J. LAVRENS BARN A. Sub littoral Gammaridea

(Amphipoda) of the

Hawaiian Islands

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 34

j . Laurens Barnard Sublittoral Gammaride a

(Amphipoda) of the

Hawaiian Islands

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

I97O
ABSTRAC T

Barnard, J. Laurens. Sublittoral Gammaridea (Amphipoda) of the Hawaiian

Islands. Smithsonian Contributions to Zoology, 34:1-286, 1970.—Gammaridean

Amphipoda from depths of 0-30 m on hard bottoms are increased from about 40 to

about 120 species, including 59 new species and 9 new genera and subgenera. About

half of the species is endemic. Nearly 70 percent of the faunule has come from

archipelagoes to the southwest of Hawaii, but only 15 percent of the faunule is of

tropicopolitan character. About 20 percent of the fauna has affinities with cool waters

of the North Pacific. This is the first significant record of cool-water species of any

marine group in Hawaii. Most of these species have a tubicolous ecology, suggesting

that nestlers are less successful in completing the long journey from the cool waters

of northern continents. All but three species with cool-water affinities have diverged

specifically from their mainland ancestors and at least two require erection of new

genera to describe their divergence. Other endemic genera of Hawaii have low

affinities with tropical Pacific faunas and have affinities with places like Antarctica,

the Caribbean Sea, and warm-temperate Australia. The tropical component of

Hawaiian Gammaridea is not impoverished by a priori standards of diversity. Few

elements expected to occur in Hawaii are missing. No evidence of interisland adaptive

radiation has been observed, but several pairs and triads of species with sibling

affinities are described as a result of successive waves of immigration of parent species.

Since no cool-water stepping stones occur between Hawaii and cool mainland shores

of the North Pacific, the divergent cool-water species of Hawaii probably reflect

morphological changes occurring after one increment of isolation. Some of these fairly

radical changes such as loss of palp articles, coalescence of urosomites, and possibly

axial reversal of dominance in gnathopods are seen to be of lower conservative value

than heretofore accorded. This upsets classifications in Atylidae, Dexaminidae,

Aoridae, and Isaeidae to a significant degree.

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

Purpose and scope of the study 1

Methods 2

Marine biogeography of the Hawaiian Islands 2

Endemic Gammaridea of the Tropics 3

Composition and general relationships of the gammaridean fauna 4

The Pantropical Indo-Pacific element 11

The Pacific tropical element 12

The cool-water northern element 13

The southwesterm warm-temperate element 16

The tropical American and Caribbean element 16

Introduced, minor and cryptic elements 16

Dispersal by rafting 17

The sweepstakes phenomenon 19

Lack of adaptive radiation in Hawaiian Amphipoda 22

Results of the one-step, long-distance isolation of cool water

immigrants to Hawaii 25

Enrichment of diversity in gammaridean genera 27

Summary and conclusions 28

Acknowledgments 29

Systematical procedure 30

Amphilochidae 30

Amphilochus Bate 30

Amphilocuhs kailua . 30

Amphilochus likelike 33

Amphilochus menehune 33

Gitana liliuokalaniae 37

Gitanopsis pele, new species 40

Cyproideinae 40

Mokuoloe, new genus 40

Mokuoloe ninole, new species 40

Ampithoidae : 44

Ampithoe akuolaka, new species 44

Ampithoe kaneohe, new species 44

Ampithoe orientalis Dana 50

Ampithoe ramondi Audouin 47

Ampithoe waialua, new species 53

Ampithoe species 54

Ampithoe (Pleonexes) poipu, new species 57

Cymadusa filosa Savigny 61

Cymadusa hawaiensis (Schellenberg) 61

Paragrubia vorax Chevreux 61

Anamixidae 61

Anamixis stebbingi Walker 61

iii
CONTENTS

Systematical procedure—Continued

 Pa£e

Aoridae 63

Aloiloi, new genus 63

Aloiloi nenue, new species 65

Aoroides Pcolumbiae Walker 68

Aoroides nahili, new species 70

Konatopus, new genus 70

Konatopus paao, new species 72

Lembos ?aequimanus Schellenberg 72

Lembos intermedius Schellenberg 76

Lembos kamanu, new species 77

Lembos leapakahi, new species 79

Lembos macromanus (Shoemaker) 82

Lembos pualani, new species 82

Lembos waipio, new species 85

Lembos species 87

Neomicrodeutopus (?) makena, new species 90

Atylidae 93

Kamehatylus, new subgenus 93

Atylus (Kamehatylus) nani, new species 93

Colomastigidae %

Colomastix kapiolani, new species %

Colomastix lunalilo, new species %

Colomastix species 100

Corophiidae 100

Corophium ?baconi Shoemaker 100

Corophium insidiosum Crawford 101

Dexaminidae 102

Paradexamine Stebbing ] 02

Wailele, new subgenus 102

Paradexamine (Wailele) maunaloa, new subspecies 103

Eophliantidae 103

Biancolina mauihina, new species 103

Eusiridae 109

Eusiroides diplonyx Walker 109

Pontogeneia pacifica Schellenberg 110

Gammaridae 115

Ceradocus hawaiensis J. L. Barnard 115

Elasmopus calliactis Edmondson 115

Elasmopus Pdiplonyx Schellenberg 115

Elasmopus ecuadorensis hawaiensis Schellenberg 119

Elasmopus hooheno, new species 121

Elasmopus molokai, new species 121

Elasmopus pectenicrus (Bate) 125

Elasmopus piikoi, new species 127

Elasmopus pocillimanus (Bate) 128

Elasmopus rapax Costa 131

Elasmopus spinidactylus Chevreux 136

Eriopisa Stebbing J3g
CONTENTS

Systematical procedure—Continued

Gammaridae—Continued Page

Eriopisa (?) hamakua, new species 138

Eriopisa laakona, new species 140

Eriopisella sechellensis upolu, new subspecies 143

Matra insignis (Chevreux) 147

Maera kaiulani, new species 147

Maera pacifica Schellenberg 150

Maera quadrimana (Dana) 152

Maera serrata Schellenberg 155

Maera species A 156

Maera species B 159

Melita appendiculata (Say) 161

Melita pahuwai, new species 161

Nuuanu, new genus 166

Nuuanu amikai, new species 166

Isaeidae ( = Photidae) 166

Chevalia aviculae Walker 166

Gammaropsis Liljeborg 170

Gammaropsis afra Stebbing 170

Gammaropsis alamoana, new species 170

Gammaropsis atlantica Stebbing 174

Gammaropsis digitata (Schellenberg) 178

Gammaropsis haleiwa, new species 178

Gammaropsis kaumaka, new species 181

Gammaropsis pali, new species 183

Gammaropsis pokipoki, new species 185

Photis aina, new species 185

Photis hawaiensis J. L. Barnard 191

Photis kapapa, new species 192

Ischyroceridae 195

Ischyrocerus (?) kapu, new species 195

Ischyrocerus oahu, new species 195

Jassa (?) lilipuna, new species 198

Parajassa angularis Shoemaker 202

Ventojassa, new genus 204

Ventojassa ventosa (J. L. Barnard) 205

Leucothoidae 205

Leucothoe hyhelia J. L. Barnard 205

Leucothoe lihue, new species 209

Leucothoe tridens Stebbing 211

Leucothoides Ppottsi Shoemaker 211

Liljeborgiidae 213

Liljeborgia heeia, new species 213

Liljeborgia laniloa, new species 216

Lysianassidae 216

Lysianassa ewa, new species 216

Ochlesidae 220

Ochlesis alii, new species 220
CONTENTS

Systematical procedure—Continued

Oedicerotidae 224

Kanaloa, new genus 224

Kanaloa manoa, new species 224

Phliantidae 226

Palinnotus Stebbing 226

Palinnotus alaniphlias, new species 227

Pleustidae 227

Parapleustes derzhavini makiki, new subspecies 227

Parapleustes (?) honomu, new species 233

Podoceridae 233

Laetmatophilus hala, new species 233

Podocerus Leach 237

Podocerus brasiliensis (Dana) 237

Podocerus hanapepe, new species 240

Podocerus talegus lawai, new subspecies 241

Sebidae 247

Seba ekepuu, new species 247

Stenothoidae 247

Stenothoe haleloke, new species 247

Stenothoe Pvalida Dana 250

Stenothoe species A 251

Stenothoe species B 251

Talitroidea 251

Hyale qffinis Chevreux 251

Hyale ayeli J. L. Barnard 255

Hyale grandicornis (Kr0yer) 255

Hyale grandicornis bishopae J. L. Barnard 257

Hyale honoluluensis Schellenberg 257

Hyale iole, new species 260

Hyale laie, new species 260

Lelehua, new subgenus 264

Hyale (Lelehua) waimea, new species 264

Hyale species (cf. rubra Thomson) 268

Parhyalella pietschmanni Schellenberg 271

Literature cited 271

Appendix 1 276

Appendix 2 277

Index 279
j . Laurens Barnard Sublittoral Gammaridea

(Amphipoda) of the

Hawaiian Islands

Introduction

Gammaridean Amphipoda of the Hawaiian chain

were last reviewed by J. L. Barnard (1955a), but that

report was based largely on a few intertidal and shallow sublittoral collections accumulated over the years

in the Bernice P. Bishop Museum. The true sublittoral

faunule had never been collected adequately and a

somewhat impoverished fauna was presumed to be

present. Through an exchange agreement between the

Bishop Museum and the Smithsonian Institution, the

writer was fortunate to spend 9 months of 1967 in the

Hawaiian chain in order to explore this sublittoral

fauna and refine the earlier taxonomy of the intertidal

species.

This report treats primarily sublittoral collections,

but new species, important new records, and various

taxonomic problems involving intertidal amphipods

are included. The littoral-sublittoral fauna is now increased from about 40 to about 120 species. Nine new

genera and 59 new species are described. Several diverse genera have been studied as thoroughly as necessary to identify their individuals in most of their instars

(Elasmopus, Colomastix, Gammaropsis, Leucothoe,

Lembos, Ampithoe, Gitanopsis, Photis, Podocerus),

but other genera remain to be so treated (Hyale,

Maera, Stenothoe, Cymadusa).

This paper will be supplemented by the publication

(Barnard, in press) of an illustrated key to the

Hawaiian shallow-water Gammaridea; keys and special discussions on identification from that paper are

therefore not included herein.

/. Laurens Barnard, Department of Invertebrate Zoology,

Smithsonian Institution, Washington, D.C. 20560.

Purpose and Scope of the Study

The effects of insular isolation on terrestrial faunas

and floras and to a degree in marine biotas are well

known in the Hawaiian Islands. The study at hand was

not formulated to replicate or reinforce any conclusions in that area of knowledge, for the taxonomy

of Indo-Pacific Amphipoda is too rudimentary for any

such treatment. This study was motivated simply by

the need to establish the faunistic composition of

Amphipoda in tropical biotic provinces.

Only one active taxonomic program on Gammaridea is being carried out anywhere in the world

tropics (Pillai and Nayar in India) and to my knowledge, except for that contemporary program, there

has never been a longtime resident gammaridean specialist in the tropics. James Dwight Dana, aboard the

Wilkes U.S. Exploring Expedition, came closest to this

categorization nearly 130 years ago.

Previous studies in the Indo-Pacific tropics have

been summarized briefly by J. L. Barnard (1965a)1

in a study of collections from atolls in Micronesia. The

tabulation of the Micronesian fauna is no more complete than was the compilation made by J. L. Barnard

(1955a) for the Hawaiian Islands. To illustrate the

paucity of knowledge on amphipods in the tropics,

one must state that the shores and shelves of Oahu

now represent the second most intensively studied tropical area (after the Madras and Ceylon coasts) in the

Indian and Pacific Oceans. The number of very rare

species for which only a few individuals are at hand -

suggests that further exploratory effort would continue

to reveal additional Hawaiian species but probably

1

 To that compilation should be added Ampithoe alluaudi

Chevreux and Pleonexes species (now named Ampithoe kulafi,

new species), omitted by error.

2

 Appendix 2.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with diminishing returns. The present fund of knowledge on Oahu corresponds to that of many nontropical

continental shores and shelves of comparable size. This

sampling of Oahu, though representing only a few

square miles, presumably is a fair example of the large

islands of the chain. One should expect little interisland speciation in a marine group of this kind but

naturally some of the other islands may have certain

species not recorded from Oahu. This circumstance is

known to occur rarely in fishes and mollusks.

The exploratory effort on Oahu seems to be comparable to that effort of J. L. Barnard (1950-67)3

 in

the warm-temperate of California, taking a midpoint

at Corona del Mar, California, from shore to 70 meters

and 50 miles in either direction. It appears to be comparable to exploration carried out in parts of the

Mediterranean (to 1925), New England (to 1918),

cold-temperate Norway (to 1895), Point Barrow,

Alaska (to 1955), South Africa (to 1925), and New

Zealand but is undoubtedly exceeded by our knowledge of the current Norwegian cold-temperate fauna,

that of cold-temperate British Isles, subarctic western

Russia to north Norway and the cold-temperate to

subarctic northern Japan Sea to Bering Sea, including

the arctic embayment of the Sea of Okhotsk, tropical

west Africa, and various antarctic and subantarctic regions. These are only subjective evaluations of the

efforts made in exploration of various provinces. They

are necessary if we are to make any conclusions about

the composition and diversity of Hawaiian Amphipoda, by balancing initial but intensive exploratory

efforts against one another on the probability that a

major share of the common species has been collected.

Methods

Samples of littoral and sublittoral flora, rocks, and

coral heads were obtained either by wading in intertidal zones or by utilizing rock dredges aboard vessels.

A few samples were obtained with the help of scuba

divers. Substrates were washed in large buckets of seawater poisoned with small amounts of formaldehyde to

kill the amphipods. Residues settling to the bottom

of the bucket were screened through a Tyler screen

of 100 mesh per inch and preserved in a sea water solution of 5 percent formaldehyde. Amphipods were removed from the samples within 2 days of the collecting

period and color notes were made, the amphipods

* Supplementing and reviewing many earlier works.

being sorted into species and then subdivided according

to color phenotypes. This process readily assisted in

the detection of several species that might have been

overlooked but several species had as many as 4 significant color variants. Rapid sorting of the samples was

possible by attention to color patterns in this artificial

medium. Color in formaldehyde remained fairly stable

for periods of 7 to 30 days after death. Notes on

identification methods utilizing color will be presented

elsewhere (Barnard, in press).

After gross sorting the specimens were represerved

in 70 percent alcohol to await morphological study.

Samples of gross dimensions were made so as to collect large quantities of amphipods (see station list in

Appendix 2). Generally an hour was spent collecting

the materials for each sample in the field and the number of specimens of amphipods usually exceeded 10,000

per sample. Aliquots of 500 to 1,000 specimens were

taken from each sample and counted by species (Appendix 2) and each sample then searched for individuals of rare species. The counts given in Appendix 2

represent the numerical proportions of each species encountered in the aliquot, the numbers occasionally having been reduced to low common denominators. Those

species having only one individual in a sample represent taxa not found in the aliquot but found elsewhere

in the sample. Besides aliquots many other specimens of

rare species were removed from the samples for taxonomic analysis and curatorial preservation in the collections of the Bishop Musuem and the Smithsonian

Institution.

Marine Biogeography of the Hawaiian Islands

A conspicuous part of the gammaridean Amphipoda

differ so strikingly in their geographic affinities from

other marine groups of Hawaii that a brief discussion

of those groups is warranted so as to document this

contrast clearly. A significant percentage (20) of

Hawaiian Gammaridea appear to have direct affinities

with warm and cold-temperate waters of adjacent continents and especially to warm-temperate California. Few other marine groups have any affinities

with America.

Ekman (1953) summarizes the Hawaiian situation

compactly. He notes in Vaughan's work on scleractinian corals their strong relationship to the western

Indo-Pacific and the scarcity of endemic genera; but

several reef-building families are missing from Hawaii.

More than a third of the warmwater echinoderm
NUMBER 3 4

species are endemic, but less than ten percent of the

genera are endemic and the others are Indo-Malayan.

None of the 70 species of warmwater starfishes lives

in America but a few echinoids and ophiuroids do

live there.

The fish fauna has smaller percentages of endemic

taxa than the invertebrates mentioned, but the west

Pacific character of the fauna is even more pronounced

in fishes than in the invertebrates.

Edmondson (1940) points out the following generalities about the marine invertebrate biogeography of

Hawaii in greater detail than Ekman: (1) corals have

their affinities with the Indo-Pacific region but are

attenuated by the absence of various families and the

absence of all but a few species of the characteristic

genus Acropora except in the leeward part of the

chain; Vaughan finds little relationship of corals to

American waters; (2) Hawaiian echinoderms have

their affinities with the Indo-Pacific, have a strong

Japanese element, but a very small group is believed

to have reached Hawaii from tropical America; (3)

gastropods also have notable affinities with Japanese

faunas though the bulk is purely Indo-Pacific; gastropods have stronger connections to tropical America

than do pelecypods, most of which are endemic to

Hawaii even though they have Indo-Pacific affinities;

(4) many of the common Hawaiian macrocrustaceans

have pan-Indo-Pacific or Japanese distributions and

there is a strong influence of Polynesian Crustacea on

the shores of tropical America but little evidence of

migration between America and Hawaii is seen (by

Rathbun).

Though students of stony corals (Scleractinia)

found little relationship between the faunas of the

American tropical Pacific and Hawaii, the most recent

study by Snuires (1959) shows that 11 of 29. snecies

of eastern Pacific reef corals also occur in the IndoPacific, and 8 of 27 coral species in the Uulf oi California occur in Hawaii. Ahermatypic corals of the

eastern Pacific, however, have a very low specific

relationship to the Indo-Pacific, for only 6 of 55

species occur to the west. A total of 18 percent of all

American Pacific tropical species also occurs in the

Indo-Pacific (13 out of 74 species). The generic relationships are strong with the Indo-Pacific and weak

with the Caribbean.

De Laubenfels (1957) notes that a weak affinity with

America is the only conspicuous relationship yet to

be seen or understood in Hawaiian Porifera, but studies in that group from the Central Pacific were cut

short by Dr. de Laubenfels' untimely death.

According to Knox (1958) about 43 percent of the

Hawaiian shallow-water polychaetes are endemic to

the islands and the relationships of the others are with

Japan and Indo-Malaya rather than with the tropical

Pacific. Knox treated only a portion of the fauna and

since that time Hartman (1966) has reviewed the

fauna. She reports 168 species of polychaetes occurring

in depths of 0-200 m, of which 42 percent are endemic,

31 percent cosmopolitan or subcosmopolitan in the

tropics, 13 percent of Pacific distribution and 14 percent of Indo-Pacific distribution. Connections with

warm-temperate North America appear non-existent.

Hawaiian Bryozoa are comprised of 46 genera and

80 species of which only 10 species and no genera are

endemic (Okada and Mawatari, 1958). Many Japanese and Californian species occur in the fauna, as yet

poorly explored. Continental provinces have many

times more species but are more thoroughly explored

(Malay province with 637 and California-Peru with

520 species).

Kohn (1959) notes that no species of eastern Pacific

Conus occur in Hawaii.

About 50 percent of the 400-500 species of inshore

Hawaiian fish are endemic but only two endemic

genera are known (Gosline, 1958). The fauna is a

balanced one and not impoverished. Apparently, no

specific connection occurs between Hawaii and the

warm-temperate of North America (Gosline and

Brock, 1960).

Endemic Gammaridea of the Tropics

Myers (1940) wrote that Hawaiian fish distributions

support the thesis of Austin Clark on echinoderms, that

the Atlantic Ocean is an inland sea biologically, meaning that the tropical Atlantic Ocean may be defined

biogeographically by the lack of certain Indo-Pacific

groups and not by the positive occurrence of unique

groups. Myers and Clark are of course discussing taxa

at the family level. In contrast, the endemic families

of Gammaridea are so poorly represented in the tropics

that a comparable analysis to that made for fishes and

echinoderms has to be concerned with genera. Table 1

has been prepared as a list of facts about the Gammaridea, of which there are more than 500 known

tropical species, or about 20 percent of the world total.

More than 150 genera, slightly more than 25 percent
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABL E 1.—Resume of tropical Gammaridea. Includes all Hawaiian species and those from

South China Sea (Imbach, in press)

Total taxa in world tropics

Pantropical endemic genera with their species

Pantropical, but genera not endemic, most species topically endemic

Indo-Pacific endemic genera with their species

Indo-Pacific nonendemic genera, but not occurring in Atlantic, with their species

West Atlantic endemic genera with their species

West Atlantic nonendemic genera, but not occurring in Indo-Pacific or east Atlantic, with their species

East Atlantic endemic genera, with their species

East Atlantic nonendemic genera, but not occurring in Indo-Pacific or west Atlantic, with their species <

Total taxa east Atlantic

West Atlantic

All Atlantic

Indo-Pacific

Total Indo-Pacific genera not occurring in Atlantic

Total Atlantic genera not occurring in Indo-Pacific

Total tropical endemic genera, world

Total species in these endemic tropical genera

Genera

160

18 with

55

32 with

24 with

• 5 with

5 with

6

 2 with

24 with

72

40

91

126

56

36

55

Species

589

57

362

36

51

5

13

2

32

185

70

238

382

77

• Plus Batea and Carinobatea occurring in east Pacific.

b

 Plus Cerapopsis occurring in Mediterranean.

« Mosdy from North Atlantic WT-CT.

of the known world genera, occur in the tropics. Very

few pantropical endemic genera occur, for most of the

pantropical genera have also spread into high latitudes

or have entered the tropics from other seas. Only 32

endemic Indo-Pacific genera are known and most of

these are monotypic. This approximates the number of

endemic abyssal genera and demonstrates how weak

Gammaridea appear to be in the tropics. This weakness

may be a function of eurythermy and highly successful

dispersal methods so that gammaridean genera, once

differentiated, tend to radiate out of the tropics. But

it also points to the fact that the Gammaridea may be

less adapted to tropical climes than to cooler waters.

This is especially marked by the large number of

tropical west African genera that are simply imports

from north Atlantic warm and cold-temperate waters

and by the large number of imports to Hawaii from

American cold waters. The tropics of west Africa is,

of course, a narrower band of environment than an

eastern continental shore and may have considerable

upwelling like that occurring along western America.

And Hawaiian thermal regimes are not purely tropical.

West tropical Africa seems to have been adequately

studied in relation to the Indo-Pacific tropics, for it

has over 70 genera already reported, while nearly 130

occur in the Indo-Pacific. The entire tropical Atlantic

has over 90 genera but the western Atlantic is very

poorly known. The situation in Gammaridea parallels

that in fishes mentioned by Myers; the tropical Atlantic

can be described in terms of what it lacks of IndoPacific genera, while a strong secondary factor is the

high fraction of genera immigrated from cooler seas to

replace missing Indo-Pacific taxa. Many of these west

African genera are represented by one or a few species

of genera otherwise centered in higher latitudes.

The Indo-Pacific also has similar cool-water imports

but a significant proportion of their records comes from

the days of amphipod taxonomy when species superficially familiar to a European taxonomist were firmly

identified with the European members. In my opinion,

these identifications remain to be confirmed.

Composition and General Relationships of the

Gammaridean Fauna

Even a rudimentary knowledge of the world of Gammaridea and especially their tropical taxa made it

possible to predict with fair success the kinds of tropical amphipods that might occur in Hawaii (Table 2),4

but the overall prediction contained two gross errors

that are of interest to biogeographers. In the first place,

* Appendix 1 cites taxa not expected in Hawaii.
NUMBER 34

TABLE 2.—Groups, or genera and species expected to occur in Hawaiian Islands

based on past exploration (3), number and kinds of species in other parts of JndoPacific tropics (/), their known ranges (2), kinds of species with known affinity for

islands or known affinity for tropical shores (4), known affinity for adjacent warmtemperate environments (7), or nearest insular or continental group (8), expected as

ectoparasites (5), or imported by humans (6)

Numbers in parentheses are codes for column 4. Codes

1 and 2 are amplified in

provides quick reference

Hawaii prior to 1967.

been predicted to occur

columns 1 and 2. Column 3

to genera known to occur in

Vote that most would have

according to reasons in column 4. The partial success in predicting additional

genera and species demonstrates the low occurrence

of generic endemicity and the very low occurrence of

sweepstakes dispersal in

occurrence of occasional

warm-temperate regions

this marine group. But the

affinities to North American

upset the success of the prediction. See Tables 3 and 4. Family names listed are

Genus or group

Acanthonotozomatidac

Ampelisea

Amphilochinae, additional

Amphilochus

Ampithot

Anamixidac

Aorid-photid, additional

Byblis

Calliopiid-cusirid, additional

Ceradocus

Cerapus

Cheiriphotis

Chelura

Chevalia

Colomastix

Corophiidae, additional

Corophium

Cymadusa

Cyproideinae

Dexaminidae

Elasmopus

Ericthonius

Eriopisa-Eriopisella-ctc.

Eusiroides

Gammaropsis

Uyale

dunella-Listriella

(1) Number

of species

in Indo- (2)

exclusive of genera in those families also expected. The

cosmopolitan index ranges from A-G as based on 7

arbitrary geographic areas quoted in Barnard (1965a,

table 1), from the Red Sea to Hawaii. The higher the

letter (G is highest), the broader is the Indo-Pacific

distribution. In column 5 the species expected to occur

in Hawaii are in addition to any already known there.

If the predicted number is lower than the actual number known to occur then the category is more diverse

than predicted.' fhe initials "NM" indicate very strong

cases for predicting the number of species stated. Note

that many were overfulfilled.

(3) Species Why expected Species

Cosmo- known in

Pacific politan index Hawaii

tropics

(prior to

1967)

3

13

0

3

4

3

3

6

3

2

1

2

1

1

2

1

5

4

2

11

21

3

3

3

8

10

2

(prior to

1967)

F 0

F 1

F 0

F 1

G 2

D 0

C 0

E 0

D 0

F 1

C 0

C 0

D 1

C 0

G 1

A 0

G 1

G 3

F 0

F 0

G 4

G 1

E 0

E 1

F 0

G 4

D 0

in Hawaii, expected Total species

by code to occur now known

numbers 1,2, in Hawaii in Hawaii

3, 4, 5, 6, 7, 8

2 1 0

1,2 3 1

— — 5

2 1 ?1

2 2 7

8

4

1,2

7

7

4,8

4,7,8

4,6

4,7

1 1

I 4

1 0

1 0

1 1

0

0

1 1

1

2,4,7 1 4

— 0 0

1,2,6 2 3

1,2 1NM 3

2,8 1 -1

1,2 2 1

1,2 4NM 10

2,6 1 1

2,4,7 1 4

2,4 1NM 1

1.2 2 7

1,2 4NM 8

4,7 1 0
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus or group

Isaea

Jassa x

Lembos

Lembopsis l

Ltucothoe

Leucothoides

Liljeborgia

Lysianassidac

Maera

Mandibulophoxus

Megaluropus

Megamphopus-Podoceropsis

Mtlita

Paragrubia

Paraphoxus

Parelasmopus

Parhyolt

Parhyalella

Photis

Platyiseknopus

Podocerus

Podoceridae, additional

Pontogeneia

Stenothoe

Synopia

Turn

Xmocheha

TABLE

(1) Number

of species

in IndoPacific

tropics

(prior to

1967)

0

1

12

1

8

1

3

16

15

1

1

1

3

1

3

3

1

2

9

1

9

2

1

4

3

1

2

2.—Continued

(2) Cosmopolitan index

D

G

A

F

E

D

F

G

A

C

A

G

E

D

F

G

E

G

A

G

D

A

G

F

A

C

(3) Species

known in

Hawaii

(prior to

1967)

0

0

2

0

0

0

0

0

3

0

0

0

()

0

1

0

1

2

0

0

0

Why expected

in Hawaii,

by code

numbers 1,2,

3, 4, 5, 6, 7,8

5

6,7

1,2

4,8

1,2

2,8

8

1,2

1,2

4,7

7

8

2

2,8

4,7

2,4,7

2,4

4

1,2

4

1,2

4

—

2,6

2,4

4,8

8

Species

expected

to occur

in Hawaii

r

:

:

>

i

1NM

1NM

2

1

1NM

1NM

2NM

1

2

1

0

2NM

1

1

1

Total species

now known

in Hawaii

0

1

8

0

3

1

2

1

7

0

0

0

2

1

1

0

1

1

3

0

3

1

1

4

0

0

0

1

 Not J. falcate (Montagu), the species expected.

1

 As represented by L. tridens (Schellenberg) formerly in genus Microdeutopus.

Unexpected Genera and Their Total Species

New genera are marked with asterisks.

Aloiloi* (1), Atylus (1), Biancolina (1), Ischyrocerus

(3), Kanaloa* (1), Nuuanu* (1), Ochlesis (1),

Palinnotus (1), Parajassa (1), Parapleustes (2),

Seba (1), Ventojassa* (1).

Genera Substituted for Familial Expectations

Amphilochidae: Amphilochus (3), Gitana (1). Aoridae: Aoroides (2), Konatopus* (1), Neomicrodeutopus (1). Cyproideinae: Mokuoloe* (1). Dexaminidae: Wailele* (1). Lysianassidae: Lysianassa

(1). Podoceridae: Laetmatophilus (1).

the estimates were based only on tropical components

since no significant occurrence of cold-water imports

was expected; in the second place, the maximal estimates based on tropical imports were expected to pinpoint, by negative occurrences, not only an impoverishment of species in common genera but the

complete absence of significant familial or generic

groups. The fulfillment and overfulfillment of these

estimates demonstrate the presence of a rich, balanced

fauna of tropical amphipods. The few missing elements

appear insignificant in the final analysis and some have

probably been replaced by unpredicted taxa. Several

of the tropical genera are surprisingly more diverse

than expected (Table 2). Appended to the tropical
NUMBER 3 4

TABL E 3.—Hawaiian marine aquatic Gammaridea and their relationships, by distribution

and morphological affinities

Name of species

*Aloiloi nenue

Ampelisca schellenbergi

(Amphilochus marionis)

*Amphilochus kailua

* Amphilochus likelike

*Amphilochus menehune

* Ampithoe akuolaka

'Ampithoe kaneohe

Ampithoe orientalis

* Ampithoe poipu

Ampithoe ramondi

*'Ampithoe waialua

* Ampithoe species

*Anamixis stebbingi

'Aoroides ?columbiae

*Aoroides nahili

'Atylus nani

*Biancolina mauihina

Ceradocus hawaiensis

Chelura insulae

'Chevalia aviculae

*Colomastix kapiolani

*Colomastix lunalilo

Colomastix pusilla

'Colomastix species

Corophium acherusicum

*Corophium Ibaconi

*Corophium insidiosum

Cymadusa filosa

Cymadusa hawaiensis

Cymadusa oceanica

Elasmopus calliactis

'Elasmopus diplonyx

Elasmopus e. hawaiensis

'Elasmopus hooheno

'Elasmopus molokai

Elasmopus pectenicrus

*Elasmopus piikoi

Elasmopus pocillimanus

Elasmopus rapax

Elasmopus spinidactylus

Ericthonius brasiliensis

'Eriopisa hamakua

'Eriopisa laakona

'Eriopisella s. upolu

Eusiroides diplonyx

Ecology Endemic

D +

D

IN +

IN +

IN +

D +

D +

D

D +

D

D +

D +

I

D

D +

N +

L +

N +

L

D

I +

I +

I

I +

D

D

D

D

D +

D +

N +

N

N +

N +

N +

N

N +

N

N

N

D

N +

N +

N +

N

Geographic affinity if appropriate

Place:

WT-CT?

Hawaii to Caribbean

Unidentifiable; from

Schellenberg, 1938

WT-California

WT-California

WT-California

North America WT

Micronesia

Philippines

Micronesia

Circumtropical

Gulf of California

WT-CT

Ceylon

Oregonian

Oregonian

Japan

Australia or Mediterranean

WT Mexico

Circumtropical; wood borer

Circumtropical

An ti boreal

Australia WT

Circumtropical (?)

Unknown

Cosmopolitan, possibly

introduced

East Pacific, Alaska-Peru

Cosmopolitan, possibly

introduced

Circumt ropical

Unknown

Unknown

Micronesia

Species:

G. vilordes

G. vilordes

G. vilordes

A. pollex, A. simulans

A. kulafia

A. kulafia, A. kaneohe

A. columbiae

A. japonicus

C. paucidentatus

C. fissilingua

C. subcastellata

C. brazieri

E. pseudqffinis

Micronesia, southern Polynesia

Galapagos

Southern Polynesia

Southern Polynesia

Circumtropical

Possibly southern Polynesia

Circumtropical-WT

Circumtropical-WT

Indo-Pacific

Cosmopolitan WT-CT

Unknown

Unknown

Seychelles

Indo-Pacific

E. e. ecuadorensis

Presumptive

E. minimus

E. s. seychellensis

Depth

in Hawaii

(m)

4

94-229

2

0-5

30

5-30

0

0-4

0

0

0

0-30

0

0-30

0-30

0

0

0

0-30

0

0-30

30

5

0

30

0

2

0

0-5

0

0

0-40

33

0

0

0-30

0

0-18

0-30

0

0

0-30

4

0-4

3-30

0-30

See footnote at end of table.
SMITHSONIAN CONTRIBUTION'S TO ZOOLOGY

TABLE 3.—Hawaiian marine aquatic Gammandea and their relationships, by distribution

and morphological affinities—Continued

Name of species Ecology Endemic

Geographic affinity if appropriate

Place: Species:

Depth

Distribution

in Hawaii

(m)

*Gammaropsis afro

*Gammaropsis alamoana

*Gammaropsis atlantica

*Gammaropsis haleiwa

*Gammaropsis kaumaka

*Gammaropsis pali

*Gammaropsis pokipoki

*Gitana liliuokalaniae

*Gitanopsis pele

Hyale affinis

Hyale ayeli

Hyale g. bishopae

Hyale honoluluensis

*Hyale iole

*Hyale laie

* Hyale (Lelehua) waimea

*Hyale species (cf. rubra)

*Ischyrocerus kapu

*Ischyrocerus oahu

*Ischyrocerus species C

*Jassa lilipuna

*Kanaloa manoa

*Konatopus paao

*Laetmatophilus hala

Lembos Jaequimanus

Lembos intermedius

*Lembos kamanu

*Lembos leapakahi

*Lembos macromanus

* Lembos pualani

*Lembos waipio

*Lembos species

*Liljeborgia heeia

*Liljeborgia laniloa

*Leucothoe hyhelia

*Leucolhoe lihue

*Leucothoe tridens

*?Leucothoides pottsi

*Lysianassa ewa

Maera insignis

* Maera kaiulani

Maera pacifica

Maera quadrimana

* Maera serrata

* Maera species A (cf. hamigera)

* Maera species B

Melita appendiculata

D

D

D

D

D

D

D

IN

IN

N

N

N

N

N

N

N

N

D

D

D

D

N

D

PIN

D

D

D

D

D

D

D

D

N

N

I

I

I

I

N

N

N

N

N

N

N

N

N

Indo-Pacific

h Oregonian-Californian

Indo-Pacific

+• Unknown

•f- Unknown

f Probably WT-CT

+- Probably Indo-Pacific

f CT

+• Caribbean

SouthernPoly nesia-M icronesia

+- WT Southern Hemisphere,

possibly Japan also

+- Japan

Micronesia; possibly race of

4- Possibly CT-WT

+• Japan

-1- PTethyan

4- Unknown, possibly race of

+ CT

-f- CT

4- Unknown, probably CT

+ CT

4- Antiboreal

+ California-Mexico WT

+ Australia WT

Southern Polynesia

4- Unknown

4- Unknown

+ Micronesia-West Africa

Indo-East Pacific

4- California

+ Micronesian race apparent

4- Unknown

+ Micronesia-Southern Polynesia

+ Micronesia-Southern Polynesia

Micronesia

+ California-WT

Micronesia

Micronesia to Caribbean

4- Indonesia; Caribbean

Indo-Pacific

+ Micronesia-Southern Polynesia

Micronesia-Southern Polynesia

Tropical Pacific

Southern Polynesia-Micronesia

Indo-Pacific

4- Indo-Pacific

Circumtropical-WT

G. thompsoni

Gitanopsis arctica

G. tortugae

H. media

H. g. grandicornis

H. chevreuxi

?H. plumulosus

"H. dollfusP' of Iwasa

//. carinata

H. rubra

I. anguipes

L. hystrix

L. species

L. jrancanni

L. concavus

L. proximo

L. proximo

L. alata

L. cubensis

L. heterodonta

M. pacifica

M. hamigera

M. othonides

30-33

0-30

2-30

2

0

0-2

2-30

4

2

0

0

0

0

0

0

0

0

2-30

0-2

30

0

2-30

0-30

18-30

0

0-4

0

0-30

2-30

0-4

0-30

0

2-30

0

0-33

33

2-30

0-30

0-18

0-30

4

0-30

0-30

0-4

30

0

0-30

See footnote at end of table.
NUMBER 3 4

TABLE 3.—Hawaiian marine aquatic Gammaridea and their relationships, by distribution

and morphological affinities—Continued

Name of species

*Melita pahuwai

*Mokuoloe ninole

* Neomicrodeutopus (? ) makena

*Nuuanu amikai

'Ochlesis allii

*Palinnotus alaniphlias

* Paradexaminc {Wailele) maunaloa

Paragrubia vorax

*Parajassa angularis

Paraphoxus centralis

*Parapleustes d. makiki

*Parapleutes (?) honomu

Parhyale hawaiensis

Parhyalella pietschmanni

*Photis aina

Photis hawaiensis

*Photis kapapa

Podocerus brasiliensis

*Podocerus hanapepe

* Podocerus t. lawai

Pontogeneia pacifica

*Seba ekepuu

Stenothoe gallensis

*Stenothoe haleloke

Stenothoe Ivalida

*Stenothoe species A

*Stenothoe species B

*Ventqjassa ventosa

Ecology

N

PIN

D

N

I

L

I

D

D

N

N

N

N

D

D

D

IN

IN

IN

N

I

IN

IN

IN

IN

IN

D

Geographic affinity if appropriate

Place:

+ Ceylon

+ Unknown, possibly Caribbear

+ Boreal?

•+• Unknown

+ Possibly Indonesian species

+ Australoafrican WT

+ Unknown

Indo-Pacific

California-Mexico WT

+ Unknown

+ Japan (Sea of)

+ Boreal

Circumtropical

+ Bermuda, Caribbean

+ Australia WT

+ Ceylon

+ Celebes

Circumtropical

+ Australia WT

+ Micronesia

+ Caribbean

+ New Zealand

Circumtropical

+ CT North Atlantic

Cosmopolitan

+ Unknown

+ Unknown

California-WT

Species:

M. zeylanica

1

P. d. derzhavini

P. batesoni, P. whelpleyi

P. dolichommata

"P. longimanus" of

Walker

Photis species of Pirlot

P. inconspicuus

P. t. talegus

P. longleyi

S. typica

S. monoculoides

Depth

DtstftbuttOTi in Hawaii

(m)

0

30

0

18

0

0

0

0-30

4-30

?

3-4

33

0

0

2-30

0

2-30

0-5

0

0-30

0

0

0

0

33

18

2

4-33

1

 Asterisks indicate first records for Hawaii. WT= warm-temperate; CT=cold-tempcrate. Ecological symbols: D=domicolous

I =inquilinous; IN=inquilinous nestler; L=lignivorous; N=nestler.

fauna is a diversity increment of more than 70 percent,

an element composed mainly of cool-water species from

American and Japanese waters. Few of these could

have been brought to Hawaii by human means.

These conditions indicate that the main IndoPacific amphipod fauna probably will be found to be

far more diverse than indicated by previous studies and

that my estimates of Hawaiian impoverishment, based

on the literature, have been too low. Probably it will be

shown that: (1) the tropical component of Hawaii is

indeed impoverished by a revised standard and open

niches may have been partially filled with cool-water

imports; (2) numerous kinds of fully tropical amphipods, though dispersed into the vicinity of the Hawaiian

Islands, fail to become established because of marginal

thermal conditions and the absence of suitable physical

and biotic substrates (e.g., acroporan corals); (3) coolwater organisms also dispersed into the vicinity of

Hawaii have been able to survive in the marginal

thermal conditions because fully tropical amphipods

have not occupied certain favorable niches.

The Hawaiian shallow-water fauna has the normal

appearance of the tropics in the presence of numerous

species of Hyale, Elasmopus, Gammaropsis, Cymadusa,

Lembos, and Maera, but these widely dispersed genera

are not always strong indicators of tropical conditions
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

for many extend abundantly through warm-temperate

regions and become reduced markedly in diversity only

within cold-temperate waters. There is a possibility

that Hyale will be shown to be impoverished in Hawaii

when the continental Indo-Pacific fauna is better

known, but for the present the other genera mentioned

above are far more diverse in Hawaii than expected

and more diverse than in any other restricted locality

so far studied in the Indo-Pacific.

Various other tropical indicators occur in Hawaii

either as members of tropical genera or as specifically

tropical species, but a large share of these also extends

far into warm-temperate regions (Table 4).

About half of the Hawaiian Gammaridea is endemic,

as far as known, to the Hawaiian chain (Table 4), but

our knowledge of many taxa to the west and south is

poor and some Hawaiian species will undoubtedly be

found there. The species of Amphilochus and Gitanopsis, for instance, are presumably associated with

scleractinians and this relationship has been previously

TABLE 4.—Distribution of Hawaiian Gammaridea by groups of geographic affinity, commencing with widespread species and terminating with endemic species having highly topical

(sic!) affinities

Noncndemic Species

A. Imported by humans (see also asterisks below as possible

candidates).

Corophium acherusicum, C. baconi, C. insidiosum.

B. CSrcumtropical and circum-warm- temper ate.

Ampithoe ramondi, Chevalia aviculae, Colomastix pusilla,

Elasmopus pocillimanus, E. rapax, Erkthonius brasiliensis*,

Melita appendkulata, Podocerus brasiliensis*, Stenothoe gallensis*, S. valida*.

C. Circumtropical only.

Chelwra insulae*, Cymadusa filosa, Elasmopus pectenicrus,

Parhyale hawaiensis.

D. Indo-Pacific.

Anamixis stebbingi, Elasmopus spinidactylus, Eusiroides diplonyx, Gammaropsis qfra, G. allantica, Hyale honoluluensis, H. ayeli, Maera insignis, Paragrubia vorax.

E. Western tropical Pacific.

Ampithoe orientalis, Lembos macromanus (and east Pacific).

F. Micronesia-Polynesia.

Elasmopus diplonyx, Lembos aequimanus, Leucothoe hyhelia,

L. trtdens, Hyale qffinis, Maera pacifica, M. quadrimana,

M. serrata.

G. Caribbean Sea and tropical America.

Amptlisca schellenbergi, Leucothoidespottsi (also Micronesia).

H. Cool waters of North America and Asia.

Aoroides columbiae, Parqjassa angularis, Ventqjassa venlosa.

Endemic Species with Affinities Classified

I. Indo-Pacific.

Cymadusa hawaiensis, C. oceanica, Elasmopus calliactis,

Erisopisella sechellensis upolu, Gammaropsis pokipoki, Lysianassa ewa, Maera species A, Maera species B, Melita

pahuwai, Paradexamine (fVailele) maunaloa, Photis aina,

P. hawaiensis, P. kapapa.

J. Western tropical Pacific.

Ochlesis alii.

K. Warm-temperate of Australia.

Biancolina mauihina, Colomastix lunalilo, Laetmatophilus hat a,

Palinnotus alaniphlias, Podocerus hanapepe, Seba ekepuu.

L. Antiboreal-Antarctica.

Colomastix kapiolani, Kanaloa manoa.

M. Caribbean Sea and tropical America.

Elasmopus ecuadorensis hawaiensis, Gitanopsis pele, Mokuoloe

ninole, Parhyalella pietschmanni.

N. Micronesia-Polynesia.

Ampithoe kaneohe, A. poipu, Elasmopus hooheno,E. molokai,

E. piikoi, Hyale laie, Lembos intermedius, L. leapakahi,

L. waipio, Liljeborgia heeia, L. laniloa, Maera kaiulani,

Podocerus talegus lawai.

O. Cool waters of North America and Asia ("J"=Asian

species).

Aloiloi nenue, Amphilochus kailua, A. likelike, A. menehune,

Ampithoe akuolaka, Ampithoe waialua, Aoroides nahili, Atylus

"wi (J), Ceradocus hawaiensis, Gammaropsis alamoana, G.

pali (?J), Ghana liliuokalaniae, Hyale g. bishopae, H. iole,

H. laie (J ), Hyale species (J), Ischyrocerus kapu, I. oahu,

I. species C, Jassa lilipuna, Konatopus paao, Lembos pualani, Leucothoe lihue, Neomicrodeutopus makena, Parapleustes

derzhavini makiki (J), P. honomu, Pontogeneia pacifica,

Stenothoe haleloke.

P. Unknown affinities.

Colomastix species, Eriopisa hamakua, E. laakona, Gammaropsis haleiwa, G. kaumaka, Hyale (Lelehua) waimea, Lembos

species, Nuuanu amikai, Paraphoxus centralis, Stenothoe

species A, Stenothoe species B.
NUMBER 34 11

unknown and unstudied in the tropics. The smallbodied taxa have apparently been ignored or overlooked previously.

The other half of Hawaiian Gammaridea has broader distributions and one-seventh of the total Gammaridea is circumtropical. A few species are distributed

between the central Pacific and the Caribbean Sea

but most of the remainder occur either throughout the

Indo-Pacific or as far west as Micronesia.

Cognates of endemic species are dominated by those

with affinities to American shores, as the remaining

half of the endemic species has cognates divided among

the Caribbean Sea, warm-temperate Australia, the

central Pacific, the Indo-Pacific, and antiboreal regions. This presumably indicates the higher frequency

of long-term interruption to continuous gene flow between Hawaii and America than between Hawaii and

places to the west and south. The migratory direction

of all taxa between America and Hawaii cannot be

elucidated. Several of the species of Amphilochus and

Gitanopsis may have moved from west to east because

more species of the genera occur in Hawaii than in

America and because their scleractinian hosts are a

stronger dominant of Hawaiian waters than of American waters. American pocilloporans also came from the

west but whether Hawaii formed their stepping stone

is debatable. In contrast, the ampbipods Parajassa,

Ventojassa, and other ischyrocerids may have come

to Hawaii from America or Asia because the family

Ischyroceridae is more typical of cool-water continents

than it is of purely tropical zones.

Species of Amphipoda coming to Hawaii from the

west must maintain more frequent genetic contact than

those from America as there is a smaller proportion

of Hawaiian species with immediate

 5 southwestern

affinities in relation to the total affinity structure of

the Hawaiian fauna than with American affinities.

Thus, 30 percent of the Hawaiian fauna is of coolwater affinity and nearly 90 percent of those species is

endemic, whereas only half of the remaining 70 percent of the Hawaiian fauna with tropical affinities is

composed of endemic species.

Taxa not expected to occur in the Hawaiian Islands

are heavily oriented to cool waters (Table 2, terminus) , primarily in warm-temperate regimes. Only Ochlesis, of 12 unpredicted genera, was previously known

in tropical waters. Few of the remaining genera are

expected to be found as common members of the ultratropical regime. A list of nine genera that are substitutes for predicted families is also given in Table 2,

and only one of these is of widespread tropical occurrence, whereas most of the others occur primarily in

nontropical waters.

The Pantropical and Indo-Pacific Element

Before the present study was commenced, the IndoPacific tropics had 297 species of gammarideans, 11 of

them endemic to Hawaii (J. L. Barnard, 1965a, Table

I and see footnote herein, p. 1). About 40 of those

species are widespread through four or more of the

seven regions that the Indo-Pacific was divided into by

Barnard. The sum of 40 species has been amplified by

recent knowledge extending ranges of a few species

into the eastern Pacific. Slightly more than half of

those species (22) has been collected in Hawaii but

several of the remainder can possibly be dismissed as

"problem" records. For example, Leucothoe furina

and L. spinicarpa, as representative of widespread

species, may be the result of taxonomic errors and

Elasmopus spinimanus may be the catchall for various

misidentifications. Two species of Synopia have not

been looked for in Hawaii, and various others like

Cyproidea ornata are difficult to find in cursory surveys. The list of missing members of the 40 is narrowed

to the following significant species, that, were they in

Hawaii, should have been collected in the surveys so far

completed: Ceradocus "rubromaculatus" 6 Cerapus

tubularis, Cheiriphotis tnegacheles, Jassa falcata,

Maera inaequipes, Lembos podoceroides, and Parelasmopus suluensis. The latter two species are confined to

the Indo-Pacific region and their absence in Hawaii

might indicate Hawaii's isolation or weakly tropical

thermal regime but the absence of the other four may

indicate primarily isolation as they also occur in the

eastern Pacific Ocean. How they may have bypassed

Hawaii is of considerable interest. Other widespread

species like Leucothoella bannwarthi, Dexaminoides

orientalis, and Jphimedia discreta have been sought in

Hawaii but the first two may occur on ascidians and

may not have been able to reach Hawaii without a

simultaneous journey of their hosts.

Significantly, only two species of Indo-Pacific tropical Ceradocus occur, the widespread C. "rubromacu5

 Referring to northeastern Micronesia and the nearest

parts of Polynesia to Hawaii.

360-034 O—70 2

* Tropical records of this warm-temperate Australian species

may be of erroneous identification.
12

latus" everywhere but Hawaii, and the endemic

C. hawaiensis. Though Cheiriphotis megacheles occurs

from the Indian Ocean eastward to the warmtemperate of California and lives sympatrically in

Mexico with Ventojassa ventosa and Parajassa angularis, the missing Cheiriphotis in Hawaii may be the

result of the presence of only two niches available foi

this morphotype; Cheiriphotis strongly resembles the

other two genera in physical appearance.

The presumed absence of Cerapus tubularis reflects

the possibility that an organism obligatorily living in a

portable cone made of sand grains has not been able to

make the long journey to Hawaii whereas organisms

with fixed soft tubes would not easily fall off of flotsam.

But C. tubularis has made the journey from Asia and

Europe to N. America or vice versa.

The absence of Jassa falcata and Maera i. inaequipes

may reinforce the heretofore weak supposition that

they have been distributed in subcosmopolitan degree

by ships rather than by natural means and that, by

chance alone, the ships coming to Hawaii have succeeded in bringing Corophium acherusicum, C. insidiosum, perhaps Elasmopus rapax, Stenothoe valida,

S. gallensis, and Podocerus brasiliensis but have failed

to bring the other 2 species.

The vacuum formed by the absence of an expected

acanthonotozomatid is filled by an ochlesid, representing a monotypic family presumed to have evolved

from acanthonotozomatid ancestors.

Members of the Calliopiidae such as Atylopsis and

Apherusa have been found in tropical and warmtemperate zones but none has been found in Hawaii.

Tropical genera like Idunella, Byblis, Tiron, and

Isaea, missing so far in Hawaii, occur on sedimentary

bottoms or on fishes and have not been pursued.

One might presume that among the list of 297 IndoPacific tropical species occurs a significant number

that, as explorations continue, will be found to be more

widespread than is now known. If not found in Hawaii,

these widespread species, when added to the present

list, will impart a stronger measure of isolation, and

perhaps impoverishment, to the Hawaiian analysis

than now shown.

The Pacific Tropical Element

Though Hawaii has many of the pantropical and

widely dispersed Indo-Pacific elements in its fauna,

they, like the restricted fauna of the central and western Pacific Ocean, have their migratory pathways to

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hawaii through the tropical islands nearest Hawaii.

The nearest archipelagos are the Line Islands to the

south, the Phoenix and Gilbert Islands to the southwest, and the Marshall Islands to the west-southwest.

Johnston Island may serve as a stepping stone for

southern Polynesian and Micronesian species dispersing to Hawaii. That small atoll "717" miles from

Honolulu may be a stepping stone for fishes (Gosline,

1955). Whereas 14 fishes of southern origin have

reached Johnston but have not passed on to Hawaii,

16 fishes otherwise endemic to Hawaii have also

reached Johnston. Gosline concludes that Johnston is

an outlier of the Hawaiian Archipelago rather than a

peripheral component of the central Pacific faunal

area. The two groups of fishes are notably subequal in

number (16 and 14) and this might indicate that

Johnston formed a perfect stepping stone except for

the probability that the pressure of migration toward

Johnston from the west and south must be higher than

that from Hawaii. In view of the impressive list of

fishes occurring in both places (86) that migratory

pressure is obviously of sufficient magnitude to extend

well beyond Johnston toward Hawaii from Polynesia.

One must presume that most of these fishes are of central Pacific rather than Hawaiian origin. How many

of those necessarily using Johnston as a stepping stone

is unknown. One has to balance the small target size

of Johnston and the large target size of the remoter

Hawaiian chain against the probability that once a fish

is entrained within a migratory journey it has a greater

chance of hitting a large target than a small, despite

the moderately greater distance of the larger target.

Phycophilous Gammaridea probably do not migrate

as active nekters, although the instigation of an accidental journey may come when a species has fully

saturated its habitat and high numbers of individuals

in a deme are carried away in flotsam by storms. I

judge that few Amphipoda have necessarily required

Johnston Island as a way-station for consolidation of

their populations before the extended journey to

Hawaii has been undertaken. But Gosline's (1955)

analysis does offer evidence to refute the target theory

in the sense that only a tiny fraction of fish taxa shared

between the Line Islands and the Hawaiian chain does

not also occur in Johnston Island. This may support

the stepping stone theory unless one can demonstrate

that Johnston has all of those common fishes because of

extremely high dispersion pressure from the direction

of the Line Islands or because many of the species have

reached Johnston in immigrations from Hawaii.
NUMBER 34

Approximately 80 percent of the Hawaiian Amphipoda have passed through the nearest tropical islands,

and we should expect that the strongest direct affinities of Hawaiian Amphipoda are therefore with those

southwestern archipelagos, but the picture is not

simple. About 75 percent of the tropical Hawaiian species that obviously had to reach Hawaii from the west

or south are composed of species not strictly central

Pacific in distribution. They are species of circumtropical character, or those of broad Indo-Pacific affinities.

