 about 150 percent as long as peduncle (43Ou3), inner slightly longer than

outer, inner with 2 stout apical spines, urosomites 1-2 simple, each with 1 dorsolateral seta,

lateral margins straight, dactyl of male gnathopod 2 overlapping palm (45Og2), palmar

excavation dominating palmar area, gnathopod 2 small, eyes retaining black pigment in

alcohol, black in life G. pokipoki

Rami of uropod 3 subequal to peduncle in length (43Zu3), outer slightly longer than inner,

inner with 1 small apical spine, urosomites 1—2 with 1 dorsolateral seta marking dorsal

boundary of bulging lateral urosomital margin, dactyl of male gnathopod 2 not overlapping

palm, palmar excavation not dominating palmar area, gnathopod 2 large, eyes becoming

clear or rust colored in alcohol, red or brown in life 6

6. Eyes horizontally ovate (44Zh) G. atlantica, form B

Eyes vertically ovate, lageniform, reniform or trapezoidal 7

7. Eyes lageniform (43Z) adults of G. atlantica, form A

Eyes otherwise 8

8. Eyes vertically ovate (like 44Oh) juveniles of G. atlantica forms A & B?

Eyes vertically reniform or trapezoidal (44Xh) 9

9. Adults 2.2 mm or less in body length, eyes brown in 2-day formaldehyde, retaining dark

brown or rust stain in alcohol, body white G. afra

Adults about 3.0 mm or longer, specimens smaller always immature, eyes red in 2-day formaldehyde, becoming clear in alcohol but body (and especially head) retaining pink-purple

stain G. afra-atlantica hybrids

•Separated from each other by a stout spine.
NUMBER 5 8 91

Gammaropsis afra Stebbing

Gammaropsis afra and G. afer, Stebbing, 1888:1097, pi. 113;

J. L. Barnard, 1970:170, fig. 108.

Eurystheus afra.—Stebbing, 1906:612.— ?K. H. Barnard,

1916:249-250, pi. 28, fig. 11; 1937:165-166, fig. 12.—

Pillai, 1957:55-56, fig. 13.—J. L. Barnard, 1961:113-114,

fig. 79.

Outer ramus of uropod 3 with 2 articles, inner plate of

maxilla 1 with 3+ medial setae, right mandibular

molar with seta, no flake, epistome unproduced, article 5 of gnathopods 1-2 lacking stout anterior spines,

article 6 of gnathopod 1 ovate and with stout defining

spine, article 2 of pereopods 3-5 ovate or pyriform,

uropod 2 lacking ventral peduncular tooth, rami of

uropod 3 not significantly longer than peduncle, urosomites 1-2 with bulging posterolateral margins marked

dorsally by a seta, gnathopod 2 of both sexes large,

dactyl fitting palm, palm oblique, excavate at proximal end only, with giant articulate spine.

Eyes brown, body clear in 2-day formaldehyde, eyes

trapezoidal or reniform on short ocular lobes. Adults

not exceeding 2.2 mm in length.

Hybrid form, with G. atlantica: reaching 4 mm in

length, eye slightly more reniform than in G. afra,

palm of gnathopod 2 much less oblique, concavity narrowest, palmar bulge quadrate instead of lobed as in

atlantica and afra, proximal inner margin of dactyl

with low quadrate process; color as in G. atlantica,

with red eyes and body-head pigment.

Almost circumtropical. Oahu, 30-33 m, on deep

Pocillopora reef and black coral base.

Gammaropsis alamoana J. L. Barnard

Gammaropsis alamoana J. L. Barnard, 1970:170-174, figs.

109, 110.

Outer ramus of uropod 3 lacking article 2, inner plate

of maxilla 1 lacking setae (bearing setular wisps only),

right mandibular molar with large seta and molar

flake, epistome with long, conical anterior cusp, article

5 of gnathopods 1-2 lacking stout anterior spines, article 6 of gnathopod 1 ovate and lacking stout defining

spine, article 2 of pereopods 3-5 ovate or pyriform,

uropod 2 lacking ventral peduncular tooth, rami of

uropod 3 not significantly longer than peduncle, urosomite 1 with shallow pair of dorsal protuberances,

gnathopod 2 of both sexes large, dactyl fitting or shorter

than palm, palm oblique, with strong defining tooth,

and broad palmar tooth, excavation flat, bearing large

spine becoming lost in terminal males, latter having

coxa 7 enlarged enormously.

Eyes black, ovoreniform, ocular lobes of medium

extent.

Oahu, widely distributed from intertidal zone to

depths of 30 m.

Gammaropsis atlantica Stebbing

Gammaropsis atlantica Stebbing, 1888:1101, pi. 114.—J. L.

Barnard, 1970:174-178, figs. 111-113.

Gammaropsis zeylanicus Walker, 1904:282-283, pi. 6, fig. 41.

Gammaropsis Gardineri Walker, 1905:929-930, pi. 88, figs.

11-14, 16, 17.

Eurystheus atlanticus Stebbing, 1906:611.—Pillai, 1957:56-

57, fig. 14.—J. L. Barnard, 1965:534-535, fig. 29.

Outer ramus of uropod 3 with 2 articles, inner plate of

maxilla 1 with 3 plus medial setae, right mandibular

molar with seta, no flake, epistome unproduced, article 5 of gnathopods 1-2 lacking stout anterior spines,

article 6 of gnathopod 1 ovate and with stout defining

spine, article 2 of pereopods 3-5 ovate or pyriform,

uropod 2 lacking ventral peduncular tooth, rami of

uropod 3 not significantly longer than peduncle,

urosomites 1-2 with bulging posterolateral margins

marked dorsally by a seta, gnathopod 2 of both sexes

large, dactyl fitting palm, palm oblique, excavate at

proximal end only, with giant articulate spine.

Eyes red, body and head clear or purplish pink in a

2-day formaldehyde, eyes lageniform on medium to

long ocular lobes or ovate on long ocular lobes. Ovate

eyes either horizontal or vertical. This species occurs in

several ocular forms with intermediate stages, the eyes

of adults showing ommatidial proliferation to form a

lageniform condition. See G. afra for remarks on

hybrids.

Almost circumtropical; Oahu, widely distributed

from intertidal depths to 30 m, mainly in depths

exceeding 5 m.

Gammaropsis haleiwa J. L. Barnard

Gammaropsis haleiwa J. L. Barnard, 1970:178-181, fig. 115.

Outer ramus of uropod 3 with 2 articles, inner plate

of maxilla 1 with 1-2 terminal setae, right mandibular

NUMBER 5 8 93

molar with seta and flake, epistome unproduced, article 5 of gnathopods 1-2 lacking stout anterior spines

(rarely with an anterior seta), article 6 of gnathopod 1

ovate and with stout denning spine, article 2 of pereopods 3-5 ovate or pyriform, uropod 2 with long

peduncular tooth between rami, rami of uropod 3 significantly longer than peduncle, urosomites 1-2 with

slight modifications like afra-atlantica, gnathopod 2 of

male small, dactyl fitting palm, palm oblique, scarcely

excavate, bearing 2 stout but not greatly enlarged

spines; urosomites 2 and 3 with posterodorsal marginal

nobs; maxillipedal palp article 3 extended in strong

adze-shaped process, article 4 elongate, tipped with

nail. Female unknown.

Eyes small and subcircular, with black central core

surrounded by a few clear ommatidia on elongate narrow ocular lobes.

Oahu, Kaneohe Bay, 2 m.

Gammaropsis kaumaka J. L. Barnard

Gammaropsis kaumaka J. L. Barnard, 1970:181, fig. 116.

Outer ramus of uropod 3 with 2 articles, inner plate of

maxilla 1 with 2 medial and 2 terminal setae, right

mandibular molar poorly known, probably without

flake but with basal rugosity, epistome unproduced,

article 5 of gnathopods 1-2 lacking stout anterior

spines, article 6 of gnathopod 1 subrectangular and

with stout defining spine, article 2 of pereopods 3-5

pyriform, uropod 2 with long peduncular tooth between rami, rami of uropod 3 not exceeding length of

peduncle, urosomites 1-2 with smooth margins,

gnathopod 2 of presumed male of medium size, dactyl

fitting palm, palm oblique, smoothly rounded, unexcavate, defined by large spine; maxillipedal palp

article 3 unproduced or lobate, article 4 short and

bearing long apical setae. Female unknown.

Eyes of medium size, irregularly ovate and slightly

tilted from horizontal axis, with black-brownish purple

central core surrounded by a few clear ommatidia on

ocular lobes of medium extension.

Oahu, Kawela Bay, intertidal.

Gammaropsis pali J. L. Barnard

Gammaropsis pali J. L. Barnard, 1970:183-184, figs. 117,

118.

Outer ramus of uropod 3 with 2 articles, inner plate of

maxilla 1 with 3+ medial setae, right mandibular

molar with seta and flake, epistome unproduced,

article 5 of gnathopods 1-2 with 2-3 stout anterior

spines, article 6 of gnathopod 1 rectangular and with

stout defining spine, article 2 of pereopods 3-5

slender, rectangular, uropod 2 lacking ventral peduncular tooth, rami of uropod 3 not significantly longer

than peduncle, urosomites 1-2 simple, gnathopod 2

of both sexes of medium size, dactyl overriding palm,

palm oblique, widely and deeply excavate but scarcely

defined, with giant articulate spine.

Eyes bright garnet-ruby red in 2-day formaldehyde,

head with similar red spots, red lost in alcohol revealing black core, antennae dark splotchy garnet, a few

body segments with brown vertical stripes, ocular lobes

of medium extent.

Oahu, Kaneohe Bay, 2 m.

Gammaropsis pokipoki J. L. Barnard

Gammaropsis pokipoki J. L. Barnard, 1970:185, figs. 119,

120.

Outer ramus of uropod 3 with 2 articles, inner plate of

maxilla 1 with 3+ medial setae, right mandibular

molar with seta, no flake but with basal rugosity,

epistome unproduced, article 5 of gnathopods 1-2

lacking stout anterior spines, article 6 of gnathopod

1 ovatorectangular, bearing stout defining spine, article

2 of pereopods 3-5 ovate or pyriform, uropod 2 lacking ventral peduncular tooth, rami of uropod 3 nearly

150 percent as long as peduncle and outer slightly

shorter than inner, urosomites 1-2 simple, gnathopod 2

of both sexes small, sexually dimorphic, dactyl overlapping palm in male, palm oblique, broadly and

deeply excavate, with giant articulate spine, female

palm very weakly and scarcely defined.

Eyes large, subcircular, with black central core in

alcohol surrounded by thick ring of small, numerous

clear ommatidia on long ocular lobe.

Oahu, 2-30 m.

FIGURE 45.—Isaeidae. Gammaropsis; all unlettered figures are gnathopod 2: H, G. haleiwa J. L.

Barnard; K, G. kaumaka J. L. Barnard; o, G. pokipoki J. L. Barnard; p, G. pali J. L. Barnard;

x, G. afra Stebbing; Y, G. alamoana J. L. Barnard; z, G. atlantica Stebbing.

383-197 O - 71 - 7
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Aep

FIGURE 46.—Isaeidae, Photis: A, Photis aina J. L. Barnard; H, P. hawaiensis J. L. Barnard; K,

P. kapapa J. L. Barnard.

FIGURE 47.—Isaeidae, Photis: A, Photis aina J. L. Barnard; H, P. hawaiensis J. L. Barnard; K,

P. kapapa J. L. Barnard.
NUMBER 5 8 95

Kg2
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Hawaiian Species of Genus Photis Kr0yer

Figure Numbers Referenced in Italics

1. Eyes composed of relatively large ommatidia, many of them noncontiguous and few of them

perfectly spherical, eyes horizontally ovate and filling ocular lobe and protruding into head

proper, ocular lobe of medium length, eyes microscopically eosin in alcohol; uropods 1-2

with minute but distinct sharp deltoid tooth on peduncle between rami; telson sharply and

evenly triangular in adult male (like 49Ptu3), but softly triangular in other stages, in male

with 2 sharp lateral cusps in place of normal nobs P. hawaimsis

Eyes composed of tiny spherical ommatidia tightly packed into small horizontally ovate or

circular eyes occupying only apex of long ocular lobes (46Ah), eyes black or clear in alcohol;

uropods 1-2 lacking peduncular tooth between rami; telson softly triangular or irregular

(with separate apex) and with nobs, unlike males of first half of couplet 2

2. Eyes black or brown in alcohol and formaldehyde, anterior coxae distally rounded weakly,

essentially without setae except a few on coxa 1, inner ramus of uropod 3 about one-fourth

as long as outer (46Ktu3) P. kapapa

Eyes red or clear in alcohol and formaldehyde, anterior coxae long rectangles with numerous

long ventral setae, inner ramus of uropod 3 about half as long as outer (46Atu3). .P. aina

Photis aina J. L. Barnard

Photis aina J. L. Barnard, 1970:185-191, figs. 121, 122.

Male gnathopod 2 small, article 2 lacking distal lobe,

hand broad, palm slightly oblique, bisinuate, defined

by quadrate corner, bearing weak hump near defining

corner and one near base of dactyl, medioproximal

face of palm with large spine, dactyl fitting palm in

young male, much shorter in terminal male; female

gnathopod 2 slightly smaller than in male, defining

corner rounded, palm with 1 weak smooth sinus, otherwise crenulate.

Pereopod 3 very stout. Eyes clear in formaldehyde.

Oahu, 2-30 m, primarily in deeper end of depth

range.

Photis hawaiensis J. L. Barnard

Photis hawaiensis J. L. Barnard, 1955:35-37, figs. 18, 19;

1970:191-192, fig. 123.

Male gnathopod 2 grotesque: article 2 bearing immense anterolateral mammilliform lobe armed with

stridulation ridges, article 3 with lobe projecting medially, article 4 strongly setose medially, article 5 with

large, heavily setose anterior keel-lobe, posteriorly with

large lobe partially hiding base of hand, latter tapering

distally, occupied by palm posteriorly, palm defined by

proximomedial tooth and small medial excavation,

dactyl equalling palm; female gnathopod 2 with

oblique palm, tooth and slight excavation, defined by

quadrate corner, hand expanded.

Pereopod 3 about normal for genus, slightly stoutened, bearing only 1 accessory cusp on dactyl.

Male coxae 3-4 with ventral stridulation ridges,

coxae 1-2 lacking ridges, all coxae scarcely setose.

Inner ramus of uropod 3 very short, less than onefourth as long as outer ramus. Eyes pale eosin or clear

in formaldehyde.

Oahu, Kaneohe Bay, shallow water.

Photis kapapa J. L. Barnard

Photis kapapa J. L. Barnard, 1970:192-195, figs. 124, 125.

Male gnathopod 2 small, article 2 with anterodistal

mammilliform lobe bearing stridulation ridges, article

4 not produced posterodistally, article 5 of medium

breadth, cup-shaped, with tumid posterior lobe, hand

expanded, palm slightly oblique, defined by large blunt

tooth, palm weakly bisinuate, with small obtuse cusp

near dactylar base, medial spine on low hump and

medial quadrate process anterior to spine, young male

with weaker defining cusp, dactyl fitting palm in

juvenile, becoming progressively shorter in terminal

male; female gnathopod 2 palm variable, slightly

oblique, s-shaped and fully crenulate or with semicircular smooth excavation in middle, dactyl fitting palm

or much shorter, palm defined by softly rounded

quadrate corner.

Pereopod 3 normal for genus.

Male coxae 1-2 with weak posterior stridulation

ridges, 4 with complex of medial and lateral ventral

ridges, 3 with weak ventral stridulation ridges. Eyes

black in formaldehyde.

Oahu, 2-30 m.
NUMBER 5 8 97

Family ISCHYROCERIDAE

FIGURES 48-50

See Isaeidae (page 88) for an introduction to this

family. Ischyroceridae are part of the isaeid tubicolous

complex and are difficult to distinguish from Isaeidae

without clear microscopic resolution of the apical hook

or apicolateral denticles on the outer ramus of uropod

3. Jassa and Ischyrocerus are relatively easy to recognize because the peduncle of uropod 3 is elongate and

the rami short; isaeids usually have the peduncle short

and the rami elongate. Occasionally ischyrocerids lose

uropod 3 after death. Parajassa and Ventojassa have

the peduncle of uropod 3 less elongate than the other

genera and one must observe the very minute apical

uncinus on the outer ramus of uropod 3. Often the oil

immersion adaptation of the microscope must be used.

But Parajassa angularis, the only Hawaiian member

of the genus, has long plumose setae on the lateral margins of the peduncles of uropods 1-2, a character

unique to that species and the biancolinid Biancolina

mauhina. One must not confuse misdirected setae of

the pleopods for uropodal setae. Adults of Ventojassa

ventosa, the only Hawaiian member of the genus, have

characteristic outer rami on uropod 3, seen even with

low power microscopy. The outer ramus forms a digit

with 1-3 short, stout, wirelike marginal setae {49

Vu3or). All isaeids have either stout lateral spines or

thin terminal setae on the outer ramus of uropod 3.

Ventojassa and Parajassa form a pair of "mimics"

often co-occurring with relatively similar color patterns (fading in alcohol) but Ventojassa has pink

antennae and Parajassa has clear antennae reflecting

the brownish purple of the body. A species of Lembos

also has pink antennae but can be distinguished by

the enlarged gnathopod 1. Shapes of ocular lobes

differ between Parajassa and Ventojassa; and Parapassa has short antennal flagella, very stout article 2 on

pereopods 1-2 and numerous plumose setae on the anterior margin of article 2 on pereopod 3. Sexes in both

species are similar to each other. Even gnathopod 2 is

relatively similar between the sexes of both species.

The almost ubiquitous Jassa falcata (Montagu)

has not yet been found in the Hawaiian Islands. It

differs from /. lilipuna and other Hawaiian ischyrocerids in the presence of an articulate but partially immersed apical hook on the outer ramus of uropod 3

besides the 2 or 3 apicolateral, reverted, large, asymmetrical sharp cusps, all of these features being visible

under medium power compound microscopy. Males

and females of /. falcata have a variety of shapes in

gnathopod 2 as the species is of extreme multiform

character (Sexton and Reid, 1951).

Jassa lilipuna, Ischyrocerus kapu, and /. oahu are

small bodied species (1-3 mm), with superficial

similarities, the heads with sharp but not acute cephalic

lobes and circular to subcircular eyes with clear peripheral ommatidia and brown cores; the antennal pairs

are subequal in length, moderately short and stout, the

accessory flagellum being grossly visible, a thick 2-

articulate piece usually standing erect from antenna 1

on article 3.

Key to Hawaiian Species of Ischyroceridae

Figure Numbers Referenced in Italics

1. Accessory flagellum a small tubercle on article 3 of antenna 1, not usually visible macroscopically, uropods 1-2 with long peduncular setae (49Pur) Parajassa

Accessory flagellum 2-3 articulate and elongate, often thick (50Jaf), uropods 1-2 without

peduncular setae {49Jur) 2

2. Outer ramus of uropod 3 armed with 1-2 (or 3) stiff lateromarginal setae besides distal nail

or hook (49Vu3or) Ventojassa

Outer ramus of uropod 3 lacking gross spines or setae, with slightly hooked apex and bearing

(including apical hook) 1-12 large or small apicolateral denticles, cusps or villi (49Pu3or,

Ju3, Ou3) 3

3. Eyes lacking black-brown pigment in alcohol, bleaching to rusty-orange—Ischyrocerus sp. C

Eyes retaining black-brown pigment cores in alcohol 4

4. Gnathopod 2 of male (female unknown) with long spike-like article 4 (50Kg2).

Ischyrocerus kapu

Gnathopod 2 with unproduced article 4 5
98

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 48.—Ischyroceridae: j , Jassa lilipuna J. L. Barnard; p, Parajassa angularis Shoemaker;

v, Ventojassa ventosa (J. L. Barnard).
NUMBER 5 8 99

5. Article 6 of pereopods 1-2 short and stout, male gnathopod 2 article 5 shielded from posterior

margin by articles 4 and 6 (50Jg2), outer ramus of uropod 3 with 2-4 large cusps (49Ju3),

coxae 3-4 longer than broad, article 2 of pereopods 3-5 broad, posteroventrally lobate on

pereopods 3-4 Jasta lilipuna

Article 6 of pereopods 1-2 long and moderately slender (50 Opl), male gnathopod 2 article 5

not shielded from posterior margin and bearing blunt lobe (50 Og2), outer ramus of uropod

3 with 4-7 small cusps (seen only under oil-immersion lens) (49 Ou3), 3-4 as broad as

or broader than long, article 2 of pereopods 3-5 narrow and not lobate posteroventrally.

Ischyrocerus oahu

Ischyrocerus (?) kapu J. L. Barnard

Ischyrocerus (?) kapu J. L. Barnard, 1970:195, fig. 126.

Only the male is known so far and it is unique in the

world family for the large spikelike projection on article 4 that seems to serve in place of a proximal palmar

tooth on gnathopod 2. The palm of gnathopod 2 has

2 distal processes, and extends the full length of article

6; the dactyl is long. The coxae are very short. The

outer ramus of uropod 3 is shorter than the inner and

bears a row of about a dozen tiny apicolateral denticles

appearing as villi. Eyes rusty purple in alcohol.

Oahu, 2-30 m, rare.

Ischyrocerus oahu J. L. Barnard

Ischyrocerus oahu J. L. Barnard, 1970:195-198, fig. 127.

The terminal male gnathopod 2 is very large, article 6

elongate and slender, the posterior margin slightly concave and moderately setose marginally and submarginally, distally the palm bearing a weak bilobation, dactyl

as long as article 6 and inner basal margin bisinuate

to fit palmar process, article 5 with posterior margin

reaching free space and bearing blunt lobe (in contrast

to /. kapu and Jassa lilipuna), young male with shorter

dactyl and stouter article 6; pereopods 1-2 with long

article 6 (in contrast to /. lilipuna) and bearing locking spines (in contrast to /. kapu); outer ramus of

uropod 3 slightly shorter than inner ramus and often

projecting laterally so as to appear even more

shortened, bearing 4-7 long, apicolateral denticles.

Eyes rusty purple in alcohol.

Oahu, 0-2 m.

Ischyrocerus sp. G

General appearance, uropod 3 and coxae like /. oahu

but eyes lacking any dark pigment in alcohol, rustyorange.

S. Oahu, 30 m.

Jassa (?) lilipuna J. L. Barnard

Jassa (?) lilipuna J. L. Barnard, 1970:198-202, figs. 128-

130.

Male gnathopod 2 is large and undergoes numerous

changes during maturation. Coxa 1 is slightly shorter

than coxa 2 but this is difficult to see without dissection

and mounting of parts. Uropod 3 is diagnostic; the

outer ramus is slightly curved distally and bears 2-3

medium-sized cusps. The species of Ischyrocerus have

extremely minute denticles.

Article 6 of pereopods 1-2 is short in /. lilipuna

whereas it is long in the other 2 species. Eyes clear in

alcohol or retaining rusty pink core under high magnification.