The nearby Micronesian and Polynesian archipelagos

seem to have contributed only about 25 percent of the

Hawaiian tropical component from their endemic

members. This calculation suggests an obvious biogeographic fact that widespread marine species are

hindered in their distribution by few barriers within

their thermal regimes. But it also equates the proportions of imports from Micronesia-Polynesia with those

from North America. Though Polynesia forms the

bridge for the dispersal of cosmopolitan species, it has

not provided a major share of Hawaiian species from

its own pool of endemics. Micronesia may actually

have a low proportion of endemics as the archipelagos

are not significantly isolated from the Indo-Pacific.

Lesser Polynesia is therefore not comparable to the

warm-temperate of North America because the species

of the latter region are essentially all classed as endemic,

not distributed anywhere in the tropics, whereas the

fauna of Micronesia-Polynesia is composed mostly of

species distributed elsewhere in the tropics. The question is at best academic but one that could be misinterpreted by overestimating the importance of the

American contribution to Hawaii in comparing its 20

percent of the input with the 20 percent input from

Micronesia-Polynesia. The significance of the American

input is that whereas the Micronesian-Polynesian element is presumably broadly preadapted to existence in

Hawaii, the cool-water American element is not. The

contrast is not completely extreme, however, for

Hawaii is not metatropical and many Polynesian elements presumably would not survive on its shores.

The real Micronesian-Polynesian element must include

all of its nonendemic components. In balancing the

mechanisms one must consider the species of the pantropical element in a different light from others. Most

of them would have reached and have become established in Hawaii under most circumstances; they occur

in and therefore originated from Micronesia-Polynesia.

The cool-water American element may have reached

13

the importance of the endemic Micronesia-Polynesia

element because that tropical element is low in diversity, whereas the American element is extremely high

in diversity.

Two conspicuous elements of the Polynesian-Micronesian fauna do not appear to have reached Hawaii.

Xenocheira, with two species in Australia, Indonesia,

and Polynesia, and the very abundant Micronesian

Lembopsis tridens (Schellenberg) (formerly placed in

the genus Microdeutopus) have not been found in

Hawaii.

The Cool-Water Northern Element

Establishment of Hawaiian species in archipelagos to

the south and west of Hawaii may be difficult to detect

in various marine groups unless one can recognize them

through generic analysis and values of probability, because Hawaii apparently is a destination and not a

stepping stone for species coming from cool-water

America. In another vein Hawaii may be far less significant as a stepping stone from west to east than the

Marquesas Islands. No Hawaiian group other than

Amphipoda, as yet studied, seems to have any significant affinities with the Americas except perhaps in the

direction from west to east (corals). Approximately 25

species of Hawaiian Amphipoda have clear affinities

with North America and if cognates of those species

occur to the west and south of Hawaii in such places

as the Line Islands or the Marshall Islands, one must

presume very strongly that Hawaii served as the stepping stone and the evolutionary ground for those taxa.

But there is little doubt that any of the cool-water

Hawaiian imports will be found in those other places,

even as far as Johnston Island. The fishes of Hawaii, as

an example of most of the known marine groups, have

no affinities with North America; they are of IndoPacific tropical origin and in possibly passing to and

from Hawaii have left little direct evidence of their

migratory directions. One may thus suggest that Hawaii

is almost completely positive as an area of biological

demand and almost completely negative as an area of

biological supply as far as Amphipoda are concerned.

The discovery of a significant fauna of cool-water

American origin in Hawaii adds another facet to the

calculations heretofore described in attempting to predict the faunal structure of Hawaiian Amphipoda. The

previous discussion on the tropical Hawaiian component has already demonstrated that no significantly
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

diverse tropical group of amphipods has failed to reach

Hawaii, although the Lysianassidae are short of species

in comparison to the coasts of India. We must consider

which of those elements in the rich American faunule

has succeeded in becoming established and which has

not, so as to suggest, perhaps, fruitful lines of ecologic

inquiry. Amphipoda of epifaunal habitat have to be

separated from the North American faunal pool because the Hawaiian mud-bottom fauna is not being

considered (Table 6).

Again, as in previous conclusions, this discussion is

under the influence of various exploratory deficiencies.

The score of Hawaiian species with American affinities has been detected primarily under an aegis of a

literature that includes only warm-temperate and coldtemperate America. Tropical America is virtually unknown. Additional Hawaiian species may have tropical

American affinities and, indeed, several of the American warm-temperate species may extend into American tropical waters. Many of these extended species, if

occurring, may not be dismissed as semiubiquitous species if they are found to occur in the tropics only in

habitats influenced by cool-water upwelling. Under

that circumstance, they would still qualify as cool-water

species.

Almost all of the Hawaiian imports from America

have an ecological relationship to marine plants but

only three of them might be considered as simple nestlers (Atylus nani, Ceradocus hawaiensis, and Pontogeneia pacifica). At least a half dozen of the species are

probably semi-inquilines, being associated with small

sessile invertebrates that may be attached to algae or

masses of dead material (AmphUochus, Gitanopsis,

Gitana, and Leucothoe). The remainder are tubicolous

amphipods, building soft tubes attached mainly to

algae or marine grasses or occasionally hard substrates.

In the Californian epifaunal Amphipoda there are

40 species of nestlers, 10 species of inquilines, and 45

species of tubicolous habit (Table 5). The proportion

of Hawaiian nestlers (about 20 percent of the total

Hawaiian species with American affinities) seems fairly

low in relation to the pool of continental nestlers. The

high proportion of Hawaiian species of tubicolous habit

(nearly 60 percent of the Hawaiian-American imports)

and inquilinous relationship (20 percent) suggest

that these kinds of taxa with obligatory, or semiobligatory, fixed positions are more successful in completing long and difficult journeys than are nestling amphipods. The latter may have a higher tendency to swim

away from an object entrained in a journey than are

amphipods with a microhoming instinct oriented to a

tube, burrow, or host. The nestler would thus be lost

during the journey more frequently as it would evacuate its vehicle, food, and place of rest. As the migratory vehicle passed through different hydrographic environments, the reaction of the nestler may be to swim

actively so as to escape unfavorable conditions, whereas

the domicoious animal may cling even tighter to its

home. If the unfavorable conditions are transitory or

not ultimately fatal, the domicoious animal might survive the full journey more frequently than the nestler.

A strong exception to this possibility may occur in

the discovery of dense populations of the nestler Hyale

species accompanying masses of Sargassum as flotsam

far from shore (J. L. Barnard 1961), but the theory is

not an all or none situation, and Hyale is not an average nestler in the sense of its enormously dense populations on that substrate. Such flotsam may also not have

passed through unfavorable hydrographic conditions.

Two of the three Hawaiian nestlers of American

affinity would have been of improbable prediction as

one should expect the candidates from the American

pool to be composed of very abundant genera in warmtemperate regions, but Atylus, especially, and Pontogeneia are rare in the Californian warm-temperate.

They are primarily centered in cold-temperate waters.

The species group from which the Hawaiian Atylus

may have come is not found farther south than middle

California. The one Hawaiian species of Ceradocus also

has affinities with the southern (Mexican) species of

Ceradocus. The genus is not abundant in Pacific warmtemperate America.

The puzzle of AmphUochus, Gitanopsis, Konatopus,

and various ischyrocerids is discussed elsewhere.

Of the remaining species, the two members of

Aoroides, the two of Gammaropsis, the one Stenothoe,

one Leucothoe, and one Ampithoe would have been

included in a prediction as they belong to genera with

high population numbers on continental shores nearest

Hawaii. But the occurrence of even one species of

Aoroides might not have been predicted because of low

generic diversity. The new species from Hawaii represents only the third Pacific member of the genus.

Continental Amphipoda that have not arrived in

Hawaii or, by the terms of the present exploration,

would not seem to be significantly abundant in Hawaii

are noted in Table 6. There is a surprising conformity

in the total number of epifaunal species in the Cali-

16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fornian warm-temperate (109) and Hawaiian (116)

regions, with a fair balance in the familial and generic

representation. Hawaii has only four fewer genera than

does the Califomian warm-temperate. There is scarcely

any morphotype or niche represented in the Califomian

pool that is not already represented in Hawaii either

through warm-temperate imports or in the tropical

component.

The Southwestern Warm-Temperate Element

There can be no species in Hawaii that also occurs

both in the warm-temperate Australian region and in

the tropics that would not be classed as a widespread

tropical-warm temperate species. No species has been

identified as discontinuously distributed between the

two extremes, but several endemic Hawaiian species

such as Biancolina mauihina, Colomastix lunalilo,

Laetmatophilus hala, Palinnotus alaniphlias, Seba

ekepuu, and Podocerus hanapepe have their closest

affinities with the antiwarm-temperate region. If one

added these species to those widespread tropical-warmtemperate species also occurring in the Australian area,

one might strike a remarkable affinity between the two

areas. But several of these endemic Hawaiian species

may be figmentarily related to the Australian regime as

they belong to groups not yet studied in the tropics or

subtropics of either the eastern Pacific Ocean or the

Indo-Pacific. It may be happenstance that they are

known in Australia, possibly as southward extensions

from tropical centers of gravity, and not yet found in

the main Indo-Pacific. They must be reserved for

future consideration.

The Tropical American and Caribbean Element

Only six Hawaiian species of Ampipoda are considered

presently to have affinities with tropical America. One

of these species, the endemic Gitan&psis pele, is a very

conspicuous cognate of the Caribbean G. tortugae and

because the Pacific side of tropical America is virtually

unexplored for amphipods we might presume that the

more direct relationship is with a taxon in the east

tropical Pacific that is either G. tortugae itself or another cognate. That a direct pathway between tropical

America and Hawaii is possible for amphipods is

shown in the occurrence of the Caribbean-E. Pacific

Ampelisca schellenbergi in Hawaii. There is no morphological distinction among various populations of

this species, and it does not qualify as a taxon that

could be transported artificially.

Leucothoides Ppottsi represents the next best case of

a direct connection between tropical America and the

central Pacific Ocean, but there is a modicum of doubt

as to whether Pacific and Caribbean populations represent the same species. Like its familial relatives it

may be carried from place to place by transport of its

possible host, a tunicate (or ?sponge), on ships, but

in Hawaii, Micronesia, and the Caribbean Sea it is

found in the open sea and only its congener L. pacifica

has been found in unpolluted Califomian harbors.

Elasmopus e. hawaiensis is a subspecies of the Galapagan E. ecuadorensis, but I have a few observations

that suggest this epigenotype is widely scattered

throughout the Pacific Ocean.

The other two species are of very dubious relationships to Caribbean taxa, but some of the cryptic species

in Table 4, like Hyale waimea, may eventually be

added to the list of Hawaiian amphipods with tropical

American affinities.

Introduced, Minor, and Cryptic Elements

More than 20 species of Hawaiian Gammaridea do

not fit into classifications mentioned in previous pages

(various parts of Table 4). Undoubtedly, many mistakes have been made in establishing the affinities of

the species heretofore mentioned, but I estimate that,

as various species are shifted in classification, the end

result, numerically, will be very similar to that now

propounded. Ten species of unknown affinity appear

to be composed of taxa of the same geographic diversity as computed elsewhere in this paper. For example,

one of the species of Eriopisa may come from cool

water and the other from the tropics, and while the

stenothoids are from cool water, the Paraphoxus may

come from tropical America.

The so-called introduced species are very difficult

to evaluate. Doty (1961) has discovered an alga,

Acanthophora species, probably introduced by shipping and already dispersed from its probable port of

entry, Pearl Harbor, to the islands of Kauai and Lanai.

Doty also gives references to earlier remarks on other

marine organisms known to be introduced and established in Hawaii by shipping and to species known to

be introduced but not established.

Numerous amphipods could qualify as candidates

for introduction by means of shipping, as many of

them form tubes easily attached to hard surfaces, and
NUMBER 34 17

many others could nestle among projections or within

cavities of thick fouling masses. Some of the tubicolous

imports have probably arrived in Hawaii on logs floating naturally from North America as well as logs towed

slowly by boats. But little is known about the ability

of amphipods to remain with ships moving at speeds

creating great water turbulence. I have recovered

specimens of Corophium acherusicum from fouling

matter on the side of the R/V Velero IV after it had

steamed at a speed of 9.5 knots for several days.

The Amphipoda immediately qualifying as those

subject to artificial importation occur commonly in

harbors throughout the world or are various congeners

of those harbor taxa. Members of Corophium, Ericthonius, Podocerus, Jassa, and Stenothoe are the

strongest candidates (Table 4, various parts). There is

little doubt that Corophium acherusicum and C. insidiosum have been artificially imported to Hawaii as

they maintain fairly strong harbor restriction, but the

presence of C. baconi (if indeed it is correctly identified) may be a natural import on logs. The highly

atypical sexual morphology of C. baconi in Hilo Harbor

suggests that the population may represent a very

narrow segment of the total genosome (the "founder

effect"). On American shores, C. baconi is a sparse

inhabitant of harbors for it apparently cannot tolerate

the same degree of pollution as its congeners, but it also

is more frequent in the open sea than the other species

of Corophium.

Ericthonius brasiliensis and Podocerus "brasiliensis" 7

are strong fouling organisms in harbors, but the former

is also extremely abundant in many open-sea localities.

One may thus debate its mode of dispersal with equal

conviction on either extreme. Both species have, so far,

been found in Hawaii mostly in artificial situations such

as harbors or on suspended floats and boards.

Stenothoe gallensis is a common tropical fouler and

S. valida is a common subtropical fouler, but neither

builds tubes as do the other species mentioned above.

They are commonly associated with masses of hydroids.

I am unsatisfied with my taxonomic appraisal of these

species in Hawaii, but if they are found to be highly

variable species with conspicuous shifts in morphology

from one region to another, then I may have identified

them correctly. So-called S. valida has been found in

Hawaii only in association with the black-coral Antipat hes irregularis off the east coast of Oahu in 33 m of

water depth. Under normal circumstances that unusual

1

 Possibly misidentified in various harbors.

habitat would suggest a misidentification of S. valida,

for the species normally inhabits harbors. The associates

of Antipathes should presumably have been sorted out

as highly characteristic species long ago, but the

Oahuan population of Antipathes is composed of

colonies moved by biologists to that locale from other

parts of Hawaii. One might thus expect that a eurytopic harbor fouling organism could find an open niche

in such an artificial situation. But S. valida has not as

yet been collected in Hawaiian harbors.

Species of Elasmopus may also be carried by shipping

even though they do not make tubes on hard substrates. Elasmopus rapax is a common inhabitant of

unpolluted harbors, and it does show a strong orientation to hard, floating substrates. I have collected it

in Newport Harbor, California, as one of the first settlers on bare, anchored, floating wooden blocks. Specimens were collected occasionally even on blocks without other macroscopic foulers; perhaps they graze on

microscopic fouling organisms. The Hawaiian population of E. rapax may be confined entirely to Pearl

Harbor; I found it nowhere in the open sea in my

1967 explorations; my 1955a identification cf it in

Honolulu Aquarium was erroneous, and I have not

been able to locate materials of it identified by me in

the open sea in my report of 1955. I probably confused it with one of the other nine species now shown

to be present on Hawaiian shores. Thus, E. rapax is

undoubtedly a strong candidate for reconsideration as

an import by artificial means.

The final case to be considered is that of Tropichelura insulae, the wood-grazer. This "wood-borer"

is circumtropical in distribution and probably, like the

well-known Chelura terebrans of cool waters, simply

enlarges the tunnels of its obligatory associates, Limnoria species, by grazing. The question as to whether

this species has been distributed around the world

by modern shipping is entirely open to question.

Chelura terebrans is also almost completely circumboreal, but the Japanese species, Nippochelura brevicauda, has a highly restricted distribution.

Dispersal by Rafting

Amphipoda reproduce without larval stages. Their

dispersal might therefore seem to be impaired and one

might expect an extraordinary degree of endemicity

and impoverishment of Amphipoda in highly isolated

places. On the contrary, we have adequate samples
18

of their distribution patterns and a sketchy knowledge

of their shallow-water dispersal so as to suggest that

even the isolated Hawaiian chain, though it might be

impoverished, could have a very low endemicity of

species and virtually no endemic genera. If Amphipoda reach the Hawaiian Islands at all, we might expect a recurrence of the same founders, and gene flow

might be relatively continuous. As a broad generality

that thesis is found to be false, though being a complex

of ideas, part of the statement remains true. A duplicate recruitment of the same founders that produce

cognate species appears to have some basis (see following chapter), but that mechanism requires long

periods of interrupted gene flow. Isolation but not

impoverishment are apparent in the Hawaiian fauna.

Dispersal frequency of amphipods into the region has

been high but often interrupted so as to play a role in

the development of many endemic species and several

novel morphotypes necessarily described in generic

terms.

Rafting by means of detached marine plants, debris,

and logs is the assumed primary dispersal mechanism of

shallow water tropical Amphipoda. Islands within a

few hundred kilometers of continents (Azores, Canaries) have few endemic species reported in their shallow

waters, thus suggesting that they have been populated

continuously by rafted organisms. Even the Galapagos

Islands appear to demonstrate this situation by incongruous fashion (report in preparation by J. L.

Barnard). For example, those equatorial islands 600

km west of Ecuador have a primarily tropical fauna of

Brachyura (reproduction mainly by larvae), whereas

their amphipods have a strong warm-temperate affinity (see Garth (1946) for brachyuran summary). Importation of tropical crab larvae must come during the

favorable but infrequent El Nino years when warm

currents reach the Galapagos from the Bay of Panama.

A few crabs have arrived via the relatively constant

and swift Humboldt-Peruvian current system and

fewer still have come from the north and west through

the meandering complex of the California current

that becomes mixed with the most anti-Galapagan

tropical counter-current.

Since amphipods do not disperse by larval means and

presumably are carried over long distances by rafting of adults, it appears incongruous that so many

warm-temperate amphipods occur in the Galapagos

Islands, whereas many species from nearby Panama

have not arrived there. Because few warm-temperate

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

crabs occur in the Galapagos, one might suggest that

differences in crab and amphipod reproduction play

a role in faunal incongruities. Because the majority of

crabs has arrived in the Galapagos as short-lived larvae,

they have to come from shores nearly adjacent to the

Galapagos during El Nino years. Currents during those

years must also bring amphipods aboard flotsam, but

the infrequency of storms in the Panamic doldrums

may constrict the quantity of rafted material available, whereas storms in warm-temperate latitudes may

cast much larger quantities of shallow-water debris into

flotation. Although the journeys of rafts from higher

latitudes may be extremely tortuous and indeed appear almost impossible from sketchy charts of currents

now known, these journeys must be of higher significance to the Galapagan amphipod fauna than journeys

during years of El Nino. The implication is also made

that the amphipods reproduce during their journeys.

But one should not dismiss the indications that the

Galapagos have several characteristics of warm-temperate environments and thus are favorably situated

for successful immigration of various warm-temperate

taxa.

Waters of the Galapagos are only sufficiently tropical

to support some impoverished patches of hermatypic

corals on particular islands and Sargassum, a good tropical indicator in the eastern Pacific Ocean, does occur

in isolated patches and might flourish more were it not

for the intensive browsing of marine iguanas. Whereas

many of the tropical crabs that have arrived from the

continent may have been able to survive despite thermal

minima below tropical limits (20°C), one may speculate whether a stronger immigration of subtropical or

warm-temperate crabs might not alter that balance

if it were possible for more of those species to reach

the islands. If the thesis about El Nino currents and

lack of storms in the Gulf of Panama were incorrect

then perhaps those tropical amphipods arriving in the

Galapagos have failed in many cases to meet the competition from warm-temperate amphipods arriving

earlier or later.

Crabs of continental west America, except in areas

of the Las Perlas Islands and near Punta Malo on the

northwest Panamic shore, must be primarily eurytopic

organisms able to withstand highly turbid conditions of

the Central American inshore waters. Those turbid

conditions seem to work against the algal flora and

thus reduce the supply of materials for rafting tropical

amphipods. Unless continental runoff decreases dur-
NUMBER 34

ing El Nino years, large rafts of terrestrial debris

brought into the sea by rivers may become infested with

amphipods once the rafts reach salt water.

Despite these numerous suppositions, the fact remains that many Galapagan amphipods are very

closely related, if not identical, to species occurring in

the warm-temperate zone of the Califomias and there

must be, or have been in fairly recent times, a ready

pathway for their transmissal. The area of upwelling

near Punta Malo possibly reflects a Panamanian warmtemperate outpost that supports northern amphipods.

It is only academic to question the particular route of

migration by calling attention to the complete absence of knowledge about antiwarm-temperate amphipods of the Peruvian-Chilean shores. If the supposed

Californian-Galapagan cognates arrived from Chilean

shores via the Humboldt current then that warmtemperate fauna must have Californian species established within it. Such knowledge would be strong support of a drifting hypothesis with a major connection

in the upwelling area of Panama or in the Galapagos

themselves.

Whereas a journey by currents from warm-temperate

North America to the Galapagos is tenuous and difficult by appreciation of present-day current charts, a

journey from warm-temperate North America to the

Hawaiian Islands appears as a direct and almost continuous possibility. Waters entrained from the California current could pass through the Hawaiian chain

as northern outliers of the North Equatorial Current.

That this direction of surface sea flow dominating the

oceanographic picture has had so little influence on the

Hawaiian chain except in the amphipodan fauna is

difficult to explain in gross terms. Most Hawaiian

marine organisms have affinities with the main IndoPacific or with Polynesia

 8 and Micronesia, but dominant surface currents coming to Hawaii from those

directions do not exist although the North Pacific

Current could bring immigrants over a long northern

route from the China Sea region. The major share

of the marine Hawaiian fauna must be a product of

survival of immigrants coming from the southwest or

west during occasional reverses in the gross current

system, or by minor and subsurface currents as yet

undiscovered. The weakly tropical Hawaiian thermal

regime thus favors those occasional immigrants traveling within warm tropical waters rather than the potential warm-temperate immigrants aimed at Hawaii via

1

 Term excludes Hawaii and New Zealand.

19

the California-North Equatorial Current system. Those

taxa originating in warm-temperate North American

waters and directed toward Hawaii must encounter

primarily lethal conditions as the cool waters become

warm; few survive the journey and fewer survive the

Hawaiian environment. But the Hawaiian amphipods

do reflect in a small way the influence on Hawaii by

the west and southward flowing waters from western

North America.

The Sweepstakes Phenomenon

The theory of sweepstakes dispersal implies that the

biota of an isolated place is a result of the vagaries of

transporting mechanisms and the timing of events.

On isolated islands this applies most harshly to those

terrestrial regimes having contraction of niches and

habitat space, but the biota established on islands is

also controlled through biotopical preadaptations.

Mere examination of a checklist of species from a

biota cannot support a statement about the absolute

failure of dispersal mechanisms to import various taxa

into an extrinsic locality. The paucity of acroporan

corals in the Hawaiian Archipelago suggests an unfavorable biotope and not necessarily the failure of

the transporting mechanism. The failure may occur

in the timing or extent of the scleractinian dispersal

in the sense that by chance alone the pocilloporans

reached the islands first and the normal dominating

mechanisms of an initial immigrant prevent a foothold

of later immigrants. But there is also the strong possibility that if the vagaries of sweepstakes dispersal had

brought acroporans first, their dominating mechanisms

would have failed to prevent establishment of pocilloporans because acroporans are outside of their optimal environment in Hawaii. Sweepstakes dispersal is

furthermore a tenuous term disguising various absurdities. It has to be applied only to those species with any

chance of reaching the locality under question. Its

topical applicability is as variable as the number of

places under question. In effect it is best applied to

groups of species with generally equivalent preadaptation to the environment under question but with a relatively low, albeit positive chance of reaching that

environment. Many of the major incongruities of

terrestrial faunistics and floristics on islands seem to

have been the result of highly improbable migrations.

The dominant avian faunas of the Hawaiian Islands

and the Galapagos Islands thus represent immigrations of high improbability as to length of journey, pre-
20

dictable direction, and frequency of occurrence. These

improbabilities may, however, be amplified by our ignorance of the frequency of events, the lack of ecological preadaptations in various other candidates that

might have populated those archipelagos, and the extent to which founder species prevent subsequent establishment of competitive taxa.

An evaluation of absurdities suggests the need to

restrict the broadest implications of sweepstakes dispersal to a pool of species with definite chances of

reaching a locality under question, no matter how slim.

Certainly some very large African land mammals could

be eliminated from a pool of potential natural immigrants to Hawaii, but a broad spectrum of groups not

so obviously unqualified for the journey makes the concept almost incomprehensible. The shore fishes of

America probably also should be included in the list

of unqualified candidates. Despite their obvious dispersal mechanisms, the single journey to Hawaii has

not occurred or the preadaptation to survival in Hawaii

does not exist in American shore fishes.

TABL E 5.—Mega-niche classification of Hawaiian and

Californian epifaunal amphipods, noting strongest affinities

of Hawaiian species in domicolous taxa

Classification

Nestling

Domicolous

[nquilinous

Inquilinous

ncstlers

Lignivorous

Others

Hawaiian

43

43

12

15

2

4

Californian i

40

45

10

11

2

0

Hawaiian Species

with Californian

Affinities

5

16

1

5

0

0

1

 Californian data mostly from J. L. Barnard (1969a).

Amphipods represent a case toward the opposite

extreme to fishes, in that many of their species have a

distinctly positive chance of reaching the Hawaiian

Islands from long distances and from localities outside

the local thermal regime. To what extent the vagaries

of transportation have affected the composition of

Hawaiian immigrants from America cannot be estimated simply by examining the pool of American and

Hawaiian species. The general absence of information

on the competitive positions of American species

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

against Micronesian species is just one of many defects

in a possible analysis. There is a strong indication that

nestling species are not as well adapted to making the

journey across a thermal boundary as are tubicolous

species, and this might account for the disproportionate

number of American tubicolous species occurring in

the Hawaiian fauna. That nestling species also encounter difficulties in completing the journey to isolated

places within their normal thermal regime is also apparent in the following analysis of some statistics that

are admittedly inadequate. The epifaunas of the IndoPacific have a higher number of nestling (115) than

tubicolous (71) species, whereas Hawaii has the numbers in approximately a 1 to 1 ratio. This suggests a

possibility that numerous nestling niches remain open

or do not exist in Hawaii, whereas those tubicolous

niches that remained open because of failures of importation from the southwest direction have been filled

with immigrants from America. It is possible, therefore,

that vagarious transporting mechanisms operating over

a relatively short period of time have failed to supply

Hawaii with a full complement of nestling amphipods

from a thermally similar area, but have supplemented

the deficiency of tubicolous immigrants from that same

direction with members from the American pool. The

deficiency of nestlers in Hawaii is marked most conspicuously by the presence of only one lysianassid in

comparison to the normal complement of five or six

species in localities of India. The contradiction that

Hawaii has a greater diversity of nestlers in its common

genera than do other Indo-Pacific localities may be explained momentarily by the differing taxonomic emphasis that has occurred in tropical studies.

If American nestling amphipods have failed to reach

Hawaii, there remains a strange coincidence that

warm-temperate America has a pool of nestlers with

similar generic and familial composition to that of

Hawaii (Table 6). The only conspicuous warmtemperate group not represented either by a congener

or a confamilial substitute is the family Bateidae. One

could partially reverse the previous supposition so as

to suggest that a full measure of nestling amphipods

immigrated to Hawaii from the southwest, whereas

a deficiency of tubicolous imports was complemented

by immigrants from America. American tubicolous

imports therefore succeeded in open niches, whereas

American nestlers, if Imported in equal frequency as

tubicolous species, were unable to compete with

normally occurring tropical nestlers.
NUMBER 3 4 21

TABLE 6.—Pool of Californian warm-temperate epifaunal amphipods

Epifaunal amphipods

Family

Calif ornian warm-temperate Hawaiian

Genus Genus

Acanthonotoiomatidac

Ampcliscidac

Amphilochidac

Ampithoidae

Anamixidac

Aoridae

Atylidae

Bateidac

Colomastigidae

Corophiidae

Dcxaminidac

Eophliantidae

Eusiridac

Gammaridae

Hyalcllidae

Hyalidae

Isacidae (Photidae)

Ischyroceridae

Panoploea (1) See Ochlcsidae

Ampelisca (2) Ampelisca (1)

Amphilochus (2) {Amphilochus 4)

Gitana (1) Gitana (1)

Gitanopsis (2) Gitanopsis (1)

None Mokuoloe (1)

Ampithoe (7) Ampithoe (7)

Cymadusa (1) Cymadusa (3)

None Paragrubia (1)

Anamixis (1) Anamixis (1)

Acuminodeutopus (1) None

Amphideutopus (1) Aloiloi (1)

Aoroides (1) Aoroides (2)

Z,«m*oj (3) Lerofow (8)

Microdeutopus (1) Neomicrodeutopus (1)

Neomegamphopus (1) Konatopus (1)

Rudilemboides (1) None

A(7/ux (1) 4(y/ttf (1)

Bated (2) None

Colomastix (1) Colomastix (4)

Cerapus (1) None

Corophium (4) Corophium (3)

Ericthonius (1) Ericthonius (1)

Polycheria (1) Paradexamine (1)

Lignophliantis (1) Biancolina (1)

Eusiroides (1) Eusiroides (1)

Pontogeneia (3) Pontogeneia (1)

Ceradocus (2) Ceradocus (1)

Elasmopus (5) Elasmopus (10)

None Eriopisa (2)

Eriopisella (1) Eriopisella (1)

Afaera (4) Afaera (7)

Megaluropus (2) None

A/<r/£ta (3) Afrfito (2)

Metaceradocus (1) None

None Nuuanu (1)

None Parhyalella (1)

Allorchestes (2) None

HyaleQ) Hya/f (8)

ParallorchesUs (1) None

ParA)w/* (1) .ParA?a/<r (1)

Cheiriphotis (1) Non e

CA»a/w (1) CAroa/ta (1)

Gammaropsis (2) Gammaropsis (7)

Megamphopus (2) ° None

PAo/ir (5) PArtu (3)

Ischyrocerus (3) Ischyrocerus (3)

Ja«« (2) Jawa (1)

Mierojassa (1) None

Parajassa (1) Parqjassa (1)

Ventojassa (1). Ventqjassa (1)
2 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 6.—Pool of Californian warm-temperate epifaunal amphipods—Continued

Epifaunal amphipods

Family

Californian warm-temperate Hawaiian

Genus Genus

Lcucothoidac

Liljeborgiidae

Lysianassidac

Ochlesidae

Oediccrotidae

Phliantidac

Pleustidac

Podoccridac

Sebidac

Stenothoidae

Leucothoe (2) Leucothoe (3)

Leucothoides (1) Leucothoides (1)

Liljeborgia (2) Liljeborgia (2)

Ensayara (1) None

Lysianassa (3) Lysianassa (1)

Sec Acanthonotozomatidac Ochlesis (1)

None Kanaloa (1)

Heterophlias (1) Palinnotus (1)

Parapleustes (2) Parapleustes (2)

None Laetmatophilus (1)

Podoeerus (2) Podocerus (3)

None Seba (1 )

Parametopella (1 ) None

Stenothoe (2) Stenothoe (5)

Stenothoides (1) None

Note.—Generic relationships to Hawaii are noted by dotted lines and direct affinities by dashes. Number of species in parentheses. Californian data mostly from J. L. Barnard (1969a).

• Morphologically similar to Gammaropsis and possibly not distinct generically.

The affinities of Hawaiian Amphipoda demonstrate

a strong preadaptation of Amphipoda from similar

thermal regimes to migrate long distances. The

problem of sweepstakes dispersal is not so much as to

identify missing phyletic units or species (see "Pantropical and Indo-Pacific Elements") but to study the

niches and transportational ecology of amphipods so

as to determine the biological stresses influencing the

present composition of the Hawaiian fauna.

Lack of Adaptive Radiation in Hawaiian Amphipoda

Approximately 20 genera of Hawaiian Amphipoda are

represented by two or more species, but there is little

evidence of local adaptive radiation in these various

species flocks. This situation is a strong contrast to the

terrestrial Hawaiian biota in which numerous cases of

adapative radiation occur among honey creepers,

snails, and fruit flies. Most of the present study of

amphipods has occurred on only one of the Hawaiian

Islands, but this does not seriously weaken the conclusion that little adaptive radiation occurs in the Amphipoda. The conclusion is supported by results in other

marine groups.

Evolution of endemic Hawaiian Amphipoda has undoubtedly occurred, but the lists of affinities in Table

3 demonstrate that cognates of the several species

in each genus are located in extrinsic regions. Those

few cases of strong interrelationships between a pair or

a triad of Hawaiian endemic species can usually be

explained by assuming the successive immigration of

a parent species.

Discussions of adaptive radiation in marine Gammaridea have not yet entered the literature, but a few

potential examples may be cited in the flock of genera

and species of haustoriids in the northwest Atlantic,

the genus Harpinia in the northeast Atlantic, a section

of the genus Ampelisca in southeast Asia, and especially the flocks of closely related species of Anonyx,

Hippomedon, and Orchomene in the Okhotsk Sea and

the genus Parawaldeckia in New Zealand. Each of

these sympatric flocks must signal the former occurrence of barriers that isolated various populations from

one another. Subsequently, the barriers must have retracted and those various isolates again became sympatric. Visualization of the mechanics of this process in

marine organisms remains highly rudimentary.

The Hawaiian fauna of Amphipoda is still relatively

youthful, but nearly half of the species is now divergent,
NUMBER 34 23

morphologically, from their presumed ancestors in

Micronesia-Polynesia and adjacent continents. No

endemic genus with more than one species occurs in

Hawaii, but several multispecific genera have pairs of

species in Hawaii with close interrelationships. These

pairs can be identified with an extrinsic cognate, but

that fact does not necessarily obviate the possibility

that one member of the Hawaiian pair did not evolve

from its sympatriot. The conclusion that these Hawaiian pairs are the result of successive immigrations of

a single parent species and do not represent topical

radiation is supported by our knowledge of the high

success of amphipods in completing journeys from unexpected sources. If they can both come long distances

as well as overcome thermal barriers, then there might

occur successive waves of a species from a nearby area

of thermal characteristics similar to those of Hawaiian

waters.

Among the following multispecific Hawaiian genera

there is little question that their various species have

diverse affinities (Table 3): Colomastix, Corophium,

Eriopisa, Ischyrocerus-Jassa, Leucothoe, Melita, Photis, and Podocerus. Some of the species of Corophium

have probably been brought into the islands by ships,

but the various species of Ischyrocerus and Jassa are

sufficiently divergent from each other as to suggest the

development of incipient subgenera.

Pairs of Hawaiian species among genera with coldwater affinities occur in Amphilochus, Aoroides, Parapleustes, and possibly Gammaropsis, Lembos, and

Stenothoe. Heretofore Aoroides has been composed of

only two continental species, one in America and one

in northern Asia. Hawaii has two species—one is possibly synonymous with the American species, and the

other is apparently a Hawaiian evolute. Both species

are perhaps endemic to Hawaii, but the wide American

distribution of A. columbiae plus the tubicolous adaptations of the genus suggest a succession of dispersals to

Hawaii of the American species rather than a Hawaiian

radiation of the genus. If the two Hawaiian members

of Parapleustes evolved from a single boreal immigrant,

there is little morphological evidence to suggest that

course. One of the species is an incipient genus in its

loss of a part of the maxillipedal palp, and its true

affinities are obscured by the change because of the

vagaries of classification within the complex of boreal

pleustid genera. Three species of Amphilochus form a

tetrad with an American species. They would probably

represent the best potential case of Hawaiian adaptive

radiation were it not for inadequate studies of the

genus in America elsewhere than the Californias. They

may be figmentary siblings in Hawaii because we have

yet not found their cognates in America. But the other

side of the question can be supported through the

ecological evidence that Hawaii is better supplied with

their scleractinian hosts than is America.

The pertinent affinities of several members of

Gammaropsis, Lembos, and Stenothoe are very

sketchy; this ignorance may conceal the presence of

three more pairs of Hawaiian siblings with cool-water

affinities.

Most of the remaining Hawaiian multispecific genera

have their strongest affinities with tropical regions.

They are Ampithoe with (5) species, Cymadusa (3),

Elasmopus (9), the complex of Eriopisa-EriopsiellaNuuanu (4), Gammaropsis (7), Hyale (8), Lembos

(8), Liljeborgia (2), Maera (7), and Photis (3).

Ampithoe kaneohe and A. poipu form a pair of

Hawaiian species with their closest affinities to each

other despite the presence of certain morphological

characters in one species that heretofore had generic

value. Ampithoe poipu is assignable to the genus

Pleonexes, but that genus simply categorizes a grade of

evolution. There is little evidence that the Micronesian

allopatriot was a precursor to either of the Hawaiian

species.

Hawaii is the center of a complex of species represented by the tropicopolitan A. ramondi, the Californian A. simulans, the Pan-American A. pollex, and

the Hawaiian A. akuolaka and A. waialua. A sixth

member from the Gulf of California is also apparent in

other material. Hawaiian Ampithoe waialua has especially close affinities with tropicopolitan A. ramondi

and may be the result of Hawaiian evolution of an

early wave of members of A. ramondi before the latter

became well established by frequent immigration.

Hawaiian A. akuolaka may have affinities with A.

simulans and A. pollex that are stronger than its affinities with A. ramondi, but all of these species form a

complex that requires detailed study. All of them could

have been products of former isolation in populations

of A. ramondi.

Cymadusa has ten species in the world fauna, three

of which occur in Hawaii. One of its members, C.

filosa, is tropicopolitan, like Ampithoe ramondi. Owing

to the variability of C. filosa in its scattered populations, plus the weak divergence of the other nine species, the question arises as to whether the other species
24

were the products of former isolation of C. filosa in

places like Hawaii, California or the Gulf of California, the Gulf of Mexico (or farther north), Bermuda,

Brazil, South Africa, and Micronesia-Polynesia. If so,

C. filosa has since become sympatric with most of its

congeners. Again, the Hawaiian pair of endemic Cymadusa is in the midst of a genus with a tropicopolitan

member that could have arrived in Hawaii in successive waves and finally, without diverging, itself become

established through frequent immigration. Both Ampithoe ramondi and Cymadusa filosa have been collected

on flotsam far at sea and thus are known to be viable

migrants.

Two or more combinations of pairs of endemic

Elasmopus occur in Hawaii. The clearest case is that

pair of E. piikoi and E. tnolokai, the latter possibly

either synonymous with or forming a cognate pair

with Polynesian E. minimus. Hawaiian Elasmopus hooheno may be added to the pair to form a triad; in

all but one character there appears to be morphological convergence between E. hooheno and E. piikoi,

but that character, a spine on pereopods 1-2, may have

special significance that will denote a different affinity

of E. hooheno when the taxonomy of the genus has

been properly evaluated. Meanwhile, E. hooheno also

forms a weak pair with E. ecuadorensis hawaiensis,

but the latter species (sensu lato) has its strongest affinities with tropicopolitan E. pectenicrus. Indeed there

is some merit in suggesting that an early wave of E. pectenicrus became isolated for a long period of time in

an archipelago like Hawaii or the Galapagos and later

became codistributed between the two archipelagos.

But the stem of E. ecuadorensis may have an even

broader distribution as I have examined a population

resembling E. ecuadorensis in flotsam collected by the

R /V Galathea. The strongest case of possible Hawaiian radiation in Elasmopus remains the pair of species

E. molokai-E. piikoi but again a definite extrinsic

geminate is known.

Gammaropsis pokipoki, an endemic Hawaiian species, may form another evolute of an early wave of a

tropicopolitan species, G. afra (or G. atlantica). Two

other Hawaiian species of the genus, G. haleiwa and

G. kaumaka, may prove to be members of a system of

this sort when their adult morphology has been

clarified.

The eight or more species of Hyale in Hawaii have

mostly diverse affinities. An interrelated pair of species

occurs in H. affinis-H. honoluluensis but they are also

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

distributed outside of Hawaii. Hyale ayeli and H. g.

bishopae represent Hawaiian endemic species probably

derived from widespread parent populations, but the

parent species have not immigrated frequently enough

to Hawaii either to form a continuous genetic connection with extrinsic regions or to have formed more

than one descendent genotype. Both of the parent epigenotypes are confined to nontropical waters.

The situation in Lembos is obscured by taxonomic

ignorance, but again a potentially widespread L. macromanus may have established a pair of descendents

in Hawaii; repeated immigration is attested to by

the presence of American-Oriental L. macromanus in

Hawaii.

Both species of Hawaiian Liljeborgia are closely

related to each other and form a triad with Polynesian

L. proximo.

Only one pair of related species occurs in Hawaiian

Maera. A very strong relationship, perhaps the strongest example yet known in Hawaii, occurs between M.

pacifica and M. kaiulani. Maera pacifica occurs both

in Polynesia and in Hawaii, whereas M. kaiulani is

presumably endemic to Hawaii.

Relationships among Hawaiian members of Stenothoe are clouded by weak taxonomy; until adults of

two species are obtained and the life history of two

others worked out, there will only remain a potential

case of a pair or triads of species for inclusion in this

discussion. That Stenothoe is dominantly a cool-water

genus does amplify this possibility of a single immigrant

having completed the long journey from America (or

cool-water Asia) and radiating solely within the

Hawaiian Islands.

The numerous cited examples of strong relationships

between various Hawaiian species and widespread

tropical species that also occur in Hawaii suggest that

the Hawaiian endemic evolved from the cosmopolitan

member at a time in the history of the archipelago

when the environment differed from that of presentday conditions. This may have occurred in the early

history of the archipelago when a single small western °

island or bank had been developed and may have been

concomitant with small differences in ocean current

systems. A species like Cymadusa filosa, possibly in

continuous migration aboard flotsam, may have arrived as a small population in a very constricted early

*The islands of Hawaii have developed sequentially from

west to east and the old western islands have now been eroded

to subaqueous level.
NUMBER 3 4 25

Hawaiian environment and have undergone fairly

rapid selection to form a species like C. oceanica. As

more space and niches became available through formation of additional islands, a second wave may have

been established that diverged into C. hawaiensis.

Finally, as the Hawaiian Archipelago assumed its present dimensions or if an increased frequency of immigration occurred in relationship to favorable changes

in current patterns, the parent species C. filosa may

have become established a third time but retained frequent genetic exchange with its extrinsic populations

or underwent little morphological change because its

optimal niche, in terms of space or food source, had

finally appeared in an evolving archipelago.

Results of the One-Step, Long-Distance Isolation of

Cool-Water Immigrants to Hawaii

The classic pattern of terrestrial evolution in Hawaii

contains the element of saltatory (or incremental) isolation, from which extremely divergent suprageneric

taxa evolve fairly rapidly by a series of steps from one

isolated biotope to the next, through island hopping,

valley hopping, and by other small-scale increments.

The genotypical end products of this incremental isolation often are so divergent as to defy any tracing of the

steps (Zimmerman, 1948). The primordial end of the

process is the recognition of the first step in the isolation of populations; perhaps a few examples occur in

Hawaiian amphipods.

Unlike members of the recent terrestrial fauna and

flora of the main high islands of the Hawaiian chain,

no amphipod of direct cool-water affinity (origin)

would have arrived in the Hawaiian chain by incremental isolation, for not since the origin of the chain

have there been any intermediate land masses or

islands along the cool-water pathway from America or

Asia. Zimmerman presumes that part of the terrestrial

Hawaiian fauna passed along the now environmentally

extinct western islands and diverged with each step.

With minor exceptions, this could not have happened

in the cool-water amphipod fauna so that many of

those species now in Hawaii that have diverged have

undergone only one increment in isolation. Some of

those species may have, of course, passed to and fro

from Hawaii to cold-water continents several times,

but this occurrence is entirely hypothetical and supported only vaguely by the triangular occurrence of

Amphideutopus - Konatopus - Neomegampropus and

possibly by Japanese Atylus: Hawaiian Kamehatylus:

American Atylus. The hypothesis cannot be entirely

dismissed without damaging the concept that certain

gammarideans are highly preadapted to making long

oceanic journeys; if they could reach Hawaii, perhaps

in other times when currents were suitable they would

arrive again on the continents after a period of isolation and natural selection in Hawaii. If so, there may

be continental evidence of this interchange, but detailed studies have not yet been made on the problem.

A more logical supposition is that a successful reverse

journey would be more difficult than the initial trip

from cool continent to island, for the amphipod that

has undergone genetic change in an island would seem

less likely to be a successful reimmigrant to a continent

where specific diversity is high and where few niches

might be open to an organism that has changed only

slightly from its continental ancestors. On the other

hand, the establishment on islands should be easier, as

niches may be open even though there are fewer in

existence. Evidence that journeys to and from the

continent to Hawaii have rarely occurred is found in

the low incidence of species flocks in amphipods of

cool-water affinity. One would expect to see many

groups of closely related species that have diverged by

a few morphological changes, but the cool-water Hawaiian amphipods are primarily of the one species per

genus category, thus indicating a single period of importation for that species, or else the occurrence of only

one available broad-scale niche for each morphotype.

Three exceptions to this situation seem to occur: (1)

in the Amphilochus complex, with three closely related species; (2) in the ischyrocerid complex with

possibly three related species; and (3) the pair of

Aoroides. The ischyrocerids, by taxonomic concepts

developed from continental studies, would fall into

three different genera, but as will be seen from following paragraphs, the changes that have occurred in

other taxa suggest that one incremental isolation occasionally results in fairly radical morphological changes.

These changes, of course, are only radical in terms of

gammaridean morphometrics. We lack any rudiment

of understanding of events in gammaridean genetics

or whether gammaridean taxonomists oversplit the

morphotypes.

The impact on the rich North American fauna by

Hawaiian amphipods of these few species is almost nil.

Probably, the warm-temperate isolation that has occurred in the northern Gulf of California is of greater

importance to the development of endemic species in
26

the Pacific North American warm-temperate region

than any developments in Hawaii, and even there in

the Gulf the total impact in a one-step isolation probably is very low (J. L. Barnard, 1969b).

The importance of these divergent Hawaiian coolwater imports is thus not as evidence of a major staging ground for evolution but simply an opportunity to

observe changes that have occurred after one increment of isolation. Most of those changes are of no

greater degree than seen between sibling sympatriots

but, in a few cases, the changes assume the degree of

generic differences heretofore considered highly

qualitative.

One cannot estimate the morphological changes that

may have occurred in these species after the initial impact of isolation on a small population, for there may

have been progressive changes associated with genetic

drift, mutative accidents, or selective factors associated

with slow changes in environment after immigration.

The most striking case of radical change occurs in

Atylns (Kamehatylus) nani. By narrow definition, this

species, with affinities either to a Japanese or to an

American cool-water species of Atylus, has changed its

familial membership from the Atylidae to the Dexaminidae. Such a change might be dismissed as evidence that the two familial concepts are worthless, but

the Gammaridea have several families and genera

characterized by a loss of parts. For instance, Beaudettia palmeri from Micronesia (J. L. Barnard, 1965a)

and a new species of Maera like M. subcarinata from

Australia (in preparation) have strongly reduced or

missing mandibular palps, yet they have very close

affinities to other living species. The viability of these

losses is a function of natural selection and may be conserved more readily in highly isolated environments

where competition from immediate phyletic relatives

is reduced or absent. The Australian Maera, however,

is sympatric and cohabitational with its sibling partner.

The simple act of synonymizing two families that

have been considered distinct for so long, on the basis

of one exception, is poor practice if one considers more

than the morphology of the groups. G. O. Sars (1895)

may have been prescient in considering the two families

as one (they were distinguished in 1906 by Stebbing).

The Hawaiian case suggests indeed that the one family

came from the other, perhaps by repeated isolation of

various atylids that commenced their dexaminid status

by loss of a palp. The Dexaminidae are multigeneric,

whereas the Atylidae are now monogeneric (recently

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

they were digeneric). The small morphological distance between the only genus of Atylidae and the many

genera of the Dexaminidae is not necessarily matched

by a functional similarity, for many dexaminids are

believed to be associated in various degrees of inquilinous obligation with ascidians, whereas atylids and a few

dexaminids are algal nestlers. There is no evidence

that Atylus nani of Hawaii has become ascidiophilous.

Thus, it may be the founder of an entirely new phyletic

line and that potential is reinforced by one additional

morphological change, the incremental coalescence of

urosomite 1 with 2. In dexaminids and atylids only

urosomites 2 and 3 are coalesced but A. nani has increased that consolidation by one segment. It therefore approximates the conditions found in the Prophliantidae, a family with one species in Polynesia and

one in Australia.10

Another radical change that has occurred in a coolwater Hawaiian immigrant is the loss of maxillipedal

palp article 4 in Parapleustes(?) honomu. Morphology of maxillipedal palps is often highly stable in various families of Amphipoda and the only morphological variation of generic value in Pleustidae has been

the presence or absence of a protrusion on article 3 of

the maxillipedal palp. The unique loss of article 4

in P.(?) honomu obscures its generic relationships because the point of attachment of article 4 has heretofore marked the presence or absence of the process on

article 3. Parapleustes(?) honomu is thus an incipient genus by morphological standards.

Ischyrocerus(?) kapu represents a third case of

radical change in morphology in a cool-water immi10

 Since this study was completed in 1967 I have had the

opportunity to study many species of Dexaminidae, Atylidae,

Prophliantidae, and Lepechinellidae in Australia and New

Zealand. These include two species of Atylus, seven species

of Par ad examine, three species of Guernea, six species of

Lepechinella, one species of Prophlias, and specimens in

USNM of Guernea (^Prinassus) nordenskioldi (Hansen)

and Dexamonica reduncans J. L. Barnard. The latter two

species have been found to be congeneric, requiring revival

of Prinassus Hansen as senior synonym of Dexamonica J. L.

Barnard, Prinassus then being attached to Guernea Chevreux

as a subgenus. The families Dexaminidae, Atylidae, Anatylidae, Prophliantidae, and Lepechinellidae have been found to

be synonymous and divisible into only two subfamilies, Dexamininae and Prophliantinae: the latter subfamily contains

Prophlias Nicholls, Haustoriopsis Schellenberg, and Guernea

Chevreux (Prinassus Hansen = Dexamonica J. L. Barnard) ;

the remaining genera are assigned to Dexamininae (paper in

preparation).
NUMBER 3 4

grant to Hawaii. Article 4 of male gnathopod 2 bears

a large spike that by analogy seems to replicate the

commonly occurring palmar tooth on article 6. This

unique occurrence in the diverse family Ischyroceridae

also marks the development of an incipient genus.

The case of Neomicrodeutopus(?) makena is

clouded by the absence of certain information about

the genus in African waters, but there is a strong possibility that the Hawaiian species has its closest relationship directly with boreal members of Microdeutopus and not through the genus Neomicrodeutopus. If

so, the Hawaiian species is marked by the loss of an

inner ramus on uropod 3 that may have been a direct

coincidence with its importation to Hawaii. There is

no evidence of incremental relationship to Acuminodeutopus on continental America. The latter genus has

the inner ramus of uropod 3 present but partially

reduced in size.

A less easily documented case of radical change occurs in the endemic Hawaiian genus, Aloiloi. This

situation has consequences on familial classification in

the Aoridae and Isaeidae ( = Photidae). From time to

time doubts have been expressed as to the phyletic

validity of the two families and Aloiloi further destroys

the concept of the Aoridae except as a taxonomic category of convenience in identification. Isaeidae have

gnathopod 2 normally larger than gnathopod 1, whereas Aoridae have the dominance reversed, with secondary sexual modifications also removed to gnathopod

1 in favor of gnathopod 2. But the Aoridae no longer

appear to have been funneled through a single line of

descent from the Isaeidae, and Aloiloi appears to form

the strongest example yet known. The closest affinities

of Aloiloi seem to lie with the isaeid genus Gammaropsis rather than with any other known genus of

Aoridae. Aorids probably consist of various kinds of

isaeids in which a radical reorganization of axial

dominance in gnathopods has occurred. In Aloiloi this

may have occurred within the confines of a single

step of isolation.

Another aorid, the Hawaiian endemic Konatopus,

stands morphologically between two continental American genera, Amphideutopus and Neomegamphopus.

The character of Konatopus minimizes the strong differences between the two continental genera and may

represent one of two alternatives: (1) the convergence

of either continental genus toward the other by the

immigration and further evolution of one or the other

in Hawaii; (2) the concept that Hawaii forms the

360-034 O—70 3

27

evolutionary apex of a triangle; that one of the continental genera diverged in Hawaii, then returned to

the continent, and evolved into the second continental

genus.

The pair of Hawaiian species of Aoroides represents

the only case known so far of a presumed double wave

of immigration of a single cool-water species to Hawaii.

Only two species of the genus have heretofore been

known to occur in the northern hemisphere, one in

Pacific America and the other in northern Asia. The

American Aoroides columbiae presumably underwent

two waves of immigration to Hawaii to result in one

species that is very distinct qualitatively although the

differences are small and in a second species that, if not

identical to the mainland population, may be found

to represent a neotenic dwarf.

The remaining cases of cool-water importation of

amphipods to Hawaii represent what might be called

normal speciation. For example, Gammaropsis alamoana is simply an incremental cognate of American

G. thompsoni without any radical morphological

changes.