Oahu, Waikiki, intertidal, on Sargassum, Ulva, and

Padina.

Parajassa angularis Shoemaker

Parajassa angularis Shoemaker, 1942:41-44, figs. 14, 15.—

J. L. Barnard, 1962:58, figs. 26, 27; 1970:202-204, figs.

131, 132.

The outer ramus of uropod 3 of Hawaiian individuals

has 2 non-articulate hooks and 2 minute apical setules

(use oil immersion lens). The peduncular setae on

uropods 1-2 are diagnostic. Eyes black.

A warm-temperate Californian species. S. Oahu,

2-30 m.

Ventojassa ventosa J. L. Barnard

Eurystheus ventosa J. L. Barnard, 1962:20-22, figs. 6, 7.

Ventojassa ventosa J. L. Barnard, 1970:204-205, figs. 133,

134.

The outer ramus on uropod 3 of Hawaiian individuals

has a slightly hooked apex with 2-3 partially immersed

setules and 1-3 stiff, wirelike lateromarginal setae visible under low-medium power on slide 2. They are

diagnostic. Eyes brown-black in formaldehyde.

A warm-temperate Californian species. Oahu,

4-33 m.
100

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pur

FIGURE 49.—Ischyroceridae: j , Jassa (?)lilipuna J. L. Barnard; o, Ischyrocerus oahu J. L.

Barnard; p, Para jassa angularis Shoemaker; v, Vento jassa ventosa (J. L. Barnard).

FIGURE 50.—Ischyroceridae: j , Jassa lilipuna J. L. Barnard; K, Ischyrocerus kapu J. L. Barnard;

o, Ischyrocerus oahu J. L. Barnard; p, Par a jassa angularis Shoemaker; v, Vento jassa ventosa

(J. L. Barnard).
NUMBER 5 8

Kg2
102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Family LEUCOTHOIDAE (cf. Anamixidae)

FIGURES 14 (part), 15 (part)

Leucothoids have a characteristic and unmistakeable

gnathopod 1 like the Anamixidae (14Lgl), with

strong carpochelation and narrow article 6. They have

ordinary mandibles and maxillae, whereas anamixids

have these replaced by a ventral keel. Leucothoid

mandibles are flabellate, lack molars, and have weak

palps. The Hawaiian anamixid has a characteristic

gnathopod 2, whereas leucothoids have 2 kinds of

gnathopod 2 (15Lg2 and Pg2). These animals are

small (1-3 mm), shiny, usually pure white or with red

vertical bands at body segment junctions and with red

eyes; but some phenotypes of Anamixis have red vertical bands in the middle parts of the segments.

Uropod 3 is elongate, with the peduncle equalling

the two rami, but the appendage often falls off during

preservation. Uropod 2 is shortened. Leucothoe has

large and visible coxa 1 but Leucothoides has a small

coxa 1 like the Amphilochidae and may be distinguished from the latter by the characteristic gnathopod 1.

Leucothoids appear to be associated as inquilines

with sessile invertebrates; some non-Hawaiian species

are known to live in the branchial cavities of ascidians.

Key to Genera of Leucothoidae

Figure Numbers Referenced in Italics

1. Coxa 1 small and partially covered by enlarged coxa 2; gnathopod 2 characteristic (15Pg2),

palm distinct, scarcely oblique, slightly concave, palp of mandible 1-articulate.

Leucothoides Shoemaker

Coxa 1 as large as coxa 2 and visible, gnathopod 2 characteristic (15Hg2), very large, article 6

ovate, palm and posterior margin of article 6 confluent, mandibular palp 3-articulate 2

2. Coxa 1 beveled and/or smooth anteroventrally, coxae 2 and 4 rounded anteroventrally.

Leucothoe Leach

Coxa 1 sharply serrate and pointed anteroventrally, coxae 2 and 4 pointed anteroventrally.

LeucothoeUa* Schellenberg

Key to Hawaiian Species of Leucothoe Leach

Figure Numbers Referenced in Italics

1. Gnathopod 1 short and stout, dactyl very short (14Lgl), article 6 only 130 percent as long

as anterior margin of article 5 L. lihue

Gnathopod 1 long, slender, dactyl long (14Hgl), article 6 about 190 percent as long as

anterior margin of article 5 2

2. Lateral cephalic lobe with sharp anterior cusp (14Th), coxa 4 short and not excavate

posteriorly, article 2 of pereopods 3-5 evenly ovate (15Tp5), pleonal epimeron 2 roundedquadrate posteroventrally or scarcely protuberant, uropod 2 extending only three fourths

along peduncle of uropod 3, anterior cephalic keel between antennae sharply pointed (not

to be confused with epistome) L. tridens

Lateral cephalic lobe evenly and broadly rounded (14Hh), coxa 4 long and deeply excavate

posterodorsally, article 2 of pereopods 3-5 broadened and especially broadly beveled on

pereopod 5 (15Lp5), pleonal epimeron 2 with large posteroventral tooth, uropod 2 extending to end of peduncle of uropod 3, anterior cephalic keel between antennae rounded.

L. hyhelia

* LeucothoeUa is a widely distributed Indo-Pacific genus that may be discovered in Hawaii.
NUMBER 5 8

Leucothoe hyhelia J. L. Barnard

Leucothoe hyhelia J. L. Barnard, 1965:489-490, fig. 5;

1970:205-209, fig. 135.

Two phenotypes appear to occur in Hawaii; the typical member has a rounded quadrate posteroventral

corner on epimeron 2 and the atypical member has

that corner slightly protuberant. Gnathopods 1—2 of

this species and L. tridens are identical. The telson is

scarcely trilobate apically.

Micronesia to Hawaii, Oahu, widely distributed

from intertidal zones to depths of 30 m.

Leucothoe lihue J. L. Barnard

Leucothoe lihue J. L. Barnard, 1970:209-211, fig. 136.

This species superficially resemble L. hyhelia in terms

of coxae, antenna 1, epimera, and pereopods 3-5 (see

description of L. hyhelia in couplet 2); but article 2 of

pereopod 5 is weakly concave on the obliquely beveled

posteroventral margin. The head weakly resembles

that of L. tridens but the cephalic lobe is obtusely angular, not sharp. Gnathopod 2 is characteristically

broadened in the middle of article 6 (in males) and

article 5 has a villus-like apical protrusion on the long

lobe. The telson resembles that of L. tridens.

Oahu, off Moku Manu Island, 33 m, on base of

black coral in mollusk association.

Leucothoe tridens Stebbing

Leucothoe tridens Stebbing, 1888:777-779, pi. 47.—Schellenberg, 1938:21-23, fig. 11.—J. L. Barnard, 1970:211,

fig. 137.

103

The telson has a weak trilobation apically. See key and

discussions with other species of the genus.

Tropical Pacific Ocean. Oahu, 2-30 m.

Genus Leucothoides Shoemaker

Leucothoides ?pottsi Shoemaker

?Leucothoides pottsi Shoemaker, 1933a: 249-250, fig. 3.—

Schellenberg, 1938:26-28, fig. 13; J. L. Barnard, 1970:

211-213, fig. 138.

This genus is easily recognizable by the small coxa 1

and the relatively small gnathopod 2, with short article

6 and subtransverse, concave palm with conical

ornaments.

Pan-American. Oahu, 0-30 m.

Family LILJEBORGIIDAE

FIGURE 51

The two Hawaiian species of this family may be distinguished from other gammarideans by a combination

of characters: the thick antenna 1 (51H) with a very

stout and well developed primary and secondary

flagella; the presence of a tiny dorsal point on pleonites

1-2 (microscopically composed of 3 points each); thin

or stout but characteristically shaped uropod 3

(51Hu3), large black eyes; and large gnathopods 1-2

(51H and Hg2), these being similar to each other and

bearing long carpal processes like those of gnathopod

2 in leucothoids. But the palms of Liljeborgia have a

complex 2 or 3 cycles of densely crowded, short, medium and long spines or setae, one cycle of spines often

being composed of hooks. See Eusiroides in the Eusiridae (page 66), and Parapleustes in the Pleustidae

(page 112).

Key to Hawaiian Species of Genus Liljeborgia Bate

Figure Numbers Referenced in Italics

Pereopods 1-2 with minute palms on article 6 and several stout distal spines (51H), dactyls of

gnathopods not flagellate, uropod 3 of medium stoutness, pleonal epimeron 3 with a simple

posteroventral tooth (51H), mandibular spine row with numerous long articulate spines, palp

article 3 of mandible shorter than article 1 L. hema

Pereopods 1-2 simple and article 6 with numerous short posterodistal setae (51Lpl), dactyls of

gnathopods flagellate (51Lgl), uropod 3 slender, pleonal epimeron 3 with small sinus above

posteroventral tooth (51Lep3), mandibular spine row with only 3 tiny sharp inarticulate

serrations (51 Lm), mandibular palp article 3 slightly longer than article 1 L. laniloa
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hg2

FIGURE 51.—Liljeborgiidae: H, Liljeborgia heeia J. L. Barnard; L, L. laniloa J. L. Barnard.
NUMBER 5 8

Liljeborgia heeia J.L.Barnard

Liljeborgia heeia J. L. Barnard, 1970:213-216, figs. 139, 140.

This species may be recognized without dissection by

observing on slide 2 the minute palms of pereopods

1-2.

Oahu, 2-30 m.

Liljeborgia laniloa J. L. Barnard

Liljeborgia laniloa J. L. Barnard, 1970:216, fig. 141.

The pereopods of this species are normal and like most

species of Liljeborgia in other seas; hence to guarantee

a solid identification, in the event other undiscovered

liljeborgias occur in Hawaii, one must confirm the

identification by observing the other characters mentioned in the key.

Intertidal, northeast Oahu, living on coral block

covered with ascidians.

Family LYSIANASSIDAE

This family has been inadequately collected in Hawaii

and should yield several more than the one species now

known. The mitten-shaped article 6 of gnathopod 2,

the presence of dense brushes of short stiff setae on

articles 5-6, the frequent occurrence of a "pineapple

texture" (term invented by the Russian Gurjanova)

on article 5, and the elongate article 3 of gnathopod 2

are unmistakeable characters of this family. Antenna 1

is short and stout and usually has a large accessory

flagellum; articles 2-3 of the peduncle are very short

and partially telescoped. The body is plain, heavy, hard

and shiny. Mouthparts and uropod 3 are highly diverse

as numerous species are ectoparasites.

Genus Lysianassa Milne Edwards

Lysianassa ewa J. L. Barnard

FIGURE 52

Lysianassa ewa J. L. Barnard, 1970:216-220, figs. 142,143.

A short diagnosis will serve to distinguish at least this

this genus from others expected: accessory flagellum

2-3 articulate; upper lip produced as vertical, rounded

lamella (seen from lateral view); mandibular palp

105

attached proximal to weak spiny or setulose molar;

maxillae and maxillipeds normally gammaridean but

inner plate of maxilla 1 non-setose on medial edge;

coxa 1 as large as coxa 2; gnathopod 1 simple but

slightly stoutened; inner ramus of uropod 2 with deep

notch; uropod 3 biramous and inner ramus usually

well developed, but peduncle of L. ewa expanded into

a laterally flat plate, rami short and uniarticulate;

telson entire or weakly excavate; pereopods simple.

Eyes are brown in formaldehyde and spirit.

Oahu, 4-18 m.

Family OCHLESIDAE

Genus Ochlesis Stebbing

Ochlesis alii J. L. Barnard

FIGURE 53

Ochlesis alii J. L. Barnard, 1970: 220-223, figs. 144,145.

Unmistakeably recognized by its general body plan and

coxae (see figure), this monogeneric family has affinities with Acanthonotozomatidae, in which some of the

coxae are distinctly pointed. Possibly members of the

latter family occur in Hawaii but have not been found;

they may occur as ectoparasites on sessile invertebrates

or motile crustaceans. Ochlesis also is probably such

an ectoparasite but its host has not been determined.

Its mouthparts are of the piercing and sucking kind

and the maxillipedal palps are absent, the only known

shallow-water genus of its kind in the suborder. Eyes

orange in formaldehyde.

Southern Oahu, 18 m, on Pocillopora reef. Rare.

Family OEDICEROTIDAE

Genus Kanaloa J. L. Barnard

Kanaloa manoa J. L. Barnard

FIGURE 54

Kanaloa manoa J. L. Barnard, 1970:224-226, figs. 146, 147.

Like all oedicerotids, Kanaloa, an endemic Hawaiian

genus, differs from other Hawaiian gammarideans in

the following combination of characters: (1) Uropod

3 peduncle elongate as in Leucothoidae, Amphilochidae; (2) Pereopod 5 much longer than pereopods 3-4;
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ur

FIGURE 52.—Lysianassidae: Lysianassa ewa J. L. Barnard.
NUMBER 5 8 107

FIGURE 53.—Ochlesidae: Ochlesis alii J. L. Barnard.
108

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

xp

FIGURE 54.—Oedicerotidae: Kanaloa manoa J. L. Barnard.
NUMBER 5 8 109

(3) Head massive, nearly as long as pereonites 1-3

together, and often galeate (54, body); (4) Accessory

flagellum absent (or extremely minute vestige present);

(5) Eyes coalesced together or contiguously so close

together as to appear coalesced from dorsal view.

Kanaloa has large yellow-orange eyes in formaldehyde, fading in spirit; an enlarged pereonite 7; and a

characteristically spiny gnathopod 2 (54g2), both pairs

of gnathopods being feeble. If the synopiid genus

Synopia also occurs in Hawaii it could be confused

with Kanaloa, but Synopia has a short peduncle of

uropod 3 and bears a macroscopic accessory flagellum.

Oahu, (?2)-30 m. Affinitives are with antiborealantarctic genera.

Family PHLIANTIDAE

Genus Palinnotus Stebbing

Palinnotus alaniphlias J. L. Barnard

FIGURE 55

Palinnotus alaniphlias J. L. Barnard, 1970:226-227, figs.

148-150.

Adult phliantids are immediately recognized by the

broad, rugose, dome-shaped or boat-shaped bodies resembling an abbreviated stegosaurid. The thorax is

dominant and the small abdomen flexed beneath the

thorax. The Hawaiian genus has only 2 urosomal segments visible dorsally; uropod 2 has only 1 ramus and

the third uropods are a small pair of flaps adhering

to the ventral surface of the telson, as if they were anal

flaps. The rostrum is obsolescent, mandibles lack palps,

maxilla 1 has a small palp and the maxilliped has only

3 palp articles. Gnathopods 1 and 2 are simple and

resemble pereopods; the medial surface of gnathopod

1 has numerous prickles and asparagoid spines.

Juveniles may resemble eophliantids in the large

head and semicylindrical bodies but the urosome

remains diagnostic for Palinnotus.

Gut often filled with a black streak of presumed

lignin material; phliantids probably feed on tough,

almost woody stipes of algae. Body ochraceous-orange

in formaldehydej eyes black.

Oahu, intertidal, among algae.

Family PHOXOCEPHALIDAE

Presumably soft bottoms of Hawaii, when explored

adequately, will yield more than the one species now

known. Phoxocephalids have a visor-like rostrum,

elongate pereopod 4, and short pereopod 5 with broad

article 2; pereopod 3 and the antennae are strongly

setose or spinose; eyes when present are ommatidial

(rather than corneal as in Ampeliscidae) and are

black. The near-related Platyischnopus of Haustoriidae

has a rostrum like an ordinary shark nose and very

elongate pereopods 4-5; this genus is also expected to

occur in Hawaii. Both families are comprised of species

forming burrows in soft sediment.

Genus Paraphoxus Sars

Paraphoxus centralis (Schellenberg)

FIGURE 5P

Pontharpinia centralis Schellenberg, 1938:15-17, fig. 8.

"Fern Island"; "Whale Island."

Family PLEUSTIDAE

Pleustidae, close to Liljeborgiidae and Eusiridae ( =

Calliopiidae), are non-tropical. Apparently these are

the first tropical pleustids to be described. One is highly

aberrant and easy to distinguish from Liljeborgia in

the clear or red eyes lacking black pigment, the uncleft

telson, the large rostrum, very small gnathopods with

slender articles and obsolescent palms, and the absence

of a large accessory flagellum. The other species is

grossly distinguished only in the uncleft telson, the

absence of strong teeth on the gnathopodal dactyls,

the vestigial accessory flagellum, and the absence of

the faint dorsal teeth on pleonites 1-2 seen in

Liljeborgia.

The two Hawaiian pleustids resemble Liljeborgia

also in the non-triturative molars. The lower lip of

pleustids is formed of tilted oval outer lobes astride

partially coalesced apically truncate inner lobes forming a broad gap between the outer lobes. This character differentiates pleustids from calliopiids but not

distinctly from Liljeborgia, which has a weakly similar

lower lip. The outer rami of pleustid uropods are

shorter than the inner, either slightly or strongly, and

they are usually strongly spinose like those of podocerids. The mandibular palp is large, with heavily

developed spine-setae on a subclavate or subfalcate

article 3.

383-197 O - 71 - 8
FIGURE 55.—Phliantidae: Palinnotus alaniphlias J. L. Barnard.
NUMBER 5 8 111

FIGURE 56.—Pleustidae: D, Parapleustes derzhavini makiki J. L. Barnard; H, P. honomu J. L.

Barnard.
112

Genus Parapleustes Buchholz

FIGURES 56, 57

Parapleustes derzhavini makiki J. L. Barnard

Parapleustes derzhavini makiki]. L. Barnard, 1970:227-230,

figs. 151, 152.

The maxillipedal palp is normally 4-articulate, article

4 being long and unguiform; pereonites and pleonites

lack dorsal carinae or teeth; pleonal epimera 1-3 with

convex posterior margins, 1-2 with lateral ridge, 1 and

3 with small posteroventral cusp, 2 with longer sharply

extended tooth, ventral margins of 1 and 3 spinose and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ventrolateral face of 2 spinose; pereopods 1-5

of normal stoutness, gnathopods 1-2 of medium size

and similar to each other, article 2 of gnathopod 1

strongly setose anteriorly (setae very large but few in

number), article 4 of gnathopod 2 with sharp posterodistal process (seen only under high power), fifth

articles shorter than sixth, posteriorly lobate, sixth

articles expanded but not as strongly as in Liljeborgia,

palms defined by group of spines and bearing another

palmar spine set and a small cusp in the middle, but

dactyls not deeply serrate; eyes formed of dark brownish purple cores surrounded by clear ommatidia.

Oahu, 3-4 m.

FIGURE 57.—Pleustidae: D, Parapleustes derzhavini makiki J. L. Barnard; H, Parapleustes (?)

honomu J. L. Barnard.
NUMBER 5 8

Parapleustes (?) honomu J. L. Barnard

Parapleustes (?) honomu J. L. Barnard, 1970:233, figs. 153,

154.

The maxillipedal palp lacks a fourth article, heretofore not characteristic of pleustids; pleonites 1-2 have

strong dorsolateral carinae in addition to the dorsal

humps that are found also on pereonites 6-7, pleonite

3 with weak partial lateral carina and no dorsal hump,

epimeron 3 serrate posteriorly; pereopods 1-5 very

robust; antennae unknown but presumably typical of

other pleustids: eyes clear in spirit.

Oahu, off Moku Manu Island, 33 m, on base of

black coral in mollusk association.

113

Family PODOCERIDAE

FIGURES 58-60

Podocerids have the normal mouthparts of the basic

isaeid plus the fleshy telson, but the pereopods are not

glandular and the body of Hawaiian members is

broad and flattened, the coxae are small, the pleon

flexes strongly under the thorax, and urosomite 1 is

very elongate. Uropod 3 is formed of a pair of small

plates without rami below the telson. They are easily

overlooked. Podocerids in preservative often lose all

pereopods and antennae, but Hawaiian species may

be clearly distinguished by other characters even

though the most distinctive characters often occur in

missing parts.

Key to Genera of Podoceridae

Figure Numbers Referenced in Italics

Urosome (58L) with only 1 pair of long uropods ( = uropod 1), other uropods composed only

of small pair of flaps below telson, pereonite 1 elongate and grossly marked off from pereonite 2

by posterior sleeve-like constriction Laetmatophilus Bruzelius

Urosome (58T) with 2 pairs of long uropods (=uropods 1-2) and bearing also pair of uropodal

flaps below telson, pereonite 1 neither elongate nor strongly marked off from pereonite 2.

Podocerus Leach

Genus Laetmatophilus Bruzelius

Laetmatophilus hala J. L. Barnard

Laetmatophilus hala J. L. Barnard, 1970:233-237, fig. 155.

Body and head covered sparsely with blunt dorsal teeth,

or weak humps in juvenile, head with 3 teeth, pereonite

1 with 2 dorsal teeth in tandem and 2 dorsolateral

teeth in tandem on each side, pereonites 2-5 each with

1 dorsal and one lateral tooth on each side, in each

case primary hump middorsal; in addition, pereonites

1-7 with humps just above coxae; head strongly extended forward and upward, ocular lobes much posterior of anteroventral cephalic corner; coxae 3-4

broadly triangular and with obtuse ventral points;

gnathopod 1 with ovate articles 5-6, palm occupying

posterior margin of hand, dactyl nearly as long as

hand; article 2 of female gnathopod 2 (male unknown) with sharp anterodistal cusp, article 4 with

medium sized posterodistal process, article 5 unlobate,

article 6 narrow basally and broadly expanding distally, palm long and oblique, defined by slight acclivity, dactyl fitting palm.

Eyes red in formaldehyde, clear in alcohol.

Oahu, 18-30 m.

Genus Podocerus Leach

Uropods 1-2 are very long, with outer rami shorter

than inner but strongly spined, telson strongly elevated

dorsally, appearing from dorsal view to have a tumid

nob, bearing a pair of spines in two species and many

spines in a third species; pleonal epimera broadly

rounded below; pereonites 6-7 very large, elongate and

heavily muscularized in P. talegus lawai, slightly

shorter in P. hanapepe and scarcely different from anterior segments in P. brasiliensis. Epistome produced

conically only in P. brasiliensis and this species also

bears in the male an enormous medial flange projecting from the coxa above the insertion of article 2; the

flange bears ridges as if it were used for stridulation

and can be seen only by dissecting off gnathopod 2

with its coxal plate and viewing it medially.

NUMBER 5 8 115

Hgl

FIGURE 59.—Podoceridae: B, Podocerus brasiliensis (Dana); H, P. hanapepe J. L. Barnard; L,

Laetmatophilus hala J. L. Barnard; T, Podocerus talegus lawai J. L. Barnard.