If any cases of radical morphological change occur

in endemic Hawaiian amphipods with tropical or antiboreal affinities, they are presently obscure. They

would occur in the endemic Hawaiian genera,

Mokuoloe, Kanaloa, and Nuuanu. The first genus has

obscure affinities, but its generic characters are simply

predictable recombinations of characters known for

other members of Cyproideinae. Kanaloa has affinities

with antiboreal genera and though very distinct is not

unusual in the presently known framework. Nuuanu is

apparently a member of the Eriopisa-Eriopisella complex that probably includes many species living in interstitial niches. Its deep body and head and deep

pereopodal serrations are unusual, but it belongs with

an ecological group of organisms yet poorly studied, and

it may not prove to be as unusual as it now appears.

Enrichment of Diversity in Gammaridean Genera

The diversity of the Hawaiian genera Elasmopus,

Gammaropsis, Hyale, Lembos, and Maera exceeds that

found heretofore in any other tropical locality. Hawaii

may represent a fairly unique situation in which niches

have been compressed by the establishment of so manyexotic species from so many directions. For example,

this may be reflected in the impoverished numbers of

individuals of Hyale in the Sargassum zone despite
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the occurrence of a high diversity of species in the

region. The unusual latitudinal and thermal position

of a group of highly isolated islands like Hawaii may

contribute to the possibility of enrichment in taxal

diversity, but this circumstance could only occur in a

group of organisms like Amphipoda with extraordinarily successful means of dispersal. The possibility

that various species of diverse genera have their immediate origins from highly distinct regions suggests

the possibility that those allopatric species had already

undergone slight shifts in their niches or habitats and

thus on their meeting in Hawaii they were already

preadapted to occupying very narrow niches. More

and more species of certain genera with high dispersive

means were thus able to become assorted without an

extinctive degree of competition. Because one may

presume that isolated and environmentally impoverished Hawaii has fewer niches than a continent in a

similar thermal regime, one must look to factors other

than compression of niches for an explanation of a

high diversity of a genus in Hawaii by continental

standards. The total number of epifaunal species is

comparable between the warm-temperate of California

and Hawaii but one may presume that the Hawaiian

fauna, with fewer genera, has several taxa in other

genera that serve as ecological substitutes for those

missing genera.

But if the Hawaiian fauna is relatively young (assumed average age of 3 million years), one might

presume that we are witnessing a late stage of faunal

evolution that includes several species in each diverse

genus now on the verge of extinction as niche adjustments become perfected by the various immigrants.

For instance, the marginally tropical thermal regime

of Hawaii may also be able to support an abnormally

high number of species in certain genera because the

sympatric taxa have different seasons of maximal competitive activity that reflect the slight thermal differences of the warm-temperate and metatropical seas

from which they came. Hawaiian thermal extremes

may interfere with optimal year round activity of the

several species but are of insufficient degree to cause

extinction. These species thus occupy, by definition,

different niches, but given the same time they have

had on continents, some of these species now surviving

in Hawaii because of thermal niche separation, may

become extinct if their ecological analogues undergo

drifting genetic changes that shift their thermal optima

to the existing Hawaiian condition. One may imagine

a large number of other components of the niche that

may also be involved in the selective adjustments of

Hawaiian immigrants. These thoughts are antithetical

to the concept that species may become more numerous with increasing age of an ecosystem.

The Hawaiian Islands should form an ideal environment for the study of interactions between

amphipodan species imported from warm-temperate

and metatropical regions, but one must hope that

great care is taken not to release such imports into

Hawaiian waters before extensive studies have been

made on those species already present.

Summary and Conclusions

1. The Hawaiian Gammaridea now comprise 120

species in 52 genera, of which 59 species and 9

genera are newly described and 21 other species are

reported for the first time from Hawaii.

2. Nearly 70 percent of the fauna has obviously come

to the islands through archipelagoes to the southwest, but only 15 percent of the fauna is of tropicopolitan character. About half of the Hawaiian

Gammaridea comprises endemic species. Small

percentages (2-6) of the fauna have affinities with

Antarctica, the Caribbean Sea, and Australian

warm-temperate regions, but a significant percentage (20) of the fauna has affinities with cool

waters of North America.

3. Amphipods of cool-water affinity in Hawaii are

primarily tubicolous species; nestling taxa apparently are infrequently successful in completing the

long journey or surviving once in Hawaii. Three

of the cool-water imports have not diverged from

their continental populations, but the remainder

have; at least two of those species have diverged

sufficiently to be assigned to distinct genera and

probably several others will eventually be assigned

to unique genera.

4. Endemic Hawaiian species of Indo-Pacific affinity

include one unique genus, whereas the other discrete Hawaiian genera have apparently come from

Antarctica or the Caribbean region or have

unknown affinities.

5. The tropical component of the Hawaiian fauna is

diverse and not impoverished by a priori standards, but those standards may have to be revised in

view of the unexpected Hawaiian diversity and

the poorly explored tropical regions. Added to the

diverse tropical component is the cool-water component. Nearly all of the expected tropical fam-
NUMBER 34 29

ilies and genera occur in Hawaii, and some are

more diverse than in any other tropical locality so

far studied.

6. Micronesia and Polynesia have contributed to

Hawaiian amphipods about equally with coolwater North America, but only about half of those

central Pacific imports to Hawaii have diverged.

The pool of amphipods endemic to the central Pacific is far less than the cool-water pool, and members of the former pool would already be preadapted to Hawaiian thermal regimes in contrast

to members of the cool-water pool. As small units,

various archipelagoes in the central Pacific should

have low endemicity owing to their close connection toward the I ndo-Pacific to the west. But their

topical influence on Hawaii is great because they

form the pathway for immigration of the majority

of tropical Hawaiian species having broad IndoPacific affinity. Thus, half of the Hawaiian endemic species has come from the Polynesian direction. This degree of endemicity should be

sufficient to demonstrate the high degree of isolation of the Hawaiian Archipelago, but the additional isolates from other regions reinforce that

knowledge considerably.

7. The low incidence of sympatric polytypy in Hawaiian amphipodan genera and the absence of any

multispecificity of endemic Hawaiian genera suggest the near lack of adaptive radiation having

occurred within the Hawaiian chain. Several pairs

of species with close sympatric relationships can

be explained alternatively as the consequence of

successive immigrations of the same ancestral

species; most of these cases have also resulted in

the establishment of one of the members of each

pair in nondivergent condition.

8. The absence of any steppingstones between Hawaii

and cool waters of the northern warm-temperate

regions suggests that Hawaiian species with warmtemperate affinities probably represent good examples of changes that can occur within one increment of isolation. Most of the speciation that

has occurred in these warm-temperate imports is

of ordinary magnitude but a few species have undergone radical morphological changes within this

framework. There is little likelihood that these

radical changes occurred stepwise by repeated to

and fro migration of populations. Though each

case represents only one or two changes and each

change may be under the control of simple genetic

events, the results seem radical because they concern morphological characters normally considered

to be conservative and therefore of familial importance. The suggestion is made that these changes

and many others may actually be highly replicative

genetic occurrences but that the morphological and

functional consequences to the organism result in

selective lethality except in very special or youthful

environments such as that of Hawaii. There is nothing novel in that idea except in pinpointing the

particular characters in Amphipoda that have

been held as conservative for many decades. If

events like the loss of a mandibular palp, the loss

of an article on a maxillipedal palp, the coalescence of two urosomal segments, the development

of a special gnathopodal tooth on an abnormally

proximal article, and the reversal in dominance

of gnathopodal pairs could occur within the framework of a single increment of isolation, then one

might prudently reexamine the affinities of various

families such as Prophliatidae, Kuriidae, Calliopiidae, Ochlesidae, and Cressidae and reexamine generic goodness in varius inquilinous families such

as Stenothoidae and Stegocephalidae now based

on palpar metamery. To consider Aloiloi, a perfectly good aorid genus under present taxonomic

practice, to have arisen from a species of Gammaropsis (Isaeidae) that may still be extant, is to remove a mystique of antiquity in our viewpoint that

all gammaridean genera are old, solidly fixed, and

conservatively developed by eras of slow genetic

change and selection.

Acknowledgments

This project owes its stimulus to discussions between

Dr. Roland B. Force, Director of Bishop Museum and

Dr. Donald F. Squires, Deputy Director of the Museum of Natural History, Smithsonian Institution, resulting in a joint Bishop-Smithsonian Fellowship.

I am especially indebted to the firm support of Dr.

Richard S. Cowan, Director of the Natural History

Museum and Dr. Force who made every resource

of their museums available to me.

At Bishop Museum Dr. Dennis H. Devaney gave me

considerable aid, and I am especially indebted to Dr.

and Mrs. John E. Randall for their deep concern and
30

commitment to the success of this work. Mr. E. H.

Bryan, Jr. helped me in many ways.

At Smithsonian, my colleagues provided numerous

solutions to my logistical problems at long distance;

without this prompt, efficient aid, I would not have

been able to complete this project within the time commitment. My thanks go especially to Dr. Raymond B.

Manning, Chairman, Department of Invertebrate

Zoology, Dr. Thomas E. Bowman, Curator of Crustacea, Dr. I. E. Wallen, Head of Office of Oceanography, Mr. Roland H. Brown, museum specialist, Mrs.

Montague Smith, and Mrs. Dessie M. Best, secretaries.

At the University of Hawaii, both at the Institute

of Marine Biology with laboratories on Coconut Island

and within the Department of Oceanography, I must

thank the following persons for their aid and encouragement: Mr. Emil Keene, Mr. Ralph L. Bowers, Mrs.

Lois Lane Kruschwitz, Dr. Philip Helfrich, Dr. and

Mrs. P. B. van Veel, Dr. Jerry Welch, Mr. James Fee,

Mr. Jack M. Levitz, and Dr. W. A. Gosline.

Dr. John Hendrickson of Sea Life Park, Makapuu

Point, gave me much encouragement and good advice.

Without the assistance of Mrs. Muriel King of

Honolulu, who provided her well-equipped vessel

Pele, this project would be barren of nearly two-thirds

of its results, for on board this ship Dr. Devaney and I

were able to collect the vast majority of the materials

included herein.

Prof. H. Caspers gave his collection of Hawaiian

Amphipoda for inclusion in this study.

Finally, I am indebted to Mr. J. M. Brodie, Director

of the New Zealand Oceanographic Institute, and

Dr. D. E. Hurly who provided a place to complete the

typescript. The drawings were made by Miss N. B.

Manowitz, Mr. L. I. Kaneshiro. Mr. M. D. Lenhoff

and the writer.

Systematical Procedure

A diversity of descriptive methods is used that varies

with the state of knowledge in various taxa. Some

species require descriptions in view of their uncertain

taxonomic positions whereas, at the other extreme, a

short diagnosis suffices for species in well-known genera

or families. Long diagnoses often characterize species

in genera of extreme diversity, and a few taxa have

long diagnoses or descriptions because of the richness

or variability in morphological detail. Many words are

eliminated by consultation of the figures.

Except in obvious cases, the Hawaiian names proposed here for amphipods stand as nouns in apposition

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

to their generic appellations. They are primarily chosen

for euphonious reasons from place names scattered

throughout the islands or from the names of important

personages or deities in Hawaiian history. Some generic

names are also proposed in this way but others are

formed of contractions containing only partially

Hawaiian roots. It is senseless to invent reasons for

choosing these names so they are not considered further.

Family AMPHILOCHIDAE

Subfamily AMPHILOCHINAE

Amphilochus Bate

The Hawaiian members of this genus have better developed molars on the mandible than attributed heretofore to Atlantic and west American species. For that

reason they approach the genus Gitanopsis and remarks

on their affinities to members of that genus are included

with each of the species.

Amphilochus kailua, new species

FIGURES 2, 3

DIAGNOSIS.—Body without dorsal cusps; lateral

cephalic lobe rounded or slightly truncate; eye very

large, clear in alcohol; antennae extending equally (including distal setae) ; gnathopods large, slender lobe

of article 5 on gnathopod 1 guarding article 6 and extending almost to palm, on gnathopod 2 extending

fully to palm; anterodistal corners of sixth articles on

gnathopods softly rounded, dactyls with medial inner

tooth but no pectinations or scales; gnathopods of the

enlarged kind, but gnathopod 2 larger than 1, hands

longer than wide, palms scarcely oblique but convex;

coxa 1 elongate, anteroventrally extended as short coniform process; mandibular molar narrow and weakly

triturative; distomedial margin of outer plates on

maxillipeds not excavate, with 6-7 slender, widely

spaced setae; palp article 1 of maxilliped with winglike

medial extension; all pereopods with about four sets

of spines on "active" margins, distal locking set (and

occasionally other sets) paired and closely placed to

next proximal set, dactyls with short distal nail and

prominent mammilliform bulb proximal to nail;

pleonal epimera 1-3 each with sinuous posterior margins and very weak posteroventral tooth or protrusion;

telson of medium length, shorter than peduncle of

uropod 3, apically rounded but coniform.
NUMBER 34 31

FIGURE 2.—Amphilochus kailua, new species, ?female, 1.3 mm, Waikiki, Sargassum: a, head; b,

palp article 1 of maxilliped; c, maxilliped; d, inner plate of maxilliped; e, upper lip. Holotype,

ovigerous female, 1.7 mm: f, head; g, lower lip; h, mandible; i, ;, antennae 1, 2; k, telson; /,

uropod 3; m, maxilla 2.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Amphilochus kailua, new species, holotype, ovigerous female, 1.7 mm, Waikiki, Sargassum: a, b, c, pereopods 2, 3, 5; d, e, gnathopods 1,2;/, coxa 6; g, mandibular molar; h, outer

plate of maxilliped; i, gnathopod 1, enlarged; ;, dactyl of pereopod 2; k, pleonal epimera 1—3.

left to right. PFemale, 1.3 mm: /, apex of palp article 3 of maxilliped.
NUMBER 34 33

HOLOTYPE.—Bishop Museum collections, catalog

number 7248, ovigerous female, 1.7 mm.

TYPE-LOCALITY.—Off Waikiki Marine Laboratory,

from Sargassum in shallow water, 11 March 1964 (3

specimens).

MATERIAL.—Type-locality, also from the alga Padina

(1). JLB Hawaii 2 (2). Fee 1 (38).

RELATIONSHIP.—The mammilliform process of the

pereopodal dactyls is unique to this species. In other

characters it comes closest to Gitanopsis vilordes J. L.

Barnard (1962c) from California, but differs from that

species in the rounded and not sharp corner on the

anterodistal end of article 6 on gnathopod 1.

Amphilochus kailua resembles G. pusilla K. H.

Barnard (1916), but G. pusilla has a tooth on the dactyls of only pereopods 1-2.

The mammilliform process on the dactyls of the

pereopods distinguishes this species from its close relative Amphilochus neapolitanus Delia Valle (J. L.

Barnard, 1962c).

DISTRIBUTION.—Hawaiian Islands.

Amphilochus likelike, new species

FIGURE 4

DIAGNOSIS.—Body without dorsal cusps; lateral cephalic lobe rounded; eye very large, ommatidia of

circumference large, those in core much smaller than

in other Hawaiian species and more numerous;

antenna 1 almost as long as antenna 2 (including distal

aesthetascs); stout lobe of article 5 on gnathopod 1

extending about halfway along article 6 and slender

lobe on gnathopod 2 extending fully along article 6;

anterodistal corner of article 6 on gnathopod 2 softly

rounded, article 4 of gnathopod 2 with no stout

posterior spines, dactyls of gnathopods not pectinate

besides major tooth, each article 2 of gnathopods 1-2

with stout posterodistal spine; coxa 1 short, softly

quadrate, bearing 2 weak distal serrations, coxae 2-3

with slight distal serrations; mandibular molar of

medium size, conical and weakly triturative; outer plate

of maxilliped distomedially excavate, weakly and

scarcely pectinate, distolateral end with large sabreshaped spine; dactyls of pereopods 1-5 simple; pleonal

epimeron 1 with slight posteroventral tooth, epimeron

2 with slightly extended and sharp corner, epimeron

3 with rounded quadrate corner; telson exceeding 50

percent length of peduncle of uropod 3, apically

rounded but coniform.

Urosome like Gitanopsis pele, new species, but inner

ramus of uropod 2 reaching three-fourths distance between distal ends of peduncle of uropods 1 and 3.

COLOR IN 2-DAY FORMALDEHYDE.—Body albinid but

eye red; eye in alcohol clear with slight rust stain in

central core.

HOLOTYPE.—Bishop Museum collections, catalog

number 7249, female, 1.6 mm.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, coralline footballs, 29 January, 1967.

MATERIAL.—JLB Hawaii 5 (2), 6 (5), 11 (4).

RELATIONSHIP.—Resembles Amphilochus kailua,

new species, most of any Hawaiian species because the

gnathopodal dactyls lack supernumerary pectinations

but differs from that species in the simple dactyls of

pereopods 1-5. The individuals of A. likelike so far

collected, though adult, are smaller than 1.7 mm and

the undeveloped outer plate of the maxilliped may

be a reflection of this small size.

Amphilochus likelike is like G. vilordes in many respects, as is A. menehune, new species, but A. likelike

has the stout spine on article 2 of gnathopods 1-2

which A. menehune does not have and which G.

vilordes has only on gnathopod 2; A. likelike differs

from G. vilordes in the shorter article 5 lobe on

gnathopod 1 and the more conical mandibular molar.

Amphilochus menehune differs from A. likelike in the

presence of stout spine (s) on the posterior margin of

article 4 on gnathopod 2.

Among Hawaiian species of the genus, the fine

pectinations on the gnathopodal palms are especially

small in A. likelike.

The absence of spines on article 4 of gnathopod 2

distinguishes this species from its close relative, Amphilochus neapolitanus Delia Valle (J. L. Barnard,

1962c).

The name is pronounced in English transliteration:

leek-ee-leek-ee.

DISTRIBUTION.—Hawaiian Islands.

Amphilochus menehune, new species

FIGURES 5, 6

DIAGNOSIS.—Body without dorsal cusps; lateral

cephalic lobe rounded; eye very large, varying from

clear in formaldehyde and alcohol to a clear ring of

ommatidia surrounding a slightly pigmented core to

eyes with fully black pigmented core; antenna 1

scarcely extending to end of peduncle of antenna 2;
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Amphilochus likelike, new species, holotype, female, 1.6 mm, JLB Hawaii 6: a, upper

lip; b, head; c, lower lip; d, coxa 3; e, outer plate of maxilliped; /, maxilla 2; g, maxilla 1, minus

palp; h, inner plate of maxilliped; i, coxa 6; j , telson and lateral wing of pleonite 6; k, I, m,

pereopods 2, 3, 5; n, accessory flagellum on article 3 of antenna 1; o, epistome and upper lip;

p, q, gnathopods 1, 2.
NUMBER 3 4 35

9

FIGURE 5.—Amphilochus menehune, new species, holotype, female, 1.8 mm, JLB Hawaii 5: a,

head; b, telson; c, accessory flagellurn; d, e, f, g, pereopods 2, 3, 4, 5; h, gnathopod 1; i, mandible;

;', dactyl of pereopod 4; k, coxa 3; /, m, maxillae 1, 2; n, half of lower lip. Female, 1.9 mm, JLB

Hawaii 6; o, head.
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 6.—Amphilochus menehune, new species, holotype, female, 1.8 mm, JLB Hawaii 5: a,

head; b, upper lip; c, maxilliped; d, accessory flagellum (arrow) ; e, mandible; /, g, gnathopod 2;

h, gnathopod 1; i, pleonal epimera 1-3, left to right. Female, 1.9 mm, JLB Hawaii 6; ;,

gnathopod 2.
NUMBER 34 37

stout lobe of article 5 on gnathopod 1 extending about

halfway along article 6 and slender lobe on gnathopod

2 extending about 90 percent along article 6, anterodistal corner of article 6 on gnathopod 2 sharply extended, article 4 of gnathopod 2 with 1-3 very stout

ventral spines; dactyls of gnathopods pectinate or bearing scales on proximal inner edges; coxa 1 short, almost

hemispherical, bearing 1 distal notch, coxae 2-3 with

medium-sized distal serrations; mandibular molar

weakly triturative, subconical and of medium size;

outer plate of maxilliped distomedially excavate and

pectinate, distolateral end with large spine; dactyls of

pereopods 1-5 simple or with very faint distal nail

marked by 1-2 weak scales; pleonal epimeron 1 with

slight posteroventral tooth, epimeron 2 with slightly

extended corner and 3 with rounded-quadrate corner; telson about 40 percent as long as peduncle of

uropod 3, apically rounded but coniform.

Urosome like that of Gitanopsis pele, new species,

but inner ramus of uropod 2 reaching only to a point

halfway between the ends of peduncles of uropods 1

and 3.

COLOR IN 2-DAY FORMALDEHYDE.—Widely variable

from completely albinid having black ocular core to

medium brown body with brown eye turning clear in

alcohol. Forms with medium brown body have the

pigment on posterior half of head, pereonites 1-6,

coxae 1-6, articles 2-5 of gnathopod 1, all of gnathopod 2 and pereopod 3; these individuals turn rusty

pink in alcohol; in formaldehyde also are individuals

already turned rusty pink; a fourth form has extremely

rare and widely scattered chromatophores over the

thorax and abdomen and occasional weak blotches of

brown or pink in various places.

HOLOTYPE.—Bishop Museum collections, catalog

number 7250, female, 1.8 mm.

TYPE-LOCALITY.—JLB Hawaii 5, off Ewa Beach,

Oahu, 30 m, Pocillopora, 29 January 1967.

MATERIAL.—JLB Hawaii 2 (3), 3 (2), 5 (8), 6 (8)

(3 color phenotypes), 8 (2), 10 (3), 13 (7), 14 (13),

15 (l).Devaney 1 (1).

RELATIONSHIP.—This species resembles the Californian Gitanopsis vilordes J. L. Barnard (1962c), but

the mandibular molar of A. menehune is more conical,

article 6 of gnathopod 2 lacks midanterior spines,

article 4 of gnathopod 2 has 1-3 posterior spines, article

2 of gnathopod 2 has a posterodistal seta and not a

spine, coxa 1 is less quadrate in shape, the lateral

cephalic lobe is more rounded, and article 6 of

gnathopod 1 smaller relative to article 5, with its posterior lobe stouter and shorter, than in G. vilordes.

The short coxa 1 and strong molarial rugosites of

A. menehune distinguish it from its close relative, A.

picadurus J. L. Barnard (1962c).

DISTRIBUTION.—Hawaiian Islands.

Gitana liliuokalaniae, new species

FIGURES 7, 8

DIAGNOSIS.—Rostrum large, lateral cephalic lobe

weak, rounded, eye large, red in 2-day formaldehyde,

clear peripherally with dim orange core in alcohol;

flagellum of antenna 1 with numeous long aesthetascs

(antenna 2 broken, upper lip unknown); palp article

2 of mandible stout, incisor normal; palp of maxilla 1

apparently 1-articulate; outer plate of maxilliped

densely serrate distomedially, not excavate, distolateral margin with long seta, palp article 1 longest but

not twice as long as article 2, articles 1 and 2 slightly

produced distomedially, dactyl as long as article 3 and

bearing 1 apical setule; gnathopods 1-2 very weak,

slender, similar to each other, simple article 5 produced

in posterodistal lobe strongly on gnathopod 2, lobe

overriding article 6, lobe weaker on gnathopod 1, article 5 scarcely longer than article 6, anterior margin of

article 6 distally constricted, dactyl pointing distad;

coxa 1 broadly rounded below, coxa 2 narrowing

evenly from anterior margin only, with 1—2 small posteroventral teeth or one tooth substituted by distomedial notch (right and left sides differ); pleonal epimera

1-3 increasingly sharply quadrate posteroventrally;

uropods 1 and 3 extending equally, inner ramus of

uropod 2 reaching end of peduncle of uropod 3; telson

half as long as peduncle of uropod 3.

HOLOTYPE.—Bishop Museum collections, catalog

number 7247, ?male, 1.2 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

Oahu, 3-4 m, small corals, fleshy algae, corallines,

8 April 1967.

RELATIONSHIP.—Only 4 other species of this genus

are known, 3 from the cold-temperate North Atlantic

and one from warm-temperate California. Gitana liliuokalaniae resembles G. calitemplado J. L.-Barnard

(1962c) from California least of any and has characters of the other three mixed together. The extraordinarily weak article 6 of the gnathopods resembles that

of G. rostrata Boeck (Sars, 1895, pi. 79, fig. 1) but

the presence of a weak lobe on article 5 of gnathopod
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

h

FIGURE 7.—Gitana liliuokalaniae, new species, holotype, male, 1.2 mm, JLB Hawaii 13: a, b,

medial gnathopods 1, 2; c, d, e, pereopods 1, 4, 5, medial; /, head; g, pleon; h, spines of article

6 on pereopod 1.

2 is more reminiscent of G. abyssicola Sars (1895, pi.

78, fig. 2) or G. sarsi Boeck (Sars, 1895, pi. 78, fig. 1).

Gitana liliuokalaniae also resembles Gitanopsis arctica

Sars (1895, pi. 77, fig. 2) in maxillipeds and mandibular palp more than it does its congeners. In fact, G.

artica is intermediate between Gitana and Gitanopsis

in that it has the 2-articulate first maxillary palp of

Gitanopsis but the weak gnathopods of Gitana. Gitana

liliuokalaniae differs from all cold-temperate species

of Gitana in the short telson.

Affinities with Gitanogeiton sarsi Stebbing (1910)

from warm-temperate Australia (Manning River

area) are strong in Gitana liliuokalaniae, and the

Hawaiian species almost forms a good link between

Gitana and Gitanogeiton. The latter monotypic genus

differs from Gitana in several fairly weak characters:

the slightly excavate outer plate of the maxillipeds,

the slightly elongate palp article 1 of the maxilliped,

and the broadened incisor of the mandibles. Gnathopods of Gitanogeiton resemble those of Gitanopsis
NUMBER 34 39

k

FIGURE 8.—Ghana liliuokalaniae, new species, holotype, male, 1.2 mm, JLB Hawaii 13: a,

mandible; b, c, coxae 4, 1; d, right mandibular molar; e, right mandibular incisor and spines; /,

outer plate of maxilla 1; g, maxilliped; h, maxilla 2; i, apex of lower lip; j , maxilla 1; k, telson.

arctica in that gnathopod 2 has a strong palm and gnathopod 1 a weak palm, and each article 5 has a strong

posteroventral lobe, stronger than that of G. liliuokalaniae. The latter has the strong palp article 2 of the

mandible seen in Gitanogeiton, but the molar in G.

liliuokalaniae is stronger, the incisor weaker, the palp

of the maxilliped is relatively longer and stouter. Unfortunately, the upper lip of G. liliuokalaniae was lost

during dissection and cannot be compared with the unusually deeply incised lip of Gitanogeiton. Gitana

liliuokalaniae seems to have its presently known

strongest affinites with Gitanopsis arctica, a species

that, because of its gnathopods, may rightly belong

with Gitana rather than with Gitanopsis despite the

occurrence of a 2-articulate palp on maxilla 1.

Gitanopsis simplex Schellenberg (1926), from

Antarctica, resembles the Hawaiian species in gnathopods, but the palm of gnathopod 2 in G. simplex is

distinct and article 6 is longer than article 5. The telson

of G. simplex is longer than it is in G. liliuokalaniae,

and apparently the peduncle of antenna 1 in the first

species is much longer than it is in the second species.

DISTRIBUTION.—Hawaiian Islands.
40

Gitanopsis pele, new species

FIGURE 9

DIAGNOSIS.—Body without dorsal cusps; lateral cephalic lobe broadly rounded; eye large, with core of

black pigment in alcohol; antennae extending subequally; gnathopods small but palms strong and

nearly transverse, stout lobe of article 5 on gnathopod

1 extending about halfway along article 6 and about

75-85 percent on gnathopod 2, sixth articles weakly

expanded distally and much longer than broad, about

half as broad as their coxae, dactyls proximally pectinate on inner edges; coxa 1 distally tapering and subconical, posteroventral corner serrate, coxa 2 with 1

small posteroventral notch; mandibular molar large,

cushion-shaped, strongly triturative; outer plate of

maxilliped with mediodistal margin slightly excavate

and pectinate (as in Gitanogeiton and Gitanopsis tortugae) ; dactyls of pereopods 1-5 simple; pleonal epimera 1-2 each with small posteroventral tooth, epimeron 3 rounded-quadrate posteroventrally; telson of

medium length, about 60 percent as long as peduncle

of uropod 3, apically rounded but coniform.

HOLOTYPE.—Bishop Museum collections, catalog

number 7251, ovigerous female, 2.7 mm.

TYPE-LOCALITY.—JLB Hawaii 10, Kaneohe Bay,

Oahu, 2 m, Pocillopora, 23 February 1967.

MATERIAL.—JLB Hawaii 10 (50).

RELATIONSHIP.—This species resembles the Caribbean G. tortugae Shoemaker (1933a, p. 248, fig. 2).

Maxilla 1 and the accessory flagellum of the 2 species

are similar among numerous other characters of even

greater importance. Gitanopsis pele differs from G.

tortugae primarily in two minor characters: (1) the

serrations of coxae 1-2 and (2) the large eyes.

Gitanopsis pusilla K. H. Barnard (1916) from South

Africa and G. vilordes J. L. Barnard (1962c) from

California differ from G. pele and G. tortugae in their

large gnathopods with article 6 wider than their coxae

and strongly expanded distally. Gitanopsis pusilloides

Shoemaker (1942) from Pacific Mexico has very short

processes on article 5 of the gnathopods and more

strongly transverse palms than do G. pele and G.

tortugae.

DISTRIBUTION.—Hawaiian Islands.

Subfamily CYPROIDEINAE

Mokuoloe, new genus

DIAGNOSIS.—Rostrum short; flagella of first and

second antennae 4 and 2-articulate respectively, bearSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ing long aesthetascs; epistome nearly flat anteriorly;

upper lip evenly incised; mandible with triturative

molar and normal 3-articulate palp; lower lip angularly excavate medially; palp of maxilla 1 uniarticulate, bearing large strap-shaped terminal setae, outer

plate with 7 spines, inner plate short and truncate;

maxilla 2 with apically truncate lobes; outer plate of

maxilliped large and exceeding end of palp article 2,

palp article 3 apicomedially produced as thumb, dactyl

stout and closing on thumb; coxa 2 larger than coxa 1;

gnathopods 1-2 of medium stoutness, subchelate, dactyl of gnathopod 1 strongly overlapping palm, of

gnathopod 2 fitting palm, article 5 of gnathopod 1

lobe obsolete, of gnathopod 2 lobe extending more than

halfway along article 6, palms of gnathopods deeply

and sharply serrate; pereopods 3-4 with slender article

2, pereopod 5 with expanded article 2; uropod 1 longest, uropods 2-3 extending successively less, outer

ramus of uropod 3 shorter than inner ramus; telson

extending almost to apex of uropod 3, boat-shaped,

triangular, entire, with lateral keel; pleonite 4 long,

weakly keeled bilaterally on dorsal surface, pleonite 5

covered by junction of 4 and 6, 6 with weak lateral

keel on each side and not vaulting over telson.

TYPE-SPECIES.—Mokuoloe ninole, new species.

RELATIONSHIP.—Peltocoxa Catta (Chevreux and

Fage, 1925) and Hoplopheonoides Shoemaker (1956)

from Florida are the cyproidins seemingly closest to

this genus—not various Australian and New Zealand

genera, but even relationships with those two Atlantic

genera are weak.

Peltocoxa Catta has a long maxillipedal palp, slender

inner plate of maxilla 1, strong mandibular lobes on

the lower lip, narrow lobes on maxilla 2, no strapshaped setae, a dorsoventrally thickened telson, inflated

article 2 of pereopod 4, small gnathopods, gnathopod

2 with transverse palm and gnathopod 1 with oblique

palm confluent with the posterior margin of article 6

and bearing only articulate spines.

Hoplopheonoides has small gnathopods with rectangular sixth articles and transverse palms, an enormous

urosomite 1 with dorsal keel, no mandibular palp, long

maxillipedal palp, and numerous other differences.

Mokuoloe ninole, new species

FIGURES 10, 11

DIAGNOSIS.—With the characters of the genus.

COLOR IN 2-DAY FORMALDEHYDE.—Eyes red, becoming clear in alcohol.
NUMBER 34 41

n

FIGURE 9.—Gitanopsis pele, new species, holotype, ovigerous female, 2.7 mm, JLB Hawaii 10:

a, lateral view of body; b, telson; c, mandible; d, head; e, upper lip; f, maxilla 2; g, spiny ridge on

article 2 of pereopods 3—5; h, half of lower lip; i, maxilliped; ;, k, gnathopods 1, 2; I, part of coxa

1; m, dactyl of pereopod 3; n, epistome and upper lip, right side.
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Mokuoloe ninole, new genus, new species, holotype, male, 1.5 mm, JLB Hawaii 6:

a, body; b, head; c, lateral gnathopod 2; d, mandible; e, urosomites "4", 5, "6" and telson

("T").
NUMBER 34

m

FIGURE 11.—Mokuoloe ninole, new genus, new species, holotype, male, 1.5 mm, JLB Hawaii 6:

a, maxilla 1; b, maxilla 1 flattened; c, maxilla 2; d, right mandibular incisor; e, maxilliped; /,

inner plate of maxilliped; g, mandibular molar; h, left coxa 1; i, maxillipedal palp articles 3-4;

;, lower lip; k, left coxa 2; /, upper lip; m, lateral gnathopod 1.

:(60-034 o—70-—
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HOLOTYPE.—Bishop Museum collections, catalog

number 7252, ?male, 1.5 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, coralline footballs, 29 January 1967.

DISTRIBUTION.—Hawaiian Islands.

Family AMPITHOIDAE

Ampithoe akuolaka, new species

FIGURES 12, 13

DIAGNOSIS OF MALE.—All pleonal epimera with lateral ridges, rounded posteroventrally, but with small

obtuse notch bearing setule; articles 5 and 6 equal to

each other in length, article 5 with truncate posterior

margin and triangular posterodistal extension slightly

overlapping article 6, article 2 with large distolateral

lobe, article 6 narrowly rectangular and slightly tapering, palm very short and obsolescent, posterior margin

of article 6 with defining spine, dactyl strongly overlapping palm; gnathopod 2 of medium size, article 2

with large anterodistal lobe, article 4 weakly coniform

distally, article 5 short, with narrow bifid, setose posterior lobe, hand longer than broad, deeply bifid on

palm, heavily setose, dactyl strongly overlapping large

tooth of palm; article 2 of pereopods 1-2 more than

twice as long as broad; pereopods 3-5 with simple sixth

articles bearing distal pair of small straight locking

spines, one smaller and more erect than other, largest

very long on pereopod 3, dactyl simple; article 2 of

pereopod 3 very broad and lacking spines or notches,

article 2 of pereopods 4-5 with 2 or 3 stout posterodistal spines; antenna 1 about as long as head and

pereonites 1-6 together, antenna 2 short, peduncular

articles short and of medium thickness, entire antenna

about as long as head and pereonites 1—4 together,

flagellum slender, almost as long as articles 4—5 together; apical lobules of lower lip very short and

tumid, lateral longer than medial; coxa 1 produced

forward; peduncular processes between rami of uropods 1—2 absent, peduncles of uropods 1-3 with lateral

setae, arranged in axial row on uropod 1, in one bundle

on uropods 2-3; eyes pale purple in alcohol.

FEMALE.—Gnathopod 1 lacking posterodistal triangular extension on article 5; gnathopod 2 small,

article 5 elongate and bearing medium-sized triangular

lobe, palm of narrow article 6 very oblique, slightly

S-shaped, defined by large spine, dactyl fitting palm.

JUVENILE (smallest available, 3.6 mm).—Similar to

female and bearing peduncular setae on uropods 1-3,

notches and lateral ridges on pleonal epimera.

HOLOTYPE.—Bishop Museum collections, catalog

number 7253, male, 5.6 mm.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, other algae, rocks, corals,

24 May 1967.

MATERIAL.—JLB Hawaii 17 (2), 18 (2).

RELATIONSHIP.—The lower lip of this species resembles that of A. pollex Kunkel (W Atlantic, NE Pacific) and gnathopod 2 of the male resembles the

juvenile male of A. pollex. Ampithoe akuolaka appears

to differ from the Pacific population of A. pollex (J. L.

Barnard, 1965b), in the shorter medial lobules on the

lower lip, the broader article 2 of pereopod 3, the

presence of stout spines on article 2 of pereopods 4-5,

the stronger palm of gnathopod 1, and the posterodistal

triangular extension on article 5 of male gnathopod 1.

The latter characteristic suggests a relationship of

A. akuolaka to A. ramondi Audouin, but the apical

hooks on the outer ramus of uropod 3 of that species

are highly reverted, no stout spines occur on the posterior margins of article 2 on pereopods 4-5, gnathopod

2 of the male apparently does not go through the

akuolaka stage, the lateral apical lobules of the lower

lip are thinner, and the female hand of gnathopod 2

is stouter. The hand of young male gnathopod 2 is like

that of Oregonian A. simulans Alderman (1936) (J. L.

Barnard, 1965b) and so are pleonal epimera 1-3 highly

similar, but male gnathopod 1 article 5 of A. simulans

is not produced posterodistal ly, the lateral apical lobule

of the lower lip is thin, pereopods 4-5 lack the posterior

spines on article 2, and the dactyl of male gnathopod 2

does not strongly overlap the palmar tooth. Possibly

the specimens represented in J. L. Barnard (1965b, fig.

18) are a fourth sibling species in the complex of A.

pollex-simulans-akuolaka.

DISTRIBUTION.—Hawaiian Islands.

Ampithoe kaneohe, new species

FIGURES 14, 15, 16, 24/

DIAGNOSIS OF MALE.—All pleonal epimera posteroventrally rounded; article 6 of gnathopod 1 longer

than article 5, posterior margin of article 5 moderately

broad but with weak, broadly coniform lobe, article

6 ovato-rectangular, of medium stoutness, palm very

oblique and defined by stout spine; gnathopod 2 with
NUMBER 34 45

h

FIGURE 12.—Ampithoe akuolaka, new species, holotype, male, 5.6 mm, JLB Hawaii 17: a,

gnathopod 1; b, dactyl of pereopod 5; c, pereopod 2; d, pleonal epimera 1-3; e, telson; f, maxilla

2; g> h, gnathopod 2, medial and lateral views. Female, 6.2 mm: i, lateral gnathopod 2.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Ampithoe akuolaka, new species, holotype, male, 5.6 mm, JLB Hawaii 17: a, b, c,

pereopods 3, 4, 5; d, uropod 3; e, mandibular palp; /, head; g, lower lip; h, urosome. Female, 6.2

mm: i, gnathopod 2, setae removed;;, anteroventral corner of head.
NUMBER 3 4 47

FIGURE 14.—Ampithoe kaneohe, new species, male, 6.0 mm, Fee 1: a, head and antennae. Holotype, male, 5.4 mm, Waikiki: b, head; c, lower lip; d, e, pereopods 4, 5; /, pleonites 1—4. Female,

5.2 mm: g, dactyl of gnathopod 2; h, article 5 of gnathopod 1; i, pereopod 3, unflattened; ;,

pereopod 3.
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 15.—Ampithoe kaneohe, new species, holotype, male, 5.4 mm, Waikiki: a, b, gnathopod 2,

lateral and medial views; c, maxilliped; d, e, apex of pereopod 4; f, telson; g, mandibular molar;

h, palmar spine of gnathopod 1; i, gnathopod 1.
NUMBER 34 49

FIGURE 16.—Ampithoe kaneohe, new species, holotype, male, 5.4 mm, Waikiki: a, uropod 3;

b, maxilla 2; c, d, coxae 5, 3; e, pereopod 2; /, g, uropods 1, 2; A, maxillipedal palp; i, maxilla 1,

inner plate shaded; ;, dactyl of pereopod 2; k, mandibular palp. Female, 5.2 mm: /, medial

gnathopod 2.
50

elongate, tapering article 6, palm and posterior margin

congruent, undefined, minutely crenulate, sparsely

setose, medioterminal face of article 6 bearing large

brush of long, apically turned, stiff setae, article 5 short

and bearing strong, slender posterior lobe, article 7

half as long as article 6, distal lobe of article 2 very

large; article 2 of pereopods 1-2 more than twice as

long as broad; pereopods 3-5 with subprehensile sixth

articles, each bearing palm closely guarded by a striate curved spine, 2 other striate spines at palmar

corner and 1 small spine near dactylar hinge, dactyls

partially striate on pereopods 3-5; antennae 1-2 long

and extending equally, antenna 2 peduncle especially

long, flagellum scarcely as long as article 5, slender,

weakly setose; lateral apical lobule of lower lip much

longer than medial and bearing apical nipple; coxa 1

produced forward; peduncular process of uropod 1

short, peduncle with sparse row of ventrolateral setae;

eyes very small, clear or with slight posterior dark pigment in alcohol.

FEMALE.—Gnathopods like those of male gnathopod

1, but article 5 lobe of gnathopod 1 truncate, that of

gnathopod 2 sharp; antennae shorter than in male,

articles 4-5 of antenna 2 shorter and flagellum as long

as articles 4-5 together.

HOLOTYPE.—Bishop Museum collections, catalog

number 7254, male, 5.4 mm.

TYPE-LOCALITY.—Off Waikiki Marine Laboratory,

11 March 1964, from Ulva, shallow water (4

specimens).

MATERIAL.—Hanauma Bay, Oahu, 29 February

1964, from Acanthophora, depth unknown. Fee 1

(24). JLB Hawaii 12 (4), 13 (6), 15 (2).

RELATIONSHIP.—The subprehensile pereopods 3-5

give this species the aspect of a Pleonexes, but that

subgenus should be restricted for those few species also

having large cornified lobes or wings on the telson.

See Ampithoe {Pleonexes) poipu below. Ampithoe

should include that taxon described by Barnard

(1965b) as Pleonexes species from Micronesia, which

I designate now as Ampithoe kulafi, new species. It and

Ampithoe kaneohe appear to be a closely related pair

in terms of subprehensile pereopods, lower lip,

gnathopod 1, and uropod 1, but A. kulafi (anagram

of Ifaluk, its type-locality) has a very different male

gnathopod 2, with subcircular article 6 bearing a transverse palm. Ampithoe kaneohe is the only known

species of the genus with male gnathopod 2 having the

palm and posterior margin of the hand congruent and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

that also has a nonrectangular gnathopod 1 with

oblique palm. Thus, it differs from Ampithoe orientalis Dana of the Hawaiian Islands in that character

plus the subprehensile pereopods 3-5.

DISTRIBUTION.—Hawaiian Islands.

Ampithoe orient a Us Dana

FIGURE 17a-i

Amphithoe (sic) orientalis Dana, 1853, pp. 937-939, pi. 64,

fig. 2.

Ampithoe orientalis.—'Stebbing, 1906, p. 641.—J. L. Barnard,

1955a, pp. 26-28, fig. 14.

MATERIAL.—JLB Hawaii 17 (1), 18 (1). Fee 1 (7).

DISTRIBUTION.—Philippine and Hawaiian Islands.

See Ampithoe species for special notes and comparisons supplementing the description of the Hawaiian

population of this species by J. L. Barnard (1955a).

Ampithoe ramondi Audouin

FIOURES 18, 19

Ampithoe ramondi Audouin.—J. L. Barnard, 1955a, pp. 28-

29 [with references).

NOTES.—Drawings of this common circumtropical

species are presented to establish means for future

comparison of the Hawaiian population (or a small

portion of it) to others in far parts of the world and

with other Hawaiian members of the genus.

Of special interest is the thickening of the setae on

the anterodistal lobes of the second articles on gnathopods 1-2. The distal locking spines of pereopods 3-5

in the juvenile are composed of a very small, thin,

straight, spirally striate spine, next proximal an erect

slightly curved, enlarged, spirally striate spine and next

proximal a straight, axially striate spine. On pereopods

3-5 of adults a pair of distal, weakly sabre-shaped,

large, semispirally striate spines occurs with the next

proximal spine, a nearly straight, axially striate

member.

Eyes clear in alcohol or very pale pink.

Uropod 1 has a lateral setal row. The inner plate

of maxilla 1 has 4 setae in adults.

MATERIAL.—JLB Hawaii 2 (1), 15 (4). The material reported by Barnard (1955a). Canton Island,

February 1958, Drs. Degener (100+ ).

DISTRIBUTION.—Circumtropical.
NUMBER 34 51

FIGURE 17.—Ampithoe orientalis Dana, and Ampithoe species; A. o, male, Fee 1: a, articles 2-4 of

lateral pereopod 3; b, lateral gnathopod 2; c, pereopod 1, setae removed; d, apex of pereopod 3.

Female, e, half of lower lip; /, lateral gnathopod 1, setae removed; g, pleonal epimera 2—3; h,

gnathopod 2; i, articles 4—5 of peduncle and flagellum of antenna 2. Ampithoe species, female,

JLB Hawaii 17: j , half of lower lip; k, pereopod 1, setae removed; /, lateral gnathopod 1, setae

removed; m, uropods 2-3 from lateral view; n, apex of gnathopod 1; o, apex of pereopod 4.

Female, JLB Hawaii 18: p, articles 4-5 of peduncle and flagellum of antenna 2.
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

h

FIGURE 18.—Ampithoe ramondi Audouin, male, 6.8 mm, Kawela Bay: a, head; b, c, pereopods 3,

5; d, telson. Female, 4.6 mm, JLB Hawaii 15: e, f, pereopods, 3, 4; g, uropod 3; h, pereopod 2.

Juvenile male, 2.7 mm, JLB Hawaii 2: i, palm of gnathopod 2;;", apex of pereopod 3.
NUMBER 34 53

FIGURE 19.—Ampithoe ramondi Audouin and Neomicrodeutopus (? ) makena, new species. A.

ramondi, female, 4.6 mm, JLB Hawaii 15: a, gnathopod 2. Male, 6.8 mm, Kawela Bay: b, c,

gnathopods 1, 2; d, dactyl of pereopod 5; e, lower lip. N. (?) makena, female, 3.9 mm, JLB

Hawaii 18: /, lateral gnathopod 1.

Ampithoe waialua, new species

FIGURES 20, 21

DIAGNOSIS OF MALE.—All pleonal epimera rounded

posteroventrally; article 6 of gnathopod 1 longer than

article 5, latter subtriangular, posterior lobe turned

slightly distalward and article 4 broadly extended

triangularly to fit posteroproximal oblique margin of

article 5, article 2 with large distolateral lobe, article

6 subrectangular, palm short and oblique, excavate,

dactyl strongly overlapping palm, large medial-defining spine strongly proximal to apex of defining cusp;

gnathopod 2 small, article 2 with mammiliform,

sparsely setose distolateral lobe, article 4 slender, posterodistally coniform, article 5 relatively long for genus,

bearing asymmetrically triangular posterior lobe, hand

expanded, palm oblique and excavate, with defining

cusp, dactyl failing to fit palm; article 2 of pereopods
54

1-2 about twice as long as broad; pereopods 3-5 with

simple sixth articles bearing distal pair of striate, nearly

straight locking spines and one other striate spine

proximal to locking pair, dactyl slightly striate; article

2 of pereopod 3 very broad and bearing 1-2 large

anterior notches armed with stout spine; antenna 1

about as long as head and first 8 body segments together, antenna 2 about two-thirds as long as antenna

1, flagellum slender and as long as articles 4-5 of

peduncle together; lateral apical lobule of lower lip

much longer than medial and bearing apical nipple,

inner lobes partially coalesced with medial lobule of

outer lobe; coxa 1 produced forward; peduncular

process between rami of uropod 1 obsolescent, sharply

and obtusely triangular, peduncle with short lateral

setae; eyes clear (adult) or with pale purple cores

(juveniles) in alcohol.

FEMALE.—Gnathopod 1 with narrower lobe on

article 2 than in male, much weaker article 4 with

small process, article 5 longer and with broader posterior lobe, palm not excavate; gnathopod 2 with narrower article 6 bearing slightly S-shaped palm defined

by spine, dactyl slightly overlapping palm; peduncle

of antenna 2 more slender than in male, peduncle of

antenna 1 reaching only to end of article 4 on peduncle

of antenna 2.

JUVENILE.—Uropod 1 with a few lateral setae and

pereopod 3 with 1-2 notches; palm of gnathopod 2

on young male smooth like that of female.

HOLOTYPE.—Bishop Museum collections, catalog

number 7255, male, 4.0 mm.

TYPE-LOCALITY.—JLB Hawaii 14, 1 mi N of Kualoa

Point, Oahu, intertidal, algae, dead coral heads, 23

May 1967.

MATERIAL.—JLB Hawaii 5(36), 6(8), 8(3), 12(3),

14(13), 15 (7).

RELATIONSHIP.—Ampithoe waialua resembles the

adolescent stages of A. simulans Alderman (J. L. Barnard, 1965b) from Pacific America. The resemblance

occurs in various gross similarities such as antennae

and gnathopods, but A. waialua has a much broader

article 2 on pereopod 3 with an anterior notch and

spine. The palm of male gnathopod 2 in A. waialua

apparently does not pass through a stage of deep

notching and splitting off of a tooth as the male ages,

but gnathopod 2 remains relatively small, the palm

becomes excavate, and a small defining cusp develops.

The hand expands so that the dactyl fails to fit the

full palmar length. Article 4 of gnathopod 2 maintains

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a posterodistal protrusion and the gnathopod generally remains in a juvenoid condition. The spines on

uropods 1-2 of A. simulans are very stout and short

and the palm of male gnathopod 1 is not excavate.

Ampithoe waialua also has a strong relationship to

A. ramondi Audouin, the only circumtropical ampithoe. Gnathopod 2 is close to that of A. ramondi except

for the absence of the anterodistal protrusion on article

6, and gnathopod 1 of A. waialua is, of course, a small

edition of gnathopod 2 in the excavate palm. The lobe

on article 2 of gnathopod 2 in male A. waialua is

smaller than that on A. ramondi, and the hand of

gnathopod 1 in both sexes of A. ramondi is very weakly

developed and scarcely subchelate. Californian populations of A. ramondi resemble A. waialua in the partial coalescence of the inner lobes on the lower lip with

the medial lobules of the outer lobes; that condition may be a result of preservation, however, as

Chevreux and Fage (1925) show the inner lobes as

free in Mediterranean material and Hawaiian members of A. ramondi have free lobes.

DISTRIBUTION.—Hawaiian Islands.

Ampithoe species

FIGURE 17j-p

MATERIAL.—J LB Hawaii 17 (1), 18 (1).

NOTES.—Tw o females of a species belonging to the

same group as A. orientalis differ from A. orientalis in

the parachelate palms of the gnathopods, slightly

longer hands, with gnathopod 2 of the same slenderness as gnathopod 1 in contrast to gnathopod 2 on A.

orientalis female that is much stouter and bears a distinct lobe on article 5; gnathopod 2 has article 5 about

10 percent shorter and article 6 about 10 percent longer

than shown for the figured gnathopod 1; articles 2 and

4 of pereopods 1-2 are even more tumid than those on

A. orientalis; antenna 2 is extremely short and stout

in comparison to that of A. orientalis, the flagellum

slightly longer than article 5 of the peduncle; uropod

3 is enlarged like small individuals of the American

A. lacertosa Bate (J. L. Barnard, 1965b) ; the apical

lobules of the lower lip are short and extend subequally

in contrast to those of A. orientalis.

Pereopods 1-5 have locking spines and dactyls like

those of A. orientalis and Figure 17o represents both

species; the distal locking spines are formed of a pair

of weak, striate sabres, the next proximal spine being

straight and axially striate.
NUMBER 3 4 55

FIGURE 20.—Ampithoe waialua, new species, holotype, male, 4.0 mm, JLB Hawaii 14: a, head;

b, telson; c, medial gnathopod 2; d, apex of medial gnathopod 1; e, apex of gnathopod 2, medial;

/, anterior notch on article 2 of pereopod 3; g, gnathopod 1. Female, 4.9 mm: h, apex of medial

gnathopod 2; i, anterior notches on article 2 of pereopod 3; j , gnathopod 1; k, gnathopod 2.
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Ampithoe waialua, new species, holotype, male, 4.0 mm, JLB Hawaii 14: a,

pereopod 3; b, c, d, uropods 1, 2, 3; e, apex of pereopod 5; /, pereopod 2; g, lower lip; h, i,

pereopods 4, 5; j , mandibular palp.
NUMBER 34 57

Uropod 1 has a large peduncular tooth like that of

A. orientalis, and the tooth of uropod 2 is obsolescent.

The eyes are medium in size, with faint purple cores

and small ommatidia, whereas the eyes of A. orientalis

are large, clear in alcohol, and have large ommatidia.

Pleonal epimera, like those of A. orientalis, are

rounded posteroventrally but bear lateral ridges.

Head with weak anteroventral excavation in Ampithoe species and strong in A. orientalis.

Anterior notches and spines on article 2 of pereopod

3 are large like those of A. waialua. Those of A. orientalis are smaller.

REMARKS.—This species is obviously distinct from

any other in the Hawaiian Islands but whether it is new

cannot be decided until males are collected. It apparently has relationships to the boreal North Pacific

group with thin, transverse (or parachelate) palm of

gnathopod 1, but Hawaiian A. orientalis also has the

same boreal aspect even though it, or a species like it,

was described originally from the Philippine Islands.

Whether the Hawaiian A. orientalis is indeed the same

species as the type described by Dana from Manila is

now debatable in view of the strong endemicity found

in Hawaiian Gammaridea. One also may wonder if

the early collecting labels were highly accurate as

several cases of incorrect labeling have been noted or

suspected in Amphipoda.

No North Pacific Ampithoe with parachelate

gnathopod 1 has yet been described (J. L. Barnard,

1965b and Gurjanova, 1951), and the Hawaiian

Ampithoe species probably is a new, endemic species.

Ampithoe (Pleonexes) poipu, new species

FIGURES 22, 23, 24a-k

DIAGNOSIS OF MALE.—All pleonal epimera posteroventrally rounded, lacking lateral ridges; article 6 of

gnathopod 1 much longer than article 5, posterior lobe

of article 5 very narrow and slightly hooked, article 4

with small posterodistal lobe, article 6 rectangular,

moderately thin, palm oblique, weak, defined by striate

spine occurring almost on posterior margin of hand;

gnathopod 2 like gnathopod 1 and not strongly enlarged, distal lobe on article 2 of medium size; article

2 of pereopods 1-2 more than twice as long as broad;

pereopods 3-5 with subprehensile sixth articles similar

to but not as strong as those of A. kaneohe, new species,

large striate sabre spine not guarding smaller palm as

closely as in A. kaneohe, other member of distal pair

very small and nearly straight, next proximal spine

large and next to that one of medium size, dactyls

strongly striate; antenna 1 much shorter than 2,

peduncle of antenna 2 short and stout, flagellum

heavily armed with setular brushes and nearly as long

as peduncle; lateral apical lobules of lower lip much

longer than medial; coxa 1 weakly produced forward;

peduncular process of uropod 1 short and blunt, that

on uropod 2 obsolescent and bluntly triangular, peduncle of uropod 1 with lateral setal row; eyes small

(clear in alcohol but specimens old) ; telson with pair

of large reverted dorsal hooks formed from normal

nobs of other ampithoes.

FEMALE.—Gnathopods 1-2 like male gnathopod 1;

antenna 2 with slightly thinner peduncle and flagellum than on male and flagellum only as long as articles

3-5 of peduncle.

HOLOTYPE.—Bishop Museum collections, catalog

number 7256, male, 9.0 mm.

TYPE-LOCALITY.—Poipu Beach, Kauai, 17 September 1959, IV. 4. d, Chir. haw., collection H. Caspers.

MATERIAL.—Eleven specimens from the typelocality.

REMARKS.—Pleonexes Bate now seems to have been

an artificial genus composed of members of Ampithoe

with dorsal telsonic hooks as extensions of the normal

nobs plus subprehensile pereopods 3-5. This is well

demonstrated with A. (P.) poipu in its affinities with

A. kaneohe, new species, that has stronger palms on

the pereopods but no dorsal hooks and with A. kulafi,

new species (Pleonexes species of J. L. Barnard 1965a),

that has stronger palms of pereopods but no hooks. The

type-species of the genus, P. gammaroides Bate (Sars,

1895, pi. 207) from the Atlantic Ocean has the same

overall aspect of A. (P.) poipu, but the gnathopodal

palms are better developed, the lower lip has no medial

lobules, the antennae are more strongly subequal to

each other, the palmar spines on pereopods 3-5 are

weaker but of similar aspect, and uropod 3 is slightly

enlarged in the female (like that of Ampithoe species

herein).

There is undoubtedly some genetic connection

among A. (P.) gammaroides and A. (P.) poipu as

seen also in the special shape of the peduncular process

on uropod 1, but A. kaneohe is equally to be considered

in this connection, for it resembles P. gammaroides

in the obsolescence (or concealment) of medial lobules
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 22.—Ampithoe (Pleonexes) poipu, new species, holotype, male, 9.0 mm, Poipu Beach,

Kauai: a, coxa 5; b, c, pereopods 1, 4; d, mandibular palp; e, f, gnathopods 1, 2. Female, 7.0 mm:

g, gnathopod 2; h, pereopod 3.
NUMBER 34 59

FIGURE 23.—Ampithoe (Pleonexes) poipu, new species, holotype, male, 9.0 mm, Poipu Beach,

Kauai: a, b, head; c, posteroventral corner on article 2 of pereopod 4; d, mandible; e, uropod 3;

/, maxilliped; g, maxillipedal palp article 4; h, i, maxillae 1, 2; j , inner plate of maxilliped: k,

lower lip.

;S60-034 O—70-
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 24.—Ampithoe (Pleonexes) poipu, new species and Ampithoe (Ampithoe) kaneohe, new

species. A. (P.) poipu, holotype, male, 9.0 mm, Poipu Beach, Kauai: a, inner ramus of uropod 3;

b, urosome; c, uropod 2; d, palm of medial gnathopod 2; e, outer ramus uropod 3; /, telson.

Miscellaneous loose pereopods in this sample, possibly of this species: g, h, ?pereopods 4, 5.

Female, 7.0 mm: i, dactyl of pereopod 3; ;, left side of telson (compare with 24 /); k, pleonal

epimera 1-3. A. kaneohe: I, left side of telson (compare with 24 j).
NUMBER 34 61

on the lower lip unlike A. (P.) poipu and also shares

the same uropod 1 process.

Pleonexes lessoniae Hurley (1954a) is a member of

the group of ampithoes bearing thin hands of gnathopod 1 with transverse palm. The palms of pereopods

3-5 resemble those of gnathopod 1. The outer rami of

uropods 1-2 lack dorsal spines and the outer lobes of

the lower lip are scarcely modified apically, bearing

only one small nipple instead of two lobes.

DISTRIBUTION.—Hawaiian Islands.

Cymadusa filosa Savigny

Cymadusa filosa Savigny.—-J. L. Barnard, 1955a, pp. 29-30,

fig. 15 [with references]; 1965a, p. 540.

MATERIAL.—J LB Hawaii 2 (3), 12 (1), 13 (8), 14

(4), 15 (5). Fee l (3).

DISTRIBUTION.—Circum tropical.

Cymadusa hawaiensis (Schellenberg)

FIOURES 25, 26

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

DIAGNOSIS OF MALE.—Pleonal epimera 1-2 with

lateral ridges, all with small rounded posteroventral

tooth, notch and setule; gnathopods 1-2 elongate,

articles 5-6 of gnathopod 1 equal to each other in

length, article 4 with large conical posterodistal extension partly guarding article 5, latter with truncate,

setose posterior lobe with quadrate corner, article 6

about as broad as 5 but basally constricted, palm very

oblique, shorter than posterior margin of hand, defined

by spine, dactyl overlapping palm, article 2 with

broadly triangular anterodistal lobe; gnathopod 2

larger than 1, lobe on article 2 low and obtuse, extension on article 4 smaller than on gnathopod 1, article

5 shorter than 6, posterior lobe slightly convex, article

5 of same size as on gnathopod 1 but less setose, article

6 large, similar in shape to that of gnathopod 1, palm

slightly more prominent, defined by large spine, dactyl

overlapping palm and closing across medial face of

hand; article 2 of pereopods 1-2 about 3 times as

long as broad; pereopods 3-5 with simple sixth articles

bearing pair of straight locking spines, full lengths of

anterior margins bearing more spines in tandem, dactyl with a few striations; article 2 of pereopod 3 of

medium breadth, pyriform, with stout posterodistal

spine and not lobate, pereopods 4-5 with narrower,

pyriform article 2 bearing row of several posterodistal

marginal spines, anterior margin of article 2 of pereopods 3-5 lined with stout short spines; antenna 1 as

long as head and first 8 body segments, antenna 2

as long as head and first 5 body segments, accessory

flagellum 2-articulate, article 2 on peduncle of antenna

1 slightly longer than article 1, article 3 short, flagellum

of antenna 2 nearly as long as peduncular articles 4—5

together; apical lobules on lower lip of medium tumidity but lateral lobules projecting much farther than

medial; coxa 1 produced forward, peduncular process between rami of uropod 1 long, that on uropod 2

obsolescent, sharp and leaflike, peduncle of uropod 1

with several ventrolateral setae, uropod 3 with many

peduncular spines, outer ramus slightly shorter than

inner, latter with 2 or more medial spines; eyes ochraceous, clear in alcohol.

FEMALE (from Schellenberg, 1938).—Gnathopods

weaker than in male, article 4 angular but not produced, gnathopod 2 like gnathopod 1, article 4

triangular.

ABERRATION.—Male, 10.3 mm, with right uropod

3 bearing extra, blunt apical spine on outer ramus considerably shorter than inner.

MATERIAL.—Hanauma Bay, Oahu, 15 May 1946 (3

males) ; Waikiki Marine Laboratory, from Padina, 11

March 1964 (1 male).

DISTRIBUTION.—Schellenberg: "North Islands";

Kaneohe Bay, Oahu.

Paragrubia vorax Chevreux

FIGURE 32h-m

Paragrubia vorax Chevreux, 1901, pp. 427-431, figs. 50-

55.—J. L. Barnard, 1955a, pp. 31-34, fig. 17 [with references]; 1965a, pp. 541-542, fig. 33.

A few figures of Hawaiian specimens supplementing those of Barnard (1955a) are presented.

MATERIAL.—JLB Hawaii 14 (3). Fee 1 (1). Hanauma Bay, Oahu, 15 May 1946 (1). Molokai, 1959,

Algen, collector H. Caspers (2).

DISTRIBUTION.—Indo-Pacific.

Family ANAMIXIDAE

Anamixis stebbingi Walker

FIGURE 27

Anamixis stebbingi Walker, 1904, pp. 259-261, pi. 3, fig.

18.—J. L. Barnard, 1965a, pp. 488-489, fig. 4.

MATERIAL.— J L B Hawai i 6 (1) , 10 (2) , 12 (1) ,

13 (2) , 15(1) .
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Cymadusa hawaiensis Schellenberg, male, 10.3 mm, Hanauma Bay: a, b, gnathopods

1, 2; e, d, pereopods 1, 4; e, antenna 2; /, telson; g, coxa 4. Male, 8.2 mm, Waikiki, Padina:

h, head.
NUMBER 34 63

FIGURE 26.—Cymadusa hawaiensis Schellenberg, male, 10.3 nun, Hanauma Bay, a, pereopod 3;

b, uropod 3, normal. Male, 8.2 mm, Waikiki, Padina: c, pleonal epimera 1-3; d, lateral apex of

gnathopod 2; e, dactyl of pereopod 4; /, lower lip; g, uropod 3, abnormal.

REMARKS.—The anteroventral cephalic corner is

either typical of Micronesian individuals, with a small

sharp tooth at the corner, or it is perfectly quadrate.

The material is in superior condition and one may see

in better detail than heretofore that the mouthparts

normally said to have become vestigial or absent have

a recognizable structure; 3 pairs of tiny vestiges possibly representing mandible and maxillae 1-2 occur

on either side of the ventral keel. The presumed

mandible appears as a tiny "palp" with a long terminal

seta, maxilla 1 appears as a small lobe dorsolateral to

a larger lobe presumed to be maxilla 2.

COLOR IN 2-DAY FORMALDEHYDE.—Either white with

ruby eye or body with wide vertical pink-red stripes on

middle of each of pereonites 1-7, becoming paler

posteriorly on pereon.

DISTRIBUTION.—Indo-Pacific tropics from Ceylon to

Hawaii.

Family AORIDAE

Aloiloi, new genus

DIAGNOSIS.—Antennae equal in length, as long as

head and 5.5 pereonites, accessory flagellum multiarticulate, article 3 of antenna 1 as long as article 1, recessment of head for antenna 2 moderate; epistome unproduced anteriorly; mandibular palp stout, dominating mandibular body; inner plate of maxilla 1 lined

medially with setae; maxillipedal palp article 4 very

short, bearing long apical spine; article 4 of gnathopod

1 not produced, article 5 short, broad, cup-shaped,

article 6 broad and long, broader than article 5, bearing oblique palm, gnathopod 1 lacking any teeth;

gnathopod 2 smaller than 1 but large for family, similar to but thinner than gnathopod 1, poorly setose

anteriorly on article 5; rami of uropod 3 equal to each

other in length and slightly shorter than peduncle.

TYPE-SPECIES.—A. nenue, new species.

RELATIONSHIP.—This genus superficially resembles

Lembos Bate, but the structure of its gnathopods,

antennae, and maxillae suggest that it may represent

a species with affinities to the genus Gammaropsis

Liljeborg of the Isaeidae. Gnathopod 2, though smaller

than gnathopod 1, has all the characters typical of

many members of Gammaropsis, including the excavate palm and giant medial spine, maxilla 1 has

setae on the medial edge of the inner plate like many

species of Gammaropsis, and article 3 of antenna 1 is

elongate.

One would have to assume that an axial reversal in

gnathopodal dominance has produced the enlarged

gnathopod 1. It also has the appearance of a gnatho-
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 27.—Anamixis stebbingi Walker, specimen 1.8 mm, JLB Hawaii 10: a, lateral view of

body. Male, 2.2 mm, JLB Hawaii 12: b, telson and lateral wings of pleonite 6; c, lateral view of

ventral cephalic margin: l=mandible, 2=maxi!Ia 1, 3 = maxilla 2; d, maxillipeds; e, gnathopod

1; f, lateral lobe of head; g, head; h, gnathopod 2.
NUMBER 34 65

pod 2 on various members of Gammaropsis. A strong

resemblance to Gammaropsis pali, new species, occurs

in Aloiloi.

This genus may assist in demonstrating that various

aorids have had diverse origins from isaeids (photids)

and do not represent a fundamental line of evolution

with a common ancestor.

Aloiloi nenue, new species

FIGURES 28, 29, 30

DIAGNOSIS.—With the characters of the genus.

DESCRIPTION.—Head with medium-sized trapezoidal lobe bearing ovatocircular eye light purple in

alcohol, a few ommatidia with pigment; anteroventral

corner of head quadrate; gland cone large and sharp;

right mandibular molar with flake; coxa 1 broad,

short, slightly extended and angular anteroventrally;

palm of gnathopod 1 with shallow excavation posterior

to obtusely conical process near dactylar base, palmar

defining corner quadrate and bearing minute ridges,

posteromedial face of hand bearing submarginal stout

spine proximal to palmar corner, spine smaller than

FIGURE 28.—Aloiloi nenue, new genus, new species, holotype, male, 4.4 mm, JLB Hawaii 13:

a, b, head; c, d, maxillae 1, 2; e, f, mandibles; g, mandibular palp; h, mandibular lobe on lower

lip; i. outer plate of maxilliped; j , urosome.
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 29.—Aloiloi nenue, new genus, new species, holotype, male, 4.4 mm, JLB Hawaii 13:

a, maxilliped; b, c, medial apices of gnathopods 1, 2; d, e, lateral gnathopods 1, 2; /, pereopod 2;

g, ventral peduncular tooth on uropod 2; h, proximal part of medial gnathopod 1.
NUMBER 3 4 67

on gnathopod 2, dactyl shorter than palm, article 5

of gnathopods 1-2 anteriorly spinose, hand of both

gnathopods with groups of stout anteromedial setae,

gnathopod 2 similar to 1 but smaller, thinner, and with

spine on hand larger and more strongly submarginal,

article 2 of gnathopods 1-2 with sharp anterodistal,

lateral cusp; pereopods 3-5 with rectangular article 2,

bearing seta or weak spine on unlobate posteroventral

corner, lacking facial spines on article 5, locking spines

straight, dactyl bearing one distal facial setule; pleonal

epimeron 1 strongly bulging posteriorly, with slightly

posteroventral tooth, epimeron 2 weakly bulging and

with obsolescent tooth, epimeron 3 with medium bulge,

strong notch and rounded posteroventral cusp; outer

rami of uropods 1-2 slightly shorter than inner, uropod

1 bearing large ventral peduncular tooth, uropod 2

with obsolescent sharp tooth, uropod 3 bearing peduncular spines and small barrel-shaped article 2 on outer

ramus; telson with apical nipple and 1 dorsolateral

spine and nob on each side; sternites lacking teeth.

COLOR.—Eyes garnet in formaldehyde (faint purple

in alcohol), article 3 of antenna 1 and terminal 4

articles of antenna! flagella bright pink.

HOLOTYPE.—Bishop Museum collections, catalog

number 7257, male, 4.4 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

FIGURE 30.—Aloiloi ntnut, new genus, new species, holotype, male, 4.4 mm, JLB Hawaii 13: a, b,

pereopods 5, 4; c, thoracic sternites; d, part of pereopod 3; e, uropod 3; /, telson; g, pleonal

epimera 1-2.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Oahu, 3-4 m, corals, fleshy algae, corallines, 8 April

1967.

DISTRIBUTION.—Hawaiian Islands.

Aoroides ?columbiae Walker

FIGURES 31, 32a-g

? Aoroides columbiae Walker, 1898, p. 285, pi. 16, figs. 7-

10.—Thorsteinson, 1941, pp. 83-84, pi. 6 figs. 65-66.—

J. L. Barnard, 1954a, pp. 24-26, pi. 22.

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

DESCRIPTIVE NOTES.—Article 3 of antenna 1 slightly

more than half as long as article 1; medium-sized and

terminal males with large spine on coxa 1, young males

lacking a coxal spine but on gnathopod 1 bearing spine

on article 6 subsequently lost by terminal males, subapex of dactyl with several stout spinules besides setae;

gland cone apically truncate; maxillipedal palp article

3 with apical protrusion; inner plate of maxilla 1 with

one long curved seta; uropods 1-2 with long, acute

peduncular cusp on ventral side; outer ramus of uropod 3 with small barrel-shaped article 2.

COLOR IN 2-DAY FORMALDEHYDE.—Body white with

a few splotches of faded brown-purple pigment, eyes

bright orange or brownish orange, remaining pale

orange in alcohol.

MATERIAL.—JLB Hawaii 3 (3), 5 (5), 6 (2), 12

(2), 13 (9),14(?2).

FIGURE 31.—Aoroides columbiae Walker, male, 2.1 mm, JLB Hawaii 12: a, gnathopod 1; b,

body; c, telson; d, maxilla 2, setae removed and marked with pits; e, apex of maxillipedal palp,

setae removed. Male, 2.3 mm: /, head; g, gnathopod 2, enlarged; h, gnathopod 1.
NUMBER 3 4

m

FIGURE 32.—Aoroides columbiae Walker and Paragrubia vorax Chevreux. A. columbiae, male,

2.3 mm, JLB Hawaii 12: a, uropod 3; b, apex of uropod 2; c, inner plate of maxilliped. Male, 2.4

mm, JLB Hawaii 3: d, gnathopod 1; e, mandibular palp. Female, 2.2 mm, JLB Hawaii 3: /,

gnathopod 1. Male, 2.1 mm, JLB Hawaii 12: g, pleonal epimera 1, 2, 3. P. vorax, male, 10.0 mm,

JLB Hawaii 13: h, head; i, apex of pereopod 3; j , apex of pereopod 5; k, uropod 3; /, apex of

outer rarnus on uropod 3; m, apex of pereopod 5.
70

REMARKS.—The Hawaiian population has minor

differences from any stages so far described from the

cold and warm-temperate regions of Pacific North

America, but it is pointless to attach any nomenclatural significance to these minor differences until the life

history of the species can be thoroughly examined. The

largest male at hand (2.4 mm long) is not as fully

developed as that described by J. L. Barnard (1954a),

but coxa 1 has undergone the expected terminal modification, article 4 of gnathopod 1 is attenuate, but articles 5 and 7 are shorter and article 6 stouter than in

the cold-temperate male; thus, the Hawaiian male resembles the younger stages of the cold-temperate populations. Young Hawaiian males lack the dense brushes

of setae on article 2 of gnathopod 1 seen in mainland

juvenile males.

DISTRIBUTION.—Cold and warm-temperate Pacific

America and the Hawaiian Islands.

A oroides nahili, new species

FIGURE 33

DIAGNOSIS.—Similar to Aoroides columbiae Walker

but differing in the presence of eyes retaining black or

brownish-purple pigment in alcohol, the asymmetrically and obtusely coniform lateral lobes on the head,

the short palp article 4 of the maxilliped and thin

spines on the inner plate, the less divergent plates of

maxilla 2, the thinner articles 5-6 of male gnathopod

2 and female gnathopods 1-2; in the terminal male

gnathopod 2 the anteroproximal turgidity of article 5 ;

uropod 2 lacking a peduncular tooth between the

rami; the fully developed uropod 3 with 1 spine in the

middle of the outer ramus and only 1 terminal seta

on article 2.

Pereonite 5 retaining diffuse brownish-purple pigment in alcohol.

HOLOTYPE.—Bishop Museum collections, catalog

number 7258, male, 2.3 mm.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, other, algae, corals, 24

May 1967.

MATERIAL.—JLB Hawaii 12 (2), 13 (7), 14 (2),

17(16).

REMARKS.—The diagnosis contains only the gross

morphological differences from A. columbiae and thus

describes the relationship. This species occurs in shallow waters from intertidal to about 5 m, whereas. A

columbiae occurs in waters from about 4 to 30 m.

DISTRIBUTION.—Hawaiian Islands.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Konatopus, new genus

DIAGNOSIS.—Article 3 of antenna 1 as long as article

1, accessory flagellum short, 2-articulate; head elongate

and deeply recessed ventrally for attachment of antenna 2; article 3 of mandibular palp stout, clavate;

inner plate of maxilla 1 with medial setae; male

gnathopod 1 like that of Microdeutopus Costa, with

elongate article 5 bearing short posterodistal tooth,

article 6 shorter and more slender than article 5, nearly

simple; gnathopod 2 much smaller than gnathopod 1,

weakly subchelate, article 5 longer than 6 and scarcely

setose anteriorly; coxae touching or overlapping serially; rami of uropod 3 only slightly longer than peduncle, outer with small barrel-shaped article 2; telson

broader than long, posteroventrally pointed, scarcely

excavate dorsally; male pereonite 1 and coxa 1 becoming extremely elongate anteroposteriorly in terminal

stage.

TYPE-SPECIES.—Konatopus paao, new species.

RELATIONSHIP.—The species on which this genus

is based might fit either Neomegamphopus Shoemaker

(1942) or Amphideutopus J. L. Barnard (1959) if

those monotypic genera were broadened for its inclusion, but in so doing one would have to join together

the two monotypic genera with the species at hand as

a third species. Konatopus paao clearly stands between

the other two genera by sharing characters of both

and may stand somewhere on the evolutionary pathway. One might visualize Neomegamphopus coming

to Hawaii from the warm-temperate of North America,

eventually differentiating and returning to North

America as an organism something like Konatopus

that later differentiated into Amphideutopus. The

converse pathway is perhaps even more probable.

Konatopus has a strongly enlarged coxa 1 in terminal

males, a character of rudimentary development in

Neomegamphopus and not occurring in Amphideutopus. Konatopus has the short uropod 3 of Neomegamphopus, but male gnathopod 1 is of the short

simplified kind seen in Amphideutopus. In fact, gnathopod 1 of Konatopus has exceeded both of the other

genera in the obsolescence of the tooth on article 5

while developing a palmar protrusion on article 6.

Neomegamphopus has gnathopod 2 nearly simple in

males and females, and gnathopod 1 in the female is

very weak, whereas Konatopus has a stronger

gnathopod 2 in males and females, while Amphideutopus has a large stout gnathopod 2 in males, with

articles 5 and 6 equal in length and strongly setose.
NUMBER 34 71

FIGURE 33.—Aoroides nahili, new species, male, 2.2 mm, JLB Hawaii 17: a, head; b, mandibular

palp; c, apex of medial gnathopod 2; d, lateral gnathopod 1; e, f, apex of maxillipedal palp;

g, maxilla 2, setae marked with pits; h, inner plate of maxilliped, setae marked with pits; i,

gnathopod 2. Female, 2.0 mm: ;, k, apices of gnathopods 2 and 1.
72

The three genera form a group of aorids distinguished from Microdeutopus and most other aorids by

the deeply recessed ventral cephalic margin and the

medial placement of maxillary setae on the inner plate

plus the clavate stout mandibular palp article 3 and

elongate maxillipedal palp with clavate article 4. The

geographic "triangulation" of the three genera suggests the extremity of their characteristics is of only

specific value, but in the sense of aorid classification

as it now stands the characters mark good genera.

Shoemaker has already pointed out how these

characters as seen in Neomegamphopus are uncharacteristic of aorids and recall characters of certain genera

of isaeids. One might suggest that a megamphopal

isaeid has undergone axial reversal of gnathopods

parallel to those of normal aorids derived from nonmegamphopal isaeids.

The elongate article 1 of the mandibular palp distinguishes Grandidierella Coutiere from Microdeutopus, Neomicrodeutopus, and Konatopus.

Konatopus paao, new species

FIGURES 34, 35

DIAGNOSIS.—With the characters of the genus.

DESCRIPTION.—Eyes ovate vertically, a dark core

surrounded by pale ommatidia (in alcohol), lateral

lobes of medium forward extent but thick and solid

from side to side as in Amphideutopus and Neomegamphopus; antennae equal in projection; epistome

rounded anteriorly; mouthparts generally as in Neomegamphopus; coxa 1 of terminal male with medial

papilla, article 2 of gnathopod 1 grossly swollen, article 4 with slight distal process, palm of article 6

with protrusion, dactyl thick; coxae 3-4 slightly excavate posteriorly; article 2 of pereopods 3-5 slender

but not completely linear, pereopod 3 of terminal male

with article 2 proximally expanded (only one pereopod

3 of a terminal male at hand, possibly aberrant);

female gnathopods 1—2 indistinguishable from each

other in general size but article 5 of gnathopod 1

longer than that of gnathopod 2; uropod 1 with long

ventral peduncular cusp, absent on uropod 2; pleonal

epimera 1-3 with bulging posterior margins, posteroventral corners with notch and rounded protrusion to

ventral margin, pleonite 4 with dorsal seta on either

side marking dorsal terminus of lateral margin extension (as in Gammaropsis atlantica), pleonite 5 with

only dorsolateral seta.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

COLOR IN 2-DAY FORMALDEHYDE.—Eyes dirty orange

or brownish-orange, body generally clear or occasionally pereonite 5 with wide vertical brownish-purple

stripe; in alcohol cores of eyes retaining brownishorange, turning darker than in life, or rarely eyes

turning pale orange.

HOLOTYPE.—Bishop Museum collections catalog

number 7259, male, 2.5 mm.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

Oahu, 3-4 m, corals, algae, corallines, 8 April 1967.

MATERIAL.—JLB Hawaii 2 (13), 3 (5), 6 (11), 8

(1),10(4),12 (2),13(43),14(1),15(1).

DISTRIBUTION.—Hawaiian Islands.

Lembos ?aequimanus Schellenberg

FIGURE 36a-*

?Lembos (Bemlos) aequimanus Schellenberg, 1938, pp. 76-

77, fig. 39.

Not Lembos {Bemlos) aequimanus.—J. L. Barnard, 1955a,

p. 34.

DIAGNOSIS OF FEMALE.—Lateral lobes of head

strongly extended forward but narrow and mamilliform, anteroventral cephalic excavation weakly

concave, anteroventral corner quadrate, scarcely extended ; eyes circular, purple cores surrounded by half

row of clear ommatidia; inner plate of maxilliped with

normal terminal spines; gnathopod 1 narrow, especially elongate, rectangular article 6 with oblique, excavate palm bearing weak distal protrusion and strong

defining cusp armed with spine, appendage weakly

setose for Lembos, article 2 with strong, sharp anterodistal cusp; gnathopod 2 like that figured for male

L. pualani, article 2 lacking anterodistal process, palm

very oblique, setation moderate; pleonal epimera 1-3

with bulging posterior margins, all with lateral ridge,

slight notch and tooth at posteroventral corner, ventral setae absent; uropod 3 with very short peduncle

and long rami as figured for L. kamanu, inner ramus

twice as long as peduncle and longer than outer ramus,

latter with small article 2 present.

DESCRIPTION.—Antennae and pereopods 3-5 missing; epistome rounded anteriorly; mouthparts generally like those for L. leapakahi, new species, but left

mandibular molar lacking "shark-tooth"; urosome and

uropods appearing similar to those of L. leapakahi, but

spines thinner, peduncle of uropod 1 with basolateral

spine and distolateral seta but not lateral setae; peduncular tooth between rami of uropod 1 shorter than
NUMBER 34 73

FIGURE 34.—Konatopus paao, new genus, new species, holotype, male, 2.5 mm, JLB Hawaii 13:

a, head; b, dactyl of gnathopod 1; c, articles 1—3 of pereopod 1; d, e, gnathopods 2, 1; /, uropod

3; g, maxillipedal palp articles 3—4; h, pleonites 2-6, left to right. Female, 2.3 mm: i, telson with

lateral view of one side offset; j , k, gnathopods 2, 1.
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 35.—Konatopus paao, new genus, new species, female, 3.0 nun, JLB Hawaii 13: a,

pereopod 5; b, pereopod 3. Female, 2.3 mm: c, head; d, articles 1-4 of pereopod 2. Male, holotype, 2.5 mm: e, mandibular palp; f, pereopod 2; g, epistome and labral complex from right

lateral view; h, inner plate of maxilla 1; i, pereopod 3, less coxa.

in other species of Hawaiian Lembos and tooth of

uropod 2 obsolescent; telson large, strongly elevated,

moderately excavate dorsally, with small pair of distolateral nobs, deep slit from lateral view and pair of

long dorsal setae (appearance like that figured for L.

kamanu). Urosomites as figured for L. pualani (see

figure group for L. macromanus).

COLOR IN ALCOHOL.—Body dull white.

MATERIAL.—JLB Hawaii 17 (1).

REMARKS.—My former identification of this species

(1955a) from Hawaii was erroneous and that material

is referred to L. pualani, new species, based now on

fully adult specimens. The female at hand has gnathopod 1 like that figured by Schellenberg; its configuration is unusual for the genus in the presence of an

excavation and weak distal protuberance. The species

is clearly distinct from all other Hawaiian members of

the genus but many characters cannot be compared

with Schellenberg's Gilbertian-Fijian species as the material at hand has no antennae or pereopods 3-5 and
NUMBER 3 4 75

FIGURE 36.—Lembos aequimanus Schellenberg and Lembos species. Lembos aequimanus, female,

4.2 mm, JLB Hawaii 17: a, head; b, lateral gnathopod 2, setae removed; c, lateral gnathopod 1;

d, e, peduncular tooth of uropods 1, 2. Lembos species, female, 4.4 mm, Fee 1: /, medial

gnathopod 2, setae removed; g, lateral gnathopod 2; h, uropod 3; i, dorsal margin of urosome.

360-034 O—7C
76

Schellenberg made no notes on the unique character

of uropods 1-2 as described herein.

DISTRIBUTION.—Gilbert, Fiji, and possibly the Hawaiian Islands.

Lembos intermedius Schellenberg

FIGURE 37

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

J. L. Barnard, 1955a, p. 34.

DIAGNOSIS OF MALE.—Lateral cephalic lobes

strongly extended forward and truncate, anteroventral

margin of head excavate, nearly flat-bottomed, then

produced to weak, obtuse anteroventral cusp; eyes subcircular, brownish purple cores in alcohol surrounded

by 1—2 layers of clear ommatidia; coxa 1 short and

broad, rectotrapezoidal, with slight extension and almost sharp anteroventral corner; gnathopod 1 of

medium to strong enlargement, article 2 of normal

stoutness, anterodistal corner with strong, thin mammilliform lobe, article 4 with distal point not free, article 5 of medium breadth, less than half as long as article

6, latter expanded, bulging anteriorly, moderately

setose medially, palm very oblique, bearing deep incision separating 2 processes, unarmed defining process

short, thus making palm oblique, dactyl slender and

scarcely overlapping palm; article 2 of gnathopod 2

with large anterodistal lobe similar to that of gnathopod 1, remainder of appendage like figure of L. pualani, article 4 with distal point not free, articles 5 and 6

equal to each other in length, palm nearly transverse,

defined by spine and slight protuberance, setation

dense; thoracic sternites 2-5 each with low, broad,

obtusely bifid ventral tooth, sternite 1 with single obtuse tooth, sternite 6 with small single cusp; pleonal

epimera 1—3 with bulging posterior margins, weak

lateral ridges and slight notch and tooth at posteroventral corners; ventral setae absent or epimeron 2

with 1 seta; uropod 3 much smaller than in other

Hawaiian species of Lembos, with extension on outer

ramus appearing like that of a second article but apparently coalesced with and as broad as article 1, peduncle with large spine.

FEMALE.—Coxa 1 slightly longer and less pointed

anteroventrally than in male; gnathopod 1 of normal

kind in Lembos, anterodistal corner of article 2 lacking process, otherwise like that of L. pualani, but palm

slightly more transverse and dactyl slightly shorter;

article 2 of gnathopod 2 with anteroventral corner

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sharply quadrate, not lobate; sternites unkeeled; pleonal epimeron 2 with several midventral setae and one

posterior seta besides normal setule in posteroventral

notch, setae absent in smallest juveniles.

DESCRIPTION.—Antenna 1 at least as long as head

and pereon together (thereafter broken off in all

specimens), antenna 2 much shorter and stouter than

in L. waipio, new species, but similar to L. pualani,

new species; epistome slightly rounded anteriorly, remainder of mouthparts like those of L. leapakahi, new

species, but shark-tooth of left mandibular molar

weakly developed; article 2 of pereopod 3 with small

posteroproximal extension, pereopods 4-5 missing on

all material; urosome and uropods 1-2 appearing

similar to figure of L. leapakahi, but uropod 3 small,

spines thinner, but peduncle of uropod 1 bearing basolateral spine and distolateral setae, no basolateral setae

on uropods 1-2, both with long tooth between rami,

peduncle of uropod 3 with 2 basolateral setae; telson

especially large, not strongly elevated, scarcely excavate dorsally, with pair of posterolateral nobs forming

slit from lateral view.

COLOR IN ALCOHOL.—Posterior part of head, anterior part of pereonite 1, and posterior part of pereonite

6 deeply purple-brown.

MATERIAL.—JLB Hawaii 10 (5), 12 (4), 13 (13),

15 (9).

REMARKS.—The male is characterized by the bidentate sternal teeth, not described by Schellenberg; by

the oblique palm of gnathopod 1 with scarcely overlapping dactyl; by the small uropod 3 lacking a second

article on the outer ramus and is recognizable by the

color pattern.

J. L. Barnard (1965a) suggested that L. intermedius

might be synonymous with L. processifer (Pirlot,

1938) (not J. L. Barnard, 1965a), but L. intermedius

is now shown to be clearly distinct The bifid sternal

teeth of the male are a major difference, but the heavy

medioterminal setation on male gnathopod 1 article 6

is also noticeable. Male and female gnathopod 2 of

L. intermedius are much smaller, thinner, and more

densely setose than in L. processifer, antenna 2 is

thinner, article 2 of pereopods 3-5 has a sharp posteroproximal extension, and the eyes are dark. The identification of Indonesian L. processifer by J. L. Barnard

(1965a) is erroneous and probably should be associated as a subspecies with L. waipio, new species,

from Hawaii.

Coxa 1 of male L. intermedius is neither as sharp
NUMBER 3 4 77

FIGURE 37.—Lembos intermedius Schellenberg, male, 4.7 mm, JLB Hawaii 12: a, uropod 3; b,

gnathopod 1; c, anterior outline of head; d, apex of outer ramus on uropod 3; e, left mandibular

molar; /, pereopod 3; g, lobe on article 2 of gnathopod 2; h, telson; i, sternal teeth. Female, 4.1

mm, JLB Hawaii 13: j , pleonal epimeron 2; k, part of urosome; /, gnathopod 1, setae removed.

nor bears as elongate setae as represented by Schellenberg (1938), but no other species at hand fits L.

intermedius as well as this one.

DISTRIBUTION.—Hawaiian Islands.

Lembos kamanu, new species

FIGURE 38

DIAGNOSIS OF FEMALE.—Lateral lobes of head

strongly extended forward and truncate or slightly

convex, anteroventral margin of head hemispherically

excavate and bearing broad, conical, obtuse anteroventral corner; eyes circular, pale purple cores surrounded

by 1 layer of clear ommatidia; inner plate of maxilliped with very broad spines on inner plate in comparison to most other Hawaiian species of Lembos:

gnathopod 1 of normal female kind in Lembos but

article 6 sharply quadrate distally, palm with weak

scallops, moderately setose, article 2 lacking anterodistal process; gnathopod 2 like that figured for male
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

k

FIGURE 38.—Lembos kamanu, new species, holotype, female, 3.6 mm, JLB Hawaii 15: a, uropod

3; b, palm of medial gnathopod 1; c, telson; d, "shark-tooth" of left mandibular molar; e, f,

pereopods 3, 5; g, medial gnathopod 1; h, pleonal epimera 1-3; i, head; j , spines on inner plate

of maxilliped; k, urosome.
NUMBER 34 79

L. pualani, article 2 lacking anterodistal process, setation moderate and setae very stout; pleonal epimera

1-3 with bulging posterior margins, no lateral ridges

and slight notch and tooth at posteroventral corners,

ventral setae absent; uropod 3 with very short peduncle and long rami, inner ramus twice as long as

peduncle and longer than outer ramus, latter with

small article 2 present.

DESCRIPTION.—Antenna 1 as long as head and first

7 body segments, antenna 2 as long as head and first

5 body segments, flagellum of antenna 2 with 4 articles and shorter than article 5 of peduncle, accessory

flagellum with 4 articles; epistome rounded anteriorly;

mouthparts except inner plate of maxilliped generally

like those of L. leapakahi, new species, including presence of "shark-tooth" on left mandibular molar; urosome and uropods appearing similar to those of L.

leapakahi but spines thinner, peduncle of uropod 1

with basolateral spine and distolateral seta but no

lateral setae; telson large, strongly elevated, weakly

excavate dorsally, with small pair of distolateral nobs,

deep slit from lateral view and pair of long dorsal

setae. Urosomite 3 very deeply depressed in relationship to urosomite 2 and unique for Hawaiian species of

Lembos; urosomite 2 with long dorsal surface as in L.

pualani but in contrast to L. macromanus.

COLOR IN ALCOHOL.—Ground color of body white

and weakly spotted with faint brownish-purple dots.

HOLOTYPE.—Bishop Museum collections catalog

number 7260, female, 3.6 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 15, 1 mi N of Kualoa Point, Oahu, intertidal, coral block with ascidians,

23 May 1967.

RELATIONSHIP.—As known in the female stage only,

this species is impossible yet to relate to extrinsic members of Lembos, but in the Hawaiian fauna it stands

out as a sibling of L. pualani in which gnathopod 1 has

a weakly scalloped palm, urosomite 3 strongly drops

below the dorsal line of urosomite 2, the main spines on

the inner plate of the maxilliped are elephantine, the

rami of uropod 3 are slightly longer, and the anteroventral cephalic tooth is thicker.

DISTRIBUTION.—Hawaiian Islands.

Lembos leapakahi, new species

FIGURES 39, 40

DIAGNOSIS OF MALE.—Lateral lobes of head strongly

extended forward and truncate, anteroventral margin

of head excavate, flat-bottomed, then produced to

strong anteroventral cusp; eyes obliquely pyriform,

brownish-purple cores in alcohol surrounded by 1-2

layers of clear ommatidia; coxa 1 rounded at anteroventral corner; gnathopod 1 very large, article 2 extremely thick, anteroventral corner unproduced, article 4 with distal point not free, article 5 very wide

and nearly 80 percent as long as article 6, latter almost

subcircular, palm equal in length to posterior margin

of article 6, palm simple, convex, defined by spine and

slight protuberance, dactyl slender and curved, fitting

palm, medial face of hand strongly setose anteriorly,

posterior margin of hand setose; article 2 of gnathopod

2 with small anteroventral mammilliform cusp, article

4 with distal point not free, article 5 longer than 6,

palm of latter scarcely oblique, defined by spine and

slight protuberance, palm much shorter than posterior

margin of article 6, articles 5 and 6 densely setose

medially and marginally except for anteroproximal end

of article 5; thoracic sternites with keel formed of

spiniform tooth on segments 2-3, obtuse keel on segments 4-5 and obsolescent or obsolete on segments 6-

7, segment 1 with tiny obtuse point; pleonal epimeron

1 with a few ventral setae and small posteroventral

notch with setule, epimeron 2 ventrally setose, rounded

posteroventrally, epimeron 3 shortened on posterior

margin, rounded posteroventrally; uropod 3 lacking

article 2 on outer ramus.

FEMALE.—Generally like male even as to gnathopods but latter slightly smaller and gnathopod 2 not as

strongly setose, ventral thoracic keel absent.

DESCRIPTION.—Antenna 1 about 95 percent as long

as head and body together, antenna 2 much shorter;

epistome flat in front, remainder of mouthparts like

those of L. websteri (Bate) (Sars, 1895, pi. 194), but

mandibular lobes of lower lip more attenuate, palp

of mandible slightly longer, left mandibular molar

with complex flat, sharklike tooth on trituration face

(see figure), base of left molar with small cuspidate

rugosity, right molar normal, rugosity at base obsolescent, inner plate of maxilla 1 smaller but its one

seta longer; article 2 of pereopod 3 with posteroproximal triangular extension, pereopods 4-5 missing on

all material; uropods 1-2 with long ventral peduncular

tooth between rami; telson large, strongly elevated,
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 39.—Lembos leapakahi, new species, female, 5.0 mm, JLB Hawaii 17: a, medial

gnathopod 1, setae removed; b, head; c, gnathopod 1. Holotype, male, 6.0 mm: d, gnathopod 1;

e, gnathopod 2; f, pleon.
NUMBER 34 81

FIGURE 40.—Lembos leapakahi, new species, holotype, male, 6.0 mm, JLB Hawaii 17: a, head;

b, side of telson; c, anterior portion of coxa 1; d, palm of gnathopod 2; e, pereopod 1; /,

dactyl of gnathopod 1; g, pereopod 3; h, telson; i, palm of gnathopod 1; j , ventral processes of

pereonites. Female, 5.0 mm: k, left mandibular molar; I, uropod 3; m, half of lower lip.

with pair of small lateral processes on each side, appressed to each other so as to form deep slit from lateral

view; juvenile epimeron 2 with setae reduced or absent.

COLOR IN 2-DAY FORMALDEHYDE.—Body completely

white or very rarely with weak brownish-purple dots.

HOLOTYPE.—Bishop Museum collections catalog

number 7261, male, 6.0 mm.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, other algae, rocks, 24

May 1967.

MATERIAL.—JLB Hawaii 5 (1), 10 (3), 11 (7), 15

(7), 17 (8).

RELATIONSHIP.—This species has affinities with the

west African L. francanni Reid (1951) and the Micronesian Lembos species (J. L. Barnard, 1965a), but

differs from both in the presence of an anteroventral

cephalic cusp. Male gnathopod 2 of L. francanni also

has the dactyl strongly overlapping the palm, not

characteristic of L. leapakahi.

DISTRIBUTION.—Hawaiian Islands.
82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lembos macromanus (Shoemaker)

FIGURES 41, 42a-m, o

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

Lembos macromanus.—J. L. Barnard, 1962a, p. 9, fig. 3.

DIAGNOSIS OF MALE.—Lateral lobes of head strongly

extended forward and truncate, anteroventral margin

of head excavate, nearly flat-bottomed, then produced

to medium-sized anteroventral cusp; eyes subcircular,

brownish-purple cores in alcohol, surrounded by 1-2

layers of clear ommatidia; coxa 1 quadratiform, anteroventral corner softly quadrate; gnathopod 1 very

large, article 2 stout, anterodistal corner produced

downward conically, article 4 with distal point not free,

article 5 broad and short, article 6 very broad, rectangular, anterior margin bulging, poorly setose, palm

transverse, bearing deep incision separating 2 processes,

defining process slender and unarmed, main process

rounded or mammilliform, in terminal stage marked

with basal suture, dactyl slender and slightly overlapping palm in terminal stage, more so in medium age

male; article 2 of gnathopod 2 with weak anterodistal

cusp, article 4 with distal point not free, article 5 much

longer than 6, densely setose medially and anteriorly,

palm oblique but short, defined by spine and slight

protuberance, setation dense; thoracic sternites with

short sharp tooth on segments 2-3, small nipple on 1;

pleonal epimera 1-3 with bulging posterior margins,

lateral ridges and notch and tooth at posteroventral

corners, ventral setate absent; uropod 3 with article 2

on outer ramus and latter shorter than inner ramus,

peduncle with large spines.

FEMALE.—Gnathopod 1 of normal female kind in

Lembos, but especially tumid in terminal stage, article

5 about two-thirds as long as 6; article 2 of gnathopod

2 lacking anterodistal lobe and article 5 not strongly

setose anteriorly; sternal keel absent.

DESCRIPTION.—Antenna 1 nearly as long as head

and body together, antenna 2 much shorter; epistome

flat in front, remainder of mouthparts like those of

L. leapakahi, new species, including left mandibular

molar "shark-tooth"; article 2 of pereopods 4-5 especially setose; urosome and uropods appearing similar

to figures for L. leapakahi, but spines thinner and

peduncles of uropods 1-2 lacking small basolateral

setae, uropods 1-2 with long peduncular tooth between

rami and distolateral peduncle of uropod 1 with 1

seta, no distal spine and 1 basolateral spine, peduncle of

uropod 3 with 2 basomedial setae; telson large but not

strongly elevated, with weak pair of posterolateral

nobs forming slit from lateral view.

COLOR IN ALCOHOL.—Body background pale buff

with numerous splotches and spots of brown, eyes

caramel brown, especially bright.

MATERIAL.—JLB Hawaii 2 (2), 8 (1), 11 (28), 12

(1), 13 (6), 15 (7),Oahu, Coconut Island, 7 September 1959, species 24, collector, H. Caspers (4).

REMARKS.—In Hawaii this species compares very

strongly with the fine original figures of Shoemaker

except that the male sternal teeth are smaller. The

slightly darker body background color in alcohol with

more numerous and larger body blotches and spots

helps one in separating females and juveniles from

those of other species all having a pure white body

background with or without spots. The pale fawncolored pigment on pereonites 2-6 of L. waipio is a

strong contrast to the dark appearance of L. macromanus, and male L. waipio has 3 sternal teeth. Lembos

pualani differs from both L. macromanus and L. waipio in the relatively smooth dorsal margin of the urosome and relatively long dorsal margin of urosomite

2; the stairstep appearance of the urosome in L.

macromanus and L. waipio is easy to see under lowpower magnification.

Uropod 3 and the telson are especially setose in

terminal stages of L. macromanus, the telson bearing

4 dorsal setae in a row on each side but several other

Hawaiian species have at least 3 setae in each row; the

apex of the outer ramus on uropod 3 is densely setose

but only relatively more so than in two other Hawaiian species; the dense setae on article 2 of pereopods

4-5 are only slightly more abundant than those found

in the terminal stage of L. pualani.

DISTRIBUTION.—Warm-temperate Pacific America,

Hawaiian Islands, and reported from the Philippine

Islands.

Lembos pualani, new species

FIGURES 42n, 43

Lembos (Bemlos) aequimanus.—J. L. Barnard, 1955a, p. 34

[not Schellenberg, 1938].

DIAGNOSIS OF MALE.—Lateral lobes of head strongly

extended forward and truncate, anteroventral margin

of head excavate, nearly flat-bottomed, then produced

to medium-sized anteroventral cusp; eyes subcircular,

brownish-purple cores in alcohol, surrounded by 1-2
NUMBER 3 4 83

FIGURE 41.—Lembos macromanus (Shoemaker), male, 6.3 mm, JLB Hawaii 13: a, head; b,

gnathopod 1; c, medial gnathopod 2, setae removed; d, lateral gnathopod 2. Female, 6.1 mm,

Coconut Island, Oahu, 1959, Caspers: e, f, lateral gnathopods 1, 2. Male, 5.0 mm: g, gnathopod

1.
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 42.—Lembos macromanus (Shoemaker) and L. pualani, new species. L. macromanus,

male, 6.4 mm, JLB Hawaii 13: a, apex on outer ramus of uropod 3; b, thoracic sternites 1-4; c,

gland cone, shaded. Male, 6.3 mm: d, uropod 3; e, palmar bulge of gnathopod 1; /, telson; g,

urosomites 1—3 (compare with 42 n); h, apex of telson; i, j , k, pleonal epimera 1, 2, 3; /, side of

telson; m, aberrant outer ramus of uropod 3; n, Lembos pualani, new species, holotype, male,

4.3 mm, JLB Hawaii 14, urosomites 1-3, compare with 42 g. o, pereopod 5 from Coconut Island,

Oahu, 1959, Caspers, specimen unknown, presumed terminal male.
NUMBER 34 85

layers of clear ommatidia; coxa 1 quadratiform, anterior and ventral margins excavate, anteroventral corner sharply attenuate; gnathopod 1 of medium enlargement, article 2 of normal stoutness, anterodistal corner

with medium-sized mammilliform lobe, article 4 with

distal point not free, article 5 of medium breadth,

about 70 percent as long as article 6, latter rectangular,

very strongly setose medially, palm slightly oblique,

bearing deep incision separating 2 processes, defining

process slender but unarmed, dactyl of medium stoutness and strongly overlapping palm; article 2 of

gnathopod 2 with strongly projecting, thin mammilliform anterodistal lobe, article 4 with distal point not

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

palm nearly transverse, defined by spine and slight

protuberance, setation dense medially but not anteriorly ; thoracic sternites lacking keel or teeth; pleonal

epimera 1-3 with bulging posterior margins, weak

lateral ridges and slight notch and tooth at posteroventral corners, ventral setae absent; uropod 3 bearing

article 2 on outer ramus, rami equal to each other in

length, peduncle with large spine.

FEMALE.—Coxa 1 evenly quadrate, with softly

rounded anteroventral corner; gnathopod 1 of normal

female kind in Lembos, but slightly stouter than in L.

waipio, new species, article 5 distinctly shorter than 6;

gnathopod 2 with no anterodistal lobe on article 2.

DESCRIPTION.—Antenna 1 as long as head, pereon,

and metasome combined, exceeding antenna 2 by only

3 (female) to 6 (male) flagellar articles; epistome

slightly rounded anteriorly, remainder of mouthparts

like those of L. leapakahi, new species, but left mandibular molar lacking "shark-tooth"; article 2 of pereopods 3—5 with small posteroproximal extension; urosome and uropods appearing similar to those of L.

leapakhai, but spines thinner and peduncles of uropods

1-2 lacking lateral setae, uropods 1-2 with long

peduncular process between rami, peduncle of uropod

1 with 1 basolateral spine, and distolateral seta, no

distal spine, peduncle of uropod 3 with 2 basolateral

setae; telson large but not strongly elevated, deeply

excavate dorsally, with weak pair of posterolateral

nobs, weakly to strongly slit from lateral view.

COLOR IN ALCOHOL.—Head and pereonites 2-6 with

several small, bright, dorsolateral brownish-purple

spots on white background, or occasionally with large

blotches.

HOLOTYPE.—Bishop Museum collections, catalog

number 7262, male, 4.3 mm.

TYPE-LOCALITY.—JLB Hawaii 14, 1 mi N of Kualoa

Point, Oahu, intertidal, algae, dead coral heads, 23

May 1967.

MATERIAL.—JLB Hawaii 12 (17), 13 (24), 14 (7),

15 (17), 17 (7).

RELATIONSHIP.—This species differs from L. waipio,

new species, in coxa 1, the more strongly setose gnathopods of both sexes, the stouter male gnathopod 1 with

shorter article 5, stouter female gnathopod 1, palm of

gnathopod 2 in both sexes shorter relative to posterior

margin of hand, rami of uropod 3 shorter and equal in

length to each other, article 2 of pereopod 5 slightly

spinier, article 6 with spines proximal to locking pair,

antenna 2 slightly stouter, mandibular molar lacking

shark tooth, and the absence of a spiniform sternal keel

on the male.

Lembos pualani differs from the Gilbertian-Fijian

L. aequimanus Schellenberg (1938) in the unmodified

palm of female gnathopod 2, the highly modified male

coxa 1, the complete absence of male sternal teeth, and

the elongate article 5 on gnathopod 1 of the female.

The Hawaiian species appears most closely related

to but differs from Californian "Lembos concavus

Stout" (J. L. Barnard, 1962a) by the weakness of

anterior setation on article 6 of gnathopod 1 and article

5 of gnathopod 2 in males and the more highly modified coxa 1.

American L. macromanus (Shoemaker, 1925) differs from L. pualani in the strongly setose article 2 of

pereopods 4-5, the presence of sternal teeth in the

male, the weak anterodistal lobe on article 2 of male

gnathopod 2, the much stouter male gnathopod 1 with

short article 5, and the weakly overlapping dactyl.

DISTRIBUTION.—Hawaiian Islands.

Lempos waipio, new species

FIGURES 44, 45

DIAGNOSIS OF MALE.—Lateral cephalic lobes

strongly extended forward and truncate, anteroventral

margin of head excavate, nearly flat-bottomed, then

produced to weak obtuse anteroventral cusp; eyes subcircular, brownish-purple cores in alcohol surrounded

by 1-2 layers of clear ommatidia; coxa 1 rounded at

anteroventral corner; gnathopod 1 of medium enlargement, article 2 of normal stoutness, anterodistal

corner scarcely produced, article 4 with distal point

not free, article 5 broad, but only half as long as article
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

m

FIGURE 43.—Lembos pualani, new species, holotype, male, 4.3 mm, JLB Hawaii 14: a, head; b,

telson; c, apex of medial gnathopod 1, setae represented by pits; d, gnathopod 1; e, f, portions

of pereopods 3, 5, medial; g, uropod 3; h, apex of pereopod 5; i, gnathopod 2, setae represented by

pits. Female, 4.2 mm: j , lateral pereopod 3; k, coxa 1; /, side of telson; m, apex of medial

gnathopod 1.
NUMBER 34 87

6, latter slightly expanded, subrectangular but slightly

tapering distally, anterior margin bulging, poorly setose, palm slightly oblique, bearing deep incision separating 2 processes, defining process slender and

armed with basoposterior stout spine, dactyl slender

and strongly overlapping palm; article 2 of gnathopod

2 with weak or medium anterodistal cusp, article 4

with distal point not free, article 5 longer than 6, palm

oblique but short, defined by spine and slight protuberance, setation moderate; thoracic sternites with

spiniform tooth on segments 2-4, obtuse keel on 5,

weaker on 1; pleonal epimera 1-3 with bulging posterior margins, weak lateral ridges and slight notch and

tooth at posteroventral corners, ventral setae absent;

uropod 3 with article 2 on outer ramus and latter

shorter than inner ramus, peduncle lacking large spine.