FIGURE 58.—Podoceridae: B, Podocerus brasiliensis (Dana); H, Podocerus hanapepe J. L. Barnard ; L, Laetmatophilus hala J. L. Barnard; T, Podocerus talegus lawai J. L. Barnard.
116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 60.—Podoceridae: B, Podocerus brasiliensis (Dana); H, P. hanapepe J. L. Barnard; L,

Laetmatophilus hala J. L. Barnard; T, Podocerus talegus lavuai J. L. Barnard.
NUMBER 5 8 117

Key to Hawaiian Species of Genus Podocerus

Figure Numbers Referenced in Italics

1. Eyes occupying bulge forming anteroventral corners of head (58B), telson with 4-8 long

dorsal setal-spines (60Bt) P. brasiliensis

Eyes occupying bulges behind anteroventral corners of head (58H, T), those corners quadrate

or rounded, telson with 2 very large dorsal setal-spines (60Tt) 2

2. Palm of gnathopod 1 transverse, hand thus triangular and distally expanded (59Hgl).

P. hanapepe

Palm of gnathopod 1 oblique, hand thus ovate (59Tgl) P. talegus lawai

Podocerus brasiliensis (Dana)

Platophium brasiliense Dana, 1853:838-839, pi. 55, fig. 9.

Platophium synaptochir Walker, 1904:296-297, pi. 8, fig. 52.

Podocerus brasiliensis.—Schellenberg, 1938:94.—J. L. Barnard 1953:87 (with references); 1955:39; 1970:237-240,

figs. 156, 157.

Hand of male gnathopod 2 extremely elongate and

large, palm occupying full posterior margin of hand

and densely setose, bearing small distal process hidden

by setae, undefined, dactyl about half as long as palm;

palm of gnathopod 1 tumidly ovate (like that of P.

talegus lawai), palm long, heavily setose and bearing

a few short spines at defining corner; article 2 of pereopods 1-2 long, strongly spinose, that of pereopods

3-5 stout, densely spinose on pereopods 3-4, that of

pereopod 3 weakly bilobate; accessory flagellum long

and thin but 1-articulate; body lacking dorsal processes.

Eyes clear or ochraceous in alcohol.

Female gnathopod 2 with very large triangular posterior extension on article 4, article 5 very small and

broadly cup-shaped, hand subcircular, palm long and

defined by 2 large spines, dactyl fitting palm; gnathopod 1 with long palmar setae but no short defining

spines.

Tropicopolitan. Oahu, mainly as fouling organism

in harbors. Possibly introduced by shipping.

Podocerus hanapepe J. L. Barnard

Podocerus hanapepe J. L. Barnard, 1970:240, figs, 158, 159.

Hand of male gnathopod 2 ovate, stout, bearing distinct palm defined by cusp and bearing 2 distal processes; article 2 of pereopods 1-2 very stout and short,

article 2 of pereopod 3 complexly lobate, anteroventral

corner of head quadrate and head strongly extended

forward, antennae short and accessory flagellum very

short but 1-articulate; body lacking dorsal ornaments.

Female unknown.

Eyes red in formaldehyde, ochraceous in alcohol.

Oahu, Waikiki Beach, intertidal, on Ulva, rare.

Podocerus talegus lawai J. L. Barnard

Podocerus talegus lawai J. L. Barnard, 1970:241-243, figs.

160, 161.

Hand of male gnathopod 2 very elongate, palm occupying full posterior margin of hand, dactyl about half as

long as palm, palm undefined but bearing 3 distal

processes, one low and minutely castellate, one rounded

and one truncatoconiform; processes often difficult to

see from marginal views and hand must be rotated and

viewed from dorsomedial position, processes rudimentary in young male; article 2 of pereopods 1-2 of

medium stoutness, article 2 of pereopod 3 simply

lobate, anteroventral corner of head slightly rounded,

head moderately extended forward, antennae long, accessory flagellum long and thick (but 1-articulate).

Eyes red in formaldehyde and ochraceous in alcohol.

Female gnathopod 2 with small article 4 bearing

weak cusp, hand stout, short, broadly ovate, palm long,

strongly defined by 2 spines and weak cusp, palm

minutely castellate.

Oahu, intertidal to 30 m.

Family SEBIDAE

Genus Seba Bate

FIGURE 61

Seba ekepuu J. L. Barnard

Seba ekepuu J. L. Barnard, 1970:247, figs. 162, 163.

The body is thin and the antennae short and stiff, and

thus sebids resemble the Colomastigidae but differ

from them in the larger coxae, the uniramous uropod 3

and the remarkably chelate gnathopods 1 and 2 (61 gl

and g2). Well developed mandibles and maxillae occur

in sebids whereas colomastigids have a palpless mandi-
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

xp
NUMBER 5 8 119

FIGURE 62.—Stenothoidae: A, Stenothoe sp. A; G, Stenothoe gallensis Walker; H, S. haleloke J. L.

Barnard; v, S. valida Dana.

FIGURE 61.—Sebidae: Seba ekepuu J. L. Barnard.
120

ble and spines replacing the incisor. The elongate

article 3 of gnathopod 2 in sebids recalls the Lysianassidae but gnathopod 2 of the latter is otherwise distinct in its mitten shape, strong, stiff fuzz and pineapple texture on article 5; article 2 of antenna 1 on

Lysianassidae is very short but is elongate in Sebidae.

The uniramous uropod 3 of sebids resembles that of

Stenothoidae but the short, even coxae distinguish

sebids from stenothoids. Eyes apparently are absent

although a dense granular tissue occupies this ocular

region in these tiny amphipods (1.0-1.5 mm long).

Oahu, Kawela Bay, intertidal, in algae.

Family STENOTHOIDAE

FIGURES 62, 63

This family is characterized by a uniramous uropod 3

composed of three segments: a peduncle and articles

1-2 of the ramus; coxa 1 is strongly reduced in size

and covered by a large coxa 2 while coxa 4 is even

more enlarged, shieldlike and lacks a posterior excavation. Uropod 3 distinguishes stenothoids from amphilochids but, in addition, the known stenothoids of

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hawaii, all in the genus Stenothoe, lack a mandibular

palp and the body of the mandible is characteristic

(see 63Gm). In numerous other stenothoid genera of

higher latitudes, the mandibular palp is very fragile

and easily broken off during dissection, so that great

care must be taken in this operation when checking

Hawaiian stenothoids for the possible occurrence of

those genera.

Only five species of Stenothoe are known from

Hawaii but more members of the family are expected

to be found. They are often associated with coelenterates, especially hydroids, and numerous possible niches

have not been explored. Stenothoe is the strongest

genus of the family in the tropics and no other may

occur in Hawaii but as a safeguard all stenothoids

should be checked against the following diagnosis of

Stenothoe:

Mandibular palp either absent or composed of 1

article; palp of first maxilla 2-articulate (often very

difficult to ascertain) ; pereopod 3 with slender, linear

article 2, while pereopods 4-5 with expanded, subovatc

article 2.

Eyes apparently red, fading rapidly in all Hawaiian

species.

Key to Hawaiian Species of Genus Stenothoe Dana

Figure Numbers Referenced in Italics

1. Dactyls of pereopods 1-5 with accessory claw (62Apl) Stenothoe sp. A

Dactyls of pereopods 1-5 simple 2

2. Peduncle (basal article) of uropod 3 shorter than next article (article 1 of ramus) (62Hu3).

S. haleloke

Peduncle of uropod 3 as long as or longer than next article 3

3. Pereopods 1-5 without spines on articles 4-6 Stenothoe sp. B

Pereopods 1-5 with spines on articles 4-6 4

4. Distal bifid process on palm of male gnathopod 2 with its branches extending equally (62Gg2),

fully adult (Hawaii only) individuals with coxa 2 unexcavate posteroventrally, coxa 3 with

anterior and posterior margins parallel, female gnathopod 2 palm unarmed. . . S. gallensis

Distal bifid process on palm of male gnathopod 2 with its branches extending unequally

(63Vg2), fully adult individuals with coxa 2 excavate posteroventrally, coxa 3 with anterior

and posterior margins divergent, especially in male, female gnathopod 2 palm with small

distal tooth (63Vg2 9) S. valida

Stenothoe gallensis Walker

Stenothoe gallensis Walker, 1904:261-262, pi. 3, fig. 19.—

J. L. Barnard, 1955:3-5 (with references).

Stenothoe crenulata Chevreux, 1908:471-475, figs. 1-3.

The taxonomy of this species and S. valida, though

often treated, is not yet clarified. So-called S. gallensis

and S. valida have been reported from around the

world but each local population seems to differ from

another and the two species, where they occur in close

proximity seem to shift to and fro morphologically in

many characters of presumed phenotypic content. In

Hawaii, those specimens that have been identified as S.

gallensis because of the structure of male gnathopod

2 (see key) differ from populations in the western

Indo-Pacific in the absence of a posteroventral excavation on coxa 2 in either sex and they therefore differ
NUMBER 5 8 121

FIGURE 63.—Stenothoidae: A, Stenothoe sp. A; o, 5. gallensis Walker; H, S. haleloke J. L. Barnard ; v, S. valida Dana.
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

from so-called adult Hawaiian S. valida in this character. But Hawaiian juvenile S. valida appears also to

have unexcavate coxa 2, so that juveniles of the two

species are inseparable. Until this difficulty and several

others not worth wordage in this brief report are better

understood, the surest way to identify the two species

is by terminal males, through their gnathopods (see

key) plus the subgeniculate and ridged terminal article

of uropod 3 in S. gallensis. That article in presumed

terminal males of S. valida is smooth, simple, and

straight. One has to presume that associated females

and juveniles are specifically identical with males in

the collections, but there may be potential error even

in this attitude. We may be dealing with a single species occurring in 2 phases, a gallensis phase with special

uropod 3 that represents phenotypes of a particular

season or econiche plus a normal valida phase.

At any rate, males of the two species are easily distinguished from other Hawaiian stenothoids in the

grossly elongate article 6 of gnathopod 2, with a densely

setose palm extending the full length, plus a long dactyl

also setose on its inner edge.

The Hawaiian S. gallensis is one of the finest tropical

members of the family, in that it is relatively large (3-5

mm) and compact, heavily chitinized, and preserves

exceptionally well for a stenothoid.

Oahu and Hawaii, primarily as fouling organism in

harbors probably associated with hydroids, possibly introduced by shipping, and probably circumtropical.

Stenothoe haleloke J. L. Barnard

Stenothoe haleloke J. L. Barnard, 1970:247-249, fig. 164.

A small red-eyed species with flecks of bluish purple

on the body in 2-day formaldehyde. Gnathopod 2 of

the male resembles that of female S. gallensis and S.

valida, but uropod 3 is diagnostic (see key). Perhaps

terminal males have not been accounted for.

Oahu, Waikiki Beach, intertidal, on Ulva.

Stenothoe valida Dana

Stenothoe validus Dana, 1853:924-925, pi. 63, figs. la-o.

Stenothoe valida.—J. L. Barnard, 1953:83-87 (with synonymy); ?197O:250-251, fig. 165.

Uropod 3 of both sexes is of the simple, straight, unornamented kind seen in females, but not males, of S.

gallensis (see the latter species, above, for remarks).

The Hawaiian population of this species has an unusually narrow article 4 on pereopods 3-5, a character

of possible subspecific value.

Oahu, off Moku Manu Island, 33 m, on base of

black coral in mollusk association.

Stenothoe sp. A, J. L. Barnard

Stenothoe sp. A, J. L. Barnard, 1970:251, fig. 166 a-i.

An undescribed species known only in juvenile form

but distinct from all other Hawaiian stenothoids in the

presence of accessory claws on the pereopodal dactyls

and the thickened and distally expanding sixth articles

of pereopods 1-5.

Southern Oahu, 18 m.

Stenothoe sp. B, J. L. Barnard

Stenothoe sp. B, J. L. Barnard, 1970:251, fig. 166 ;.

An undescribed species known only in juvenile form

but distinguished from all other Hawaiian stenothoids

in the absence of any spines on articles 4-6 of pereopods 1-5. There may be several undescribed Hawaiian

stenothoids in this condition as juveniles, so the category is probably of value only as a catch-all.