FEMALE.—Gnathopod 1 of normal female kind in

Lembos, article 5 nearly as long as 6; gnathopod 2

article 2 lacking anterodistal lobe or that lobe obsolescent ; sternal keel absent.

DESCRIPTION.—Antenna 1 as long as head and body

together, antenna 2 much shorter; epistome flat in

front, remainder of mouthparts like those of L. leapakahi, new species, including left mandibular molar

"shark-tooth"; article 2 of pereopods 3-5 with small

posteroproximal extension; urosome and uropods appearing similar to figures for L. leapakahi, but spines

thinner and peduncles of uropods 1-2 lacking small

basolateral setae, uropods 1-2 with long peduncular

tooth between rami, and distolateral peduncle of uropod 1 with 1 seta, no spine, and 1 basolateral spine,

peduncle of uropod 3 with 1 basolateral seta; telson

large but not strongly elevated, with weak pair of

posterolateral nobs, with lateral slit only in terminal

stages.

COLOR IN ALCOHOL.—Body white except for weak

brown dorsal stripes on pereonites 3-6 or 3-5, eyes

brown.

HOLOTYPE.—Bishop Museum collections, catalog

number 7263, male, 6.1 mm.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, "coralline footballs," 29 January 1967 .

MATERIAL.—JLB Hawaii 3 (10), 5 (1), 6 (14), 14

(2).

RELATIONSHIP.—This species differs from L. intermedius Schellenberg (1938) in the strongly overlapping dactyl on male gnathopod 1, the rounded, asetose

anteroventral corner of coxa 1, and the presence of

simple instead of bidentate sternal teeth on the male.

It differs from L. aequimanus Schellenberg (1938)

in the presence of sternal teeth on more than one

segment in the male, in the rounded palm of female

gnathopod 1, the obtuse anteroventral corner of coxa

1, and the stouter hands of the gnathopods in both

sexes.

Lembos waipio differs from American L. macromanus (Shoemaker, 1925) in the longer dactyl of male

gnathopod 1 and the presence of the posterior spine

on the hand, the stouter, shorter, less setose gnathopod 2, the sparseness of setae on article 2 of pereopods 4-5, the thinner female gnathopod 1, and the

presence of more than 2 sternal teeth on the male.

The Hawaiian species differs from Indonesian L.

processijer (Pirlot, 1938) in the strongly overlapping

dactyl on male gnathopod 1, the thinner peduncle of

antenna 2, the presence of 3 strong sternal teeth in the

male (instead of 2), the dark eyes, slender article 2

of gnathopod 2 lacking a strong anterolateral excavation, the thinner articles 5-6 of gnathopod 2 in both

sexes, the small female gnathopod 1, and the lack of a

strong anterodistal cusp on article 2 of female gnathopod 1.

J. L. Barnard (1965a) identified L. processifer from

Micronesia, but that identification may now be seen

to be erroneous and the Micronesian specimens should

be associated with L. waipio, perhaps as a subspecies

distinguished by the presence of only 2 sternal processes

on the male and a smaller cusp on article 2 of male

gnathopod 2. My 1965a identification was supported

by a supposition that young males of Lembos might

have dactyls fitting palms whereas terminal males

might have dactyls overlapping palms but that circumstance does not justify the Micronesian identification of L. processifer in light of the high diversity of

species in Hawaii that suggests other parts of the tropics

may also have a higher than now expected diversity

in this genus. There seems to be little doubt now that

Pirlot's and J. L. Barnard's materials were both comprised of terminal males.

DISTRIBUTION.—Hawaiian Islands.

Lembos species

FIGURE 36/-»

DIAGNOSIS OF FEMALE.—Lateral lobes of head

strongly extended forward and truncate, anteroventral

margin of head excavate, flat-bottomed or slightly concave, then produced to strong anteroventral cusp (like
88

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 44.—Lembos waipio, new species, holotype, male, 6.1 mm, JLB Hawaii 6: a, body; b, c,

apices of gnathopods 2, 1. Female, 3.8 mm: d, gnathopod 1.
NUMBER 34 89

FIGURE 45.—Lembos waipio, new species, holotype, male, 6.1 mm, JLB Hawaii 6: a, b, article 2

of pereopods 3, 5; c, telson; d, uropod 3; e, pleonal epimera 1-3; /, lower lip; g, mandibular

palp articles 2-3; h, side of telson; i, article 2 of pereopod 4; j , k, articles 5-7 of pereopods 3, 4;

/, sternites of pereon. Female, 3.8 mm: m, tooth of left mandibular molar.

figure for L. leapakahi) ; eyes circular, dark purple

cores surrounded by a layer of clear ommatidia; inner

plate of maxilliped with very broad spines in comparison to other species of Hawaiian Lembos (like those of

L. kamanu) ; gnathopod 1 of normal female kind in

Lembos, but palm with weak excavation near posterior

side, forming weak denning cusp armed with spine,

appendage moderately setose, article 2 with weak anterodistal process; gnathopod 2 like that figured for

male of L. pualani, article 2 with quadrate anterodistal

corner, setation moderate; pleonal epimera 1-3 with

bulging posterior margins, all with lateral ridge and

small notch and tooth at posteroventral corner, ventral

setae absent; uropod 3 with short peduncle and short

rami about 1.3 times as long as peduncle, outer slightly

shorter than inner and bearing small article 2.

DESCRIPTION.—Antennae and perepods 3-5 missing;

epistome rounded anteriorly; mouthparts except inner

plate of maxilliped generally like those for L. leapakahi,

new species, but left mandibular molar lacking "shark-
90

tooth"; urosome and uropods appearing similar to

those for L. leapakahi but spines thinner, peduncle of

uropod 1 with basolateral spine, distolateral seta but no

lateral setae; telson large, strongly elevated, deeply excavate dorsally, with weak pair of posterolateral nobs,

deep slit from lateral view and generally appearing as in

figure for L. pualani; urosomal dorsal margin normal

as in L. pualani.

MATERIAL.—Fee 1(1).

REMARKS.—This individual differs from L. pualani

in the slightly excavate palm of female gnathopod 1,

but the palm is not scalloped as in L. kamanu. The

specimen may be an aberrant member of L. pualani.

DISTRIBUTION.—Hawaiian Islands.

Neomicrodeutopus (?)makena, new species

FIGURES 19/, 46, 47

DESCRIPTION OF MALE.—Head with truncate, trapezoidal ocular lobe bearing subcircular eye with broad

dark core; head poorly recessed for attachment of

antenna 2; mandibular palp very stout, articles 1-3

almost all subequal to each other in length, article 3

broadly clavate; epistome rounded anteriorly; lower

lip and maxilla 1 like those of Microdeutopus anomalus (Rathke) (Sars, 1895, pi. 191), maxilliped also like

that species, inner plate with 3 blade-spines and 2

coupling hooks on back, outer plate with short bladelike semiserrate (3-4 serrations) spines, becoming

longer apically as plumose, blunt spine setae (2), palp

article 4 unguiform but partially formed of 1 stout and

2 slender setae (antennae missing); coxae short, rectangular or rounded-quadrate, coxa 1 rounded anteroventrally, gnathopod 1 like that of Microdeutopus,

article 2 without distal lobes, article 5 very broad and

long and bearing double posteroventral tooth, most

posterior tooth longest, article 5 short, stout, posterior

margin bulging but truncate, palm very short and undefined, dactyl shorter than article 6; gnathopod 2 of

medium stoutness and length, article 5 longer than 6,

unlobate, weakly setose anteriorly or posteriorly, article

6 rectangular, palm nearly transverse but much shorter

than posterior margin of hand, defined by pair of spines

and posterior margin of hand with 2 more spines in

tandem, dactyl fitting palm; pereopods 1-2 of medium

slenderness, but article 2 more than twice as long as

broad, articles 5-6 about subequal in length and longer

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

than article 4, dactyl shorter than article 6; pereopods

3-5 slender, coxa 5 broadly rounded anteroventrally

but coxae 5-7 very short and 6-7 almost equally and

weakly bilobate, article 2 rectangular or tending

toward ovateness in pereopod 5, article 5 with 2 or 3

lateral spines; pleonal epimera 1-3 with protuberant

but truncate posterior margins and slight posteroventral incision bearing stout seta, ventral margins naked;

uropods 1-2 with short stout spines on all rami and

peduncles, outer ramus of uropod 2 significantly

shorter than inner ramus, uropod 1 with ventral peduncular tooth between rami; uropod 3 uniramous,

peduncle medially expanded and very thin, ramus 1.3

times as long as peduncle, 2-articulate; telson with

strong dorsal protrusion on each side, quadratocircular

from dorsal view, posteriorly truncate.

FEMALE.—Gnathopod 1 not strongly enlarged, hand

narrow, palm with angular protrusion and defined by

1 large spine, with 2 proximal and 1 distal adjacent

spines in tandem.

HOLOTYPE.—Bishop Museum collections, catalog

number 7264, male, 4.7 mm.

TYPE-LOCALITY.—Makapuu Point, Oahu, April

1967, on Zoanthus, collector, Dr. Jerry Welch.

MATERIAL.—The type-locality and JLB Hawaii 18

(2).

RELATIONSHIP.—This species differs in numerous

points from the type-species of the genus, N. cabindae

Schellenberg (1925), from west Africa and may not be

phyletically connected to that genus, but several points

about both cannot be compared owing to missing parts

on N. makena and descriptive omissions on N. cabindae. The antennae of N. makena are missing, and we

do not know the state of articles 1-3 and the accessory

flagellum; in N. cabindae the accessory flagellum is

vestigial and 1-articulate and article 3 of antenna 1 is

half as long as the interequal articles 1-2; antennae

1-2 are subequal in length to each other. The extent of

ventral cephalic recession for the attachment of antenna 2 is unknown in iV. cabindae and may be a significant generic character.

Neomicrodeutopus (?) makena differs from N.

cabindae in the stout mandibular palp with equal articles 2-3, the slender gnathopod 2, the narrow article 2

of pereopods 3-5, the configuration of pleonal epimera

and their nakedness, the short dactyl of gnathopod 1,

and the short ramus of uropod 3, bearing 2 articles.
NUMBER 3 4 91

FIGURE 46.—Neomicrodeutopus (?) makena, new species, holotype, male, 4.7 mm, Makapuu

Point, Oahu, 1967: a, b, c, pereopods 3, 4, 5; d, telson; e, mandibular palp; /, right mandibular

molar; g, apex of outer lobe on lower lip (tilted on its side); h, inner plate of maxilla 1, shaded;

i, uropods 1-2;;', pleonal epimera 1-3.

360-O34 O—70 7
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 47.—Neomicrodeutopus (?) rnakena, new species, holotype, male, 4.7 mm, Makapuu

Point, Oahu, 1967: a, head; b, coxa 1; c, apex of pereopod 5; d, uropod 3; e, gnathopod 1; /,

pereopod 1; g, lateral view of telson (tilted on its side); h, gnathopod 2.

Neomicrodeutopus elongata (Chevreux), formerly

in the genus Grandidierella Coutiere but shifted to

Neomicrodeutopus by Schellenberg (1938), differs

from N. makena in the narrower ocular lobe, the very

weak, nearly naked mandibular palp, and elongate

ramus on uropod 3; again, there may be generic differences not now known. The condition of the mandibular palp in both of the Atlantic species is strong indication of that proposition.

If this is a genus distinct from Neomicrodeutopus, it

does appear simply to be a species of Microdeutopus

without inner ramus on uropod 3 rather than having

gone through an intermediate stage such as that represented by Acuminodeutopus J. L. Barnard that has

a shortened inner ramus on uropod 3. Neomicrodeutopus makena does not appear to have any direct relationship to Acuminodeutopus.

DISTRIBUTION.—Hawaiian Islands.
NUMBER 3 4

Family ATYLIDAE »

Atylus Leach

Katnehatylus, new subgenus

DIAGNOSIS.—Atylus with small atylid rostrum and

dorsal sculpture of A. levidensus J. L. Barnard (1956) ;

all segments of urosome coalesced; mandibular molar

triturative but palp absent; accessory flagellum vestigial; lower lip lacking inner lobes; inner plate of

maxilla 1 with 2 terminal setae and no apical cusp,

outer plate normally spinose, palp 2-articulate; maxilla

2 normal; maxilliped normal; gnathopods small but

subchelate and atylid; pereopods simple; anterior

coxae intersimilar in length and surface area, none

grossly enlarged; uropods normal, uropod 3 with

short peduncle, rami of medium elongation; telson of

medium length and deeply cleft.

TYPE-SPECIES.—Kamehatylus nani, new species.

RELATIONSHIP.—The breadth of the diagnosis reflects the problem that Kamehatylus is, by definition, a

member of the family Dexaminidae. The loss of the

mandibular palp is the major distinction between atylids and dexaminids. There seems to be little point in

maintaining a distinction between these families except

at the subfamilial level, for Kamehatylus is not the

only genus to bridge the familial concepts. Anatylus

Bulycheva (Anatylidae, later Anatylinae in the Dexaminidae) and Atylus japonicus Nagata (1961a) also

form intergrades, the former having a nontriturative

mandibular molar and the latter possibly having lost

the mandibular palps like Kamehatylus.

One might presume from the congruity of appearance that Kamehatylus nani is a descendent-of either,

or is a member of a species complex including the

Japanese Atylus japonicus and the Oregonian A. levidensus J. L. Barnard (1956). The latter has mandibular palps, whereas Nagata notes "broken" palps of

A. japonicus; they are a closely interrelated group

that may also include A. collingi Gurjanova (see 1951).

Kamehatylus nani appears to be more closely related to A. japonicus than to the other species in the

fact that fewer gross differences occur between those

two species than among the others. If A. japonicus has

indeed lost the mandibular palps then the sequence

seems eminently clear, for K. nani differs only in these

other characters: (1) the short peduncle of antenna 1;

" See footnote on page 26.

93

(2) the loss of one seta and the apical cusp on the inner

plate of maxilla 1 (A. japonicus has 3 setae and a

cusp); (3) the short telson; (4) the development of

2 lateral spines on the urosome (like many dexaminids) ; and (5) the complete coalescence of all urosomites (in all other atylids and dexaminids only urosomites 2-3 are coalesced). Kamehatylus nani also

resembles A. japonicus in the tapering coxa 1 but also

differs in a slightly smaller rostrum.

Kamehatylus nani differs from A. levidensus in

characters 2-5 but differs in addition by: (6) the

slightly more elongate article 3 of antenna 1; (7) the

slightly shorter article 4 of antenna 2; (8) the apical

tapering of coxa 1; (9) the slight elongation and narrowing of article 2 on pereopods 3-5; (10) the loss of a

small cusp on article 2 of pereopod 5; (11) the loss of

a conical protuberance on coxa 3. Thus, the connections between A. levidensus and the other two species

are more remote than between A. japonicus and K.

nani. A logical progression of events would be the occurrence of a common ancestor to A. japonicus and A.

levidensus with K. nani descending from A. japonicus.

Neither A. japonicus nor A. levidensus could be the

precursor to the other, but A. levidensus could be a remote ancestor of K. nani (but not vice versa.)

Atylus japonicus has one character of complete divergence, the elongate peduncle of antenna 1 that is

reflected weakly in the slightly elongate article 3 on K.

nani, but the latter otherwise has a short peduncle.

Gnathopods of A. japonicus also are more slender

than in the other two species, and this damages the

concept of a direct progression between A. japonicus

and K. nani.

Atylus (Kamehatylus) nani, new species

FIGURES 48, 49

DIAGNOSIS.—With the characters of the subgenus.

COLOR IN 2-DAY FORMALDEHYDE.—Body mottled

with orange, rust, and ochre on a buff-yellow background, eyes red.

HOLOTYPE.—Bishop Museum collections, catalog

number 7265, female, 2.8 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, algae, rocks, corals, 24

May 1967.

REMARKS.—This species has undergone changes

analogous to those seen in Micronesian Beaudettia palmeri J. L. Barnard (1965a) that also represents a case
94

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 48.—Atylus (Kamehatylus) nani, new subgenus, new species, holotype, female, 2.8 mm,

JLB Hawaii \1: a, body; b, c, mandibles, left normal view, right obverse; d, lower lip; e, maxilla

1; /, maxilliped; g, maxilla 2; h, i, articles 2—3 of pereopods 4,3.
NUMBER 3 4 95

FIGURE 49.—Atylus (Kamehatylus) nani, new subgenus, new species, holotype, female, 2.8 mm,

JLB Hawaii 17: a, medial gnathopod 2; b, medial gnathopod 1; c, medial pereopod 5; d, telson;

e, apex of lateral pereopod 1; /, uropod 3.

of radical change in characters of familial importance.

Those characters therefore lose such value by themselves in a typological framework, but the cases are of

considerable interest in viewing the incipience of

changes that in other phyletic lines have resulted in presumably old, diversified, and genetically remote complexes. Beaudettia palmeri is believed to be a relatively

direct descendent of a species of Elasmopus in which

the mandibular palp has been lost and various changes

in uropod 3 and the telson have occurred. The two

cases demonstrate a thesis that certain changes occurring in the evolution of Amphipoda do not necessarily

involve a slow transformation from one state to another; thus a mandibular palp may either be present

(and 3-articulate) or absent, and a telson either deeply

cleft or entire. In some families (Stenothoidae) the

mandibular palp may be lost in stages, one article at a

time, but Beaudettia and Kamehatylus seem to demonstrate that a radical loss also occurs. That such species are not simply blind alleys of evolution appears to

be supported by a diverse occurrence of dexaminids

that are simply atylids without mandibular palp, which
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

then have become partially adapted to an inquilinous

life. The bright colors of Kamehatylus, that blend together as a camouflage resembling the background

color of Sargassum, suggest that the Hawaiian subgenus, like its atylid precursors, is living freely rather

than in an inquilinous niche.

DISTRIBUTION.—Hawaiian Islands.

Family COLOMASTIGIDAE

Colomastix kapiolani, new species

FIGURE 50

DIAGNOSIS OF MALE.—Epistome bluntly conical,

head with large conical anterior keel between antennae

as seen from ventral view; eyes medium in size, red in

formaldehyde, and clear or slightly pink-rust in alcohol; lateral cephalic lobe rounded; antenna 1 weakly

spinose and cuspidate, flagellum with 4 articles, latter

2 articles vestigial, antenna 2 with medial spines on

articles 3-4 of peduncle, flagellum 4-articulate; mandibular spine row simple and normal, molar typical of

C. pusilla, maxilla 1 similar to that of C. subcastellata

Hurley, maxilla 2 strongly bilobate; inner plates of

maxilliped split deeply, remainder of maxilliped like

that of C. lunalilo, new species; coxae simple, ovatorectangular, lacking points; gnathopod 1 normal, with

article 5 slightly longer than 6, but article 2 strongly

swollen anterodistally; article 2 of gnathopod 2 with

small anteroventral hump, article 5 with narrow, setose

posterior lobe, setae simple, article 6 very elongate,

palm occupying entire posterior margin of article 6,

dactyl overlapping palm and reaching back to proximal margin of lobe on article 5, palm with hump near

dactylar hinge, deep excavation and large conical cusp,

then long flat margin defined proximally by small cusp,

dactyl simple; pereopods 1-5 normal, each bearing

one locking spine on article 6; pleonal epimera 1-3

broadly rounded ventrally and posteroventrally;

uropods 1-3 extending subequally, outer ramus of

uropod 1 slightly shorter than inner ramus, outer ramus

of uropod 2 about two-thirds as long as inner ramus,

and outer ramus of uropod 3 about three-fourths as

long as inner ramus; telson unequally trilobate apically,

of medium length.

JUVENILE MALE.—Article 6 of gnathopod 2 with

short dactyl fitting palm and weak defining cusp, palmar process near dactylar hinge quadrate, undivided;

some setae of lobe on article 5 suddenly narrowed in

middle like setae shown by Hurley (1954b). Female

unknown.

HOLOTYPE.—Bishop Museum collections, catalog

number 7267, male, 1.3 mm.

TYPE-LOCALITY.—JLB Hawaii 5, off Ewa Beach,

Oahu, 30 m, Pocillopora, 29 January 1967.

MATERIAL.—JLB Hawaii 5 (1), 6 (1), 10 (2).

RELATIONSHIP.—According to Hurley's (1954b)

key to the species of this genus, C. kapiolani has its

closest affinities with C. fissilingua Schellenberg (1926)

(K. H. Barnard, 1932), but the latter differs from C.

kapiolani in the equal trilobation of the telson, the

anteriorly pointed coxae 1, 3, 4, and 5, the posteriorly

pointed coxae 6-7, the equal rami of uropod 1, and

the extremely thin article 2 of pereopods 3-5.

Colomastix subcastellata Hurley (1954b), from

New Zealand, also closely resembles C. kapiolani,

but has lateral scallops on the telson, a smaller hump

on article 2 of gnathopod 1, and equal rami of uropod

1. These differences may be of subspecific value only.

Colomastix castellata K. H. Barnard (1932) has a

multicastellate telson and equal rami on uropods 1

and 2.

DISTRIBUTION.—Hawaiian Islands.

Colomastix lunalilo, new species

FIGURES 51, 52

DIAGNOSIS OF PRESUMED MALE.—Epistome conical,

head with large conical anterior keel between antennae, eyes large, brown in formaldehyde and alcohol,

lateral cephalic lobe rounded; antenna 1 with lateral

and medial, ventral edges spinose on articles 1, 2, and

3, distoventral apex on lateral side pointed on articles

1, 2, and 3, flagellar article 1 cuspidate distodorsally,

article 2 of flagellum vestigial; antenna 2 also with

spines and cusps on articles 3-5 (see figures), flagellum 4-5 articulate and attached asymmetrically to

article 5 of peduncle; mandibular spine row simple

and normal, molar typical of C. pusilla, maxillae 1-2

relatively typical of C. subcastellata Hurley, inner

plates of maxilliped coalesced fully, outer plates each

with apical spine, article bearing outer plate not

broadly alate laterally, basal article shell-like, forming

immense posteriorly directed envelope, remainder of

maxilliped attached to basal article in geniculate

fashion; coxae 1-5 with nipple-like anterior points;

gnathopod 1 normal, with elongate article 3, article

6 subequal in length to article 6 and apically setose
NUMBER 3 4 97

FIGURE 50.—Colomastix kapiolani, new species, holotype, male, 1.3 mm, JLB Hawaii 5: a, body,

b, uropod 3; c, telson; d, inner plate of maxilliped; e, head. Male 1.0 mm: /, maxilla 2; g, lateral

gnathopod 2; h, maxilla 2. Specimen from JLB Hawaii 10: i, lateral view of maxilliped; j , basal

shell of maxilliped.
98

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

9

FIGURE 51.—Colomastix lunalilo, new species, holotype, male, 1.9 mm, JLB Hawaii 13: a, body;

b, apex of gnathopod 1; c, gnathopod 2; d, gnathopod 1; e, lateral view of maxilliped; /, mandible;

g, head.
NUMBER 3 4 99

FIGURE 52.—Colomastix lunalilo, new species, holotype, male, 1.9 mm, JLB Hawaii 13: a,

outer plate and palp of maxilla 1; b, edge of inner ramus of uropod 3; c, mandible with molar

turned to left; d, telson and uropod 3, dorsal; e, f, medial antennae 1 and 2; g, apex of telson; h,

maxillipedal palp articles 3 and 4; i, uropod 3, offset to left are high power and oil immersion

views of outer ramus edge sculpture; ;, maxilla 2; k, mandibular molar; I, ventral view of head.

Male, 1.8 mm: m, maxilla 1; n, maxilliped; o, articles 6-7 of gnathopod 2 from right side.
100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(dactyl not distinguished from setae) ; article 5 of

gnathopod 2 with broad setose posterior lobe, setae

simple, article 6 elongate, dactyl shorter than palm,

palm shorter than posterior margin of article 6 and

defined by large cusp, palm with low truncate process,

dactyl with obtuse inner cusp; pereopods 1-2 unornamented; pleonal epimera 1-3 broadly rounded ventrally and posteroventrally; uropods 1-2 extending

equally, rami all equal to each other in length, uropod

3 extending slightly beyond uropods 1-2, outer ramus

about one-third as long as inner (variable, from

slightly less than in juvenile to slightly more than

one-third in adult) ; telson triangular, of medium

length, exceeding in length peduncle of uropod 3.

JUVENILE MALE.—Gnathopod 2 of normal undifferentiated morphology (see figure).

DESCRIPTION.—Palp of maxilla 1 uniarticulate,

with deep midterminal notch defined by sharp distolateral extension, medial margin finely but deeply

serrate, outer plate with 1 apical and 2 proximal simple

spines, between those, 2 ridges or spines with flabellate,

serrate and beaded margins, inner plate subrectangular; maxilla 2 shriveled, but lobes weakly separated.

HOLOTYPE.—Bishop Museum collections, catalog

number 7266, ?male, 1.9 mm.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

Oahu, 3-4 m, corals, algae, corallines, 8 April 1967.

MATERIAL.—Three specimens from the type-locality.

RELATIONSHIP.—Colomastix brazier i Haswell

(1880) from east Australia and C. magnirama Hurley (1954b) are the only other members of the genus

with a strongly reduced outer ramus on uropod 3.

Pereopods 1-2 of C. magnirama have a large cusp on

article 5, complex mandibular spines, a much shorter

outer ramus of uropod 3 than seen on C. lunalilo, a.

broader telson, unequal rami of uropod 1 in both

sexes, a shorter article 5 and narrower lobe on gnathopod 2, and no points on the coxae.

Colomastix brazieri has article 5 of gnathopod 1

much longer than article 6, an outer ramus of uropod

3 half as long as the inner ramus and a shorter article

5 and narrower lobe of gnathopod 2 than in C. lunalilo. The latter appears to be closer to C. brazieri than

to any other species, but C. brazieri needs further

elucidation.

Colomaxtix pusilla Grube, a cosmopolitan sympatriot of C. lunalilo in Hawaii, differs from the latter in

its red eyes changing to colorless in alcohol, in the

absence of sharp and narrow nipple-like points on the

coxae, the shorter telson, the blunter epistome, the

shorter gnathopod 1, and in males a distinctly different gnathopod 1 than occurs in female C. pusilla and

males of C. lunalilo. The rami of uropod 3 in C. pusilla

are equal to each other in length.

DISTRIBUTION.—Hawaiian Islands.

Colomastix species

MATERIAL.—JLB Hawaii 6(1).

REMARKS.—This undissected individual, with

gnathopod 2 in juvenile condition, is distinguished

from other Hawaiian species in (1) the presence of

only 1 apically hooked setal spine on the apex of

gnathopod 1; (2) the absence of a hump on article 2

of gnathopod 1; (3) uropod 1 is shortened, the inner

ramus reaching only to the middle of the rami of

uropod 2 and the outer ramus being about two thirds

as long as the inner ramus; (4) the rami of uropod 2

extend equally; (5) the outer ramus of uropod 3 is

about three fourths to four fifths as long as the inner

ramus; (6) the telson appears simple (not adequately

seen) ; (7) pereopods 1-5 lack a locking spine; (8) the

eye has 9 ommatidia but 3 of the middle ones are extremely tiny.

The individual fits C. pusilla more closely than it

does the other Hawaiian species but differs in the unequal rami of uropods 1 and 3. The unusual condition

of gnathopod 1 may represent a stage of transformation from the normal setal brush to the advanced male

stage like that of C. pusilla in which a distinct, stubby

dactyl occurs on a shortened gnathopod 1.

Family COROPHIIDAE

Corophium ?baconi Shoemaker

FIGURE 53

?Corophium baconi Shoemaker, 1934, pp. 356-359, fig. 1;

1949, p. 82, fig. 5g, h.

MATERIAL.—JLB Hawaii 11 (4).

REMARKS.—No adult male is available but female

characters match those of C. baconi, the only Pacific

species of the genus in which the segments of the

urosome are coalesced and formed into a shallow cuplike plate with upturned rims bearing a slit to mark

the junction between urosomites 1 and 2, with uropods

1-2 attached ventrally rather than in lateral notches.

Dactyls of gnathopods 1-2 have only 1 and 2 inner

teeth.
NUMBER 3 4 101

9

FIGURE 53.—Corophium baconi Shoemaker, female, 2.3 mm, JLB Hawaii 11: a, medial antenna

2; b, medial antenna 1; c, gnathopod 2. Female, 1.5 mm: d, dorsal urosome; e, dorsal head; /,

lateral head; g, pereopod 1.

Young females and the ovigerous female have a

spine-seta pattern on the urosome similar to that shown

by Shoemaker (1934), but the spines are very stout in

Hawaiian specimens. Antenna 2 of the large female

has an extra proximal ventral spine on article 4 adjacent to the normal spine, for a total of 4 spines

instead of 3 in normal individuals. The short article 5

of pereopods 1-2 and the elongate posteriorly straight

and poorly armed article 6 of gnathopod 2 are

illustrated.

DISTRIBUTION.—Bering Sea to Peru.

Corophium insidiosum Crawford

FIGURE 54

Corophium insidiosum Crawford, 1937, p. 615, fig. 2a-g.—

Shoemaker, 1947, p. 53, figs. 6, 7.

MATERIAL.—Hilo, Hawaii, 20 June 1959, ClunioPolster, collector H. Caspers (70+).

Besides the normal male and female phenotypes at

least a dozen other phenotypes are present in a single

collection of this species from Hilo. The normal male

antenna 2 (Figure 54c) lacks large ventral spines on

articles 4 and 5, but males occur with a single proximoventral spine on article 4 and none on article 5 or with a

pair of midventral spines on article 4 and one ventral

spine on article 5. The medioproximal margin of article

1 of antenna 1 (from dorsal view) typically would have

2-3 spines, but males in the Hilo collection have either

2, 1, or no spines, occasionally one of the spines being

extremely and abnormally proximal. Females also have

these kinds of antenna 1 except that they occasionally

have 3 of those spines, one of which is extremely proximal. The ventral margin of article 1 of antenna 1 typically would have 2 ventral and 1 ventrodistal spines

and this is typical for Hilo females, but males often

lack the 2 ventral spines. Individuals often have one

formula on the left antennae and another formula on

the right antennae.

Female antenna 2 is often typical (Figure 54e),

but frequently either right or left antennae 2 of adult

ovigerous females look like those of males with the
102

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 54.—Corophium insidiosum Crawford, Hilo. Antenna 2: a, abnormal female; b,

abnormal female; c, normal male; d, abnormal male; e, normal female; /, abnormal female.

Antenna 1: g, male and female lateral; h, male dorsal; i, male lateral; ;, female, dotted spine

infrequent, male occasionally with only 1 spine.

large and small distal cusps in place of a single spine.

The ventral surface of article 4 may carry 3 single

spines, 2 single spines, or 2 pairs of spines. Female antennae 2 without the male configuration may also vary,

sometimes having a single ventral spine, sometimes 2,

occasionally 3, but usually having the normal 2 pairs

and a single.

Some of the female aberrancies equivocate the conditions typical of Corophium uenoi, from Japan and

California, in which female antenna 2 has spines occurring only singly instead of in pairs. Nevertheless,

I identify these specimens with C. insidiosum. Apparently, C. insidiosum, a widely distributed and transported species, has become highly variable in the

Hawaiian Islands, typical of other species having

reached island niches or special environments. Another example is the masculine females of Gammaropsis in Micronesia (J. L. Barnard, 1965a).

DISTRIBUTION.—Cosmopolitan, especially in harbors.

Family DEXAMINIDAE

Paradexamine Stebbing

Wailele, new subgenus

DIAGNOSIS.—Antenna 1 much longer than antenna

2; mandibles with spine row and lacinia mobilis on both

sides, molar triturative; lower lip with well-developed

separate inner lobes and strong mandibular processes

on outer lobes; inner plate of maxilla 1 naked, short,

palp 1-articulate and short, only reaching halfway

along outer plate, bearing 1 long apical seta; outer

plate of maxilla 2 much longer than inner plate, latter

with 2 apical setae; maxillipedal palp 4-articulate, not

exceeding outer plate, article 4 not unguiform, instead

short, stout, and bearing 2 apical setae, inner plates

obsolescent, scarcely evident, outer plates very large

and spinose; gnathopods with article 5 shorter than

article 6 and cupshaped, palms transverse; pereopods

with long dactyls but not subchelate; coxa 5 with surface area greater but plate not longer than anterior

coxae; telson cleft to base.

TYPE-SPECIES.—Paradexamine (Wailele) maunaloa.

RELATIONSHIP.—Paradexamine (Wailele) maunaloa has been compared with 7 species of Paradexamine

studied in Australian and New Zealandian collections

(paper in preparation). The variability in mouthparts

is higher than previously elucidated in this genus and

suggests that Dexaminoides Spandl is a synonym of

Paradexamine. The Micronesian identification of

Dexaminoides orientalis Spandl by J. L. Barnard

(1965a) apparently represents a distinct, new species of

Paradexamine and to some extent connects the Hawai-
NUMBER 34 103

ian Par ad examine to the mainstream of the genus in

Australia. Wailele is thus described as a subgenus of

Paradexamine rather than being accorded full generic

rank. Wailele differs from the type-species of Paradexamine and from most of the known species of that genus

in the strong reduction of the palp on maxilla 1 and in

the complete obsolescence of the inner plates of the

maxilliped. The Micronesian Paradexamine (=Dexaminoides) lessens the categorical importance of the

differences in Wailele because of the partial reduction

of the inner plate of the maxilliped. A new species of

Paradexamine from New Zealand (in press) also has a

reduced inner plate of the maxilliped.

Wailele maunaloa, new species

FIOURES 55, 56

DIAGNOSIS.—With the characters of the genus.

DESCRIPTION.—Lateral cephalic lobe rounded, eye

black-brown in formaldehyde and alcohol; article 1 of

antenna 1 with anteroventral setose hump; distal spines

on pereopods 1-5 blunt; article 2 of pereopod 3 with

posteroventral lobe, absent on pereopods 4-5; article

2 of pereopod 5 broader than on pereopods 3-4, but

pereopod 5 otherwise shorter than 4; pleonal epimera

1-2 with slight lateral ridges, posteroventral corners of

all epimera sharply produced, least on epimeron 1,

posterior edges raggedly and minutely serrate, most

evenly on the epimeron 3; pleonites 1-4 with small

sharp dorsal tooth, pleonites 3-4 with dorsolateral

tooth also; rami of uropod 1 equal in length to each

other, outer ramus of uropod 2 shorter than inner.

HOLOTYPE.—Bishop Museum collections, catalog

number 7268, female, 1.7 mm.

TYPE-LOCALITY.—JLB Hawaii 15, 1 mi N of Kualoa

Point, Oahu, intertidal, block of ascidians, 23 May

1967.

MATERIAL.— Five specimens from the type-locality.

DISTRIBUTION.—Hawaiian Islands.

Family EOPHLIANTIDAE

Biancolina mauihina, new species

FIGURES 57, 58

DIAGNOSIS.—Eyes composed of about 14-18 ommatidia, red in formaldehyde, yellow or clear in alcohol; antenna 1 about as long as head and pereon

together, flagellum with about 10 articles, antenna 2

short, flagellum and peduncle not readily distinct from

one another, flagellum with about 3 articles; maxillipedal palp article 4 very short, hemispherical, bearing

a large terminal spine, mediodistal end of article 2

obtusely conical, outer plate with 3 very stout apical

spines, inner plate with 2 elongate spine-setae; pereopods 3-5 prehensile, article 6 strongly widened, bearing

sinuous palm armed and lined with 1 curved striate

spine, palm defined by minute spine and setule; outer

rami of uropods 1-2 about 60 percent as long as inner

rami, peduncles strongly setose; rami of uropod 3 relatively stout; telson short and apically notched.

HOLOTYPE.—Bishop Museum collections, catalog

number 7269, ?sex, 1.8 mm.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, other algae, rocks, corals,

24 May 1967.

SPECIMEN NO. 2.—In general agreement with the

holotype but uropods 1-2 very distinct: uropod 1 with

outer ramus grotesquely gnarled (perhaps from preservative) ventrally bearing a flat hyaline lobe adjacent

to a long shovel-spine and apparently its distal half

formed of a flat but sharp spine, thus outer ramus exceeding inner ramus; outer ramus of uropod 2 with

long, hooked distal spine reaching almost to apex of

inner ramus, latter with medium-sized spine and small

spine. Fee 1 (1 individual, 1.6 mm long).

REMARKS.—The two specimens may belong to different species or may be the opposite sexes of a single

species; I have not been able to determine sex in either

specimen; neither specimen has brood lamellae; penial

processes are not evident.

The holotype differs from B. australis Nicholls

(1939) in the short outer rami of uropods 1-2, the

stouter and shorter article 4 of the maxillipedal palp,

the relatively longer antenna 1 and relatively thinner

antenna 2, stouter rami of uropod 3, and the prehensile

pereopods 3-5.

In the short outer rami of uropods 1-2 the holotype

resembles B. algicola Delia Valle (1893, from the

Mediterranean Sea) but differs in the prehensile pereopods 3-5 and in the distal notch on the telson.

Uropods 1-2 of specimen no. 2 are uniquely distinct.

1 have searched in vain for more individuals of this

genus through thousands of intertidal amphipods.

Gnathopod 1 of B. mauihina has one giant midposterior seta and one medial seta on article 2. Gnathopod

2 lacks either seta.

DISTRIBUTION.—Hawaiian Islands.
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 55.—Paradexamine (Wailele) maunaloa, new subgenus, new species, holotype, female, 1.7

mm, JLB Hawaii 15: a, body; b, telson; c, inner and outer plates of maxilliped, side opposite to

d; d, maxilliped; e, f, g, pereopods 3,4, 5.
NUMBER 34 105

FIGURE 56.—Paradexamine (Wailele) maunaloa, new subgenus, new species, holotype, female, 1.7

mm, JLB Hawaii 15: a, b, distal ends of gnathopods 1, 2; c, pereopod 2; d, lower lip less right

outer lobe; e, left mandible from obverse side; /, right mandible; g, upper lip; h, pleonal epimeron

3, posterior margin: i, maxilla 2. Juvenile, 1.0 mm: j , uropod 3. Juvenile, 1.3 mm: k, maxilla 1;

I, maxillipedal palp articles 2, 3, 4.
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 57.—Biancolina mauhina, new species, holotype, ?sex, 1.8 mm, JLB Hawaii 17: a, body;

b, apex of gnathopod 1, c, maxilla 2; d, uropod 3; e, maxilliped; f, g, right and left mandibles,

obverse sides; h, maxilla 1; i, apex of gnathopod 2. ?Sex, 1.6 mm, Fee 1: ;", lower lip; k, apex of

pereopod 5.
NUMBER 3 4 107

9

FIGURE 58.—Biancolina mauhina, new species, ?sex, 1.6 mm, Fee 1: a, head, anterodorsal view;

b, anteroventral view of upper lip in middle embraced by mandibles; c, uropod 2; d, uropod 1,

dorsal; e, uropod 1, ventral. Holotype, ?sex, 1.8 mm, JLB Hawaii 17: /, urosome, dorsal; g,

apex of pereopod 5; h, maxilliped; i, upper lip (on side); j , coxa 6; k, coxa 1.

360-034 0—70 8
108

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

d

FIGURE 59.—Eusiroides diplonyx Walker, female, 6.2 mm, JLB Hawaii 14: a, body; b, lateral

gnathopod 2; c, lateral gnathopod 1; d, palm of medial gnathopod 2; e, sculpture on articles 4—6

of pereopods 3-5.
NUMBER 34

Family EUSIRIDAE

Eusiroides diplonyx Walker

FIGURES 59, 60, 61

?Eusiroides Caesaris Steb., var. Walker, 1904, p. 264, pi. 4,

fig. 22.

Eusiroides diplonyx Walker, 1909, pp. 333-334, pi. 43, fig.

4.—Pirlot, 1936, pp. 302-303, figs. 126-128.—Schellenberg, 1938, p. 35.

NOTES ON HAWAIIAN MATERIAL.—Only Pirlot has

adequately described his Indonesian material. Walker's

109

(1904) Ceylonese material and Seychellesian (1909)

type were accompanied with a few notes and figures

sufficient only to recognize this species grossly by the

low number of serrations on epimeron 3 and the sabreshaped, striate locking spine of pereopods 1-2.

The Hawaiian material differs from that of Pirlot in

the stronger setation on mandibular palp article 3,

the inner plate of maxilla 2, and the apex of the outer

plate on the maxilliped. Palp article 2 on the maxilliped has a ventrally pointing linear crest produced

apically as a subfalcate lobe not definitely shown on

Pirlot's drawing. Article 2 of pereopods 3-5 is slightly

FIGURE 60.—Eusiroides diplonyx Walker, female, 6.1 mm, JLB Hawaii 14: a, head; b, maxilla 1.

Female, 6.2 mm: c, lower lip; d, right mandibular incisor; e, article 2 of mandibular palp,

flattened; /, mandible; g, maxilla 2; h, accessory flagellum and calceoli; t, upper lip; ;, maxilliped.
110

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 61.—Eusiroides diplonyx Walker, female, 6.2 mm, JLB Hawaii 14: a, b, c, uropods 1, 2,

3; d, telson; e, lateral pereopod 5; /, sculpture on medial side of article 4 on gnathopod 1; g,

pleonal epimera 1-3; h, apex of pereopod 1.

more slender but slightly more lobate posteroventrally

than in Pirlot's material.

MATERIAL.—JLB Hawaii 5 (2), 6 (2), 13 (2), 14

(6).

Notes on Ronco J. L. Barnard (1965a) : This genus

was erected for a Micronesian species resembling Eusiroides but having cleft upper lip and a few thin palmar

spines on the gnathopods. Ronco sosa appears to be a

derivative of Eusiroides diplonyx with poorly developed gnathopods, incised upper lip, and smooth epimeron 3. The stout locking spine of pereopods 1-2 is

a strong clue to this relationship, plus the condition of

the telson. The inner plate of maxilla 2 on Ronco

has lost all but a few of the numerous setae occurring

on E. diplonyx. I consider the changes that have occurred in Ronco sosa to be of the magnitude as those

found in Atylus (Kamehatylus) nani and suggest the

reduction of Ronco to subgeneric status under

Eusiroides.

DISTRIBUTION .—Indo-Pacific.

Pontogeneia pacifica Schellenberg

FIGURES 62, 63, 64

Pontogeneia pacifica Schellenberg, 1938, pp. 35-37, fig. 17.—

J. L. Barnard, 1955a, p. 5.

This species seems to form part of a triad with the

Caribbean P. longleyi Shoemaker (1933a) and the

Califomian P. "minuta" of J. L. Barnard (1959), with

characters of each of those species appearing in P.

pacifica. The rostrum and article 2 of pereopod 5 of

P. longleyi and the long coxae of P. "minuta" are combined in P. pacifica. The posterior lobe on article 5 of

gnathopod 2 in P. pacifica is intermediate in size between the weak lobe of P. longleyi and the strong lobe

of P. "minuta." Mandibular palp article 3 of P. pacifica is much better developed than in the other species.

Uropod 3 is similar in both sexes but the antennae

differ considerably. Male antenna 1 is as long as the

head and first 7.5 body segments, that of the female
NUMBER 3 4 111

only as long as the head and first 6 body segments.

Male antenna 2 is as long as the head and first 9 body

segments, that of the female only as long as the head

and first 8 segments. Male antennal peduncles have

stairstep notches and both antennae bear calceoli not

found in females.

Eyes are dark purple in formaldehyde but become

pale purple broad cores surrounded by a layer or two

of clear ommatidia in alcohol.

MATERIAL.—JLB Hawaii 14 (2), 15 (17), 17 (2).

Fee 1 (32).

DISTRIBUTION.—Hawaiian Islands.

FIOURE 62.—Pontogeneia pacifica Schellenberg, female, 5.0 mm, Fee 1: a, proportions of uropod

2; b, head; c, d, apices of lateral gnathopods 1, 2; e, urosome; /, uropod 1; g, microscopic skin

texture; h, dactyl of pereopod 5; i, pleonal epimera 1-3. Male, 4.7 mm: ;, apex of gnathopod

1, setae removed; k, pleonal epimera 1-3; /, outline of head.
112

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 63.—Pontogeneia pacifica Schellenberg, male, 4.7 mm, Fee 1: a, base of antenna 2,

lateral; b, base of antenna 1, lateral. Female, 5.0 mm: c, base of antenna 2, medial; d, e, coxa

and article 2 of gnathopods 1, 2; /, g, h, i, pereopods 1, 3, 4, 5; ;, coxa 4. Male, 4.1 mm: k,

base of antenna 1, medial.
NUMBER 3 4 113

FIGURE 64.—Pontogeneia pacifica Schellenberg, male, 4.7 mm, Fee 1: a, mandibular palp; b, c,

maxillae 1, 2; d,e, mandibles; /, lower lip; g, upper lip; h, maxilliped. Female, 5.0 mm: i, article

2 of pereopod 5, shaded; j , telson; k, uropod 3.
114

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

n

FIGURE 65.—Ceradocus hawaiensis J. L. Barnard, male, 5.0 mm, JLB Hawaii 6: a, small kind of

gnathopod 2; b, pleon; c, dactyl of pereopod 4; d, posteroventral corner on article 2 of pereopod

5; e, maxilla 1; /, article 4 of gnathopod 2; g, telson; h, posteroventral corner on article 2 of

pereopod 3; i, maxilla 2; ;, uropod 3; k, body; I, anteroventral corner of head; m, articles 2, 3, 4,

5, of medial gnathopod 2; n, article 3 of antenna 2 peduncle. Juvenile 3.3 mm: o, lateral

gnathopod 2.
NUMBER 3 4 115

Family GAMMARIDAE

Ceradocus hawaiensis J. L. Barnard

FIGURE 65

Ceradocus hawaiensis J. L. Barnard, 1955a, pp. 5-8, figs. 2, 3.

MATERIAL.—JLB Hawaii 6 (9).

REMARKS.—Excellent material of this species permits figuring a whole specimen.

DISTRIBUTION.—Hawaiian Islands.

Elasmopus calliactis Edmondson

FIOURE 66

Elasmopus calliactis Edmondson, 1951, pp. 189-191, fig. 3.—

J. L. Barnard, 1955a, p. 8.

Elasmopus rapax.—J. L. Barnard, 1955a, p. 11, specimens

from Honolulu Aquarium only, not Costa.

DIAGNOSIS OF TERMINAL MALE.—Accessory flagellum 3-4 articulate; eyes large and clear in alcohol,

ommatidia all distinct, eyes red in life, lacking black

pigment; mandibular palp article 3 long and weakly

falcate; articles 5-6 of gnathopod 1 equal to each other

in length; article 2 of gnathopod 2 not distally lobate,

anterodistal corner setose and slightly protuberant,

article 3 with normal lobe, article 6 elongate, palm

oblique and equal to posterior margin of article 6, palm

defined by large tooth and bearing middle tooth and

spinose adze-shaped process near dactylar hinge, margin between defining and middle process flat, rounded

between middle and distal process, posterodistal end

of article 6 on pereopods 1-2 with 1 irregularly shaped

chisel spine, slightly pointed or bifid apically, other

locking spine slightly curved, dactyl with subapical

constriction bearing 1 seta and 2 setules; locking spines

of pereopods 3-5 normal; pereopods 3-5 bearing long

setae on articles 2-6, posterior margin of article 2 of

regular dimensions and weakly serrate; pleonal epimera 1-2 with lateral ridge, posteroventral corners

sharply produced, posterior margins sparsely serrate,

epimeron 3 quadrate, slightly serrate posteriorly, ventral margins of epimera with pairs of spines; urosomal

segments simple; rami of uropods 1-2 with terminal

spines of medium length; inner ramus of uropod 3 extending nearly as far as outer ramus, broadly truncate

and spinose, outer ramus also broadly truncate, spinose

and bearing small article 2 concealed from dorsal

view; telson long, each apex broadly and deeply excavate, but excavation hidden by subterminal row of

5-6 spines.

FEMALE.—Gnathopod 1 slightly stouter than in male,

gnathopod 2 a miniature edition of male gnathopod 2.

JUVENILE MALE AND JUVENILE.—Palmar humps on

gnathopod 2 very weak, telson similar to adult but less

spinose (4 spines per lobe), inner ramus of uropod 3

slightly shortened; epimera similar to adult.

MATERIAL.—The type material: on Calliactis, 20

April 1949. Honolulu Harbor, with Calliactis on

Dolium, 4 October 1948 (3). Honolulu Aquarium, on

hermit crab, 26 October 1949 (25). Ewa, off Oahu,

9 August 1950, Dolium-Calliactis with Dardanus asper

association, 75-100 feet, collection, Tinker (10+).

Honolulu Aquarium, from cement overflow gutter exit

of fish tanks, 28 February 1950 (16, this sample misidentified as E. rapax by J. L. Barnard, 1955a).

REMARKS.—My comments (1955a) on the alternate

variety of E. rapax are erroneous; that material belongs

with E. calliactis and is readily identified by the clear

eyes, absence of the fourth tooth on the palm of gnathopod 2, and the shape of the telson. The black eyes of

E. rapax do not bleach clear in alcohol. The distal process on the palm of gnathopod 2 in E. calliactis is

strongly spinose in terminal adults and thus my (1965a)

key is in error. Besides relationships with E. holgurus

J. L. Barnard (1962b), E. rapax and E. piikoi, Elasmopus calliactis has a resemblance to E. pseudaffinis

Schellenberg (1938) (J. L. Barnard, 1965a) from

Micronesia. That species has black eyes, very weak

armament on the palm of gnathopod 2, a female with

normal gnathopod 2, and a telson like adult E. rapax,

with apically extended medial lobes bearing small

terminolateral spines.

DISTRIBUTION.—Oahu, apparently found exclusively

with a hermit-crab, sea-anemone association.

Elasmopus ?diplonyx Schellenberg

FIGURES 67, 68

? Elasmopus diplonyx Schellenberg, 1938, p. 54, fig. 26.

DIAGNOSIS OF MALE.—Accessory flagellum 3-articulate; eyes of medium size, brown in formaldehyde,

turning purplish-brown in spirit, ommatidia distinct

diough middle ones partially covered by pigment or

darkened; mandibular palp article 3 short and deeply

falcate; articles 5 and 6 of gnathopod 1 subequal in

length; article 2 of gnathopod 2 not lobate, article 3

weakly lobate, article 6 elongate, palm oblique, strongly

setose, bearing small, spinose distal protuberance, palm
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 66.—Elasmopus calliactis Edmondson, female, 7.4 mm, Oahu, 20 April 1949: a, head; b,

telson; c, apex of medial gnathopod 2; d, gnathopod 1, minus setae; e, pereopod 4; /, mandibular

palp; g, uropod 3; h, apex of pereopod 1. Male, 8.8 mm: i, medial gnathopod 2; ;, gnathopod 1.

Juvenile, 5.2 mm: k, palm of medial gnathopod 2.
NUMBER 3 4 117

9

FIGURE 67.—Elasmopus diplonyx Schellenberg, female, 4.6 mm, Devaney 1: a, head; b, medial

gnathopod 2; c, uropod 3; d, apex of pereopod 2; e, telson; /, mandibular palp; g, gnathopod 1.

Male, 3.3 mm: h, medial gnathopod 2; i, lateral gnathopod 2.
118

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 68.—Elasmopus diplonyx Schellenberg, female, 4.6 mm, Devaney 1: a, b, c, pereopods

3, 4, 5; d, apex of pereopod 3; e, pleonal epimeron 3; /, lower lip. Juvenile male: g, h, apices of

pereopods 2, 4.
NUMBER 3 4 119

scarcely defined by acclivity and spines, inner face of

article 6 with locking ridge for end of dactyl, ridge

with 2 spines and 2 setae, dactyl slender and slightly

overriding palmar face; posterodistal end of article 6

on pereopods 1-2 with one striate, slightly sabre-shaped

spine and 1 small, normal spine, dactyl constricted and

bearing a very stout blunt spine and 2 setules; locking

spines of pereopods 3-5 straight but striate, main dactylar spines inserted into dactyl but also joined basally

to dactyl margin by chitinous membrane; pereopods

3-5 lacking long setae, article 2 of pereopods 3-5 ovate,

but posteroventral margin strongly sloping on pereopod

5, pereopod 3 with normal serrations, but pereopods

4-5 deeply castelloserrate almost as strongly as on

pereopod 4 of E. pectenicrus; pleonal epimera 1-2

with lateral ridge and sharply produced posteroventral

corners, posterior margins sparsely serrate, epimeron 3

quadrate but with slight, sharp protrusion posteroventrally and weak posterior serrations, epimera lacking

setae, ventral spines occurring singly; rami of uropods

1-2 with long terminal spines; inner ramus of uropod

3 about 80 percent as long as outer ramus, apex truncate and spinose, outer ramus beveled and truncate,

spinose and bearing small hidden article 2; telson of

medium length, apices excavate and sinuous or raggedly serrate, each subapex with 2 spines.

FEMALE.—Gnathopod 2 like male but palm lacking

distal spinose protuberance and not as strongly setose.

JUVENILES.—Posteroventral tooth of pleonal epimera

1-3 slightly more protuberant than in adult, lacking

spines; eye small; major spine of dactyl on pereopods

1-2 sharp but stout, major locking spine of article 6

more strongly sabre-shaped than in adult; accessory

flagellum 2-articulate, membrane of spine on dactyl

of pereopods 3-5 scarcely evident; palm of juvenile

male with fewer setae than adult and armed with short

stout spine-setae; each lobe of telson with only 1 spine;

inner ramus of uropod 3 tapering, with one long apical

seta, article 2 of outer ramus relatively large; serrations

of pereopods 4-5 fewer and smaller than in adult but

relatively deep even on youngest individuals having

only 2-3 notches per article 2.

MATERIAL.—Devaney 1 (8).

REMARKS.—Schellenberg did not mention castellations on article 2 of pereopods 4-5; if they do not occur

in the Micronesian material than the Hawaiian populations should be described as a new species.

DISTRIBUTION.—Marshall, Gilbert, and Hawaiian

Islands.

Elasmopus ecuadorensis hawaiensis Schellenberg

FIGURE 69

Elasmopus ecuadorensis hawaiensis Schellenberg, 1938, pp.

54-55, fig. 27.

DIAGNOSIS OF MALE.—Accessory flagellum 2-articulate ; eyes medium in size, as dark as those of E. pocillimanus; mandibular palp normally falcate (as figured

for E. piikoi) ; article 5 of gnathopod 1 subequal to

article 6 in length; articles 2 and 3 of gnathopod 2 lacking lobes, article 5 short, bearing narrow but relatively

short, setose posterior lobe, article 6 elongate and

strongly tapering, palm and posterior margin of hand

confluent, densely setose, distal end of palm bearing

strong, obtuse protrusion, medial face of hand with

ridge defined proximally by spine near end of dactyl,

latter reaching less than halfway along palm; posterodistal end of article 6 on pereopods 1-2 with 1 striate

sabre-spine and one unstriate nearly straight smaller

spine, dactyl subapically constricted and bearing 1 stout

free spine and 2 weak setules, locking spines of pereopods 3-5 normal; pereopods 3-5 lacking long setae,

their lengths in same proportion to figures for E. diplonyx, article 2 of pereopod 3 similar to that species

but article 2 of pereopod 4 with oblique and straight

posterior edge and subquadrate posteroventral corner,

article 2 of pereopod 5 narrowly lobate posteroventrally, posterior margins weakly serrate; coxa 6 with unusually broad and slightly truncate anterior lobe;

pleonal epimera 1-2 with lateral ridge, posteroventral

corners sharply but weakly produced, posterior margins

sparsely serrate (notched), epimeron 3 with weakly

quadrate posteroventral corner bearing small notch and

several posterior notches, epimera with single spines

and no setae; urosomal segments simple; rami of uropods 1-2 with elongate spines but not unusually so;

inner ramus of uropod 3 nearly three-fourths as long

as outer, apices of both rami truncate and spinose, outer

with very small article 2 hidden among spines; telson

very short, as broad as long, apices slightly concave,

with medial portions extended as rounded lobes, bearing a long and short spine laterally on each apex.

FEMALE.—Gnathopod 2 like that shown for E.

molokai; accessory flagellum 3-articulate.

YOUNG MALE.—Gnathopod 2 bearing distinct palm

defined by 1 marginal spine and medial ridge bearing

spine similar to adult, setae present mainly on posterior

margin of hand, palm slightly excavate and bearing

several small spines, distal protrusion present.
120

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 69.—Elasmopus ecuadorensis hawaiensis Schellenberg, male, 4.9 mm, Hilo: a, telson.

Male, 5.0 mm: b, lateral gnathopod 1, setae removed; c, d, dactyls of pereopods 2, 5; e, half

of telson; f, g, article 2 of pereopods 4, 5; h, uropod 3; i, palm of medial gnathopod 2; ;, lateral

gnathopod 2; k, pleonal epimeron 3. Female, 3.7 mm: I, aberrant telson. Male, 3.6 mm,

Molokai: rn, hand of medial gnathopod 2.
NUMBER 3 4 121

MATERIAL.—Hilo, Hawaii, 20 June 1959, Clunio

Polster, collector H. Caspers (10). Molokai, Hawaii,

1959, algae, collector H. Caspers (1).

REMARKS.—This species has the same formula of

locking spines on pereopods 1-2 as does E. hooheno,

one sabre-shaped and one nonsabre-shaped but the

nonsabre-shaped spine of E. hooheno is of the corkscrew kind; the main locking spine of pereopods 3-5

in E. hooheno is a sabre while that of E. e. hawaiensis

is not.

Male gnathopod 2 resembles that of E. pectenicrus,

but the locking spines on pereopods 1-5 of the latter

are all straight; male pereopod 4 of E. e. hawaiensis

has a normal article 2.

DISTRIBUTION.—Hawaiian Islands.

Elasmopus hooheno, new species

FIOURE 70

DIAGNOSIS OF MALE.—Accessory flagellum 2-articulate; eyes medium in size, not as dark as in E. pocillimanus, with occasional central ommatidium visible,

pigment in alcohol brownish-purple; mandibular palp

normally falcate (as drawn for E. piikoi); article 5 of

gnathopod 1 about 80 percent as long as article 6;

anterodistal end of article 2 on gnathopod 2 with

slightly beveled corner bearing large spine (s), article

3 weakly lobate, article 5 short, bearing narrow but

relatively short, setose posterior lobe, article 6 elongate

and slightly tapering, palm oblique and about equal

in length to posterior margin of article 6, defined by

medium-sized conical cusp, distal end bearing spinose,

slightly scalloped protrusion, middle of palm with

truncate cusp, dactyl slender, normal, fitting palm;

posterodistal end of article 6 on pereopods 1-2 with 1

large striate sabre-spine and 1 straight horizontally

striate ("corkscrew") spine, dactyl subapically constricted, minutely striate, bearing stout seta and 2 setules; locking spines of pereopods 3-5 consisting of 1

slight sabre and 1 straight axially striate spine, dactylar

seta with weak flange; pereopods 3-5 bearing a few

long setae on articles 2, 4, 5, posterior margin of article

2 of regular dimensions and weakly serrate; coxa 6 with

unusually broad and slightly truncate anterior lobe;

pleonal epimera 1-2 with lateral ridge, posteroventral

corners sharply produced, posterior margins sparsely

serrate, epimeron 3 with small cusp at posteroventral

corner similar in size to a few posterior serrations, posterior margin nearly straight and posteroventral corner

grossly quadrate, epimera bearing ventral spines or

pairs of spines, no setae; rami of uropods 1-2 with

elongate terminal spines but not unusually so; inner

ramus of uropod 3 about two-thirds as long as outer,

apex of inner ramus truncate and spinose, apex of outer

spinose and bearing minute article 2; telson short,

squamiform, lobes weakly excavate apically, each bearing 1 long, 1 short spine.

FEMALE.—Gnathopod 2 like that of E. molokai, but

article 6 slightly stouter and palm with several small

stout spines, occasionally posteroventral corner of epimeron 3 with notch obsolescent.

HOLOTYPE.—Bishop Museum collections, catalog

number 7270, male 4.3 mm.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, other algae, rocks, coral,

24 May 1967.

MATERIAL.—Five specimens from the type-locality.

RELATIONSHIP.—This species has approximately

the same locking spine formula on pereopods 1-2 and

telson as does E. spinidactylus Chevreux, but the dactyls lack castellae and the second articles of pereopods

3-5 are of the regularly ovate, unnotched kind. Male

gnathopod 2 is very different from that of E. spinidactylus.

Perhaps this species has its closest relationship to

E. molokai because of the total number of similarities

(pereopods, telson, e.g.), but the main locking spine

on pereopods 1-2 of E. molokai is not a sabre, the eyes

are more weakly pigmented, the palm of male gnathopod 2 has no middle tooth, and article 2 lacks large

spines.

Terminal males of E. hooheno probably have not

been described and may, by indicative relationships,

have more strongly serrate and setose pleonal epimera

than seen in the present subterminal males.

DISTRIBUTION.—Hawaiian Islands.

Elasmopus molokai, new species

FIGURES 71, 72

DIAGNOSIS OF MALE.—Eyes dark but ommatidia

distinct throughout, large and few in number, pigment weak and purple to light brown in alcohol; mandibular palp article 3 normally falcate; article 5 of

gnathopod 1 about 80 percent as long as article 6;

article 2 of gnathopod 2 with large distolateral lobe,

article 3 broadly lobate, article 5 short, bearing narrow,

poorly setose posterior lobe, article 6 elongate, subrec-
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 70.—Elasmopus hooheno, new species, holotype, male, 4.3 mm, JLB Hawaii 17: a,

pereopod 5; b, medial gnathopod 2; c, dactyl of pereopod 5; d, medial gnathopod 1, setae

removed; e, uropod 3; /, telson; g, dactyl of pereopod 1; h, pleonal epimeron 3.

tangular but slightly tapering distalfy, palm oblique,

equal in length to posterior margin of article 6, defined

by large cusp and bearing sharp, subfalcate protrusion

near dactylar hinge armed with 1-2 spines, margin between processes flat and excavate, dactyl slender,

curved, fitting palm; posterodistal end of article 6 on

pereopods 1-2 with 1 irregularly shaped chisel spine,

slightly bifid apically or with small point, other locking

spine small, slightly sabre-shaped, dactyl subapically

constricted and bearing one stout free spine and 1-2

weak setules; locking spines of pereopods 3-5 normal;

pereopods 3-5 bearing a few long setae on articles 3-5

and on article 2 of pereopods 4-5, posterior margin of

article 2 of regular dimensions and weakly serrate; pleonal epimera 1-2 with lateral ridge, posteroventral

corners sharply produced, posterior margins sparsely

serrate, epimeron 3 with strong ventroposterior serrations, ventral margins of epimera bearing pairs composed of a spine and long seta; urosomal segments

simple; rami of uropods 1-2 with unusually elongate

terminal spines; inner ramus of uropod 3 about as

broad and three-fourths as long as outer ramus, apex
NUMBER 3 4 123

FIGURE 71.—Elasmopus molokai, new species, female, 3.3 mm, JLB Hawaii 6: a, b, gnathopods

1, 2. Male, 2.9 mm: c, gnathopod 2. Holotype, male, 3.1 mm: d, gnathopod 2; e, head. Male, 2.5

mm: /, pleonal epimeron 3.

of inner ramus with 2-3 spines, apex of outer ramus

with 3-4 spines and small article 2; telson short, apices

broadly sinuous, armed with 2-4 apical and subapical

spines and 1 pair of lateral setules on each side.

FEMALE.—Gnathopod 2 typical of the genus and

other characters like those of male.

JUVENILE MALE.—Distal palmar process of gnathopod 2 less falcate than in terminal male, denning cusp

occurring more distally with dactyl overlapping it,

excavation narrower and a subsidiary defining cusp

occurring proximally, this cusp gradually disappearing

360-034 0—70—9

in adult; otherwise like adult but with fewer spines

and setae elsewhere.

JUVENILE (2.0 mm).—Setae on article 2 of pereopods and epimera absent; epimeral spines when occurring set in pairs or singly; articles 3-4 of perebpods 3-5

with only a few setae; article 1 of antenna 1 with long

stout ventrodistal spine; accessory flagellum 2-articulate; inner ramus of uropod 3 with either 1 or 2 stout

spines, 1 elongate; telson with 2-3 spines on each lobe.

HOLOTYPE.—Bishop Museum collections, catalog

number 7271, male, 3.1 mm.
124

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

d

FIGURE 72.—Elasmopus molokai, new species, holotype, male, 3.1 nun, JLB Hawaii 6: a,

pereopod 5; b, uropod 3; c, dactyl of pereopod 3; d, mandibular palp. Male, 2.5 mm: e,

gnathopod 2; /, dactyl of pereopod 1. Female, 3.3 mm: g, telson.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, coralline footballs, 29 January 1967.

MATERIAL.—JLB Hawaii 3 (15), 5 (25), 6 (22),

8 (6), 10 (6), 11 (1), 12 (2), 13 (3), 14 (1). Fee

1 (?1).

RELATIONSHIP.—This species has sympatric relationships with E. pocillimanus (Bate) and in juvenile

stages superficially might be confused with that species

except for the following differences: (1) in all stages

E. pocillimanus eyes are so darkly pigmented that in

alcohol only peripheral ommatidia are visible, whereas

they show through the lighter pigment centrally in

E. molokai; occasionally the pigment of E. molokai

appears very dark but microscopic examination demonstrates the appearance of central ommatidia; (2)

the short telson with sinuotruncate apices bearing
NUMBER 3 4 125

slightly subapical spines; (3) the presence of a few

setae on epimera and pereopods 3-5 in adults; (4) the

replacement of striate sabre-spines on pereopods 1-2

with a chisel-shaped and a simple spine.

Distinction in juvenile stages from E. piikoi, new

species, are very difficult to make and are discussed

under the latter heading.

Allopatric relationships occur with E. minimus

Chevreux (1908) ; E. molokai may be synonymous with

E. minimus from Gambier Archipelago; the latter is

based on a male resembling the juvenile male of E.

molokai. The telson of E. minimus appears also to be

shortened from Chevreux's lateral drawing.

Elasmopus molokai differs from E. rapax in the same

characters mentioned for E. piikoi.

DISTRIBUTION.—Hawaiian Islands.

Elasmopus pectenicrus (Bate)

FIGURES 73, 74

Moera pectenicrus Bate, 1862, p. 192, pi. 34, fig. 8.

Elasmopus serrula Walker, 1904, pp. 277-278, pi. 8, fig. 37.

Elasmopus pectenicrus.—J. L. Barnard, 1955a, pp. 8-10, fig.

4 [with references].

DIAGNOSIS OF MALE.—Accessory flagellum about

4-articulate; eyes medium in size, clear in alcohol;

mandibular palp normally falcate; article 5 of gnathopod 1 longer than article 6, latter softly rectangular,

palm nearly transverse, medial faces of articles 5-6 and

posterior margin of article 5 densely setose; article 2 of

gnathopod 2 not lobate, article 3 normally lobate, article 5 short, bearing narrow, setose posterior lobe, article

6 elongate and tapering, palm fully oblique and not

marginally defined from posterior edge of article 6,

palm S-shaped, medial face with small cusp defining

palmar extent, distal end with small protuberance at

posterior end of transverse margin, small process at

medial hinge line near base of dactyl, one short ridge

on medial face near end of dactyl, latter slender, evenly

curved, not reaching medioproximal cusp, palm

densely setose its full length; posterodistal end of article 6 on pereopods 1-2 with long straight, and short

straight spine, long spine slightly twisted and bent

distally, sharp, dactyl stout, distal constriction bearing

normal stout seta and 2 curved setules, basal part of

constriction with small sharp, distally angular keel;

locking spines of pereopods 3-5 normal; pereopod 3

with stout, subrectangular article 2 with minute posterior serrations, posteroventral lobe scarcely extended,

pereopod 4 with posteroventral margin excavate along

strongly extended lobe and deeply castelloserrate,

pereopod 5 with broadly hemispherical, minutely serrate posterior margin scarcely extended ventrally,

pereopods 3-5 with a few short setae on articles 3-6,

none on article 2 (except setules) ; pleonal epimera 1-2

with lateral ridge and small sharp posteroventral tooth,

posterior margins with several normal serrations, epimeron 3 bluntly quadrate, posteroventral tooth absent or

obsolescent, posterior margin with several normal serrations, ventral margins of all epimera with spines in

groups of 1, 2, and 3, no setae; urosomal segments

simple; rami of uropods 1-2 with medium to elongate

terminal spines; inner ram us of uropod 3 only twothirds as long as outer ramus, both relatively short for

genus, truncate, apically spinose, outer lacking article

2; telson of medium length, lobes apically truncate and

armed with 5-6 spines each.

FEMALE.—Pleonal epimeron 3 with 2 sharp medium

serrations on posteroventral corner; gnathopod 2 of

medium size, palm even, oblique, lined with short

stout spines, defined by 1 marginal and 1 medial spine,

inner margin of dactyl wavy; article 2 of pereopod 5

slightly narrower than in male, posteroventral margin

slightly beveled on pereopod 4, not excavate, castelloserrations slightly smaller than in male; setae of pereopods rare; article 6 of gnathopod 1 relatively longer

and larger than in male.

JUVENILE (smallest available, 5.3 mm).—Pleonal

epimeron 3 like that of female; article 2 of pereopods

4—5 unmodified except for presence of coarse castelloserrations, much coarser on pereopod 5 than in adult;

ommatidia dispersed, few in number, distinctly bigeminate, eye otherwise of size similar to adult; each

telsonic lobe with only 4 spines; gnathopod 2 lacking

marginal defining spine seen in female, dactyl strongly

castelloserrate on inner margin; major locking spine of

pereopods slightly sharper than in adult, apex slightly

curved.

MATERIAL.—Same as in Barnard (1955a); material

herein figured or analyzed again: Pearl Harbor, drydock, 15 April 1948, collector C. H. Edmondson;

Oahu, 24 January 1944, 5-foot depth (also Pearl Harbor, and forming part of Station 2 material reported

in Barnard, 1955a).

REMARKS.—The strong castelloserrations occur on

pereopod 4, not pereopod 5 of the adult male; my

(1955a) statement was in error; juveniles, however,

have strong serrations on pereopod 5.

DISTRIBUTION.—Tropicopolitan.
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

k

FIGURE 73.—Elasmopus pectenicrus (Bate), male, 10.6 mm, Oahu, 5', 24 January 1944: a, head;

b, gnathopod 1, lateral; c, pereopod 2; d, uropod 3; e, pleonal epimeron 3; /, mandibular palp.

Juvenile female, 5.3 mm: g, dactyl of gnathopod 2; h, dactyl of pereopod 2; i, j , k, bases of

pereopods 3, 4, 5. Female, 6.3 mm: /, pleonal epimeron 3; m, telson; n, base of pereopod 4.
NUMBER 3 4 127

9

FIGURE 74.—Elasmopus pectenicrus (Bate), male, 10.6 mm, Oahu, 5', 24 January 1944: a, apex

of pereopod 5; b, gnathopod 2, medial; c, gnathopod 2, lateral; d, e, f, pereopods 3, 4, 5; g,

palmar processes and inner edge of dactyl on gnathopod 2; h, coxa 1; i, apex of pereopod 2.

Elasmopus piikoi, new species

FIGURES 75, 76

DIAGNOSIS OF MALE.—Accessory flagellum about 3-

articulate; eyes medium in size, not as dark as in E.

pocillimanus, with occasional central ommatidium

visible, pigment in alcohol often turning to dark purple

or brownish purple; mandibular palp normally falcate; article 5 of gnathopod 1 about 87 percent as

long as article 6; anterodistal end of article 2 on

gnathopod 2 with pendant lobe on medial and lateral

side, slight anterior protrusion above lobe, article 3

broadly lobate, article 5 short, bearing narrow but

relatively short, setose posterior lobe, article 6 elongate

and strongly tapering, palm very oblique and much

longer than posterior margin of article 6, defined by
128

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 75.—Elasmopus piikoi, new species, holotype, male, 5.0 mm, JLB Hawaii 13: a, head;

b, c, inner plates of maxillae 1; d, left palp of maxilla 1; e, uropod 3; f, telson; g, mandibular palp

article 3, setae removed; h, gnathopod 1, setae removed; i, j , apices of inner and outer rami on

uropod 3. Male, 4.1 mm: k, gnathopod 2, medial; /, pleonal epimeron 3.
NUMBER 3 4 129

FIGURE 76.—Elasmopus piikoi, new species, holotype, male, 5.0 mm, JLB Hawaii 13: a, b,

gnathopod 2, lateral and medial views; c, d, apices of pereopods 2, 3; $, coxa 6 (tilted on its

side); f, right palp of maxilla 1.
130

small cusp, distal end with truncate or slightly bilobate

process bearing a few spines, midpalmar margin with

large truncatoconiform process, dactyl slightly overriding palm and fitting facial recess defined by spine,

inner margin of dactyl with large process, margin thus

bisinuate, dactyl tending to be stout; posterodistal end

of article 6 on pereopods 1-2 with 1 irregularly shaped

chisel spine, slightly bifid apically or with small point,

other locking spine small, slightly sabre-shaped, dactyl

subapically constricted and bearing one stout free spine

and 1-2 weak setules; locking spines of pereopods 3-5

normal; pereopods 3-5 bearing several long setae on

articles 3-5 and on article 2 of pereopods (3) 4—5,

posterior margin of article 2 of regular dimensions and

weakly serrate; coxa 6 with unusually broad and

slightly truncate anterior lobe; pleonal epimera 1-2

with lateral ridge, posteroventral corners sharply produced, posterior margins sparsely serrate, epimeron 3

with evenly quadrate posteroventral corner, ventral

margins of epimera bearing pairs of short spines, no

setae; urosomal segments simple; rami of uropods 1-2

with elongate terminal spines but not unusually so;

inner ramus of uropod 3 about three-fourths as long as

outer ramus, apex of inner ramus slightly tapering and

bearing 1 small seta or several spines (right and left

differing on same animal occasionally), apex of outer

ramus spinose and bearing small article 2; telson elongate, lobes apically excavate and bearing 3 terminal

spines each.

FEMALE.—Gnathopod 2 like that of E. molokai;

pleonal epimeron 3 like that of young male; inner

ramus of uropod 3 broadly truncate and spinose;

pereopods 3-5 lacking setae (no female at hand nearly

as large as males).

YOUNG MALE.—Gnathopod 2 with more spines on

middle and distal palmar processes than in adult but

processes smaller, dactyl weakly sinuate; pleonal epimeron 3 with posteroventral tooth like epimera 1-2,

spines on ventral margin occurring singly in each set;

inner ramus of uropod 3 with strongly spinose apex;

telson with 2 spines on each lobe.

JUVENILE.—Pleonal epimera like young male but

often lacking spines; inner ramus of uropod 3 like

adult male, tapering and bearing one apical seta or

one seta and one spine; telson with 2 spines on each

apex.

HOLOTYPE.—Bishop Museum collections, catalog

number 7272, male, 5.0 mm.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

Oahu, 3-4 m, corals, algae, corallines, 8 April 1967.

MATERIAL.—JLB Hawaii 2 (32), 3 (1), 12 (12),

13 (14),14 (25), 17 (2).

RELATIONSHIP.—A sympatric resemblance occurs

with E. pocillimanus (Bate) and E. molokai, new

species, and differentiation of juveniles from similar

stages of the latter species is difficult, except for the

short telson of E. molokai. A judgment of this character is difficult to make without dissection until considerable experience with the material has been obtained. Gnathopods of the adult male are clearly different, and the presence of epimeral setae on adult

female E. molokai is characteristic. The pair of sabrespines on pereopods 1-2 of E. pocillimanus is definitive

of that species in all stages.

Elasmopus piikoi gnathopod 2 of the young male

resembles that of E. calliactis Edmondson, but the eyes

devoid of black pigment distinguish all stages of

the latter species.

Adult males of E. piikoi differ from E. rapax Costa

(J. L. Barnard, 1962b) in the absence of the fourth

palmar process on gnathopod 2 that forms a long

conical lateral tooth. Males, females, and large juveniles of E. rapax have several sharp serrations on epimeron 3. The telson of E. rapax has spines only on the

apicolateral notch, and the medial projecting portion

is smooth. Article 2 on the outer ramus of uropod 3

is absent in E. rapax.

Elasmopus holgurus J. L. Barnard (1962b) from

California differs from E. piikoi in the absence of a defining tooth on the palm of male gnathopod 2,

although the facial ridge and spine into which the

dactylar apex fits are present in both species.

DISTRIBUTION.—Hawaiian Islands.

Elasmopus pocillimanus (Bate)

FIGURES 77, 78

Moera pocillimanus Bate, 1862, p. 191, pi. 34, fig. 7.

Elasmopus pocillimanus.—Stebbing, 1906, pp. 443-444;

Chevreux and Fage, 1925, p. 246, fig. 257.—Schellenberg,

1938, p. 56, fig. 28.

DIAGNOSIS OF TERMINAL MALE.—Accessory flagellum about 3-articulate; eyes very large, only peripheral

ommatidia visible through heavy black pigment; mandibular palp article 3 extremely falcate for genus;

article 5 of gnathopod 1 about 80 percent as long as

article 6; articles 2 and 3 of gnathopod 2 not lobate,
NUMBER 3 4 131

article 5 short, bearing narrow, setose posterior lobe,

article 6 very elongate, subrectangular, palm and posterior margin of article 6 confluent but defined by

weak, obtuse cusp, palm near dactyl with slight protrusion bearing several marginal and submarginal

spines, medial palmar face bearing deep hollow defined proximally by weak spine, dactyl stout, curved,

overlapping palm to close into recess; posterodistal end

of article 6 on pereopods 1-2 with 2 large, sabre-shaped,

striate locking spines, dactyl constricted very distally,

bearing 1 large, sharp straight seta and 2 strap-shaped

thinner setae; locking spines of pereopods 3-5 composed of one sabre and one normal, both slightly striate; pereopods 3-5 lacking any long setae, posterior

margin of article 2 of regular dimensions and weakly

serrate; pleonal epimera 1-2 with lateral ridge, posteroventral corners sharply produced, posterior margins

sparsely serrate, epimeron 3 with evenly quadrate posteroventral corner, ventral margins of epimera bearing

pairs of short spines, no setae; urosomal segments simple; rami of uropods 1-2 with unusually elongate terminal spines; inner ramus of uropod 3 as broad and

about seven-eighths as long as outer ramus, apex of

inner ramus with 3 spines, apex of outer ramus spinose

and bearing small article 2; telson slightly elongate,

lobes apically extended into subovate unarmed plates,

each lobe with one midlateral spine and pair of setules.

YOUNG MALE.—Gnathopod 1 stouter than in terminal stage, article 5 shorter; palmar margin of gnathopod 2 with palmar protrusion poorly defined but

equally spinose, medial hollow rudimentary, dactyl

thin; pleonal epimeron 3 with strong posteroventral

tooth like epimera 1-2.

FEMALE.—Not in Hawaiian collections.

JUVENILES.—Inner ramus of uropod 3 shortened

and bearing 1 apical seta, article 2 of outer ramus relatively much larger than in adult, apparently remaining

same size in juvenile and adult while article 1 increasing in size; pleonal epimeron 3 with small posteroventral tooth, ventral spines on epimera when present

occurring singly; each telsonic lobe with asymmetrical

apical extensions, medial point reaching farthest, spine

on each lobe essentially terminal; eyes nearly as dark as

in adult, but slightly smaller relative to head; pereopods

1-2 also bearing 2 striated sabre-spines as in adult and

pereopods 3-5 with one such spine; terminal spines of

uropods 1-2 relatively as long as in adult.

MATERIAL.—JLB Hawaii 5 (2), 6 (1), 17 (1).

REMARKS.—Article 3 of the maxillipedal palp has a

distal protrusion ornamented like a pin cushion.

DISTRIBUTION.—Cosmopolitan in warm seas.

Elasmopus rapax Costa

FIGURES 79, 80

Elasmopus rapax Costa.—J. L. Barnard, 1955a, pp. 10-12,

fig. 5 [in part, not figs. 5b, g, material from Honolulu

Aquarium=£. calliactis] [with references]; 1962b, pp.

94-96, figs. 16, 17.

DIAGNOSIS OF HAWAIIAN MALE.—Accessory flagellum about 4-articulate; eyes medium in size, not as

dark as in E. pocillimanus, with occasional central ommatidium visible through purplish-brown pigment in

alcohol; mandibular palp normally falcate; article 5 of

gnathopod 1 as long as article 6, latter heavily setose

medially; articles 2-3 of gnathopod 2 not lobate, article 5 short, bearing narrow setose posterior lobe,

article 6 elongate and tapering, palm very oblique and

longer than posterior margin of article 6, extent defined by small tooth on medial face, distal end of palm

with large lateral tooth, short truncate medioproximal

tooth and broad shallow spinose distomedial protrusion, dactyl slender but shorter than palm and strongly

bent basally; posterodistal end of article 6 on pereopods

1-2 with one nearly straight unstriate spine minutely

constricted and/or pointed distally and one short normal spine, dactyl slender, distal constriction bearing 1

stout seta and 2 curved setules; locking spines of pereopods 3-5 normal; pereopods 3-5 bearing long setae on

articles 2-6, posterior margin of article 2 of regular

dimensions and weakly crenulate; pleonal epimera

1-2 with lateral ridge and small sharp, posterodistal

tooth, posterior margins densely serrate but serrations

of normal depth, epimeron 3 deeply serrate ventroposteriorly, one strong serration forming quadrate

posteroventral corner; ventral margins of epimera with

mixture of spines and long setae occasionally occurring

as singles or triads, frequently as pairs; urosomal segments simple; rami of uropods 1-2 with long terminal

spines; inner ramus of uropod 3 subequal to. outer

ramus in length, both broad and flat, apices broadly

truncate and spinose, outer ramus lacking article 2;

telson of medium length, apices extended as medial

lobes, bluntly coniform, laterally each lobe carrying 3

spines in notch.
132

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 77.—Elasmopus pocillimanus (Bate), male, 5.1 mm, JLB Hawaii 6: a, uropod 3; b,

apex of outer ramus on uropod 3; c, d, e, pereopods 3, 4, 5; /, telson; g, head; h, mandibular

palp; i, pereopod 1; ;, accessory flagellum. Male, 3.3 mm: k, article 2 on outer ramus of uropod 3;

I, pleonal epimeron 3 (upside down); m, lateral gnathopod 1, setae removed. Juvenile, 1.7 mm:

n, telson; o, pleonal epimeron 3 (upside down); p, uropod 3.
NUMBER 34 133

FIGURE 78.—Elasmopus pocillimanus (Bate), male, 5.1 mm, JLB Hawaii 6: a, lateral gnathopod

1; b, c, medial gnathopod 2; d, apex of pereopod 4; e, head; /, apex of pereopod 1; g, apex of

maxillipedal palp; h, pleon; i, coxa 4. Male, 3.3 mm: ;, medial gnathopod 2.
134

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 79.—Elasmopus rapax Costa, juvenile, 3.8 mm, Pearl Harbor, 1948: a, spine on article

6 of pereopod 1. Male, 8.3 mm: b, lateral gnathopod 2; c, lateral gnathopod 1; d, medial

gnathopod 2; e, telson. Female, 6.0 mm: /, head; g, medial gnathopod 2.

FEMALE.—Gnathopod 2 small, palm even, bearing

small spines and defined by medial triad and 1 marginal stout spine, each telsonic lobe with 2-3 spines,

apices sharp; pereopods and epimera lacking setae.

JUVENILE (smallest available, 3.8 mm).—Each telsonic lobe with 2 spines, apices sharp, epimera with

only short spines as singles, pairs and occasional triads,

but epimeron 3 already strongly serrate as in adult;

rami of uropod 3 narrower than in adult, but truncate

apex of inner ramus already with 7 spines.

MATERIAL.—Same as J. L. Barnard (1955a), minus

material from Honolulu Aquarium; material figured

herein: Pearl Harbor, 28 April 1948, collector C. H.

Edmondson.

REMARKS.—Distinguished in various stages by absence of article 2 on outer ramus of uropod 3 and

sharply serrate epimeron 3, but smallest juveniles

unstudied.

DISTRIBUTION.—Cosmopolitan in warm seas and occasionally in cold-temperate regions.
NUMBER 34 135

FIGURE 80.—Elasmopus rapax Costa, male, 8.3 mm, Pearl Harbor, 1948: a, b, c, pereopods 3, 4,

5, in whole or part; d, uropod 3; e, apex of pereopod 1; /, mandibular palp; g, pleonal epimeron 3.
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Elasmopus spinidactylus Chevreux

FIGURES 81, 82

Elasmopus spinidactylus Chevreux, 1908, pp. 486-

489, figs. 9-10.—Walker, 1909, pp. 336-337.—

Schellenberg, 1938, p. 55.—Shoemaker, 1942, p.

13.—J. L. Barnard, 1965a, p. 504.

DIAGNOSIS OF HAWAIIAN FEMALE AND MEDIUM-AGED

MALE.—Accessory flagellum about 3-articulate; eyes

medium in size, not as dark as those of E. pocillimanus,

with occasional central ommatidium visible, pigment

in alcohol often turning to dark purple or brownish

purple; mandibular palp article 3 normally falcate;

article 5 of gnathopod 1 about 80 percent as long as

article 6; articles 2 and 3 of gnathopod 2 without

anterodistal lobes, article 5 short, bearing narrow posterior lobe, article 6 elongate and scarcely tapering,

palm and posterior margin of article 6 confluent, palm

FIGURE 81.—Elasmopus spinidactylus Chevreux, female, 4.3 mm, Molokai: a, b, c, pereopods 3, 4,

5, in whole or part; d, distolateral end of article 5 on pereopod 1; e, telson; f, apex of pereopod 1;

g, apex of pereopod 3; h, palm of gnathopod 2, mediolateral view; i, lateral gnathopod 2.
NUMBER 34 137

FIGURE 82.—Elasmopus spinidactylus Chevreux, female, 4.3 mm, Molokai: a, head; b, gnathopod

1; c, pereopod 2; d, mandibular palp; e, pleonal epimeron 3. Another female: /, uropod 3; g,

telson. Juvenile, 2.3 mm: h, palm of gnathopod 2, medial view.

undefined, distal end slightly protuberant and bearing

3-5 pairs of spines, remainder of palm and posterior

half of medial surface of article 6 strongly setose, dactyl slender; posterodistal end of article 6 on pereopods

1-2 with 1 large, striate sabre-spine and 1 small normal

spine, dactyl constricted and bearing stout seta and

2 setules, proximal inner margin with 2 (juveniles)

to 4, occasionally 5, large to small sharp, broad teeth;

locking spines of pereopods 3-5 normal but dactyls

like pereopods 1-2; article 2 of pereopods 3-5 narrow,

subrectangular, that of pereopod 5 becoming slightly

ovate, posterior margins weakly serrate and posteroventral corners each with deep quadrate notch, article

2 lacking long setae and only distal articles of pereopod

5 with several long setae; pleonal epimera 1-2 with

lateral ridge and sharply produced posteroventral corners, posterior margins sparsely serrate, epimeron 3

also with sharp posteroventral tooth, ventral margins

of epimera with a few single spines, rarely paired; urosomal segments simple; rami of uropods 1-2 with

medium-length terminal spines; inner ramus of uropod

3 small, rami gaping, inner ramus reaching only about

halfway along outer ramus, apex with 2 spines, outer

edge with 2 more singles, outer ramus deeply stair-
138

stepped, apically truncate, article 2 not observed; telson short, apices broadly truncate, each armed with

3 long spines.

JUVENILE.—Inner ramus of uropod 3 pointed, with

2 apical setules, telson with 2 spines on each lobe;

palm of gnathopod 2 shorter than posterior margin

of article 6, weakly sinuous, broad distal protusion

spinose, proximal sinuosity distinct from posterior margin of article 6 and defined by medial spine on ridge;

pereopods 1-5 with as few as 2 accessory teeth on

dactyls.

Terminal male with large palmar protrusion on

gnathopod 2, as described by Chevreux, not present

in Hawaiian collections at hand.

MATERIAL.—Hanauma Bay, Oahu, on Microdictyon, 29-3-64 (4); USNM 262506, Molokai (?"Haleolone PI (Caspers)"? ix 59 Algen Prob (6) (place

name unknown and date illegible).

REMARKS.—The gross teeth on the pereopodal dactyls distinguish this species from all others in Hawaii.

Probably various races of this species throughout its

range, like many other Hawaiian species, will be found

but cannot be distinguished at present.

DISTRIBUTION.—Chagos Islands, eastward through

Micronesia, Hawaii, Glipperton Island, SE Polynesia;

and in the Caribbean Sea on the shores of Venezuela.

Eriopisa Stebbing

This genus, Eriopisella Chevreux and Netamelita

J. L. Barnard appear to represent a polyphyletic group

of species possibly derived from melita ancestors by

reduction in size and complexity of male gnathopod 2

but at one extreme (type-species of Eriopisa), article

2 of the outer ramus on uropod 3 becomes very elongate and flat and in the other extreme (type-species

of Netamelita) the article is lost as it is in most species

of Melita. The eriopisella line generally has species

with small article 2 on the outer ramus, like 2 species

of Eriopisa, and the medial setae on the inner plates

of the maxillae have been lost. Netamelita may not be

distinct from Eriopisella, even though article 2 of the

outer ramus on uropod 3 is lost, and gnathopod 2 is

very strongly reduced to a size scarcely larger than

gnathopod 1. The importance of these characters is

difficult to evaluate in view of the alternative presence

of article 2 on the outer ramus of uropod 3 in Melita

and the diversity of other character differences among

the various species of the 3 eriopisa genera. Thus,

article 3 of the mandibular palp in Eriopisa capensis

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

K. H. Barnard (1916) has become reduced, E. garthi

J. L. Barnard (1952) has maxillae with intermediate

setation, and E. elongata (Bruzelius, see Sars, 1895)

has extremely reduced coxae and retains a lateral

cephalic notch.

When European taxonomists established and refined

the concepts of Eriopisa and Melita, the various tropical and warm-temperate species were unknown. Numerous gradations are now seen between the two

extremes. The new Hawaiian species to be described

confuses the generic classification further. It has normal Eriopisa-Melita mandibular bodies and maxillae,

but has the short article 2 on the outer ramus of uropod

3 and resembles Netamelita more than the other genera

in the transverse anterolateral cephalic margins. It is

as clearly distinct phyletically from the type-species of

Eriopisa as are Netamelita and most members of

Eriopisa and Eriopisella; but it is retained in Eriopisa

as a matter of convenience on the basis of maxillae 1-2.

We are thus in a position of recognizing defects in

the present classification before more than a fraction

of the known species has been discovered; further

alignment should be reserved until exploration has

revealed a majority of the species (Gurjanova, 1965).

Eriopisa (?) hamakua, new species

FIGURES 83, 84

DIAGNOSIS OF MALE.—Eyes represented by 8-9 clear

facets weakly resembling ommatidia in front of light

orange pigment splotch in alcohol; lateral cephalic

lobes weakly protruding and lacking anteroventral

notch; antenna 1 as long as head and body together,

accessory flagellum 2-articulate, flagellum of antenna 2

as long as peduncular article 5 and three-fourths of

article 4 together; mandibular palp long, article 3

subfalcate and strongly setose on concave margin; inner plates of maxillae 1-2 medially setose; coxae overlapping each other, coxae 1-2 longer than broad, 3-4

about as broad as long, coxa 1 rectangular, not produced or extended anteroventrally: article 4 of gnathopod 1 not inflated, article 5 much longer than broad,

very densely setose posteriorly and laterally but unlobate, article 6 slender, rectangular, palm nearly transverse, dactyl short but fitting spinose palm; gnathopod

2 much larger than 1, article 5 short, stout, slightly

inflated, article 6 very elongate and slender, palm from

lateral view appearing to occupy entire posterior margin of hand, densely setose, dactyl about half as long
NUMBER 34 139

h

FIGURE 83.—Eriopisa hamakua, new species, holotype, male, 4.0 mm, JLB Hawaii 2: a, head;

b, medial gnathopod 2; c, lateral gnathopod 1; d, left lateral urosome, uropods removed; e,

mandibular palp article 3; /, apex of medial gnathopod 1; g, pereopod 3; h, lateral gnathopod 2.
140

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

9

FIGURE 84.—Eriopisa hamakua, new species, holotype, male, 4.0 mm, JLB Hawaii 2: a, part of

pereopod 5; b, part of pereopod 4; c, half of telson; d, pleonal epimera 1-3; e, uropod 2; /, combrow of uropod 2; g, uropod 1.

as hand and slightly sinuous along inner margin, palm

mediodistally with 3 spines and bearing defining bulge

with 2 spines at apical extent of dactyl, palm medially

with weak excavation; pereopods 1-2 with slender

article 2; article 2 of pereopods 3-5 rectangular and

relatively slender, posteroventral corners with sharp

tiny cusp but not lobate, remainder of pereopodal

articles slender; uropod 1 with large distolateral peduncular spine, outer ramus of uropod 2 slightly

shorter than inner ramus, peduncle of uropod 2 with

comb-row of spines on mediodistal end; (uropod 3 unknown) ; pleonal epimera 1-2 with small sharp posteroventral tooth and nearly straight posterior margins,

epimeron 3 with large obtusely rounded posterior extension bearing sharp cusp on posterodorsal corner,

oblique upper margin with 3 setules, ventral margin

spinose; pleonite 4 with small dorsolateral setule, 5

with dorsolateral spinule on each side; telson thin

dorsoventrally, of medium length but slightly broader

than long, fully cleft, lobes less tilted than in E. laakona,

new species, each apex with obtusely conical projection

guarded on each side by 3 spines, middlemost spine

of each triad longest; branchiae broadly ovate and

pyriform, with narrow necks.

HOLOTYPE.—Bishop Museum collection, catalog

number 7273, male 4.0 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 2, open sea off W end

of Pearl Harbor, Oahu, 4-5 m, sand, encrustations, 29

January 1967.

RELATIONSHIP.—The absence of uropod 3 does not

seriously detract from the placement of this species in

Eriopisa, but relationships with several other species

are clouded; however, the shape of pleonal epimeron

3 is so far unique in the genus.

Gnathopods have a slight relationship to those of E.

philippensis (Chilton) and E. seurati Gauthier, but

article 4 of gnathopod 1 is not swollen.

The right mandibular molar of E. hamakua has a

seta and a flake.

DISTRIBUTION.—Hawaiian Islands.

Eriopisa laakona, new species

FIGURES 85, 86

DIAGNOSIS OF ?MALE.—Eyes absent, lateral cephalic

lobes nearly vertically truncate and not protruding,

lacking anteroventral notch; antenna 1 as long as head

and pereon together, accessory flagellum 2-articulate,

flagellum of antenna 2 as long as article 5 and half of

article 4 together; mandibular palp long, slightly

thickened, article 3 subfalcate and strongly setose; inner plates of maxillae 1-2 medially setose; coxae overlapping each other, coxae 1-2 longer than broad, 3-4

broader than long, coxa 1 rectangular, not produced

or extended anteroventrally; article 4 of gnathopod 1

not inflated, article 5 longer than broad, posteriorly

setose but unlobate, article 6 stout, rectangular, palm

nearly transverse, dactyl short but fitting spinose palm;
NUMBER 3 4 141

gnathopod 2 larger than 1, article 5 of similar shape,

but article 6 elongate, palm extremely oblique and defined from posterior margin of article 6 by 2 spines, 1

of them medial, dactyl slightly overlapping palm;

pereopods 1-2 with slender article 2; article 2 of pereopods 3-5 changing from rectangular to ovatopyriform

consecutively, but relatively slender, posteroventral

corners with tiny sharp cusp but not lobate, remainder

of pereopodal articles slender; uropod 1 with large distolateral peduncular spine, outer ramus of uropod 2

slightly shorter than inner, mediodistal peduncular

margin of uropod 2 with comb-row of spines, article 2

of outer ramus on uropod 3 about 40 percent as long as

article 1, forming slender cone lacking spines, inner

ramus with sharp apex; pleonal epimera 1-2 with small,

sharp posteroventral tooth and nearly straight posterior

margins, epimeron 3 with large posteroventral tooth

and slightly bulging posterior margin; pleonites 4-5

with 1 dorsolateral articulate spine on each side; telson thick dorsoventrally, short, broader than long,

fully cleft, lobes slightly tilted away from horizontal

axis to form tent as in Hyale, each apex with obtuse

k

FIGURE 85.—Eriopisa laakona, new species, holotype, ?male, 2.0 mm, JLB Hawaii 17: a, head;

b, coxa 1 (tilted on its side); c, coxa 2; d, side view of telson; e, half of telson, dorsal; f, medial

antenna 1; g, mandibular palp; h, apex of pereopod 1; /', pleonal epimeron 3 and urosome; j ,

inner plate of maxilliped; k, pleonal epimera 1—3. Specimen from JLB Hawaii 12: /, anterior

margin of head.
142

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 86.—Eriopisa laakona, new species, holotype, ?male, 2.0 mm, JLB Hawaii 17: a, pereopod

5; b, apex of maxillipedal palp; c, uropod 3; d, medial gnathopod 2; e, part of pereopod 4; /,

medial gnathopod 1; g, pereopod 1.
NUMBER 34 143

projection guarded by spine on each side, lateral spine

shortest; branchiae ovate and pyriform, with narrow

necks.

HOLOTYPE.—Bishop Museum collections, catalog

number 7274, ?male, 2.0 mm.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, algae, rocks, corals, 24

May 1967.

MATERIAL.—JLB Hawaii 12 (1), 17 (3).

RELATIONSHIP.—Eriopisa seurati Gauthier, from

Tunisia, also has the short article 2 on the outer ramus

of uropod 3 like the Hawaiian species, but has numerous differences in mandibular palp, pereopods, gnathopods, head, telson, and otherwise on uropod 3.

Eriopisa garthi J. L. Barnard from Baja California,

E. elongata (Bruzelius) from the North Atlantic, and

the subterranean E. philippensis (Chilton) have the

elongate kind of article 2 on the outer ramus of uropod

3. Eriopisella capensis (K. H. Barnard, 1916) (see

Gurjanova, 1965) from South Africa has a short article 2 on uropod 3, but has rounded epimera, acutely

produced coxa 1, blunt inner ramus on uropod 3, a

short, rectolinear, apically setose article 3 on the

mandibular palp, and a stout, lobate article 5 on

gnathopod 2.

The four known species of Eriopisella differ from

E. laakona in the absence of medial setae on the inner

plates of the maxillae, but resemble it, like Eriopisa

seurati, in the shortness of article 2 on the outer ramus

of uropod 3. Each of them has numerous other differences from E. laakona. Those species are Eriopisella

pusilla Chevreux, E. sechellensis Chevreux, E.

nagatai Gurjanova, and E. capensis (K. H. Barnard).

Eriopisa laakona differs from E. chilkensis Chilton

(1921) in the subfalcate palp article 3 of the mandible,

the shorter article 2 on the outer ramus of uropod 3,

the larger tooth on epimeron 3, the narrower article 2

of pereopod 5, and the narrower hand of gnathopod 1.

Eriopisella nagatai Gurjanova (1965) has a very

broadly inflated palm on gnathopod 1, normally narrowed telsonic lobes, a further shortened article 2 on

the outer ramus of uropod 3, trapezoidal and anteroventrally extended coxa 1, and a grossly lobate article

2 on pereopod 5.

DISTRIBUTION.—Hawaiian Islands.

Eriopisella sechellensis upolu, new subspecies

FIGURES 87, 88

DIAGNOSIS.—Male and female similar; eyes of medium size, brownish-purple in alcohol and with several

clear peripheral ommatidia; lateral cephalic lobes

scarcely protruding and rounded anteroventrally, lacking anteroventral notch; antenna 1 as long as head

and first 9 body segments together, accessory flagellum

short, 1 or 2-articulate, article 2 very small; antenna 2

about 40 percent as long as antenna 1; mandibular

palp long, article 3 longer than 2, rectolinear and bearing 2 apical setae; inner plates of maxillae 1-2 setose

only apically, inner plate of maxilla 1 with only 3 setae,

palp article 1 elongate; coxae overlapping each other,

coxae 1-2 broader than long, coxae 3-4 about as broad

as long, coxa 1 strongly extended forward anteroventrally; article 4 of gnathopod 1 not inflated, article

5 longer than broad, weakly setose posteriorly but unlobate, article 6 rectangular, palm oblique, slightly

convex, defined by 3 long marginal spines in tandem

and 1 submarginal medial spine, dactyl fitting palm;

gnathopod 2 scarcely larger than 1, article 5 longer

than 6, bearing posterior lobe sloping distally and extended slightly along article 6, latter like that of

gnathopod 1; pereopods 1-2 with slender article 2,

middle of article 6 bearing compact group of posterior

spines, dactyl with small accessory tooth; article 2 of

pereopods 3-5 changing from rectangular to slightly

pyriform consecutively, posteroventral corners unproduced, quadrate or subquadrate, remainder of pereopodal articles slender, dactyl not bifid, but with hooked

distal nail and with constriction bearing seta and sharp

guarding process; uropod 1 with large distolateral peduncular spine, outer ramus of uropod 2 shorter than

inner ramus, peduncle of uropod 2 with comb-row of

spines on distomedial end; article 2 of outer ramus on

uropod 3 about 30 percent as long as article 1, rectangular, inner ramus with blunt apex armed with 2

spines; pleonal epimera 1-3 each with sharp posteroventral tooth and bulging or straight posterior margins,

epimeron 2 with several ventrolateral setae in stepped

row; pleonite 4 with dorsolateral setule on each side;

telson thin dorsoventrally, of medium length, as broad

as long, lobes scarcely tilted from flat plane, each lobe

with obtuse distolateral apex bearing spine and setule

on medial slope; branchiae ovate and pyriform, with

narrow necks.

HOLOTYPE.—Bishop Museum collections, catalog

number 7275, male, 2.4 mm.

TYPE-LOCALITY.—JLB Hawaii 12, Kaneohe Bay,

Oahu, 3-4 m, corals, algae, corallines, 4 March 1967.

MATERIAL.—JLB Hawaii 6 (1), 12 (7).
144

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 87.—Eriopisella sechellensis upolu, new subspecies, holotype, male, 2.4 mm, JLB

Hawaii 12: a, dorsal edge of urosomites 1-3; b, coxa 3; c, apex of pereopod 4; d, inner plate of

maxilla 1; e, gnathopod 1; /, pereopod 2; g, pereopod 3; h, apex of pereopod 2; i, pereopod 5;

;', apex of pereopod 1; k, gnathopod 2.
NUMBER 3 4 145

FIGURE 88.—EHopisella sechellensis upolu, new subspecies, holotype, male, 2.4 mm, JLB Hawaii

12: a, head; b, apex of medial gnathopod 2; c, telson; d, lower lip; e, apex of maxillipedal palp;

/, part of right mandible; g, maxilla 2; h, apex of peduncle on uropod 2 to show comb-row; i,

pleonal epimera 1-3; j , uropod 3; k, mandibular palp; /, uropods 1-2.
146

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 89.—Maera insignis (Chevreux), female, 6.8 mm, JLB Hawaii 12: a, lateral gnathopod

2; b, palm of medial gnathopod 2. Male, 7.7 mm: c, articles 2-5 of medial gnathopod 2; d,

palm of lateral gnathopod 2; e, body; f, uropod 3; g, telson.
NUMBER 34 147

RELATIONSHIP.—This subspecies resembles closely

the typical subspecies Eriopisella s. sechellensis (Chevreux, 1901), but the hands of the gnathopods are

slightly narrower, less ovate and more rectangular, the

lateral cephalic lobes are more strongly rounded, and

the dactyls of pereopods 3-5 are not bifid.

Eriopisella pusilla Chevreux (see Chevreux and

Fage, 1925), from the Mediterranean Sea, though resembling E. sechellensis in many ways, has a more

strongly hemicircular cephalic lobe and vestigial eye

dot, noncontiguous coxae 5-7, slender body, and imperfectly bifid dactyl on pereopod 5.

Eriopisella capensis (K. H. Barnard, 1916) has article 3 of the mandibular palp shorter than article 2, a

much elongate palm on gnathopods 1-2, more evenly

triangular article 5 on gnathopod 2, rounded pleonal

epimera 1-3, shorter antenna 1, and narrower anterior

process on coxa 1.

Affinities with Eriopisella (Netamelita) cortada (J.

L. Barnard, 1962b) are strong, but E. s. upolu differs

from that Californian species in the presence of article

2 on the outer ramus of uropod 3, the sharp, not

rounded apices of the telsonic lobes, the presence of a

posteroventral tooth on pleonal epimeron 1, the narrow

article 2 of pereopod 5, and strongly extended coxa 1.

DISTRIBUTION.—Hawaiian Islands.

Maera insignis (Chevreux)

FIGURE 89

Elasmopus insignis Chevreux, 1901 pp. 406-412, figs. 24-31.

Maera insignis.—Schellenberg, 1938, pp. 50-52, fig. 24.—

J. L. Barnard, 1955a, pp. 12-13.

MATERIAL.—JLB Hawaii 8 (1), 12 (23), 13 (7), 14

(9+) , 15 (18+), 17 (5+) , 18 (l).Fee l (27).

DISTRIBUTION .—Indo-Pacific.

Maera kaiulani, new species

FIGURES 90, 91

DIAGNOSIS OF FEMALE.—Lateral cephalic lobes softly

quadrangular, anterolateral cephalic corner produced

as narrow sharp tooth, eyes subreniform, orange in alcohol; upper lip lobate as in "Linguimaera" (Pirlot,

1936) ; coxa 1 weakly extended and sharp anteroventrally, anterior coxae lacking posteroventral notch;

gnathopod 1 of medium stoutness, article 6 ovatorectangular, palm oblique and convex, slightly shorter than

posterior margin of article 6, defined by tiny sharp cusp

and 3 spines, article 5 with several medial rows of

diverse setal spines, article 4 lacking posterodistal cusp;

gnathopod 2 occasionally of different sizes on either

side, very large, article 2 with medium-size anterodistal

lobe, article 3 with similar lobe, article 4 weakly extended posterodistally on large gnathopod, sharp on

small gnathopod, article 5 very short, with long tumid

posterior lobe or less tumid on small gnathopod, article

6 very large, subrectangular, slightly expanded distally,

palm nearly transverse, but weakly rounded and defined by large, sharp tooth-bearing spine, tooth next to

small sinus, palm spinose and slightly scalloped and

with one slight excavation near anterior end, dactyl

fitting palm, small gnathopod with smooth but spinose

palm and weak defining tooth; article 2 of pereopods

3-5 changing from narrowly ovatorectangular on

pereopod 3 to subovate on pereopod 5, posteroventral

lobes of medium extent; dactyls of pereopods 1-5 with

strong distal constriction bearing sharp defining corner

and armed with stout seta-bearing flange, sharp nail

with large acute accessory tooth, facial setule, constriction also bearing stout setule appressed to margin of

nail, locking spines straight and slightly pectinate;

pleonal epimera 1-3 with rounded posterior margins

and small to medium-sized posteroventral tooth, ventrally spinose, epimera 1-2 with lateral ridge, pleonites

dorsally smooth; uropod 3 scarcely exceeding extent of

uropods 1-2, rami flat, of medium breadth, apically

truncate and spinose, inner ramus slightly shorter than

outer, latter with thin article 2; telson deeply but not

fully cleft, lobes with strongly fixed medial gap, each

apex deeply and sharply bifid and bearing 2 spines

nearly as long as telson.