Oahu, Kaneohe Bay, 2 m.

Superfamily TALITROIDEA

Families HYALIDAE, HYALELLIDAE

FIGURES 64-68

This is the only portion of the Gammaridea so far ordered into a superfamily, and the Talitroidea are

herein treated as equivalent to other families.

The essentially or fully uniramous uropod 3, combined with a strongly triturative mandibular molar,

lacking a palp, and the absence of an accessory flagellum characterize Hawaiian talitroids. The regularly

quadrate anterior coxae and strong molar distinguish

talitroids from stenothoids; the latter also have a

strongly 2-articulate ramus on uropod 3 and weak lobes

on the maxillipeds.

Sebidae have a mandibular palp, nontriturative

molar, coalesced urosomites 2-3 (but Parhyalella of

Talitroidea might be confused in this regard), an accessory flagellum, and grossly chelate gnathopods.

Female brood lamellae of aquatic talitroids have

curl-tipped setae with apices appearing tangled.
NUMBER 5 8 123

FIGURE 64.—Talitroidea—Hyalidae and Hyalellidae: L, Hyale waitnea J. L. Barnard; P, Parhyalella pietschmanni Schellenberg.

Key to Aquatic Talitroidea

Figure Numbers Referenced in Italics

1. Uropod 3 with small scalelike inner ramus (68Hu3) Parkyale Stebbing

Uropod 3 lacking inner ramus 2

2. Telson cleft (68Ht and Lt), article 5 of male gnathopod 2 masked between articles 4 and 6,

unlobed (67Gg2)..'. HyaU Rathke

Telson entire (68Pt), article 5 of male gnathopod 2 bearing long lobe separating articles 4

and 6 (67Yg2) 3

3. Pleonites 1-2 with large dorsal tooth each, maxilla 1 with small palp.

Hyalella azteca (freshwater swamp)

Pleonites 1-2 smooth dorsally, maxilla 1 lacking palp Parhyalella Kunkel

Genus Hyale Rathke

One of the most commonly occurring genera in the

tropics and subtropics, Hyale is a genus of nestlers most

often found in the tropics with Sargassum. Hawaii has

at least 7 species and more probably will be found

when better collections are made. The several species

do not seem to be as numerically abundant as they are

on continents, as no collections made by J. L. Barnard

(1970) showed them dominating certain kinds of samples as they do in Panama and California.

Like most preserved amphipods the eyes slowly

change after initial preservation, so that collections of
124

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 65.—Talitroidea—Hyalidae and Hyalellidae: A, Hyale ayeli J. L. Barnard; o, H. grandicornis bishopae J. L. Barnard; H, H. honoluluensis Schellenberg; i, H. iole J. L. Barnard; L, H.

late J. L. Barnard; p, Parhyalella pietschmanni Schellenberg; w, H. (Lelehua) waimea J. L.

Barnard; Y, Parhyale hawaiensis (Dana).
NUMBER 5 8 J25

several years age may have the eyes resembling egg hands of male gnathopod 1 and female gnathopods

yolks. All but one Hawaiian species has black to purple i_2 a re very useful. One contrasts a pair (or 3) poseyes in alcohol (1 to 2 months). Pereopodal dactyls *»«;«„„„+„ • * j r /

. , , . • r i • J r • ii

 teno r seta e aSamst a spread of numerous setae (more

and locking spines are very useful in identifying all ,

 v

stages of most of the species, but females of 2 Hawaiian

 tha n 6) ' ° the r character s

 not mentioned but also of

species are grossly indistinguishable (see below). The

 dlagnostic value are the shape of uropod 3 (thick or

location of the major distal spine on the peduncle of thin) and the shape of telson from lateral view and

uropod 1 and the number of posterior setae on the from flattened dorsal view.

Key to Hawaiian Species of Genus Hyale

Figure Numbers Referenced in Italics

1. Eyes red in formaldehyde, clear in alcohol, palp article 4 of male maxilliped with long whiplike terminal seta (64Lxp) H. waimea

Eyes with purple or brown pigment in alcohol, palp article 4 of maxilliped normally setose

(64Pxp) 2

2. Article 6 of pereopods 1-5 with small or medium undifferentiated distal locking spine (68Ip5),

and medium sized mid-setule on dactyl, major peduncular spine of uropod 1 on mediodistal

margin, gnathopod 1 with numerous posterior setae on article 6 3

Article 6 of pereopods 1-5 with large striate distal locking spine (of corkscrew kind) (68Ep5)

weak or no distal setule on dactyl, major peduncular spine of uropod 1 on laterodistal

margin, gnathopod 1 with only 2 or 3 posterior setae on article 6 4

Article 6 of pereopods 1-2 with small or no poorly striate distal locking spine next to larger

striate spine (68Rp5) weak distal setule on dactyl, major peduncular spine of uropod 1 on

laterodistal margin, gnathopod 1 with numerous posterior setae 6

3. Antenna 2 in adult with dense brushes of setules (651h), article 5 of gnathopod 1 with broad

posterior lobe (661 gl), coxa 4 with weak posterior acclivity, ramus of uropod 3 with only

terminal spines (68Iu3) .• H. iole

Antenna 2 in adult lacking dense brushes of setules, article 5 of gnathopod 1 with narrow

posterior lobe (66Ggl), coxa 4 with sharp, attenuate posterior cusp, ramus of uropod 3

with one spine on side margin besides terminal spines (68Gu3)... .H. grandicornis bishopas

4. Palm of male gnathopod 1 defined by large cusp, palm of male gnathopod 2 with strong

distal process H. tot*

Palm of male gnathopod 1 bearing weak denning hump or none, palm of male gnathopod 2

simple 5

5. Palm of male gnathopod 2 distinct from posterior margin of hand (67F), posterior coxal

acclivities (coxae 1-3) close to ventral margins H. affinis

Palm of male gnathopod 2 confluent with posterior margin of hand and dactyl extending its

full length (67H), posterior coxal acclivities near middle of posterior margins.

H. honoluluensis

6. Outer rami of uropods 1-2 lacking spines on dorsal margin, antenna 2 with small but distinct

brushes of posterior setules, coxae 1-3 with sharply quadrate, slightly attenuate posterior

acclivities (66Igl) H- «>«'«

Outer rami of uropods 1-2 with spines on dorsal margins, antenna 2 lacking posterior setular

brushes, coxae 1-3 with posterior acclivities obsolescent and softly rounded (66Ygl).

Hyale sp.

Hyale affinis Chevreux Male gnathopod 1 with expanded hand, anterodisu i * • m. iono KTM tnr r 01 00 c i_ 1 tally tumid, palm oblique and bearing large spine, deHyale affinis Chevreux, 1908:503-506, figs. 21-22.—Schel- ' r

 , .

lenberg, 1938:67-68.—J. L. Barnard, 1955:14 (in part, fining corner softly rounded, setose, dactyl reaching

not fig. 6); 1970:251-255, fig. 167 e-h. ma i n spine, posterior margin of hand with 2-3 setae

Antenna 2 simple.

 i n l clum P5 sPin

y P a l m o f mal e gnathopod 2 deeply

Coxae 1-4 with quadrate posterior acclivities es- recessed from short posterior margin of hand, latter

pecially close to ventral margins. Coxa 4 with strongly bearing 1 spinule, palm defined by 2 spines.

sinuous excavation. Female gnathopods 1-2 with article 6 slender, rec383-197 O - 71 - 9
FIGURE 66.—Talitroidea—Hyalidae and Hyalellidae; gnathopod 1: A, Hyale ayeli J. L. Barnard;

T, H. affinis Chevreux; o, H. grandicornis bishopae J. L. Barnard; H, H. honoluluensis Schellenberg; i, H. iole J. L. Barnard; L, H. laie J. L. Barnard; P, Parhyalella pietschmanni Schellenberg;

R, H. rubra (Thomson); w, H. {Lelehua) waimea J. L. Barnard; v, Parhyale hawaiensis (Dana).
NUMBER 5 8 127

tangular, palm oblique, simple, posterior margin of

hand with pair of setae.

Pereopodal locking spines formed of 1 large and 1

small spirally striate members in tandem, dactyl with

small distal setule and microcastellations.

Uropod 1 with major peduncular spine distolateral.

Ramus on uropod 3 with only terminal spines. Eyes

purple in alcohol.

Southern Polynesia; Laysan; Kauai; Oahu, intertidal.

Hyale ayeli J. L. Barnard

Hyale ayeli J. L. Barnard, 1955:14-15, fig. 7; 1970:255,

figs. 168, 169.

Male and female antenna 2 with weak to strong

posterior setal brushes on peduncle and base of

flagellum.

Coxae 1-3 with sharply quadrate, slightly attenuate

posterior acclivities; coxa 4 with simple excavation.

Male gnathopod 1 with narrow rectangular hand,

palm oblique and defined by spines at rounded defining

corner, dactyl fitting palm, posterior margin of hand

with wide spread of numerous setae; spiny palm of

male gnathopod 2 oblique, equal to posterior margin of

hand, weakly defined by softly rounded defining corner,

dactyl fitting palm, in terminal male palm with weak

distal quadrate protrusion, absent in young male.

Female gnathopods 1-2 with article 6 slender, rectangular (like male gnathopod 1), posterior margins

of hand with wide spread of numerous setae.

Pereopodal locking spines differing on pereopods

1-2 and 3-5; on pereopods 1-2 with large distal spirally

striate (weakly) single spine, in pereopods 3-5 with

largest spine strongly striate spirally and bearing a

more distal small, axially-spirally striate spine; dactyl

with small distal seta.

Uropod 1 with major peduncular spine distolateral.

Uropods 1-2 with outer rami lacking dorsal spines.

Ramus of uropod 3 with only terminal spines. Eyes

brown-black.

Oahu, intertidal.

Hyale grandicornis bishopae J. L. Barnard

Hyale bishopae J. L. Barnard, 1955:16-17, fig. 8.

Hyale grandicornis bishopae J. L. Barnard, 1970:257, fig.

170.

Antenna 2 stout but simple.

Coxae 1-3 with well developed, slightly to strongly

attenuate posterior acclivities. Coxa 4 with simple

excavation.

Male gnathopod 1 with stout rectangular hand, palm

oblique and short, defined by 2 spines at defining

corner, posterior margin of hand with spread of several setae; spiny palm of male gnathopod 2 oblique

and equal to posterior margin of hand, weakly defined

by 2 spines.

Female gnathopods 1-2 with article 6 stout, rectangular (like male gnathopod 1), posterior margin

of hand with spread of several setae.

Pereopodal locking spines small and simple, dactyl

with large middle seta.

Uropod 1 with major peduncular spine mediodistal

but scarcely enlarged.

Ramus of uropod 3 with at least one subterminal

dorsal spine disjunct from terminal set. Eyes

brown-black.

Oahu, intertidal.

Hyale honoluluensis Schellenberg

Hyale honoluluensis Schellenberg, 1938:69-71, figs. 35b, c.—

J. L. Barnard, 1955:18; 1965:520, figs. 22, 23; 1970:

257-259, figs. I67a-d, 171.

Antenna 2 simple.

Coxae 1-3 with a strong quadrate posterior acclivity.

Coxa 4 with sinuous excavation.

Male gnathopod 1 with expanded hand, anterodistal

margin tumid, palm oblique, bearing striate spine,

softly rounded defining corner with 2 spines, dactyl

not reaching main spine; spiny palm of terminal male

gnathopod 2 occupying full posterior margin of hand,

dactyl very long and overlapping free margin of hand,

younger male with shorter palm defined by 2 spines

but palm not recessed and lacking posterior spinule

seen in H. affinis.