MALE.—Gnathopod 2 slightly larger than largest of

female, equal to each other on either side of animal.

JUVENILE DEVELOPMENT, 1.9 MM.—Accessory flagellum with 2 very long and 1 short articles, flagellum of

antenna 2 with 4 articles; inner plate of maxilla 1 with

3 long and 2 short setae, same as adult; gnathopod 2

with large cusp on article 4, article 5 long and poorly

setose, palm oblique, smooth, spinose, defined by slight

protrusion and several spines and setae; article 2 of

pereopods 3-4 slender, that of pereopod 5 basally expanded and tapering distally; inner ramus of uropod 2

reaching only two-thirds along outer ramus of uropod

3; outer ramus of uropod 3 over twice as long as peduncle, inner ramus slightly less than half as long as

outer ramus, thus resembling Melita, uropod 3 exceeding extent of uropods 1-2 by more than length of
148

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 90.—Maera kaiulani, new species, holotype, female, 6.3 mm, JLB Hawaii 2: a, body; b,

palm of medial gnathopod 2 (right member); c, cusp on article 4 of right gnathopod 2; d, palm of

lateral gnathopod 2 (left member). Juvenile, 1.9 mm: e, base of pereopod 4. Male: /, palm of

lateral gnathopod 2.
NUMBER 3 4 149

FIGURE 91.—Maera kaiulani, new species, juvenile, 1.9 mm, JLB Hawaii 2: a, medial gnathopod

2; b, dactyl of pereopod 5; c, uropod 3; d, base of pereopod 5. Male: e, head; /, articles 2-6 of

medial gnathopod 2. Holotype, female, 6.3 mm: g, apex on outer ramus of uropod 3; h, telson;

i, uropod 3; ;, mandibular palp; k, hand of medial gnathopod 1, setae mostly removed; /, articles

2-5 of lateral gnathopod 2; m, dactyl of pereopod 3; n, inner plate of maxilliped, setae removed.
150 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

telson; telson with only middle spine present on each

apex.

JUVENILE, 2.5 MM.—Inner ramus 80 percent as long

as outer ramus on uropod 3 but rami exceeding uropod

2 by more than length of telson, lateral telsonic spine

about half as long as medial, article 2 of pereopod 3

slender, but that of pereopods 4-5 becoming expanded.

YOUNG ADULT, 4.5 MM.—Rami of uropod 3 like

those of adult, with inner ramus nearly as long as

outer, uropod 3 exceeding uropod 2 by half length of

telson.

HOLOTYPE.—Bishop Museum collections, catalog

number 7276, female, 6.3 mm.

TYPE-LOCALITY.—JLB Hawaii 2, off W end of

Pearl Harbor, Oahu, 4-5 m, encrustations from sand

reef, 29 January 1967.

MATERIAL.—Twelve specimens from the typelocality.

REMARKS.—This species has its closest affinites with

M. pacifica Schellenberg, a species known from the

Gilbert, Fiji, and Hawaiian Islands. Maera kaiulani

differs from M. pacifica in the absence of dark ocular

pigment, the deeply and sharply bifid telsonic lobes,

the slightly shorter defining tooth on gnathopod 2, the

scalloped palms, narrower article 2 of pereopods 3-4

in adults, and the absence of a small inner acclivity on

male gnathopod 2 dactyl (mostly absent in M. pacifica

from Hawaii).

The developmental steps undergone by uropod 3

during maturation of this species are remarkable, for

in early stages this species resembles the genus Melita

in having a short inner ramus. The mouthparts of M.

kaiulani and M. pacifica are almost indistinguishable;

the mandibular body and molar are very massive, and

the inner plates of the maxillipeds bear apicolateral

cusps giving the distal end an excavate appearance.

DISTRIBUTION.—Hawaiian Islands.

Maera pacifica Schellenberg

FIGURES 92, 93

Maera pacifica Schellenberg, 1938, pp. 42^*5, figs. 19, 20.—

J. L. Barnard, 1965a, p. 511.

DIAGNOSIS OF MALE.—Lateral cephalic lobes softly

rounded, scarcely quadratiform, anteroventral cephalic

corner produced as narrow sharp tooth; eyes subcircular, bright to brownish-purple in alcohol; upper lip

lobate as in "Linguimaera" (Pirlot, 1936); coxa 1

weakly extended, setose and rounded anteroventrally,

coxa 2 very narrow anteroposteriorly, anterior coxae

lacking posteroventral notch; gnathopod 1 of medium

stoutness, article 6 ovatorectangular, palm oblique and

convex, equal to posterior margin of article 6, defined

by tiny sharp cusp and spine, article 5 with several

medial rows of diverse setal spines, article 4 lacking

posterodistal cusp; gnathopod 2 occasionally of

slightly different size on each side, very large, article

2 with medium-sized anterodistal lobe, article 3 with

similar lobe, article 4 unextended posterodistally, article 5 very short, with long tumid posterior lobe, article

6 very large, subrectangular, slightly expanded distally,

palm transverse, defined by large tooth-bearing spine,

tooth adjacent to small sinus, palm spinose and very

weakly scalloped and with 1 medium-sized excavation

near middle, dactyl fitting palm and occasionally with

inner acclivity fitting excavation; article 2 of pereopods 3-5 ovate, posteroventral lobes broad; dactyls of

pereopods 1-5 with strong distal constriction bearing

sharp defining corner and armed with stout setabearing flange, sharp nail with large acute accessory

tooth, facial setule, constriction also bearing stout setule appressed to margin of nail, locking spines straight

and slightly pectinate; pereopods 4-5 stout, article

4 of pereopod 3 very tumid; pleonal epimera 1-3 with

slightly convex posterior margins and small to mediumsized posteroventral tooth, that of epimeron 1 especially

sharp, epimera ventrally spinose, 1-2 with lateral

ridge; pleonites dorsally smooth; uropod 3 not or

slightly exceeding extent of uropods 1-2, rami flat, of

medium breadth or narrow, apically truncate and

spinose, inner ramus very slightly shorter than outer,

latter with thin article 2; telson deeply but not fully

cleft, with strongly fixed medial gap, lobes with

medium-sized or small medial cusp but not bifid, armed

with 3-5 spines on each lobe, at least 3 spines as long

as telson.

FEMALE.—Gnathopod 2 slightly smaller than in

male, palm lacking incision and slightly more strongly

scalloped; coxa 2 of normal dimensions.

VARIATIONS.—Occasional individuals have one or

two of the six members of pereopods 3-5 wfth narrow

article 2 like M. quadrimana, but most of these appear

to be the result of regeneration of legs accidentally lost.

MATERIAL.—JLB Hawaii 6 (?2), 13 (4), 14 (6),

17 (4). Fee 1 (2).

REMARKS.—The Hawaiian population of this

species has a stronger anteroventral cephalic tooth,
NUMBER 3 4 151

FIGURE 92.—Maera pacifica Schellenberg, female, 5.7 mm, Fee 1: a, b, head; c, apex of outer

ramus on uropod 3; d, coxa 1; e, half of telson; /, palm of gnathopod 2; g, gnathopod 1, setae

removed. Male, 5.0 mm: h, palm of gnathopod 2; i, uropod 3; ;', telson; k, gnathopod 2.

smaller incision adjacent to defining tooth on gnathopod 2 palm, stronger medial telsonic cusps in small

males and large females, longer telsonic spines and a

more tumid article 4 of pereopod 3 than in the materials from southern Polynesia and Micronesia.

Several characters comparable to or slightly different from the figures of M. kaiulani, new species,

are the presence of only 1 defining spine on the palm

of gnathopod 1, the more strongly setose coxae, the

additional sets of spines on articles 5-6 of pereopods

1-2, the similar dactyls of pereopods 1-5, and the

shorter antennae 1-2. Maxilla 2 of M. pacifica in

Hawaii has setae occurring as far down the inner

margin as one-third of its length, the mandibular palp

is more tumid and more setose than in M. kaiulani,

and the maxillipeds are similar between the two species.

DISTRIBUTION.—Southern Polynesia, Micronesia,

Hawaiian Islands.
152 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 93.—Maera pacifica Schellenberg, female, 5.7 mm, Fee 1: a, b, c, pereopods 3, 4, 5; d,

lateral apex of article 5 on pereopod 2; e, pereopod 2; /, uropod 3; g, pleonal epimera 1-3; h,

Maera quadrimana (Dana)

FIGURES 94, 95

Gammarus quadrimanus Dana, 1853, pp. 955-956, pi. 65,

fig. 9.

Maera quadrimana.—Schellenberg, 1938, pp. 45-48, figs. 21-

22.—J. L. Barnard, 1955a, p. 13; 1965a, pp. 511-512, fig.

17.

DIAGNOSIS OF HAWAIIAN MALE.—Lateral cephalic

lobes softly quadrate, anteroventral comer of head

produced as narrow sharp tooth; eyes subcircular,

brownish-purple in alcohol; upper lip anteriorly lobate

as in "Linguimaera" (Pirlot, 1936); coxa 1 weakly extended, setose and rounded anteroventrally, coxa 2 very

narrow anteroposteriorly, anterior coxae lacking posteroventral notch; gnathopod 1 of medium stoutness,

article 6 ovatorectangular; palm oblique and convex,

equal in length to posterior margin of article 6, defined

by tiny cusp and spine, article 5 with several medial
NUMBER 3 4 153

FIGURE 94.—Maera quadrimana (Dana), male, 6.4 mm, Waimanalo, Oahu, 17 April, 1951:

a, b, c, pereopods 3, 4, 5. Male, 3.3 mm, JLB Hawaii 5: d, half of telson; e, palm of lateral

gnathopod 2; f, g, parts of pereopods 4, 5. Female, 5.0 mm, Waimanalo: h, palm of medial

gnathopod 2.
154 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 95.—Maera quadrimana (Dana), male, 6.4 mm, Waimanalo, Oahu, 17 April, 1951: a,

telson; b, palm of medial gnathopod 2; c, uropod 3; d, anterior outline of head; t, coxa 1; /,

gnathopod 2. Male, 3.3 mm, JLB Hawaii 5: g, palm of right gnathopod 2.

rows of diverse setal spines, article 4 lacking posterodistal cusp; gnathopod 2 very large, article 2 with

medium-sized anterodistal lobe, article 3 with similar

lobe, article 4 slightly extended posterodistally, article

5 very short, with long tumid posterior lobe, article 6

very large, subrectangular, slightly expanded distally,

palm transverse, defined by large tooth-bearing spine,

tooth adjacent to sharp sinus, palm divided into 3

diverse truncate, spinose processes separated by 2

sinuses, dactyl fitting palm and lacking acclivity; article 2 of pereopods 3-5 narrowly ovatorectangular, with

slightly extended, angular posterodistal corners, each

with posteroproximal extension; dactyls of pereopods

1-5 with strong distal constriction bearing sharp defining corner and armed with stout seta-bearing flange,

sharp nail with large acute accessory tooth, facial setule,

constriction also bearing stout setule appressed to margin of nail, locking spines straight and slightly pectinate; articles 4-6 of pereopods 3-5 as in M. pacifica,

article 4 of pereopod 3 tumid, of pereopod 4 but especially of pereopod 5 grossly extended posterodistally;

pleonal epimera 1-3 with slightly convex posterior margin and small to medium-sized, sharp posteroventral

tooth, epimera ventrally spinose, 1-2 with lateral

ridges; pleonites dorsally smooth; uropod 3 not exceeding extent of uropods 1-2, rami flat, of medium

breadth, apically truncate and spinose, inner ramus

very slightly shorter than outer, latter with thin article

2 hidden among spines; telson deeply but not fully

cleft, with strongly fixed medial gap, lobes with mediodistal corner unproduced, armed with 4—5 spines,

mostly spines as long as or longer than telson.

FEMALE.—Gnathopod 2 slightly smaller than in

male, palm slightly scalloped and convex, bearing only

incision next to defining tooth; coxa 2 like that of male.

SMALL MALE OF JLB HAWAII 5.—With left male

gnathopod 2 like that originally figured by Dana and

Schellenberg in which palmar teeth all projecting

equally except defining tooth and largest middle

palmar teeth only about twice (instead of thrice) as

broad as smallest; pereopods 4-5 like those of M. pacifica and not the original M. quadrimana, but pereopods 4-5 slightly more slender than in M. pacifica;

telson with strong medial cusp on each lobe; inner
NUMBER 34 155

ramus of uropod 3 only two-thirds as long as outer

ramus, right gnathopod 2 small and plain as if undergoing regeneration.

YOUNG INDIVIDUAL OF JLB HAWAII 14.—Rami of

uropod 3 elongate, equal to each other in length.

MALE OF KAWELA BAY.—Article 2 of gnathopod 2

with anterior spines instead of setules, pereopod 5 like

pereopod 4 of M. pacifica, with broadly rounded posteroventral corner of article 2.

MATERIAL.—JLB Hawaii 5 (3), 14 (1). Waimanalo, Oahu, 17 April 1951 (4). Kawela Bay, Oahu,

2-7 September 1948 (4).

REMARKS.—The material at hand is insufficient to

link the small form of JLB Hawaii 5, with "normal"

gnathopods and abnormal pereopods, to the large form

of Waimanalo having abnormal gnathopods and

normal pereopods.

DISTRIBUTION.—Micronesia, southern Polynesia,

Hawaii.

Maera serrata Schellenberg

FIOURES 96, 97

Maera inaequipes serrata Schellenberg, 1938, pp. 41-42, fig.

18.—J. L. Barnard, 1965a, p. 510.

NOMENCLATURE.—Schellenberg described this taxon

as a subspecies of M. inaequipes, but it and the latter

are sympatric and should either be recognized as conspecific phenotypes/ecotypes or recognized as distinct

species. Though one may presume one phenotype of

a species and not the other may exist in a geographic

area, the absence so far of any record typical of M. i.

inaequipes in Hawaii suggests that M. serrata is a distinct species occurring both in southern Polynesia and

in Hawaii. Because the M. serrata of Hawaii probably

has a distinct character worthy of subspecific appellation, a quadrinomial would be required to signal that

distinction. For the present, I hesitate to establish the

Hawaiian population as subspecifically distinct from

the population in southern Polynesia, but such trinomial procedure would now be simplified in that M.

serrata is elevated to full specific rank. If the males

described below are fully terminal then the Hawaiian

population should be recognized nomenclaturally.

DIAGNOSIS OF MALE.—Lateral cephalic lobes softly

rounded, anteroventral cephalic tooth sharp and thin;

eyes subcircular, pale purple cores in alcohol; upper

lip lobate as in "Linguimaera" (Pirlot, 1936) ; coxa

1 sharply but weakly extended anteroventrally, coxa

360-034 O—70 ,11

2 narrow anteroposterioriy, anterior coxae lacking

posteroventral notch; gnathopod 1 of medium stoutness, article 6 ovatorectangular, palm oblique and

convex, palm slightly shorter than posterior margin of

article 6, defined by sharp acclivity and spine, article

6 with several medial rows of diverse setal spines, article 4 lacking posterodistal cusp; gnathopod 2 very

large, article 2 with large downward pointing process

of medium breadth, article 3 weakly lobate, article 4

sharply extended posterodistally, article 5 very short,

with long tumid posterior lobe, article 6 very large,

subrectangular, slightly expanded distally, palm transverse, defined by large tooth bearing spine, tooth adjacent to small sinus, palm spinose and weakly scalloped

and bearing weak excavation in middle, dactyl fitting

palm and bearing strong inner acclivity fitting excavation; article 2 of pereopods 3—5 narrowly ovatorectangular, posterior margins nearly straight, posteroventral

lobes narrow, short, sharply cuspidate; dactyls of pereopods 1-5 with strong distal constriction bearing

sharp defining corner and armed with thin seta, sharp

but stunted nail bearing small acute accessory tooth,

facial setule, constriction also bearing stout setule partially appressed to margin of nail, locking spines small,

straight or slightly curved; pleonal epimera 1-3 with

slightly convex posterior margins, each with sharp posteroventral tooth, ventrally spinose, epimeron 2 with 1

accessory serration and 1 notch; pleonites dorsally

smooth; uropod 3 slightly exceeding extent of uropods

1-2, rami flat, narrow, apically truncate and spinose,

equal to each other in length, outer with tiny article

2 hidden among spines; telson deeply but not fully

cleft, with strongly fixed medial gap, lobes thin, apically

bifid, lateral cusp longer than medial, each lobe with

2 long spines not fully as long as telson and 1 short

spine; posterior margin of urosomite 3 very sharp in

comparison to other species of Hawaiian Maera.

FEMALE.—Gnathopod 2 similar to that of male but

excavation in middle weaker.

JUVENILE.—Article 4 of gnathopod 2 with sharp

posterodistal cusp; pleonal epimeron 3 with sharp

posteroventral corner and 1 or 2 weak posterior serrations marked with setule; rami of uropod 3 very thin

and elongate but inner shorter than outer; each telsonic lobe with only 2 spines, one spine as long as

telson.

MATERIAL.—JLB Hawaii 12 (5), 13 (3), 14 (3),

15(2).
156

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 96.—Maera serrata Schellenberg, female, 5.4 mm, JLB Hawaii 12: a, b, c, pereopods 3,

4, 5; d, inner plate of maxilliped; e, head; /, palm of gnathopod 2; g, dactyl of pereopod 3; h,

gnathopod 1. Male, 4.3 mm, JLB Hawaii 13: i, palm of gnathopod 2. Juvenile, 2.9 mm, JLB

Hawaii 12: j , pleonal epimeron 3.

REMARKS.—Male gnathopod 2 of Schellenberg's Maera species A

material has a much deeper excavation in the middle

of the palm separating off a strong subquadrate tooth

between it and the sinus near the defining tooth.

DISTRIBUTION.—Southern Polynesia, Micronesia,

and the Hawaiian Islands.

FIGURES 98, 99

A single female has affinities with M. hamigera Haswell as represented by J. L. Barnard (1965a) from

Micronesia, but gnathopod 2 is so much better de-
NUMBER 3 4 157

FIGURE 97.—Macro, serrata Schellenberg, female, 5.4 mm, JLB Hawaii 12: a, posterior corner of

urosomite 3 (upside down); b, c, d, posteroventral corners of article 2 on pereopods 3, 4, 5; e,

uropod 3; /, telson. Juvenile, 2.9 mm: g, gnathopod 2; h, uropod 3; i, half of telson.

veloped than in the Micronesian population that a

name is reserved until the male can be found and

described.

DIAGNOSIS.—Head with normal anteroventral

notch; eye narrowly triangular, pigment in alcohol

blood red but dense and appearing very dark from

distance; coxa 1 sharply extended forward and slightly

attenuate at anteroventral corner; anterior coxae with

posteroventral notch; gnathopod 1 slender, article 6

thin, subrectangular but slightly expanded distally,

palm oblique and defined on medial surface with 5

spines, medial surface of slightly swollen article 5 with

many rows of diverse setal spines; article 4 with posterodistal cusp; gnathopod 2 elongate, slender, article

4 with posterodistal cusp, 5 with broad, setose posterior

margin, article 6 slightly tapering distally, palm short,

oblique, deeply excavate proximally and bearing protrusion with large spine near distal end, defined by
158

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 98.—Maera species A (cf. M. hamigera), female, 6.5 mm, JLB Hawaii 5: a, head; b, c,

d, pereopods 3, 4, 5, in whole or part; e, uropod 3; f, article 2 on outer ramus of uropod 3 with

setae apically cut off; g, telson; h, mandibular palp; i, coxa4;j, pleonal epimera 2, 3.

small cusp and medium spine, bearing large and medium medial spine and medium lateral spine on proximal

portion of palmar hollow, dactyl slender, fitting palm;

article 2 of pereopods 3-5 rectangular to subrectangular, of medium expansion, posterior margins with

weak but regular serrations, posteroventral lobes shallow, pereopods otherwise of normal thinness; dactyls

of all pereopods constricted distally, apical nail marked

with suture, dactyl bearing one long, and one short

setal-spine at constriction and one facial seta; pleonal

epimera 1-2 weakly bulging and bearing small, posteroventral cusp, epimeron 2 with one posterior serration, epimeron 3 nearly straight posteriorly and deeply

and sharply serrate; pleonites dorsally smooth; uropod

3 elongate and greatly exceeding extent of uropods

1-2, rami flat, narrow, apically truncate, spinose, outer

ramus with small article 2 bearing 1 short and 2 long

setae; telson long, fully cleft, each lobe with 2 spines
NUMBER 3 4 159

FIGURE 99.—Maera species A (cf. M. hamigera), female, 6.5 mm, JLB Hawaii 5: a, medial

gnat hop od 1; b, apex of gnathopod 2, medial; c, pleonal epimeron 3 and urosome; d, apex of

pereopod 5; e, medial gnathopod 2.

on medial margin, distal spine at deep notch, each Maera species

apex deeolv excavate, with sharo bordering CUSDS and

B

apex deeply excavate, with sharp bordering cusps and

bearing 2 spines and setule, lateral margins each with

2 setules and 1 basal spine.

MATERIAL.—JLB Hawaii 5 (ovigerous female, 6.5

mm).

FIGURE 100

A single female differs from other species of Hawaiian Maera notably in the absence of a lateral cephalic

notch and in other characters, but the lack of a male
160

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 100.—Maera species B, female, 3.8 mm, JLB Hawaii 17: a, uropod 3; b, pleonal epimera

1-2 and part of 3; c, half of telson; d, apex of pereopod 1; e, defining corner of gnathopod 2 palm;

/, medial gnathopod 2; g, coxa 1; h, medial gnathopod 1; x, j , head; k, apex of outer ramus on

uropod 3; I, pleonal epimeron 3 and urosome.
NUMBER 34

161

prevents appellation; affinities may be with M.

othonides Walker.

DIAGNOSIS.—Head lacking normal anteroventral

notch, lateral cephalic lobe broadly convex and produced as cusp at anteroventral corner; eye irregularly

subcircular, brownish-purple in alcohol; coxae short,

coxa 1 obtusely extended anteroventrally, anterior

coxae lacking posteroventral notch; gnathopod 1 of

medium stoutness but long, article 6 ovate, palm

oblique and defined by slight acclivity and marked with

pair of submarginal medial spines, article 5 with several

medial rows of diverse setal spines, article 4 lacking

posterodistal cusp; gnathopod 2 elongate, of medium

stoutness, article 4 with posterodistal cusp, 5 with

medium-broad, poorly setose posterior margin, article

6 broad, margins nearly parallel, palm short, oblique,

evenly convex, armed with numerous short spines, defined by weak cusp and stout submarginal spine, dactyl

slender, fitting palm; pereopods 3-5 unknown, missing;

dactyls of pereopods 1-2 constricted distally, apical

nail marked with suture, dactyl bearing one seta at

sutural margin, one facial setule and one proximosutural seta in notch; pleonal epimera 1-3 weakly bulging posteriorly, each with small posteroventral cusp,

1-2 with lateral ridge; pleonites dorsally smooth;

uropod 3 elongate and greatly exceeding extent of

uropods 1-2, rami flat, narrow, apically truncate and

spinose, but outer ramus with small protrusion representing vestigial article 2 coalesced with article 1; telson

long, fully cleft, medial and lateral margins of each

lobe smooth, apices deeply excavate, points sharp, excavation with one spine, lateral apex with spine, setule

between spines.

MATERIAL.—JLB Hawaii 17 (female, 3.8 mm, bearing one giant embryo filling brood pouch).

Melita appendiculata (Say)

FIGURES 101, 102

Gammarus appendiculatus Say, 1818, p. 374.

Gammarus fresnelii Audouin, 1826, p. 93, pi. 11, fig. 3.

Melita fresneli.—J. L. Barnard, 1955a, pp. 13-14 [with references].

MATERIAL.—J. L. Barnard Hawaii 3 (1), 5 (1), 6

(3), 13 (42).

DISTRIBUTION.—Cosmopolitan in warm seas.

Melita pahuwai, new species

FIGURES 103, 104

DIAGNOSIS OF MALE.—Head with deep lateral slit

below subnasiform lateral lobes; eyes narrowly ovatopyriform, retaining brownish-purple pigment in alcohol; flagellum of second antenna 1.3 times as long

as article 5; coxa 1 not anteroventrally produced;

article 6 of gnathopod 1 distally expanded, anterior

margin swollen, palm short and oblique, bearing 3

stout medial spines and 5-6 thin lateral spines, then

distally excavate for reception of large dactylar bulge,

beyond that produced again, dactyl thick, bulging

basally, fitting palm, distally unguiform and bearing

accessory setal spine; gnathopod 2 with unlobed articles 2-3, article 4 with posterodistal cusp, article 5

with narrow, setose posterior lobe, article 6 elongate,

expanding distally like young male M. palmata,

dactyl longer than palm and overriding medial

face, palm short and scarcely oblique, but rounding onto posterior margin of article 6 and minutely

spinose, medial face of hand with axial channel

toward posterior side, channel defined anteriorly by

sharp ridge bearing distal spines and by shorter

sharp ridge proximally; channel filled with pectinoplumose setae; all pereopodal dactyls with distal double constriction and nail, with stout marginal

seta and small facial setule, locking spines simple; pereopods 3-5 of regular form but relatively short and

thick, dactyls also thick; outer ramus of uropod 3 with

1 article only; telson with sharp apices, lobes gaping,

each lobe with 2 medial spines, one long subterminal

medial spine guarded by sharp cusp, small apicolateral

spine, and 3 basolateral spines, plus setules subapically;

pleonal epimera 1-3 each with small posteroventral

tooth, no other serrations; only urosomite 2 with small

lateral cusp guarding 2 spines, cusp obtusely angular.

Mouthparts like M. palmata as figured by Sars (1895,

pi. 179).

FEMALE.—Gnathopod 1 with simple bulging palm,

unbulging article 6, basally unthickened dactyl; gnathopod 2 small, palm evenly oblique and defined by 2

spines on medial face and bearing several slender

marginal spines; posteroventral teeth of epimera

blunter than in male; cusp on urosomite 2 sharply

attenuate, bearing 2 spines (or 1 in juvenile); telson

with few spines in slightly different arrangement (see

figures); coxa 6 with anterior hook bearing gland.

HOLOTYPE.—Bishop Museum collections, catalog

number 7277, male, 4.5 mm.
162

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 101.—Melita appendiculata (Say), male, 4.8 mm, JLB Hawaii 6: a, lateral pleon; b,

body, c, dorsal pleon; d, inner ramus of uropod 3, shaded; e, medial side of apex of gnathopod 2.
NUMBER 34 163

FIGURE 102.—Melita appendiculata (Say), female, 4.7 mm, JLB Hawaii 6: a, articles 2-3 of

pereopod 5; b, anteroventral corner of head; c, medial gnathopod 1. Male, 4.8 mm, JLB Hawaii

6: d,e, dactyls of pereopods 3, 2; f, anteroventral corner of head; g, h, maxillae 1, 2; i, posteroventral corner of article 2 on pereopod 3; j , right gnathopod 2.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, algae, rocks, corals, 24

May 1967.

MATERIAL.—JLB Hawaii 17(3), 18(2).

RELATIONSHIP.—Closely related to Melita koreana

Stephensen (1944), a species with similar male gnathopod 2 palm lacking a distinct channel on medial face

of hand but strongly setose thereon. Melita koreana

has 3 spines but no cusp on each dorsolateral side of

pleonite 4, and male gnathopod 1 dactyl is a stout,

short coniform piece not reaching the end of the palm,

the latter simply convex. The telson lacks mediobasal

spines and articles 3-6 of pereopods 3-5 are more

slender than in M. pahuwai.

Melita laevidorsum Stephensen (1944), from Japan,

also has 3 spines and no cusp on each side of pleonite

4, no basomedial spines on the telsonic lobes, slender

pereopods 3-5, and a longer flagellum on antenna 2

than does M. pahuwai, but gnathopod 1 is more like

that of the Hawaiian species than is gnathopod 1 of M.

koreana. Gnathopod 2 of the male, possibly not fully

developed, has a simple, sloping, spinose palm with

many medial setae and a spinose medial denning ridge.

Melita pahuwai has its closest affinities with M. zeylanica Stebbing (1904) from Ceylon. The latter has a

similar gnathopod 2, apparently similar ornaments on

pleonite 4, has medial spines on the telsonic lobes but

has larger coxae than does M. pahuwai, and male

gnathopod 1 article 6 is evenly rectangular, not distally

expanded and has a simple palm like the female of M.

pahuwai. Article 4 of pereopods 3-5, especially pereopod 5 of M. zeylanica, is much stouter than in M. pa-
164

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 103.—Melita pahuwax, new species, holotype, male, 4.5 mm, JLB Hawaii 17: a, head;

b, apex of gnathopod 2, medial; c, gnathopod 1; d, apex of pereopod 5; e, gnathopod 2; /, half

of telson; g, pleonal epimera 1-3; h, apex of gnathopod 1; i, uropods 1-2 and part of urosome.

Female, 3.7 mm: ;, half of telson; k, coxa 6.
NUMBER 34 165

FIGURE 104.—Melita pahuwai, new species, holotype, male, 4.5 mm, JLB Hawaii 17: a, apex

of gnathopod 1, medial; b, peduncle and accessory flagellum of antenna 1; c, pereopod 2; d,

ornamentation of pleonite 4; e, f, pereopods 3, 5, whole or part; g, coxa 6. Female, 3.7 mm:

h, apex of gnathopod 1; i, gnathopod 1, setae removed; ;', uropod 3; k, medial gnathopod 2,

setae removed; /, posteroventral corners of pleonal epimera 1-3.
166

huwai and article 6 is very short on pereopod 5 but not

on pereopods 3-4.

The west American representatives of M. nitida

Smith (Shoemaker, 1935b) have long coxae, a simple

palm on male gnathopod 1 and a weak, poorly setose

gnathopod 2, palm very oblique, but with dactyl overriding the surface to close against a spinose locking

ridge. Pleonite 4 has only 3 dorsolateral spines on each

side.

DISTRIBUTION.—Hawaiian Islands.

Nuuanu, new genus

DIAGNOSIS.—Eyes weakly developed; lateral cephalic lobe with deep notch; antenna 1 geniculate

between articles 1 and 2, accessory flagellum 3-articulate; mandibular palp article 3 subfalcate, strongly

setose and longer than article 1, molar triturative;

lower lip apparently lacking inner lobes, inner plates

of maxillae 1-2 densely setose medially, palp of maxilla

1 with 2 articles; maxillipeds basic; gnathopod 1 small

and slender but subchelate, palm nearly transverse;

gnathopod 2 of male large (female unknown); coxa

4 strongly excavate posteriorly; second articles of

pereopods 3-5 with strong castelloserrations; metasome with weak dorsal teeth, urosome with a few dorsal

spines not in bundles, urosomites separate; uropod 3

short, scarcely extending beyond uropods 1-2, inner

ramus short and scalelike, outer with 2 articles, article

2 short; telson short, deeply cleft.

TYPE-SPECIES.—Nuuanu amikai, new species.

RELATIONSHIP.—This genus belongs in the EriopisaMelita group with shortened and scalelike inner ramus

of uropod 2. It differs from Eriopisella Chevreux and

Netamelita J. L. Barnard in the medially setose inner

plates of the maxillae, from Melita Leach in the short

uropod 3, from Melitoides Gurjanova and Netamelita

in the absence of inner lobes on the lower lip, the 2-

articulate outer ramus of uropod 3, and from Eriopisa

Stebbing in the short uropod 3 and presence of weak

dorsal sculpture. The geniculate antenna 1, with article

2 bending strongly past the lateral extension of article

1, is also a character distinguishing Nuuanu from other

Gammaridae mentioned above, the telson apparently

is unusually short for this group, and the conspicuous

castelloserrations of pereopods 3-5 are signficant. The

3-articulate accessory flagellum exceeds that of the

other genera except Melita by at least 1 article.

The dorsal sculpture, though weak, and the long

coxae suggest closest affinities with Melita more than

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with Eriopisa and mandibular palp article 3 is scarcely

more swollen than in M. palmata (Montagu) (Sars,

1895, pi. 179).

Nuuanu amikai, new species

FIGURES 105, 106

DIAGNOSIS.—With the characters of the genus.

DESCRIPTION.—Lateral cephalic lobe broad, strongly

squared off and bearing deep notch, eyes composed

of about 9 widely scattered ommatidia weakly stained

orange in alcohol and formaldehyde; article 5 of

gnathopod 1 elongate, palm defined by 2 series of

stout spines, one series apically bifid, dactyl stout, short,

curved; gnathopod 2 with sharp cusp on article 4, article 5 cup-shaped, posterior lobe of medium breadth,

article 6 large and elongate, palm scarcely defined from

setose posterior margin of article 6, dactyl less than

half as long as article 6, slipping over finely setose

medial face of article 6 and locked by mediodistal

ridge bearing 2 spines, lateral palmar margin protuberant and spinose; article 2 of pereopods 3-4

similar, weakly concave posteriorly, articles 4-6 of pereopod 4 stouter than on pereopod 3, article 2 of pereopod 5 very broad, posteriorly convex and slightly

concave ventrally, pereopod 5 shorter than pereopod

4; uropod 2 with distomedial comb-row on peduncle;

pleonal epimeron 1 with sharp posteroventral corner,

epimeron 2 strongly extended ventrally and posteroventrally attenuate, epimeron 3 of more regular form,

with straight posterior margin and weak posteroventral

tooth; metasomites 1-2 each with small posterodorsal

tooth, metasomite 3 smooth, urosomite 1 with weakly

frilled dorsal margin, urosomites 2-3 each with 1

dorsolateral spine and urosomite 3 with sharp but small

posterolateral wings.

HOLOTYPE.—Bishop Museum collections, catalog

number 7278, male, 4.2 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 3, off Ewa Beach,

Oahu, 18 m, Pocillopora, bryozoans, 29 January 1967.

Family ISAEIDAE (=PHOTIDAE)

Chevalia aviculae Walker

FIGURE 107

Chevalia aviculae Walker, 1904, pp. 288-290, pi. 7, fig. 50,

pi. 8, fig. 50.—J. L. Barnard, 1962a, pp. 17-20, fig. 5 [with

references].

MATERIAL.—JLB Hawaii 2 (5), 3 (17), 5 (8), 6

(10), 8 (150+),1 5 (3), 17 (3).
NUMBER 3 4 167

FIGURE 105.—Nuuanu amikai, new genus, new species, holotype, male, 4.2 mm, JLB Hawaii 3:

a, body; b, mandible; c, mandibular palp; d, maxilla 2; e, maxilliped; /, inner plate of maxilliped;

g, half of lower lip; h, telson; i, upper lip.
168

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 106.—Nuuanu amikai, new genus, new species, holotype, male, 4.2 mm, JLB Hawaii 3:

a, head and epistome-labral complex below (shaded); b, maxilla 1; c, part of pereopod 4; d,

pereopod 5;e,f, gnathopod 1; g, h, gnathopod 2; i, medial antenna 1; ;, uropod 3; k, urosome and

pleonal epimeron 3.
NUMBER 34 169

FIGURE 107.—Chevalia aviculae Walker, male, 3.1 mm, JLB Hawaii 3: a, body; b, mandibular

palp; c, head; d, raxni of uropod 1; e, uropod 3; /, palm of right gnathopod 2; g, telson; h, apex of

pereopod 5; i, right gnathopod 1, setae removed;;, urosome.
170

REMARKS.—The right mandibular molar has a large

smooth accessory flake and a long seta. Eyes are red in

formaldehyde and clear in alcohol. The palm of

gnathopod 2 is transverse and nearly protuberant in

Hawaiian individuals.

DISTRIBUTION.—Circum tropical.

Gammaropsis Liljeborg (—Eurystheus Bate)

REMARKS.—Members of this genus in Hawaii demonstrate a meeting ground of cold-temperate and

tropical pathways. Gammaropsis atlantica, G. afra,

and G. pokipoki represent the tropical members having

an article 2 on the outer ramus of uropod 3 and the

absence of a molarial flake. Gammaropsis alamoana,

with obvious derivation from the cold-temperate Pacific G. thompsoni Walker, lacks article 2 on the outer

ramus of uropod 3 like other cold-temperate species

(of the North Atlantic Ocean), but has a molarial flake

on the right mandible. Gammaropsis pali represents the

intermediate condition, a molarial flake combined with

an article 2 on the outer ramus of uropod 3, and the

tropical G. digitata from Canton Island, the reverse

condition.

Gnathopod 1 usually has a defining spine except in

G. pali, and uropod 2 lacks a ventral peduncular tooth

between the rami except for a small tooth in G. digitata

and a large one in G. haleiwa.

Gammaropsis afra Stebbing

FIGURE 108

Gammaropsis afra Stebbing, 1888, p. 1097, pi. 113.

Eurystheus afer.—Stebbing, 1906, p. 612.—?K. H. Barnard,

1916, pp. 249-250, pi. 28, fig. 11.—K. H. Barnard, 1937,

pp. 165-166, fig. 12.—Pillai, 1957, pp. 55-56, fig. 13.—

J. L. Barnard, 1961, pp. 113-114, fig. 79.

DIAGNOSIS (relative to forms of G. atlantica).—

Lateral cephalic lobes poorly projecting, eye weakly

reniform, but not distinctly lageniform or ovate; accessory flagellum thick and 3-articulate in adults (1.9

mm) ; gnathopod 2 with palmar axis less oblique than

in form A of G. atlantica, with small subquadrate palmar concavity, large defining tooth, palmar bulge with

2 humps, dactyl especially short and thick compared

with form B of G. atlantica, and proximal inner margin with 2 small conical processes, article 6 about 1.75

times as long as broad, articles 2 and 6 with very few

setae. Female gnathopod 2 of the fully developed conSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dition as in form A of G. atlantica and spine of palm

especially enlarged.

COLOR IN 2-DAY FORMALDEHYDE.—Body white, eyes

brown.

MATERIAL.—JLB Hawaii 5 (15). Devaney 1 (19).

Hybrid or form intermediate with G. atlantica: Eye

slightly more reniform than in typical individuals; palm

much less oblique than any form of atlantica or afra,

concavity narrowest, palmar bulge quadrate, proximal

inner margin of dactyl with low quadrate process; color

as in form B of G. atlantica; body length reaching 4 mm

in contrast to 2.2 mm maximum size of G. afra.

MATERIAL.—JLB Hawaii 5(10).

REMARKS.—The metasome, urosome, uropods, and

telson resemble those of G. atlantica (Figure 111), but

owing to the small size of the adults of the typical form

the spines are less frequent. Thus, the outer ramus of

uropod 3 lacks the subterminal marginal spine and

article 2 of the outer ramus is relatively much larger

than in G. atlantica or in the "hybrid" form. The outer

ramus of uropod 2 is shortened as it is on uropod 1 of

G. atlantica and has only 1 marginal spine; the outer

ramus of uropod 2 has 2 marginal spines. Mouthparts

are less setose than in G. atlantica.

Gammaropsis afra and G. atlantica are linked

through form B of G. atlantica and the hybrid form

here assigned for convenience to G. afra; gnathopods

are almost perfectly intergraded but the hybrid form

has them more extremely coarsened than in any other

form and G. atlantica form A has them occurring in the

most graceful condition.

DISTRIBUTION.—East Atlantic Ocean, eastward to

Hawaii.

Gammaropsis alamoana, new species

FIGURES 109, 110

DIAGNOSIS.—Lateral cephalic lobes weakly extended, apex in adults obtaining sharp cusp; eyes ovate

in young, pigment core small, clear ommatidia fully

surrounding pigment, eyes subreniform in adult, pigment core large, posteriorly lacking clear marginal ommatidia; epistome with long acute cusp; right mandibular molar with flake; inner plate of maxilla 1

conicovermiform, bearing setular wisps but otherwise

naked; gnathopod 1 with article 6 subovate, palm

poorly defined; gnathopod 2 with article 5 broadly

conical and elongate, increasingly so in adults, lacking

posterior lobe, posterior margin flat, anterior margin
NUMBER 3 4 171

FIGURE 108.—Gammaropsis afra Stebbing, male, 1.9 mm, JLB Hawaii 5: a, b, head; c, d, e,

pereopods 1, 3, 5; /, gnathopod 1; g, h, gnathopod 2, lateral and medial views. Male hybrid, 2.5

mm: i, head. Male, hybrid, 3.0 mm: ;, gnathopod 2. Female, 2.1 mm: k, gnathopod 2.

360-034 O—70 12
172

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 109.—Gammaropsis alamoana, new species. Heads and eyes: a, b, male, holotype, 3.2

mm, JLB Hawaii 13; c, male, 3.6 mm; d, e, female, 2.2 mm, JLB Hawaii 8. Male holotype: /,

gnathopod 1; g, right mandibular molar, arrow pointing to flake; h, uropod 3; i, coxa 3; ;, telson

and dorsal view of urosomites 1-3; k, pleon. Male, 3.6 mm: /, lateral view of telsonic apex; m,

epistome; n, pereonites 5-7 and pleonites 1-4.
NUMBER 3 4 173

FIGURE 110.—Gammaropsis alamoana new species, male, 3.6 mm, JLB Hawaii 13: a, gnathopod

2, medial; b, gnathopod 2; c, gnathopod 1; d, palmar sculpture of gnathopod 1. Holotype, male,

3.2 mm, e, gnathopod 2, medial; /, g, h, pereopods 3, 4, 5; i, inner plate of maxilla 1, shaded; ;',

pereopod 2; k, I, gnathopod 2. Female, 4.5 mm: m, n, gnathopod 2. Juvenile, 1.5 mm: o,

gnathopod 2.
174

lacking spines, palm oblique with strong defining tooth,

excavation flat and shallow, protrusion near finger

simple in juvenile, acquiring one or two major protuberances in adults, dactyl fitting palm in juveniles

and females, but becoming shortened in adult males,

large spine of palm lost in adult males; article 2 of

pereopods 3-5 ovate, coxa 7 becoming enlarged enormously in terminal male, coxa eventually hiding

pleonal epimeron 1; epimera 1-3 with evenly convex

posterior margins, small incision and tooth at posteroventral corners; outer ramus of uropod 3 lacking article

2; telson nearly square, dorsoventrally excavate; urosomite 1 with broad shallow dorsal excavation bordered

on each side by blunt protrusion, each side of excavation bearing 1 large seta and 2 minute setules, urosomite 2 with slight dorsal excavation but lateral

protrusions obsolescent, 1 seta present on each side.

COLOR IN 2-DAY FORMALDEHYDE.—Eyes blackbrown, body almost clear white.

HOLOTYPE.—Bishop Museum collections, catalog

number 7279, male, 3.2 mm.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

Oahu, 3-4 m, corals, algae, corallines, 8 April 1967.

MATERIAL.—JLB Hawaii2 (6), 3 (1), 5 (3), 8 (26),

10 (13), 12(1), 13 (10), 14 (5).

RELATIONSHIP.—This species is a cognate of Gammar opsis thompsoni (Walker) of the cold and warmtemperate northeastern Pacific Ocean (Shoemaker,

1931, 1942, 1955 and J. L. Barnard, 1959). Gammaropsis alamoana differs from its cognate in the weaker

bulge of epimeron 3, the shorter spines of the telson,

the weaker dorsal armaments of the urosome, the

presence of a distinct defining cusp on gnathopod 2 of

females, and the shortening of the dactyl on gnathopod

2 of terminal males.

DISTRIBUTION.—Hawaiian Islands.

Gammaropsis atlantica Stebbing

FIGURES 111, 112, 113

Gammaropsis atlantica Stebbing, 1888, p. 1101, pi. 114.

Gammaropsis zeylanicus Walker, 1904, pp. 282-283, pi. 6,

fig. 41; 1909, p. 339.

Gammaropsis Gardineri Walker, 1905, pp. 929-930, pi. 88,

figs. 11-14, 16, 17.

Eurystheus atlanticus.—Stebbing, 1906, p. 611; 1908, pp.

86-87, pi. 40B.—Tattersall, 1922, pp. 10-11, pi. 1, figs.

17-20.—K. H. Barnard, 1937, p. 164.—Pillai, 1957, pp.

56-57, fig. 14.—J. L. Barnard, 1965a, pp. 534-535, fig. 29.

DIAGNOSIS OF MALE FORM A.—Lateral cephalic

lobes projecting to medium extent, eye lageniform in

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

juveniles and adults from 1.8 to 4.5 mm in body length,

juveniles of 1.2 mm with vertically oriented ovate eye

lacking dorsoposteriorly extended ommatidia; accessory flagellum thin and 6-articulate in adults; gnathopod 2 with palmar axis oblique, large palmar concavity

near defining corner, with large spine on medial surface

near concavity, palmar bulge near finger hinge forming 3 low humps each minutely undulate, palmar

defining corner scarcely produced, dactyl slightly

shorter than palm, article 6 of gnathopod 2 about 1.5

times as long as broad; terminal male with heavy

setation on anteromedial margins of articles 2 and 6.

FEMALE.—Gnathopod 2 fully developed as in Figure

112/, with crenulate palm and slight process at defining corner. Juveniles lacking process at comer.

COLOR IN 2-DAY FORMALDEHYDE.—Eyes red or

orange, head purple or white but bleaching white in

alcohol.

DIAGNOSIS OF MALE FORM B.—Lateral cephalic lobes

projecting very strongly, coniform, eyes in youthful

members (3.1 mm) ovate horizontally and confined to

ocular lobe, in terminal adults ommatidia proliferating

in dorsoposterior direction toward forehead and eye

becoming lageniform; accessory flagellum thick and

not exceeding 4 articles; gnathopod 2 with palm less

oblique than form A, concavity shallower and narrower

but defining cusp much larger, palmar bulge with 1

or 2 humps only, articles 2 and 6 with very few setae.

FEMALE.—Frequently maintaining juvenile gnathopod 2 into old age (4.0 mm), but accessory flagellum

slender and often 5-articulate.

COLOR.—Same as form A but majority of specimens

maintaining purple-pink head and anterior body

stain in alcohol.

MATERIAL.—JLB Hawaii 2 (14), 3 (23), 5 (6),

6 (10), 8 (1), 10 (20), 12 (4), 13 (18). Makapuu

Point, Oahu, April 1967, Zoanthus (1), collectors

Ralph Bowers and Dr. Jerry Welch.

DISCUSSION.—Form B is like materials reported on

by Walker (1905), Pillai (1957), and J. L. Barnard

(1965a), whereas form A represents material in the

other references. If the two forms are indeed phenotypes of a single species, they are readily distinct in

Hawaii in the male at all stages. Young males and

females of form B are recognizable by the horizontally

ovate eyes, and adult males are recognizable by the

gnathopod 2 palm and the lack of a dense brush of

setae on article 2. Form A appears to be the more

advanced in that juvenile eyes are vertically ovate
NUMBER 3 4 175

n

FIGURE 111.—Gammaropsis atlantica Stebbing, male, 4.0 mm, JLB Hawaii 10: a, anterior end

of body; b, palp article 3 of maxilliped, sockets with setae removed; c, head; d, pleonal epimera

1-3, left to right; e, right lateral view of telson; /, right lateral view of epistome and upper lip

complex; g, obverse view of right mandibular molar; h, maxilla 2; i, uropod 3; ;', inner plate

of maxilla 1. Heads, k, I, m, n: k, male 3.1 mm, form B, Hawaii 2; /, male 4.4 mm, form B,

Hawaii 2; m, intersex, 3.4 mm, Hawaii 3; n, juvenile, 1.2 mm, Hawaii 5. Female, 4.5 mm, Hawaii

10: o, urosome.
176

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 112-—Gammaropsis atlantica Stebbing, male, 4.0 mm, JLB Hawaii 10: a, b, c, d,

pereopods 1, 3, 4, 5; e, telson. Female, 4.5 mm: /, medial gnathopod 2. Juvenile, 2.6 mm: g,

gnathopod 2. Juvenile, 1.8 mm: h, gnathopod 2. Male, 4.4 mm, Hawaii 2, form B: i, gnathopod 2.
NUMBER 3 4 177

FIGURE 113.—Gammaropsis atlantica Stebbing, male, 4.0 mm, JLB Hawaii 10: a, b, gnathopod

2, lateral and medial views; c, gnathopod 1; d, medial gnathopod 2. Male, 3.1 mm, form B,

JLB Hawaii 2: e, gnathopod 2, setae removed. Male, 4.4 mm, form B, JLB Hawaii 2: f, medial

gnathopod 2.

and transformation to lageniform state occurs very

early. Form B may represent a retarded phase or holding stage in the maturation cycle, whereas form A may

be a "pelagic phase" with early and continuous reproduction. But form A may be one kind of hybrid of

G. atlanticus with G. afra. Another hybrid-like form is

reported under the latter species.

If the two forms represent sibling species, they are

unusual in that they occur throughout the tropical

Indo-Pacific together, from at least India to Hawaii.

At every locale each form has a minutely to grossly

distinct morphology from its nearest neighbors, and

they might be accorded subspecific appellations.

Strangely, G. afra must also be a sibling species as

gnathopod 2 is often almost identical to various allopatriots of G. atlantica (e.g., afra of South Africa

and atlantica of India). But in each locale the two

species may be distinguished as their morphologies

shift in various directions so as not to overlap

sympatrically.

Gammaropsis pacific a (Schellenberg, 1938) from

various islands in Micronesia and southern Polynesia

appears to be distinct from G. atlantica in the dactyl

of male gnathopod 2 overlapping the palmer defining

corner, but Barnard (1965a) found definite individuals of G. atlantica in the same islands (Marshalls)

as Schellenberg found his G. pacifica. Also, Barnard's

female G. pacifica from Micronesia probably is referable to G. atlantica or to the "hybrid" afra-atlantica

mentioned under G. afra. The diversity in Hawaiian

collections suggests to me that Megamphopus abbotti

Barnard (1965a) from Micronesia also belongs with

the G. afra-atlantica complex; it has gnathopod 2 of

afra, but the eyes are of intermediate character, and

the accessory flagellum is reduced to 2 articles. The

latter is the criterion for the Megamphopus "concept,"

but Barnard (1962a and elsewhere) has already questioned the validity of that slim difference. Megamphopus abbotti should be brought into the Gammaropsis

complex but retained as a distinct species. Undoubted-
178 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ly, the widely scattered archipelagos of Micronesia and

Polynesia provide fertile ground for speciation or at

least strong differentiation by founder populations enjoying less stressful competition than that occurring

in more diverse environments. Unfavorable mutations

recurrent in populations inhabiting diverse environments may be better suited for and become dominant

in the small scale and poorly diverse environments of

isolated atolls or small island groups.

Hawaiian populations of both G. afra and G. atlantica have the inner plate of maxilla 1 fully setose on

the medial edge; the tiny individuals of G. afra have

only 3 setae but they fully embrace the medial margin.

DISTRIBUTION.—Eastern Atlantic Ocean eastward to

Hawaii.

Gammaropsis digitata (Schellenberg) from Canton

Island

FIGURE 114

Eurystheus digitatus Schellenberg, 1938, pp. 84-86, fig. 44.—

J. L. Barnard, 1965a, pp. 535-536, fig. 30.

DIAGNOSIS OF MALE.—Lateral cephalic lobes strongly extended, narrow, antenna 2 moderately recessed

along ventral margin; eyes obliquely ovate and elongate, rusty pink in alcohol; epistome unproduced; right

mandibular molar with strong flake and seta; inner

plate of maxilla 1 with about 5 long medial and 2

short terminal setae; antennae 1-2 equal to each other

in length, relatively stout, nearly as long as head and

pereon together, flageila nearly 40 percent as long as

peduncles, accessory flagellum 4-articulate; coxa 1

bluntly angular at anteroventral corner, coxae otherwise normal and not enlarged; article 6 of gnathopod

1 ovate, not expanded, article 5 bearing 1 thin anterior

seta; palm of gnathopod 2 basically transverse, with

large tooth becoming split off posterior margin of

article 6 and becoming more strongly gaped with increase in age, crotch of tooth with large articulate

spine, reduced to a seta and finally lost in adults, distal

end of palm with stout falciform protuberance, dactyl

strongly curved at base and maintained in a transverse

position; article 2 of pereopods 3—5 slender and weakly

ovatopyriform, not rectangular; pleonal epimera 1-3

with slight posteroventral notch and tooth, posterior

bulges of medium extent; urosomites 1-2 each with 1

dorsolateral seta on each side but otherwise simple;

outer ramus of uropod 2 lacking article 2; telson deeply excavate dorsally; uropod 2 with slight peduncular

tooth between rami.

FEMALE.—Palm of gnathopod 1 evenly convex and

not bearing slight sinuosity seen in male; gnathopod 2

small, palm unexcavate, bearing stout spines.

MATERIAL.—Canton Island, February 1958, Drs.

Degener (4).

DISTRIBUTION.—Micronesia, Southern Polynesia,

and the Hawaiian Islands.