Female gnathopods 1-2 with article 6 slender, rectangular, palm oblique, simple, posterior margin of

hand with pair of setae.

Pereopodal locking spines composed of 1 large and

1 small spirally striate members in tandem, dactyl with

small distal seta and microcastellations.

Uropod 1 with major peduncular spine distolateral.

Ramus of uropod 3 with only terminal spines.

Females of this species and H. laie are grossly

indistinguishable.
128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Wur
FIGURE 68.—Talitroidea—Hyalidae and Hyalellidae: A, Hyale ayeli J. L. Barnard; E, H. laie

J. L. Barnard; o, H. grandicornis bishopae J. L. Barnard; H, Parhyale hawaiensis (Dana) ; i,

Hyale iole J. L. Barnard; L, H. waimea J. L. Barnard; p, Parhyalella pietschmanni Schellenberg;

R, Hyale rubra (Thomson).

FIGURE 67.—Talitroidea—Hyalidae and Hyalellidae; gnathopod 2: A, Hyale ayeli J. L. Barnard;

F, H. affinis Chevreux; G, H. grandicornis bishopae J. L. Barnard; H, H. honoluluensis Schellenberg; i, H. iole J. L. Barnard; L, H. laie J. L. Barnard; R, H. rubra (Thomson) ; w, H. (Lelehua)

waimea J. L. Barnard (also urosome); Y, Parhyale hawaiensis (Dana).
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Eyes garnet-ruby in formaldehyde, dark purple in

alcohol.

Indo-Pacific; Hawaiian Islands, shallow water.

Hyale iole J. L. Barnard

Hyale iole J. L. Barnard, 1970:260, fig. 172.

Male and female antenna 2 with extremely thick posterior setal brushes on peduncle and base of flagellum.

Coxae 1-3 with sharp and attenuate posterior acclivities. Coxa 4 with simple excavation.

Male gnathopod 1 with rectangular hand, palm

oblique, short, defined by large spine at rounded defining corner, posterior margin of hand with long

spread of numerous setae; spiny palm of male gnathopod 2 oblique, equal in length to posterior margin of

hand, weakly defined by softly rounded corner.

Female gnathopods 1-2 with article 6 slender, rectangular (like male gnathopod 1), posterior margin

with medium spread of posterior setae.

Pereopodal locking spines formed of 1 medium and

1 small axially striate spines in tandem, dactyl with

large seta in middle.

Uropod 1 with major peduncular spine mediodistal

and very large.

Ramus of uropod 3 with only terminal spines. Eyes

brown-black.

Maui, Kahalui Harbor.

Hyale late J. L. Barnard

Hyale laie J. L. Barnard, 1970:260-264, figs. 173, 174.

Antenna 2 simple.

Coxae 1-3 with strongly quadrate posterior acclivity.

Coxa 4 with sinuous excavation.

Male gnathopod 1 with expanded hand, anterodistally tumid, palm oblique and defined by large triangular cusp, dactyl fitting palm, spiny palm of male

gnathopod 2 oblique, defined from short posterior

margin of hand by hump and pair of spines, bearing

large blunt distal process, dactyl fitting palm, hand

with 2 anterior spines.

Female gnathopods 1-2 with article 6 slender, rectangular, palm oblique, simple, posterior margin of

hand with pair of setae.

Pereopodal locking spines composed of 1 large and

1 small spirally striate members in tandem, dactyl with

small distal seta and microcastellations.

Uropod 1 with major peduncular spine distolateral.

Ramus of uropod 3 with only terminal spines. Eyes

brown-black.

Oahu and Kauai, intertidal.

Hyale sp. (cf. rubra Thomson)

Hyale sp. (cf. rubra Thomson) J. L. Barnard, 1970:268-269,

fig. 178.

Antenna 2 simple.

Coxae 1-3 with posterior acclivity obsolescent and

softly rounded. Coxa 4 with slightly sinuous excavation.

Male gnathopod 1 with thin, tapering rectangular

hand, palm very oblique and short, defined by 2 spines,

dactyl short and fitting palm, posterior margin of hand

with wide spread of numerous setae; spiny palm of

male gnathopod 2 oblique and equal to posterior margin of hand, weakly defined by spines, dactvl fitting

palm.

Female unknown.

Pereopodal locking spines composed of 1 small,

weakly striate distal and 1 large spirally striate member in tandem, dactyl with small distal seta.

Uropod 1 with major peduncular spine distolateral.

Ramus of uropod 3 with only terminal spines. Eyes

damaged and color unclear.

Maui, Kahalui Harbor.

Hyale (Lelehua) waimea J. L. Barnard

Hyale(Lelehua) waimea J. L. Barnard, 1970:264-268, figs.

175-177.

This species is the only member of Hawaiian Hyale

with colorless eyes in alcohol; the others retaining some

purple or brown color in that medium. The subgenus is

distinguished in the male by the presence of a very

long terminal seta on palp article 4 of the maxilliped.

Coxa 4 has only a small dorsal point in the posterodorsal excavation and otherwise coxae 1-3 have essentially no posterior acclivities or cusps. Gnathopod 1 has

many posterior setae on the narrow, rectangular article

6; male gnathopod 2 has a deep hemispherical incision

on the palm and a hump on the inner edge of the

dactyl. Pereopodal dactyls have a medium distal setule

on the inner edge, a facial setule, and a thin sharp

spine-tooth appressed to the dactylar terminus; the distal locking spine is weakly striate and small and is

adjacent to a larger smoother spine. Pereopod 3 has a

widely inflated nasiform article 4 with posterior spines,
NUMBER 5 8 131

and articles 4—5 of male pereopod 4 are like the normal

pereopod 5 of the male. The main peduncular spine

of uropod 1 occurs laterally, the ramus of uropod 3 is

unusually short and bears only terminal spines, and

the telson is uniquely narrow. Female gnathopod 2 is

similar to gnathopod 1 but much less setose.

Southern Oahu, 30 m.

Genus Parhyale Stebbing

Parhyale hawaiensis (Dana)

Parhyale inyacka (K. H. Barnard).—J. L. Barnard, 1955:23,

fig. 12 (with references).

Parhyale hawaiensis (Dana).—Shoemaker, 1956:351-356,

figs. 3-4 (with references).—J. L. Barnard, 1965:521—

523, fig. 24.

Among the aquatic talitroids this species differs from

all others in the presence of a small scale representing

the inner ramus of uropod 3. The uropod is dissected

and mounted dorsal side up so it can be viewed with

a compound microscope. Eyes black, turning deep

purple in alcohol.

Circumtropical in shallow waters.

Genus Parhyalella Kunkel

Parhyalella pietschmanni Schellenberg

Parhyalella pietschmanni Schellenberg, 1938:71-74, figs. 36,

37.—J. L. Barnard, 1970:269, figs. 179, 180.

A large species, up to 11 mm in length, P. pietschmanni

is also one of the most beautiful Hawaiian amphipods,

and like the Hawaiian beach-hopper Orchestia platensis Kroyer, would form excellent introductory laboratory material in a study of amphipods. The animal

preserves well and is easy to dissect.

The adult male has very stout peduncles on antenna

2 but those of the female are thin. The eyes in alcohol

are dark purple throughout but garnet in formaldehyde. Urosomite 2 is dorsally telescoped into urosomite 1 and does not freely reach the dorsal body

margin. Urosomite 3 is shaped like the prow of a ship,

and uropod 3 projects from it like a bowsprit.

Oahu, beach wrack.

Literature Cited

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1936. Some New and Little Known Amphipods of California. University of California Publications in

Zoology, 41:53-74, figures 1-51.

Barnard, J. L.

1953. On Two New Amphipod Records from Los Angeles Harbor. Bulletin Southern California Academy of Sciences, 52:83-87, plate 15.

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1959. 3. Generic Partition in the Amphipod Family £Iheluridae, Marine Wood Borers. Pacific Naturalist, 1

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1916. Contributions to the Crustacean Fauna of South

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1862. Catalogue of the Specimens of Amphipodous Crustacea in the Collection of the British Museum,

London, iv+399, plates 1, la, 2-58.

.Caiman, W. T.

1910. On Two New Species of Wood-Boring Crustacea

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Chevreux, E.

1901. Crustac6s amphipodes. Mission scientifique de M.

Ch. Alluaud aux lies Sechelles (Mars, Avril, Mai,

1892). Memoires de la Sociite Zoologique de

France, 14:388-438, figures 1-65.

1908. Amphipodes recueillis dans les possessions francaises de POceanie par M. le Dr. Seurat, Directeur

du Laboratorie de Recherches Biologiques de Rikitea (lies Gambier), 1902-1904. Memoires de la

Sociite Zoologique de France, 20:470-527, figures

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Chevreux, E., and L. Fage

1925. Amphipodes. Faune de France, 9:1-488, figures

1-438.

Crawford, G. I.

1937. A Review of the Amphipod Genus Corophium,

with Notes on the British Species. Journal of the

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Dana, J. D.

1853-55. Crustacea. Part II. United States Exploring Expedition, 14:689-1618, atlas of 96 plates.

Edmondson, C. H.

1951. Some Central Pacific Crustaceans. Occasional

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Pillai, N. K.

1957. Pelagic Crustacea of Travancore, 3: Amphipoda.

Bulletin of the Central Research Institute, University of Travancore, 5:29—68, figures 1—18.

Pirlot, J. M.

1936. Les Amphipodes de l'expedition du Siboga.

Deuxieme partie: .. . II : .. . 3: .. . Ill : . . .

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146.

Sars, G. O.

1895. Amphipoda. An Account of the Crustacea of Norway with Short Descriptions and Figures of All

the Species, 1 :i—viii-f-1-711, plates 1-240, supplementary plates 1—8.

Schellenberg, A.

1938. Litorale Amphipoden des Tropischen Pazifiks.

Kongliga Svenska Vetenskapsakademiens Handlingar, 16:1-105, figures 1-48.

Sexton, E. W. and D. M. Reid

1951. The Life-history of the Multiform Species Jassa

falcata (Montagu) (Crustacea Amphipoda) with

a Review of the Bibliography of the Species.

Journal of the Linnean Society of London, Zoology, 42:29-91, plates 4-30.

Shoemaker, C. R.

1925. The Amphipoda Collected by the United States

Fisheries Steamer Albatross in 1911, Chiefly in the

Gulf of California. Bulletin of the American Museum of Natural History, 52:21-61, figures 1-26.

1933a. Two New Genera and Six New Species of Amphipoda from Tortugas. Carnegie Institution of Washington, Papers, 28:245-256, figures 1-8

(Publication 435).

1933b. Amphipoda from Florida and the West Indies.

American Museum Novitates, 598:1-24, figures

1-13.

1934. Two New Species of Corophium from the West

Coast of America. Journal of the Washington

Academy of Sciences, 24:356-360, figures 1-2.

1942. Amphipod Crustaceans Collected on the Presidential Cruise of 1938. Smithsonian Miscellaneous

Collections, 101 (11): 1-52, figures 1-17.

1947. Further Notes on the Amphipod Genus Corophium

from the East Coast of America. Journal of the

Washington Academy of Sciences, 37:47-63, figures 1-12.

1949. The Amphipod Genus Corophium on the West

Coast of America. Journal of the Washington Academy of Sciences, 39:66-82, figures 1-8.

1956. Observations on the Amphipod Genus Parhyale.

Proceedings of the United States National Museum, 106:345-358, figures 1-4.

Stebbing, T. R. R.

1888. Report on the Amphipoda Collected by H.M.S.

Challenger During the Years 1873-76. In Great

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Index

Main page references to Hawaiian genera and species are entered in boldface.

Omitted are subjects in tables, lists, keys, and figures, and in the discussion of uropod

3 on pages 18-19.

abditus, Ericthonius, 61

acherusicum, Corophium, 59

?aequimanus, Lembos, 50, 51

afer, Gammaropsis, 91

affinis, Hyale, 125, 127

afra, Eurystheus, 91

Gammaropsis, 91

aina, Photis, 96

alamoana, Gammaropsis, 90, 91

alaniphlias, Palinnotus, 109

alii, Ochlesis, 105

Aloiloi, 41

nenue, 41

akuolaka, Ampithoe, 35

amikai, Nuuanu, 85

Ampelisca, 22

schellenbergi, 22

Amphilochus, 14, 16, 28

kailua, 28

likelike, 28

marionis, 25

menehune, 28

Ampithoe, 2, 6, 34, 36, 37

akuolaka, 35

kaneohe, 29, 34, 35, 36

orientalis, 34, 35, 37

poipu, 34, 36

ramondi, 36

waialua, 37

Ampithoe sp. 35, 37

Anamixis, 40, 102

stebbingi, 40

angularis, Parajassa, 52, 97, 99

Antipathes, 3

Aora, 2, 48

Aoroides, 2, 41

californica, 41

columbiae, 41, 49

nahili, 41

appendiculata, Melita, 70, 85

atlantica, Gammaropsis, 91

atlanticus, Eurystheus, 91

Atylus, 61

nani, 61

aviculae, Chevalia, 88

ayeli, Hyale, 127

Pbaconi, Corophium, 59

Bemlos macromanus, 51

Biancolina, 52

mauihina, 52, 97

bishopae, Hyale grandicornis, 127

black-coral, see Antipathes

brasiliense, Platophium, 117

brasiliensis, Ericthonius, 16, 61

Podocerus, 113, 117

Byblis, 2, 16

californica, Aoroides, 48

,Calliactis, 74

calliactis, Elasmopus, 3, 14, 16, 70, 74,

75

centralis, Paraphoxus, 109

Pontharpinia, 109

Ceradocus, 2, 12, 68, 70, 77

hawaiensis, 70, 77, 85

Cheiriphotis, 88

Chelura insulae, 54

Chevalia, 88

aviculae, 22, 88

Colomastix, 16, 54

kapiolani, 55

lunalilo, 55

pusilla, 55

Colomastix sp., 55

columbiae, Aoroides, 41,49

Corophium, 2, 14, 59

acherusicum, 59

Pbaconi, 59

insidiosum, 59

crenulata, Stenothoe, 120

Cymadusa, 37

filosa, 37, 38

hawaiensis, 38

oceanica, 38

Cyproideinae, 29

Dardanus, 3, 74

derzhavini makiki, Parapleustes, 112

diplonyx, Elasmopus, 74, 85

Eusiroides, 66

Dolium, 3, 74

ecuadorensis, hawaiensis, Elasmopus,

74, 75

ekepuu, Seba, 117

Elasmopus, 2, 68, 70, 77

calliactis, 3, 14, 16, 70, 74, 75

Pdiplonyx, 74, 85

ecuadorensis, hawaiensis, 74, 75

hooheno, 74

insignis, 77

molokai, 75

pectenicrus, 70, 74, 75, 85

piikoi, 75

pocillimanus, 75

rapax, 70, 74, 75, 76

serrula, 75

spinidactylus, 76

Eophliantidae, 21

Ericthonius, 61

abditus, 61

brasiliensis, 61

Eriopisa, 68, 77, 85

hamakua, 76

laakona, 17, 76

Eriopisella, 68, 76, 77, 85

sechellensis upolu, 76

Eurystheus, 2, 89

afra, 91

atlanticus, 91

ventosa, 99

Eusiroides, 66, 103

diploynx, 66

ewa, Lysianassa, 105

falcata, Jassa, 97

filosa, Cymadusa, 37, 38

fresneli, Melita, 85

gallensis, Stenothoe, 120, 122

Gammaropsis, 2, 41, 89, 90

afer, 91

afra, 91

alamoana, 90,91

atlantica, 91

Gardineri, 91

haleiwa, 89, 90,91

kaumaka, 90, 93

pali, 41,89, 93

pokipoki, 89, 93

zeylanicus, 91

Gammarus quadrimanus, 84

Gardineri, Gammaropsis, 91

133
134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gitana, 25

liliuokalaniae, 25

Gitanopsis, 12, 28

pele, 28

grandicornis bishopae, Hyale, 127

Grubia hawaiensis, 38

hala, Lactmatophilus, 16, 113

haleiwa, Gammaropsis, 89, 90,91

haleloke, Stenothoe, 122

hamakua, Eriopisa, 76

hamigera, Maera, 84

hanapepe, Podocerus, 113, 117

hawaiensis, jCeradocus, 70, 77, 85

Cymadusa, 38

Elasmopus ecuadorcnsis, 74, 75

Grubia, 38

Parhyale, 131

Photis, 96

heeia, Liljeborgia, 105

honoluluensis, Hyale, 127

honomu, Parapleustes (?), 113

hooheno, Elasmopus, 74

Hyale, 2, 6, 123

affinis, 125, 127

ayeU, 127

bishopae, 127

grandicornis bishopae, 127

honoluluensis, 127

iole, 130

laie, 127, 130

rubra (see Hyale sp.)

(Lelehua) waimea, 130

Hyale sp. (cf. rubra), 130

hyhelia, Leucothoe, 103

inaequipes serrata, Maera, 84

insidiosum, Corophium, 59

insignis, Elasmopus, 77

Maera, 70, 77

insulae, Chelura, 54

Tropichelura, 54

intermedius, Lembos, 49, 51

inyacka, Parhyale, 131

iole, Hyale, 130

Ischyrocerus, 97

(?)kapu,97,99

oahu, 97,99

Ischyrocerus sp. C, 99

Jassa, 97

falcata, 97

(?) lUipuna, 97, 99

kailua, Amphilochus, 28

kaiulani, Maera, 84

kamanu, Lembos, 49, 51

Kanaloa, 12, 16, 18,105, 109

manoa, 105

kaneohe, Ampithoe, 29, 34, 35, 36

kapapa, Photis, 96

kapiolani, Colomastix, 55

kapu, Ischyrocerus (?), 97, 99

kaumaka, Gammaropsis, 90,93

Konatopus, 41

paao, 49

laakona, Eriopisa, 17, 76

Laetmatophilus, 113

hala, 16,113

laie, Hyale, 127, 130

laniloa, Liljeborgia, 105

Iawai, Podocerus talegus, 113, 117

leapakahi, Lembos, 49, 51

Lelehua (subg.), 130

Lembos, 2, 6, 14, 16,41,49

?aequimanus, 50, 51

intermedius, 49, 51

kamanu, 49, 51

leapakahi, 49, 51

macromanus, 49, 51

pualani, 49, 50, 51,52

waipio, 49, 52

Lembos sp., 52

Leucothoe, 2, 12,102

hyhelia, 103

lihue, 103

tridens, 103

Leucothoella, 102

Leucothoides, 102, 103

?pottsi, 103

lihue, Leucothoe, 103

likelike, Amphilochus, 28

lilipuna, Jassa (?),97,99

liliuokalaniae, Gitana, 25

Liljeborgia, 66, 103, 109

heeia, 105

laniloa, 105

lunalilo, Colomastix, 55

Lysianassa, 105

ewa, 105

macromanus, Bemlos, 51

Lembos, 49, 51

Maera, 2, 12, 16,68, 77

hamigera, 84

inaequipes serrata, 84

insignis, 70, 77

kaiulani, 84

pacifica, 84, 85

quadrimana, 84

serrata, 84

Maera sp. A, 84

Maera sp. B, 85

makena, Neomicrodeutopus (?), 52

makiki, Parapleustes derzhavini, 112

manoa, Kanaloa, 105

marionis, Amphilochus, 25

mauihina, Biancolina, 52, 97

maunaloa, Paradexamine, 61

Melita, 68, 77,85

appendiculata, 70, 85

fresneli, 85

pahuwai, 77, 85

menehune, Amphilochus, 28

Microdeutopus, 49, 52

Moera pectenicrus, 75

pocillimanus, 75

Mokuoloe, 29

ninole, 29

molokai, Elasmopus, 75

nahili, Aoroides, 41

nani, Atylus, 61

nenue, Aloiloi, 41

Neomicrodeutopus, 52

(?) makena, 52

ninole, Mokuoloe, 29

Nuuanu, 68, 77,85

amikai, 85

oahu, Ischyrocerus, 97,99

oceanica, Cymadusa, 38

Ochlesis, 105

alii, 105

Orchestia, 2

platensis, 131

orientalis, Ampithoe, 34, 35, 37

paao, Konatopus, 49

pacifica, Maera, 84, 85

Pontogeneia, 66

pahuwai, Melita, 77,85

pali, Gammaropsis, 41, 89, 93

Palinnotus, 10, 109

alaniphlias, 109

Paradexamine, 61

maunaloa, 61

Paragrubia, 38, 39

vorax, 37, 38

Para jassa, 97

angularis, 52, 97,99

Paranamixis, 39

Paraphoxus, 109

centralis, 109

Parapleustes, 103, 112

derzhavini makiki, 112

(?) honomu, 113

Parhyale, 131

hawaiensis, 131

inyacka, 131

Parhyalella, 122, 131

pietschmanni, 131

pectenicrus, Elasmopus, 70, 74, 75, 85

Moera, 75

pele, Gitanopsis, 28

Photis, 88,96

aina, 96

hawaiensis, 96

kapapa, 96

pietschmanni, Parhyalella, 131

piikoi, Elasmopus, 75

platensis, Orchestia, 131

Platophium brasiliense, 117

synaptochir, 117

Platyischnopus, 109

Pleonexes, 36
NUMBER 5 8

pocillimanus, Elasmopus, 75

Moera, 75

Podocerus, 10, 113

brasiliensis, 113, 117

hanapepe, 113, 117

talegus lawai, 113, 117

poipu, Ampithoe, 34, 36

pokipoki, Gammaropsis, 89, 93

Pontharpinia centralis, 109

Pontogeneia, 2, 66

pacifica, 66

Ppottsi, Leucothoides, 103

pualani, Lembos, 49,50, 51,52

pusilla, Colomastix, 55

quadrimana, Maera, 84

ramondi, Ampithoe, 36

rapax, Elasmopus, 70, 74, 75, 76

rubra, Hyale (see Hyale sp.)

schellenbergi, Ampelisca, 22

135

Seba, 117

ekepuu, 117

sechellensis upolu, Eriopisella, 76

serrata, Maera, 84

Maera inaequipes, 84

serrula, Elasmopus, 75

spinidactylus, Elasmopus, 76

stebbingi, Anamixis, 40

Stenothoe, 120

crenulata, 120

gallensis, 120, 122

haleloke, 122

Stenothoe sp. A, 122

Stenothoe sp. B., 122

?valida, 120, 122

validus, 122

synaptochir, Platophium, 117

Synopia, 16, 109

talegus lawai, Podocerus, 113,117

Talitrus, 2

tridens, Leucothoe, 103

Tropichelura, 54

insulae, 54

upolu, Eriopisella sechellensis, 76

?valida, Stenothoe, 120, 122

validus, Stenothoe, 122

Ventojassa, 97

ventosa, 97,99

ventosa, Eurystheus, 99

Ventojassa, 97, 99

vorax, Paragrubia, 37, 38

waialua, Ampithoe, 37

Wailele, 61

waimea, Hyale (Lelehua), 130

waipio, Lembos, 49, 52

zeylanicus, Gammaropsis, 91

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