Gammaropsis haleiwa, new species

FIGURE 115

DIAGNOSIS OF YOUNG MALE.—Lateral cephalic lobes

long and narrow, anteroventral margin of medium

recessment for antenna 2, lobes rounded-subconical

apically; eyes small, subcircular, a black core surrounded by clear, small ommatidia; epistome rounded

anteriorly; right mandibular molar with flake; inner

plate of maxilla 1 bearing 1 long apical seta and 1

small setule; maxilliped differing from Hawaiian congeners in flabellate expansion of apical spines on

inner plate of maxilliped, presence of adz-shaped

process on article 3 of palp and in elongation of article

4, subunguiform and bearing short nail and 2 or 3

setules all shorter than article 4; antennae slender,

antenna 1 as long as head plus first 9.5 body segments,

antenna 2 three-fourths as long as 1, accessory flagellum 4-articulate; coxa 1 obtuse anteroventrally, coxae

otherwise normal and not enlarged; article 6 of gnathopod 1 ovatorectangular, not expanded, palm defined

by small spine and bearing large spine like that of

gnathopod 2, article 5 of gnathopods 1-2 lacking

anterior spines; gnathopod 2 small, scarcely broader

than gnathopod 1, palm oblique, even, defined by

medium spine and bearing large medial spine in middle, dactyl slender and fitting palm; pereopods 3-4

(5 unknown) like those of Aoroides columbiae figured herein including presence of stout, slightly hooked

submarginal spine on article 5, article 2 thus thin but

weakly ovatopyriform; pleonal epimera 1-3 with moderately bulging posterior margins and weak posteroventral notch and tooth; urosomite 1 with 1 dorsolateral

seta on each side marking slight anterior turn in lateral margin, urosomite 2 similar but dorsal part of

lateral margin with 3 nobs, dorsolateral margin of

urosomite 3 also nobbed; both uropods 1 and 2 with

long ventral peduncular tooth between rami, uropod

1 also with distomedial sharp tooth on peduncle; rami

of uropod 3 about 1.5 times as long as short peduncle,

outer with small article 2; telson moderately excavate
NUMBER 3 4 179

FIGURE 114.—Gammaropsis digitata (Schellenberg), female, 3.5 mm, Canton Island: a, part of

pereopod 5; b, gnathopod 2, setae marked by pits; c, part of pereopod 3. Male, 4.0 mm: d, head;

e, telson; /, gnathopod 1; g, uropod 3; h, apex of maxillipedal palp; i, pleonal epimeron 3 and

urosome; ;, part of gnathopod 2.
180

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 115.—Gammaropsis haleiwa, new species, holotype, Pmale, 3.5 mm, JLB Hawaii 10:

a, b, gnathopods 1, 2, medial; c, coxa 1; d, head; e, inner plate of maxilla 1; f, inner plate of

rnaxilliped; g, apex of maxillipedal palp; h, pleonal epimeron 3: i, urosome, lateral.
NUMBER 34 181

dorsally and bearing only 1 lateral seta on each side,

no spine.

HOLOTYPE.—Bishop Museum collections, catalog number 7280, ?male, 3.5 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 10, Kaneohe Bay,

Oahu, 2 m, Pocillopora, 23 February 1967.

REMARKS.—The mandibular palp is slender like that

of G. maculatus (Johnston) ( = G. erythrophthalmus

(Liljeborg) in Sars, 1895, pi. 198) and the lobes of

maxilla 2 are broader than in Sars' representation.

Affinities of this species outside of Hawaii cannot

be determined until we are certain that adult males

have been described. In Hawaiian waters this species

differs from its congeners in numerous characters, some

of possible generic value such as those of the maxillipeds, but it differs in nongeneric characters by a combination of narrow cephalic lobes, small, circular eyes

with black core, elongate rami of uropod 3 with 2-

articulate outer ramus, the presence of a long peduncular tooth on uropod 2, and the occurrence of nobs

on urosomal margins.

DISTRIBUTION.—Hawaiian Islands.

Gammaropsis kaumaka, new species

FIGURE 116

DIAGNOSIS OF YOUNG MALE.—Lateral cephalic lobes

of medium elongation, anteroventral margin of medium recessment for antenna 2, lobes rounded-subconical apically; eyes of moderate size, irregularly ovate,

slightly tilted from horizontal plane, a black core surrounded by clear ommatidia; epistome rounded anteriorly ; right mandibular molar appearing to lack flake

but possibly bearing rugosity; inner plate of maxilla

1 bearing 2 short apical setae and 2 long medial setae;

maxillipedal palp and inner plate of normal condition

for genus, major spine-teeth of inner plate short conical

sabres, palp article 3 lacking process, article 4 short,

with long apical setal spines; antennae short and stout,

equal to each other in length, accessory flagellum

2-articulate; coxa 1 obtuse anteroventrally, coxae

otherwise normal and not enlarged; article 6 of gnathopod 1 subrectangular, not expanded, palm defined by

small spine, article 5 of gnathopods 1-2 lacking anterior

spines; gnathopod 2 of medium size, possibly in juvenile condition, palm oblique, rounding onto posterior

margin of hand and defined by stout spine, dactyl fitting palm; pereopods 3—4 (5 unknown) like those of

Aoroides columbiae figured herein, second articles

stouter and subpyriform; pleonal epimera 1-3 with

moderately bulging posterior margins and weak posteroventral notch and tooth; urosomites simple; both

uropods 1 and 2 with long ventral peduncular tooth

between rami; outer ramus of uropod 3 as long as

peduncle, with barrel-shaped article 2, inner ramus

slightly shorter and much thinner than outer ramus,

bearing 1 apical spine; telson moderately excavate

dorsally and bearing stout spine and nob on each side.

HOLOTYPE.—Bishop Museum collections, catalog

number 7281, ?male, 1.7 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 17, Kawela Bay,

Oahu, intertidal, Sargassum, other algae, rocks, corals,

24 May 1967.

RELATIONSHIP.—Affinities of this species outside of

Hawaii cannot be determined until definitely adult

males have been described, but in its juvenile condition,

G. kaumaka is clearly distinct from all other Hawaiian

species of Gammaropsis. The presence of a peduncular

tooth on uropod 2 links this species with G. haleiwa,

new species, the only other Gammaropsis in Hawaii so

far having that character, but G. kaumaka differs from

G. haleiwa in the normal articles 3-4 of the maxillipedal palp, the slightly shorter lateral cephalic lobes

with more irregular eyes, the smooth margins of the

urosomites, the short rami of uropod 3, the

medially setose inner plate of maxilla 1, the normal

spines on the inner plate of the maxilliped, and the

presence of stout spines on the telson.

Gammaropsis kaumaka also resembles G. pokipoki,

new species, but has shorter rami on uropod 3, less

deeply inserted antenna 2, an unexcavate palm of

gnathopod 2, short article 5 on gnathopod 2, a normally rounded and broad pleonal epimeron 3, and the

process on uropod 2.

Gammaropsis kaumaka also is difficult to distinguish

superficially from G. pali, new species, but differs in

the presence of the peduncular process on uropod 2,

the blunter spines on uropods 1-2, the broader article

2 of pereopods 3-4 (?5), the unexcavate palm of

gnathopod 2, the slightly more deeply inserted antenna

2, the shorter inner ramus of uropod 2 with its single

apical spine, and the absence of anterior spines on

article 5 of gnathopods 1-2.

Antenna 1 bears 4 long and 1 short flagellar articles

and antenna 2 bears 3 flagellar articles.

DISTRIBUTION.—Hawaiian Islands.
182

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 116.—Gammaropsis kaumaka, new species, holotype, male, 1.7 mm, JLB Hawaii 17:

a, b, parts of pereopods, 4, 3: c, head; d, pleonal epimeron 3; e, telson and uropod 3 from lateral

aspect; /, apex of maxillipedal palp; g, lateral uropod 2; h, telson; i, j , medial gnathopods 1, 2;

k, mandibular palp, setae removed; I, uropod 3; m, antenna 1; n, inner plate of maxilla 1.
NUMBER 3 4 183

Gammaropsis pali, new species

FIGURES 117, 118

DIAGNOSIS OF FEMALE AND JUVENILE MALE.—Lateral cephalic lobes of medium extension, narrow, anteroventral margin slightly insinuate; eyes dark in alcohol,

irregularly oval or trapezoidal, set obliquely in ocular

lobes, anterior and ventral margins clear of pigment;

epistome unproduced; right mandibujar molar with

flake; inner plate of maxilla 1 with about 5 long medial

setae; antennae 1-2 subequal to each other in length,

about as long as head and pereonites 1-6 together,

flagella about half as long as peduncles, accessory

flagellum about 4-articulate; coxa 1 sharply angular at

anteroventral corner, coxae otherwise normal and not

enlarged; article 6 of gnathopod 1 subrectangular, not

expanded, palm lacking defining spine, article 5 of

gnathopods 1-2 with 2-3 stout anterior spines; palm

of gnathopod 2 oblique and short, dactyl overriding

palm, palm with locking bu,lge near finger, then slightly

excavate and with slight protrusion before becoming

widely excavate, defining corner scarcely projecting,

FIGURE 117.—Gammaropsis pali, new species, holotype, female, 4.3 mm, JLB Hawaii 10: a,

lateral view of body; b, lateral view of telson; c, telson; d, inner plate of maxilla 1; e, apex of

gnathopod 2; /, apex of pereopod 1; g, uropod 1; h, urosome and margins of pleonal epimera 1-3,

to left.
184

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 118.—Gammaropsis pali, new species, holotype, female, 4.3 mm, JLB Hawaii 10: a,

medial gnathopod 2; b, medial gnathopod 1; e, pereopod 4; d, head; e, telson.

medial surface with enormous spine; article 2 of pereopods 3-5 slender and subrectangular; pleonal epimera

1-3 with slight posteroventral notch and tooth, not

bulging strongly; urosomite 1 with one lateral seta,

urosomite 2 with about 4 subrnarginal lateral setae, no

teeth or notches; outer ramus of uropod 3 with a small

article 2; telson only weakly excavate dorsally.

COLOR IN 2-DAY FORMALDEHYDE.—Eyes bright garnet-ruby, head with similar red spots, antennae dark

garnet splotchy, a few body segments with brown vertical stripes.

HOLOTYPE.—Bishop Museum collections, catalog

number 7282, ovigerous female, 4.3 mm.

TYPE-LOCALITY.—JLB Hawaii 10, Kaneohe Bay,

Oahu, 2 m, Pocillopora, 23 February 1967.

MATERIAL.—JLB Hawaii 10 (11), 14 (1).

RELATIONSHIP.—This species belong with the G.

afra-atlantica group of the genus because the outer

ramus of uropod 3 has a minute second article, but

because few species of Gammaropsis have been

decidedly examined for this character, one cannot

determine how extensively distributed is the character

nor what relationships it may signify. No male of

G. pali is as mature as, nor as well developed as, the

adult females collected, so that relationships with males

of other species are cloudy.

Gammaropsis pali differs from G. afra and G. atlantica in the shapes of the eye and cephalic lobe, in the

simple urosomites 1-2 having only submarginal setae,

in the presence of a large molarial flake on the right

mandible, the thin rectangular article 2 of pereopods

3-5, and in the stout anterior spines on the wrists of

the gnathopods.

The new species resembles females of G. semidentatus (K. H. Barnard, 1916) from South Africa but differs in the narrow article 2 of pereopods 3-5.

Affinities with G. japonica (Nagata, 1961b) occur in

view of the narrow article 2 of pereopods 3-5 on females and the shape of gnathopod 2, but G. pali differs

from G. japonica in the absence of small teeth on

urosomites 1-2, the nearly straight, unbulging posterior

margin of epimeron 3, and in the unproduced epistome.

DISTRIBUTION.—Hawaiian Islands.
NUMBER 3 4 185

Gammaropsis pokipoki, new species

FIGURES 119, 120

DIAGNOSIS.—Lateral cephalic lobes strongly extended and thick, bluntly triangular in front, antenna 2

deeply recessed along ventral cephalic margin; eyes

large, subcircular, with a black core surrounded by

clear ommatidia; epistome rounded anteriorly; right

mandibular molar with long seta, both molars with

basal rugosities but no flakes; inner plate of maxilla 1

with 4-5 medial setae; antenna 1 slightly longer than

antenna 2, about as long as head and first 8 body segments together, flagella more than three-fourths as

long as peduncles, accessory flagellum 3-articulate;

coxa 1 bluntly angular at anteroventral corner, coxae

otherwise normal and not enlarged; article 6 of gnathopod 1 an elongate trapezoid, not expanded, article 5

of both pairs of gnathopods smooth anteriorly; gnathopod 2 small, palm oblique in both sexes, short, deeply

excavate in male and aberrant females, bearing enormous spine, palm defined by conical protrusion and excavation bounded distally by similar protrusion, dactyl

slender and overextending palm, female palm normally

scarcely excavate; article 2 of pereopods 3-5 expanded

and pyriform; pleonal epimera 1-3 with slight posteroventral tooth, posterior margins strongly bulging;

urosomites 1-2 each with 1 dorsolateral seta, otherwise

simple; rami of uropod 3 about 1.5 times as long as

peduncle, outer with small article 2; telson only weakly

excavate dorsally.

HOLOTYPE.—Bishop Museum collections, catalog

number 7383, female, 2.6 mm.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, coralline footballs, 29 January 1967.

MATERIAL.—JLB Hawaii 5 (4), 6 (2), 10 (6),

12 (?1).

RELATIONSHIP.—This species has its closest affinities

with the G. atlantica-afra complex and G. pali, for it

differs from Hawaiian G. alamoana in the setose inner

plate of maxilla 1 and in the presence of a second

article on the outer ramus of uropod 3. Gammaropsis

pokipoki has few qualitative characters to distinguish

it from the 3 species mentioned above, but nevertheless

stands out sharply when encountered in various samples. The body and antennae are long and thin, the

head shallow and very smooth, and the large circular

eye with black core surrounded by clear ommatidia

immediately marks the species from all its congeners

except G. haleiwa, new species, (which see for relationships) . Gammaropsis atlantica and G. ajra have no

dark pigment in the eyes and have more complex and

larger gnathopods and weakly ornamented urosomites

1-2.

Gnathopod 2 of G. pokipoki resembles that of G. pali

very closely but the head, eyes, and antennae are very

different and G. pali has a distinct molarial flake,

thicker body, shorter rami of uropod 3, and narrow

second articles on pereopods 3-5.

Female and juvenile individuals of G. pokipoki

closely resemble their counterparts of Konatopus paao,

for the heads and eyes have strong similarities, but

careful examination shows gnathopod 1 of Konatopus

to be very slightly larger than gnathopod 2, and the

eyes retain reddish or pale brown pigment instead of

black.

DISTRIBUTION.—Hawaiian Islands.

Photis aina, new species

FIGURES 121, 122

DIAGNOSIS OF MALE.—Head with long, nearly clavate ocular lobes; eyes round or horizontally subovate,

located terminally in ocular lobes, clear in formaldehyde and alcohol, composed of small, spherical, tightly

packed ommatidia; antennae equal to each other in

length, as long as head and pereon together, article 3

of antenna 1 equal to article 1, article 2 about 1.5 times

longer than either; coxa 1 elongate, rectangular, with

slightly protuberant anteroventral corner, coxae 2-4

also of similar elongate rectangular condition, coxa 5

with narrow, slightly tapering anterior lobe (abnormal

for majority of species in Photis), posterior lobe thus

relatively wide compared with anterior lobe, all coxae

strongly setose ventrally and lacking stridulation

ridges; gnathopod 1 with articles 5 and 6 equal to each

other in length, article 5 broadly truncate posteriorly,

article 6 of normal ovateness, palm with strong medial

submarginal spine not precisely at defining corner,

palm minutely crenulate, dactyl long and slender, with

small teeth and crenulations on inner edge; gnathopod

2 small, article 2 lacking anteroventral lobe, article 2

small, thick, article 4 not produced posterodistally,

article 5 of medium breadth, cup-shaped, narrowly lobate posteriorly; article 6 broad, about 80 percent as

broad as long, palm slightly oblique, bisinuate, defined

by quadrangular corner, bearing weak hump near defining corner and weak hump distally near base of

dactyl, medioproximal face of hand with large spine,
186 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 119.—Gammaropsis pokipoki, new species, holotype, female, 2.6 mm, JLB Hawaii 6:

a, mandibular palp; b, telson; c, apex of gnathopod 2; d, uropod 3; e, right mandibular molar;

/, pleonal epimeron 3 and urosome. Male, 2.2 mm: g, head. Female, 3.2 mm, JLB Hawaii 10:

h, apex of gnathopod 2, medial.
NUMBER 3 4 187

FIGURE 120.—Gammaropsis pokipoki, new species, male, 2.2 mm, JLB Hawaii 6: a, head; b, c,

medial gnathopods 2, 1. Holotype, female, 2.6 mm: d, half of lower lip; e, f, g, pereopods 2, 3, 5;

h, lobes of maxilla 2, setae removed.

360-034 O—1C
188 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dactyl fitting palm in medium male, much shorter

than palm in terminal male, dactyl stout, bearing 3-4

medium-sized teeth on inner margin, pereopods 3-5

relatively short and stout for genus, article 2 of pereopod 3 very broadly pyriform, anteriorly and posteriorly

setose, article 5 almost quadrate, article 4 with similar

appearance but obliquely cut proximally, article 6

stout, rectangular, bearing 1 large stout and 1 small

stout distal spines, dactyl geniculate, armed with bilateral pair of sharp accessory cusps, pereopod 4 very

similar to pereopod 3 but article 2 narrower, articles

4-6 slightly longer, dactyl also with accessory cusp,

pereopod 5 with small posteroproximal cusp on article

2, articles 4 and 6 elongate, article 5 short, bearing

weak distal lockipg spine and setule, dactyl straight,

but bearing 2 setules and slight tooth on inner margin

and pair of small accessory cusps on outer; uropods 1-2

lacking ventrodistal peduncular tooth, inner ramus of

uropod 1 lacking marginal spines, otherwise rami and

peduncles with a few spines; outer ramus of uropod 3

about 60 percent as long as peduncle, thus, peduncle

elongate and outer ramus short for genus, article 2

well developed and barrel-shaped, apical spines long,

inner ramus half as long as outer, thus, as long as that

of subgenus Cedrophotis; telson subtriangular, apically

obtuse, bearing dorsolateral nob on each side plus

various setae and setules; epimeron 1 with several

anteroventral setae, posteroventral corner rounded,

epimeron 2 with 1 posteroventral notch, and epimeron

3 with 2 notches above rounded, protruding, posteroventral corner.

Mouthparts like those of P. reinhardi Kroyer (Sars,

1895, pi. 202), but mandibular palp longer, right molar

with flake and seta, and maxillipedal palp article 4

armed with numerous apical setal-spines.

FEMALE.—Gnathopod 2 slightly smaller than in

male, palmar defining corner rounded, palm with 1

weak smooth sinus, palm otherwise minutely crenulate.

HOLOTYPE.—Bishop Museum collections, catalog

number 7284, male, 2.5 mm.

TYPE-LOCALITY.—JLB Hawaii 3, off Ewa Beach,

Oahu, 18 m, Pocillopora, bryozoans, 29 January 1967.

MATERIAL.—JLB Hawaii 2 (34), 3 (88), 5 (11),

6 (40), 8( 180).

RELATIONSHIP.—This species resembles those members of Photis with long ocular lobes like P. dolichommata Stebbing (1910) and P. lamellifera Schellenberg

(1928). The latter species is distinguished by the

shorter article 5 of gnathopod 1, being only half as

long as article 6. Photis aina differs from P. dolichommata in the slightly stouter pereopods 3-5, shorter antennae, longer article 5 of gnathopod 2, broader article

6, lack of spines on the inner ramus of uropod 1, and

the slightly longer inner ramus of uropod 3. Photis aina

and P. dolichommata resemble each other in the unusual shape of coxa 5.

Photis aina has weak affinities with P. longicaudata

(Bate and Westwood) (Sars, 1895, pi. 203, fig. 1) as

gnathopod 2 has approximately the same outline, but

this general shape is fairly common in the genus, especially in juveniles of various species with stronger

ornamentation in terminal stages. The ocular lobes are

much shorter and /or broader in P. longicaudata than

in P. aina, but several decades ago those ocular lobes

of the former species were considered extreme. Coxa 5

of P. aina is distinct from the normal shape in P.

longicaudata.

The so-called P. longicaudatus (Bate and Westwood) of Walker (1904) and K. H. Barnard (1916),

which probably represents a distinct species, also apparently has the normal coxa 5, and P. longimanus

Walker has a normal coxa 5, presumably. Gnathopod 2

of the male is far more ornamented in those CeyloneseSouth African species than it is in P. aina; perhaps the

terminal male of P. aina has not been discovered.

Photis species of Pirlot (1938) from the Celebes

has a normal coxa 5, slender articles 2-6 of pereopods

3-5, a strong anteroventral lobe on article 2 of gnathopod 2, and weak coxal setation.

In Hawaii, this species differs from P. hawaiensis

J. L. Barnard (1955a) and P. kapapa new species, in

the strong setation of the coxae, the long inner ramus,

and, relative to the peduncle, the short outer ramus

of uropod 3, and the stoutness of pereopods 3-5. Male

gnathopod 2 of P. hawaiensis has a very oblique palm

and large mammilliform process on article 2, and the

oval eyes fully occupy the lateral cephalic lobes. Male

gnathopod 2 of P. kapapa has a much broadened palm,

a process on article 2, and like P. hawaiensis, but unlike P. aina, has stridulation ridges on one or more

coxae and on article 2 of gnathopod 2.

Photis aina resembles the deep-water eastern Pacific

subgenus Cedrophotis (J. L. Barnard, 1967) in the long

inner ramus of uropod 3 and the relatively long peduncle, but the upper lip of Cedrophotis is much more

deeply incised, and the inner lobes of the lower lip are

partially coalesced. Probably, P. aina shows the unsuitability of the Cedrophotis concept and demonstrates
NUMBER 3 4 189

FIGURE 121.—Photis aina, new species, holotype, male, 2.5 mm, JLB Hawaii 3: a, b, c, d,

pereopods 3,4, 5, 1; e, gnathopod 1; /, lateral gnathopod 2; g, medial gnathopod 2.
190

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 122.—Photis aina, new species, holotype, male, 2.5 mm, JLB Hawaii 3: a, b, head, lateral

and dorsal; c, palm of medial gnathopod 2; d, palm of medial gnathopod 1; e, f, dactyls of

pereopods 5, 3; g, urosome; h, pleonal epimera 1-3. Male, 3.0 mm, JLB Hawaii 2: i, palm of

lateral gnathopod 2. Female, 2.6 mm, JLB Hawaii 3: j , mandibular palp minus setae; k, coxa 4;

/, medial gnathopod 2; m, apex of maxillipedal palp; n, uropod 3 and telson.
NUMBER 3 4 191

that in isolated environments, various species of Photis

independently develop simplifications or aberrations.

DISTRIBUTION.—Hawaiian Islands.

Photis hawaiensis J. L. Barnard

FIGURE 123

Photis hawaiensis J. L. Barnard, 1955, pp. 35-37, figs. 18,

19.

DIAGNOSIS OF MALE.—Head with medium-sized ocular lobes, eyes horizontally ovate, filling ocular lobe

and projecting into head proper, ommatidia much

larger than in other Hawaiian photises and partially

discontiguous, all not perfectly spherical, eyes pale

eosin in alcohol when viewed under high power; antennae 1-2 as long as head and pereon together, article

3 of antenna 1 equal to article 1 or slightly shorter,

article 2 scarcely longer than either; coxa 1 elongaterectangular and with 1 long and several short setae,

coxae 2-4 broader than 1 and thus relatively less elongate, essentially asetose, coxae 3-4 with ventrolateral

stridulating ridges, coxa 3 with set of ventromedial

ridges also, coxae weakly rounded ventrally, coxa 5

with normally broad anterior lobe; gnathopod 1 with

FIGURE 123.—Photis hawaiensis J. L. Barnard, male, 3.3 mm, Kaneohe Bay, 1937: a, uropod 3;

b, telson; c, head; d, medial gnathopod 2; e, uropod 1; /, medial gnathopod 1, setae marked

with pits. Female, 2.2 mm: g, h, dactyls of pereopods 5, 4; i, uropod 3 and telson from lateral

view (upside down);;, gnathopod 2.
192 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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

5 broadly truncate posteriorly, article 6 normally but

broadly ovate, palm weakly convex (drawing of Barnard, 1955a, poor), undefined, bearing 1 medial spine

near middle, dactyl long, fitting palm theoretically,

with small teeth and crenulations on inner edge;

gnathopod 2 very large, article 2 with immense anterodistal mammilliform hump armed with stridulating

ridges, article 3 quadrate from lateral view and bearing medial keel projecting mediad (shown folded

down in drawing), article 4 medially setose (setae

shown by pits only in figure), article 5 very broad, unlobate medioposteriorly, but bearing large, setose posterolateral lobe and anterior rounded setose keel, setae

very heavy, article 6 broad but tapering distally, palm

occupying full posterior margin of hand and defined by

medioproximal tooth, palm distally with 2 sinuses and

2 cusps, dactyl as long as hand, inner edge smooth except for weak distal tooth and setae; pereopods 1-2

with article 4 especially turgid anterodistally; pereopod

3 fairly stout for genus but not as stout as that of P.

aina, new species, dactyl with 1 sharp accessory cusp,

pereopod 4 similar to 3 but article 2 narrower, pereopod 5 dactyl with 1 slender accessory cusp; uropods

1-2 with very small sharp leaflike ventrodistal tooth

between rami, inner ramus of uropod 1 lacking marginal spines, otherwise rami of uropods 1-2 with 1

marginal spine each, apices of each with jewel-like

nails; outer ramus of uropod 3 of variable length relative to peduncle, but about three-fourths as long as

that peduncle, article 2 well developed and barrelshaped, apical spines long, inner ramus 20-25 percent

as long as outer ramus; telson triangular, bearing deep

anterolateral notch on each side to form shape of leaf,

the 2 acclivities of the notches apparently representing

nobs normally found in females or juveniles; pleonal

epimera all with softly rounded posteroventral corners,

epimeron 2 with 2 weak ventral setae.

FEMALE.—Gnathopod 2 smaller than that of male,

article 2 with medium-sized mammilliform lobe, article

5 with narrower posterior lobe, article 6 expanded,

palm slightly oblique, with weak sinus near dactyl,

then cusp bordered by deep sinus followed by transverse sinuous margin with quadrate defining corner

and bearing small, medial, submarginal spine, dactyl

slightly shorter than palm, with 2 inner teeth.

MATERIAL.—Same as J. L. Barnard (1955a).

DISTRIBUTION.—Hawaiian Islands.

Photis kapapa, new species

FIGURES 124, 125

DIAGNOSIS OF MALE.—Head with long narrow

ocular lobes, eyes round or horizontally ovate, with

black, brown, or purplish-brown core in formaldehyde

or alcohol, eyes occupying distal ends of ocular lobes,

ommatidia small and closely packed; antennae 1-2

equal to each other in length, as long as head and

pereonites 1-5 together, article 2 equal to article 1,

article 2 about 1.3 times longer than either; coxa 1 of

medium elongation, subrectangular, anteroventral corner not protuberant, coxae 2-4 broadly rectangular,

coxae 1-2 with weak stridulation ridges on posterior

margin, coxa 3 with ridges on anteroventral margin,

4 with small lateral ones posteroventrally and large

medial ones ventrally, coxae all scarcely setose, coxa

5 with normally broad anterior lobe; gnathopod 1 with

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

5 broadly truncate posteriorly, article 6 of normal

ovateness, palm lacking any special thick spine but

bearing short setal spine in expected position, this

spine similar to neighboring palmar spines (and relatively large in juveniles only), palm minutely crenulate,

dactyl long and slender, with small teeth and crenulations on inner edge; gnathopod 2 small, article 2 with

very large, saclike anterodistal lobe bearing stridulation

ridges, article 4 unproduced posterodistally, article 5 of

medium breadth, cup-shaped but with tumid posterior

lobe, article 6 expanded, about 80 percent as broad as

long, palm slightly oblique, defined by large blunt,

gaping tooth, palm weakly bisinuate, with small obtuse

cusp near dactylar base and medial spine on low hump

near defining tooth, medial palmar face with quadrate

tooth anterior to main spine, dactyl much shorter than

palm, with crenulations and 2 inner teeth; pereopods

3-5 normal in stoutness for genus, article 2 of pereopod

3 pyriform, scarcely setose, article 6 with 1 large and 1

small distal spines, dactyl slightly geniculate, with bilateral pair of accessory cusps, pereopod 4 similar to

3, but article 2 much narrower, articles 4 and 6 slightly

longer, dactyl also with 2 accessory cusps, pereopod 5

with weak posteroproximal projection, articles 4 and

6 not strongly elongate and article 5 not strongly

shortened, article 6 bearing 2 weak distal locking

spines, dactyl nearly straight and with only 1 thin,

appressed accessory cusp and 1 setule; uropods 1-2

lacking ventrodistal peduncular cusps between rami,
NUMBER 3 4 193

inner ramus of uropod 1 naked, outer ramus with 1-2

spines, spines thinner than in P. aina, rami with jewellike apical nails, outer ramus of uropod 3 nearly 90

percent as long as peduncle, article 2 well developed

and barrel-shaped, apical spines weak, inner ramus

about one-third as long as outer; telson nearly hemicircular, apex obtusely convex, each side with small nob;

posteroventral corner of pleonal epimera 1 and 3

rounded, that of epimeron 2 rounded-quadrate.

Mouthparts generally similar to those of P. aina,

new species, but article 4 of maxillipedal palp lacking

lateral seta.

FEMALE.—Gnathopod 2 of two kinds, smaller than

that of male, palm slightly oblique and either weakly

S-shaped and fully crenulate or bearing semicircular

smooth excavation in middle, juvenile palm like that

of former kind, dactyl failing to fit palm, defined by

softly rounded quadrate corner.

FIGURE 124.—Photis kapapa, new species, holotype, male, 2.7 mm, JLB Hawaii 3: a, lateral coxa

4; b, gnathopod 2; c, upper lip; d, gnathopod 1; e, pereopod 1. Male, 2.4 mm, JLB Hawaii 2:

/, apex of pereopod 3;g,h, pereopods 5, 3.
194 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

JUVENILE MALE.—Dactyl fitting palm and defining

tooth small, hand not strongly expanded and distal

palmer tooth small and narrow.

ABERRANT FEMALE.—Large, 3.3 mm ovigerous female from JLB Hawaii 13, with noticeably small

gnathopods in juvenile state, dactyl as long as palm

but palm like that figured for 2.6 mm female of JLB

Hawaii 3 and with much larger quadrate tooth on

medial palmar face.

HOLOTYPE.—Bishop Museum collections, catalog

number 7285, male 2.7 mm.

TYPE-LOCALITY.—JLB Hawaii 3, off Ewa Beach,

FIGURE 125.—Photis kapapa, new species, holotype, male, 2.7 mm, JLB Hawaii 3: a, pereopod 4;

b, coxa 5; c, telson and uropod 3; d, head; e, pleonal epimera 1-3. Male, 2.3 mm: /, hand of

lateral gnathopod 2. Male, 2.4 mm, JLB Hawaii 13: g, palm of medial gnathopod 2. Male, 2.4

mm, JLB Hawaii 2: h, palm of medial gnathopod 2. Female, 3.8 mm, JLB Hawaii 5: i, palm of

medial gnathopod 2. Female, 2.6 mm, JLB Hawaii 3: j , palm of medial gnathopod 2. Juvenile,

1.7 mm, JLB Hawaii 3: k, hand of medial gnathopod 2.
NUMBER 34

Oahu, 18 m, Pocillopora, bryozoans, encrustations, 29

January 1967.

MATERIAL.—JLB Hawaii 2 (13), 3 (38), 5 (4), 6

(6),8(7),10(9),12(4),13(3).

RELATIONSHIP.—This species appears similar to

Photis species of Pirlot (1938) from the Celebes, a

taxon that Pirlot considered synonymous with Walker's

(1904) so-called P. Iongicaudata (Bate) from Ceylon.

Adult males of Photis kapapa have a more strongly extended defining tooth on the palm of gnathopod 2,

and weaker and less sharp distal protrusion near the

base of the dactyl than does Photis species, and bear

stridulating ridges on gnathopod 2 and coxae.

DISTRIBUTION.—Hawaiian Islands.

Family ISCHYROCERIDAE

Ischyrocerus (?) kapu, new species

FIGURE 126

DESCRIPTION.—Aspect generally like that of Jassa

lilipuna, new species, but coxae all shorter, coxa 1 as

long as 2 and coxa 5 as long as 4, thus, assignable provisionally to Ischyrocerus; eyes, head, mandible, maxillae, maxillipeds, uropods 1-2, telson all similar to those

of /. lilipuna; pereopods 3-5 and antennae unknown,

missing from all specimens; gnathopod 1 with only one

defining spine on palm, dactyl very long and slender,

overlapping palm; coxa 1 trapezoidal, coxae 2-4 much

shorter than broad; male gnathopod 2 with broadly

lobate articles 2-3, article 4 immensely produced

posterodistally to form substitute palmar defining tooth,

article 5 triangular and enclosed posteriorly by article

4, article 6 elongate, apically broadened, lacking defining tooth on palm, latter occuping entire posterior

margin of article 6, distal end of palm with 2 large

processes separated by deep, flat-bottomed excavation,

proximal tooth subconical, distal process spinose and

asymmetrically triangular, dactyl very long and strongly

overlapping palm and process of article 4; article 6 of

pereopods 1-2 longer and more slender than article 6

of /. lilipuna, bearing only 2 posterior setae, no locking spines, dactyl long and simple; rami of uropod 3

much longer than on /. lilipuna, outer with about 12

very minute serrations visible generally on oil immersion lens only, inner ramus longer than outer and

bearing 1 apical spinule.

HOLOTYPE.—Bishop Museum collections, catalog

number 7286, male, 1.4 mm.

195

TYPE-LOCALITY.—JLB Hawaii 10, Kaneohe Bay,

Oahu, 2 m, Pocillopora, 23 February 1967.

MATERIAL.—JLB Hawaii 3 (1), 6 (1), 10 (male,

1.4 mm, figured).

REMARKS.—The immense process of article 4 on

gnathopod 2 distinguishes this species from other members of Ischyrocerus, although several species of that

primarily boreal genus have small processes occurring

on that article. Perhaps the process has generic value

and this species would represent an endemic Hawaiian

genus derived from a boreal member of Ischyrocerus.

The coxae do not correspond to those of Microjassa,

and the absence of material with pereopods and antennae prevents me from establishing the true affinities of

the species.

DISTRIBUTION.—Hawaiian Islands.

Ischyrocerus oahu, new species

FIGURE 127

DESCRIPTION.—Following characters like Jassa lilipuna, new species; head, eyes, antennae, uropods 1-2,

mandible and stout palp, maxillae, maxillipeds, telson;

coxae of male very short, especially coxae 3-4 broader

than long, but coxa 2 enlarged and longer than either

1 or 3, ventral margins straight, coxa 1 nearly quadrate

anteroventrally, coxa 5 as long as 4 or slightly longer;

gnathopods of female like those of Jassa lilipuna, new

species, but lacking comb-row on dactyls; article 5 of

gnathopod 1 about three-fourths as long as article 6,

latter slightly more broadened than in /. lilipuna;

gnathopod 2 of terminal male with weakly but broadly

lobate article 2, article 3 grossly lobate and spinose,

article 4 unproduced, article 5 short but produced to

free, blunt posterior lobe unlike other Hawaiian

ischyrocerus-jassa species, article 6 greatly elongate and

midposteriorly setose its full length, bearing weak

bilobate distal process, palm and posterior margin of

article 6 confluent and slightly concave, dactyl long,

about as long as article 6, inner margin trisinuous, with

2 weak basal protrusions; gnathopod 2 of young male

with much shorter, simple, stout dactyl, short and relatively stouter article 6 with nearly straight posterior

margin, but with weakly bilobate distal protrusion

forming palm, article 3 smooth; pereopods 1-2 normally slender compared with /. lilipuna, article 6 and

dactyl long, but with 3 stout posterior spines and terminal seta, distalmost 2 spines coupled with setule, no

locking spines per se; pereopod 4 narrow, scarcely ex-
196

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

s

FIGURE 126.—Ischyrocerus kapu, new species, holotype, male, 1.4 mm, JLB Hawaii 10: a, uropod

3 and telson from lateral view; b, lateral gnathopod 2; c, coxa and article 2 of gnathopod 2;

d, medial gnathopod 1; e, articles 3-5 and part of 6 on gnathopod 2; /, pereopod 2. Specimen,

JLB Hawaii 6: g, apices of outer and inner rami on uropod 3, left to right.
NUMBER 3 4 197

FIGURE 127.—Ischyrocerus oahu, new species, holotype, male, 1.2 mm, JLB Hawaii 10: a,

gnathopod 1; b, c, uropod 3; d, medial gnathopod 2; e, pleonal epimera 2,3;/ , pereopod 1; g,

pereopod 4 ; h, coxa 4; i, j , coxae 6, 5. Young male, JLB Hawaii 17: k, medial gnathopod 2 ;

I, uropod 3 and telson from lateral view.
198

panded article 2 lacking posteroventral lobe and

armament similar to /. lilipuna, but dactyl lacking

comb-row, anterodistal end of article 6 with 1 subterminal spine and pair of locking spines, medial spine

smaller than lateral (only latter drawn in figure) ; rami

of uropod 3 longer than on /. lilipuna, but outer ramus

often projecting laterally so as to appear even shorter,

but outer ramus distinctly shorter than inner, bearing

4-7 small denticles.

Only the slight disproportionate enlargement of coxa

2 suggests a classificatory difference of /. oahu from

other members of Ischyrocerus; in terms of male gnathopod 2 palm and dactyl, I. oahu is as close to the very

common boreal /. anguipes Kroyer as to any other.

Male I. oahu differs from that species in the slightly

shorter coxae 3-4, slightly larger coxa 2, absence of

a process on article 4 of gnathopod 2, bifidation of the

palmar process, reduction in setosity of pereopods, and

decrease in width of article 2 on pereopod 4 (3 and 5

unknown in 7. oahu), and in the general equalization

of male antennae 1-2. A resemblance especially in

gnathopod 2 to 7. commensalis Chevreux, 7. dezhnevi

Gurjanova, 7. krascheninnikovi Gurjanova, and 7.

chamissoi Gurjanova occurs, but 7. oahu differs from

all except the latter, according to the key in Gurjanova

(1951), by the relatively short rami of uropod 3. The

equal antennae in the male of 7. oahu differentiate it

from 7. chamissoi. One must disregard male gnathopod

2 differences until all of these species have been thoroughly studied and the various stages of development

elucidated.

HOLOTYPE.—Bishop Museum collections, catalog

number 7287, male, 1.2 mm.

TYPE-LOCALITY.—JLB Hawaii 10, Kaneohe Bay,

Oahu, 2 m, Pocillopora, 23 February 1967.

MATERIAL.—JLB Hawaii 10(4), 15( 1), 17 (9).

RELATIONSHIP.—Although gnathopod 2 has more

of the characteristics of an Ischyrocerus than does that

of Jassa lilipuna and Ischyrocerus kapu, the relative

difference between the lengths of coxae 1-2 suggests

affinities of 7. oahu with various species of Microjassa.

But coxae 3^4 are very short, and coxa 5 is even

slightly longer than 4, quite in contrast to the type-species of Microjassa, M. cumbrensis (Stebbing and

Robertson). Jassa lilipuna is not firmly assignable to

Microjassa for the same reason, even though coxae

2-4 are elongate as in M. cumbrensis. But several species of Ischyrocerus have coxae 2-A elongate, with

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

coxa 5 short, even though coxa 4 is not excavate for the

reception of coxa 5 as in the type-species of Microjassa.

DISTRIBUTION.—Hawaiian Islands.

Jassa (?) lilipuna, new species

FIGURES 128, 129, 130

DIAGNOSIS.—Lateral cephalic lobe coniform, ventral margin of lobe and head sinuous; accessory flagellum 2-articulate, article 1 elongate, article 2 very small;

article 3 of peduncle of antenna 1 as long as article 1;

epistome weakly and bluntly produced; palp article 3

of mandible extremely clavate; inner plate of maxilla

1 simple; maxilla 2 with about 4 setae on distal half

of medial margin; palp article 4 of maxilliped stout,

apically setose; coxa 1 of male extending ventrally

about half as far as coxa 2; latter bilobate in terminal

male; coxa 5 nearly as long as coxa 4, coxae 6-7 short;

gnathopod 1 of both sexes and gnathopod 2 of female

similar, article 5 cup-shaped and shorter than 6, latter

ovate, palm and posterior margin about equal, palm

defined by 2 spines, dactyl with several flat inner serrations and lateral comb; gnathopod 2 of male initially

with article 6 becoming greatly elongate while dactyl

remaining short, palm almost simple and confluent

with posterior margin of article 6, then posterior margin developing scant cusp bearing spine, finally in

adult, dactyl elongating to full posterior length of

article 6, midpalmar tooth becoming proximal

medium-sized thumb, palm near dactylar hinge formed

of 2 processes, 1 of medium size, 1 smaller and hemispherical, articles 2-3 weakly lobate, article 4 swollen,

article 5 small, cup-shaped, not reaching free margin;

pereopods 1-2 short, moderately stout, article 6 with

4 large, blunt posterior spines, 2 of them forming a

locking pair; pereopods 3-5 with anterior margin of

article 6 bearing large pair of locking spines and 1 other

large distal spine, dactyls all bearing facial comb;

outer ramus of uropod 3 with 2-3 terminal, scarcely

reverted cusps, inner ramus with 1 partially immersed

distal spinule; telson pointed, bearing a pair of stout

dorsal spines and a pair of setules.

OTHER FEATURES.—Mandibular molar large but

weakly triturative, bearing molarial flake; eyes large

and clear in alcohol.

HOLOTYPE.—Bishop Museum collections, catalog

number 7288, male, 1.9 mm.

TYPE-LOCALITY.—Waikiki, Oahu, in ocean seaward

of aquarium, from Ulva, 11 March 1964.
NUMBER 3 4 199

FIGURE 128.—Jassa lilipuna, new species, male, 1.8 mm, Waikiki: a, mandibular palp; b, telson;

c, uropod 3; d, maxilla 1; e, mandibular molar showing flake; f, prebuccal mass, left lateral; g,

inner plate of maxilliped; h, mandible (palp removed); i, urosome; ;, maxilliped. Female, 1.4

mm: k, coxa 5; /, lower lip; m, maxilla 2.
200

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 129.—Jassa lilipuna, new species, female, 1.4 mm, Waikiki: a, b, gnathopods 1, 2. Male,

1.3 mm: c, gnathopod 2. Male, 1.8 mm: d, e, f, g, pcreopods 2, 3, 4, 5; h, coxa 3; i, gnathopod 1.
NUMBER 34 201

FIGURE 130.—Jassa lilipuna, new species, female, 1.4 mm, Waikiki: a, head; b, apex of pereopod

5; c, pleonal epimera 1-3, left to right, and coxa 7. Male, 1.8 mm: d, head; e, f, antennae 1,2 ;

g, accessory flagellum; h, i, gnathopod 2, medial and lateral views.
202

MATERIAL.—Nine specimens from the type-locality

on Ulva and 20 specimens from Sargassum and 4 from

Padina.

REMARKS.—Females have a maximum of 3 large

eggs.

RELATIONSHIP.—Jassa lilipuna fits the genus Jassa

much better than it does Microjassa even though coxa

1 is slightly shorter than coxa 2, for coxa 5 is as long

as coxa 4, and coxae 2-4 are not greatly enlarged as

in the type-species, M. cumbrensis (Stebbing and

Robertson), M. macrocoxa Shoemaker, and M. litotes

J. L. Barnard. The debatable species is M. litotes J. L.

Barnard (1954b) that was later shifted to Ischyrocerus

and then returned to Microjassa (J. L. Barnard,

1969a). Microjassa litotes may indeed deserve a genus

of its own, for it differs from the other microjassas and

Jassa in the absence of any palmer tooth on male

gnathopod 2 and coxa 5 is not as short as originally

believed. But M. litotes differs from Ischyrocerus and

Jassa in the posterodorsally excavate coxa 4 and from

Parajassa in the large accessory flagellum.

The presence of the proximal palmar tooth on male

gnathopod 2 is a good generic character used by

Stebbing (1906) to differentiate Jassa and Ischyrocerus. Thus, /. lilipuna differs from Parajassa in

accessory flagellum and from Microjassa in coxae 2-5.

The description of a new species in Jassa poses

problems in terms of Sexton and Reid's (1951)

lengthy dissertation on the multiform character of

Jassa falcata (Montagu). They synonymized many

species, and one might gather from the diversity of

form demonstrated in /. falcata that the few remaining

species Sexton and Reid did not mention might also

fit into the /. falcata complex. Jassa lilipuna does have

its closest relationship with /. falcata according to

Stebbing's (1906) key if one pursues the necessary

synonymy. According to Sexton and Reid's description of general characters in /. falcata, ]. lilipuna

would differ in: the sharp cephalic lobe, the poorly

angular coxa 1, the moderately long bilobed coxa 2 in

terminal male, the absence of a small cusp on the

posteroventral corner of pleonal epimeron 3, the palm

of gnathopod 1 lacking angular definition, the smallness of female gnathopod 2, the weakly expanded and

poorly produced article 4 of pereopods 1-2 (in /.

falcata article 4 is so produced as to cover the anterior

edge of article 5), and the longer peduncle of uropod

3.

The extremely small size, smooth shiny body, and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

overall appearance of /. lilipuna suggest very little

direct relationship with /. falcata, but indicate that

/. lilipuna and /. falcata have had diverse phylogenies

and simply appear together into the same genus as a

result of convergence in a few characters.

Californian populations of /. falcata have a partially

immersed but articulate spine on the apex of the outer

ramus on uropod 3, a character necessarily seen with

extremely high power microscopy. Its worldwide presence in all populations of all species of Jassa has not

been documented, but it may form an important character that would demonstrate the discontinuity between /. lilipuna and /. falcata and possibly necessitate the removal of /. lilipuna from Jassa.

DISTRIBUTION.—Hawaiian Islands.

Parajassa angularis Shoemaker

FIOURES 131, 132

Parajassa angularis Shoemaker, 1942, pp. 41-44, figs. 14, 15.-

J. L. Barnard, 1962a, p. 58, figs. 26, 27.

Described from Magdalena Bay, Baja California by

Shoemaker and recorded by Barnard from the northern margins of southern California, this species extends

the full length of the northeastern Pacific warmtemperate province. The specimens described from one

end of the range differ from those at the other end

significantly, but it is too early to suggest that these

extremes, or those populations from Hawaii, deserve

any nomenclatural distinction. Specimens from Hawaii

fit those from the northern end of the warm-temperate

more than those from the southern end. A cline may

occur in that range and both northern and Hawaiian

specimens are slightly smaller than that specimen figured by Shoemaker from the southern warm-temperate

locality.

The Magdalenian types have the palmar defining

tooth on gnathopod 2 occurring much farther toward

the proximal end of article 6 and standing more

erectly at right angles to the axis of the appendage than

in Californian specimens. Most Hawaiian individuals

resemble Californian specimens in this character, but

a few of the largest (3.5 mm) have that tooth approaching the Magdalenian condition. Magdalenian

specimens also have a distinct scalelike tooth on the

apex of the outer ramus of uropod 2, not seen in other

populations.

Ramal apices of uropod 3 in California specimens

have one hooked spine each, whereas in Hawaiian
NUMBER 34

203

FIGURE 131.—Parajassa angularis Shoemaker, male, 2.1 mm, JLB Hawaii 2: a, lateral view of

body. Female, 3.2 mm: b, c, outer and inner rami of uropod 3; d, e, pereopods 3,5;/ , head and

epistome-labral process (stippled behind antenna 2 base); g, apex of pereopod 3; h, telson and

uropod 3; i, accessory flagellum on article 3 of antenna 1; ;", pereopod 2. Anotfier specimen: k,

apex of inner ramus on uropod 3.

specimens the outer ramus has 2 nonarticulate hooks

and 2 minute setules, and in Magdalenian material

the outer ramus has 3 small nonarticulate hooks.

The inner ramus of Hawaiian specimens has a jewellike spine nearly immersed and coalesced with the apex,

and some specimens seem to have only a pit that fits

the shape of that jewel spine; possibly the spine has

been lost during preservation. The inner ramus of

Magdalenian specimens has a thin apical spinule like

Californian specimens.

360-034 O—70 14

COLOR IN 2-DAY FORMALDEHYDE.—Antennae clear

but reflecting brown of body; head and pereonites 1-3

purple, remainder of body brown or gold; gnathopod

2 mostly covered with brown pigment; eggs pale buff.

MATERIAL.—JLB Hawaii 2 (16), 8 (7).

HAWAIIAN FAUNISTICS.—This species and Ventojassa ventosa were not suspected to be present in the

Hawaiian fauna prior to this study. They are enigmatic. Ventojassa ventosa, heretofore put in the

Isaeidae as a member of Eurystheus, is now shown to
204

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 132.—Parajassa angularis Shoemaker, female, 3.2 mm, JLB Hawaii 2: a, urosome; b,

mandibular molar enlarged and flattened to show molar flake; c, apex of gnathopod 2, medial; d,

dactyl of gnathopod 1; e, dactyl of gnathopod 2; /, apex of gnathopod 1; g, mandibular palp;

h, dactyl of pereopod 5.

be a monotypic ischyrocerid with unknown relationships, but Parajassa angularis belongs to a genus with

4 species all previously recorded from nontropical

shores (Europe, South Georgia Islands, Tristan da

Gunha, and the Californias). Unless Parajassa has been

grossly overlooked in Indo-Pacific tropical waters, it

should mark the cool water influence in the Hawaiian

fauna. Cool waters would include those of warmtemperate regions in this context.

Parajassa angularis and Ventojassa ventosa resemble

each other strongly in color and superficial morphology

and occur together in gross samples; it is of considerable interest that together they have crossed the wide

ocean gap between North America and Hawaii. The

direction of this movement is, of course, unknown and

could be of complex status. Hawaii could possibly be

the evolutionary ground for one of the species which

then returned to the mainland—or perhaps both—

and finally there is a possibility that Ventojassa ventosa is a product of the parajassid line of evolution;

they are the only ischyrocerids known to me with any

uropodal setae, although Ventojassa is a poor example

in having only 1-3 week setae on the outer ramus of

uropod 3.

DISTRIBUTION.—Hawaii and warm temperate

California-Mexico.

Ventojassa, new genus

DIAGNOSIS.—Ischyrocerid with normal-sized overlapping coxae, coxa 1 not reduced, coxa 5 as long as

4 and much longer than coxa 6; gnathopod 2 larger

than 1; pereopods 3-5 simple, second articles ovate,

trapezoidal or slightly broadened, not narrowly linear;

fourth articles of approximately equal breadth; palp

articles 5 and 6 gnathopod 1 subequal in length; outer

plate of maxilla 1 with 9 or more spines; accessory

flagellum elongate, 2 or 3-articulate; gnathopod 2

similar in both sexes with oblique palm defined by a

cusp and with 1-2 other palmar teeth, and 2 large

palmar spines, but not like that of Jassa male with

enormous proximal palmar tooth; outer ramus of
NUMBER 3 4 205

uropod 3 in adults with 1-3 mediomarginal setae,

apex slightly hooked, and bearing 2 (occasionally 3)

immersed setules.

TYPE-SPECIES.—Eurystheus ventosa J. L. Barnard

(1962a).

RELATIONSHIP.—In juvenile stages, the unique

species of this genus resembles Ischyrocerus Kroyer

and Jassa Leach mainly in uropod 3, but in adults the

outer ramus acquires 1-3 marginal setae not found in

the other 2 genera. Second gnathopods of males are

intermediate between those of Ischyrocerus and Jassa,

and the genus has so many resemblances to Parajassa

Stebbing, especially to P. angularis Shoemaker (1942),

that one might consider them as morphological mimics.

Yet a major character, the vestigial accessory flagellum,

distinguishes Parajassa, and numerous fine details distinguish P. angularis from V. ventosa. The latter

species is transferred from the family Isaeidae. At the

time of its establishment, J. L. Barnard considered it

probable that E. ventosa should be ranked with the

Ischyroceridae, and a study of uropod 3 and the mandibles now indicates its proper position.

Ventojassa ventosa (J. L. Barnard)

FIGURES 133, 134

Eurystheus ventosa J. L. Barnard, 1962a, pp, 20-22, figs. 6, 7.

DIAGNOSIS.—With the characters of the genus.

DESCRIPTIVE NOTES (and see figures).—Lateral

cephalic lobes short but triangular, appearing superficially rounded, but anterior margin with sharp flangelike extension becoming sharper in adults; accessory

flagellum either 2 or 3-articulate, terminal article very

small; gland cone long and sharp; epistome sharp and

projecting, of medium length; mandibular molar with

typical ischyrocerid flake; article 6 of gnathopod 1

strongly expanded distally; gnathopod 2 palm with 3

short teeth, 1 tooth defining palm; second articles of

pereopods 1-2 slender, article 4 slender and unproduced anterodistally, anterior margin of article 2 on

pereopod 3 scarcely setose, article 6 with 2 or 3 large

spines or sets of spines near dactyl; epimeron 3 with

strong posteroventral notch; uropods 1-2 without

lateral peduncular setae; lower lip, maxilla 2 and

maxilliped like Ischyrocerus anguipes Kroyer (Sars,

1895, pi. 209), inner plate of maxilla 1 with 1 seta;

sexes alike, gnathopod 2 of female large like that of

male; juveniles 1.0-1.5 mm long, lacking setae on

outer ramus of uropod 3; telson usually with only 1

pair of lateral hooks even in adult but occasional specimens with 2 pairs or 1 on one side, 1 pair on other.

COLOR IN 2-DAY FORMALDEHYDE.—Antennae reddish-pink; head and pereonites 1-3 muddy purple,

remainder of body bronze but fading toward posterior

end; gnathopod 2 white, with sparse splotches of

brown; embryos in eggs orange; eyes dark brown or

black, changing to definite black in alcohol.

MATERIAL.—JLB Hawaii 2 (69), 3 (7), 5 (3), 8

(14).Devaney 1 (1).

REMARKS.—The descriptive notes distinguish this

species primarily from Parajassa angularis Shoemaker

(1942), a sympatriot both in Hawaii and Baja California. The two species resemble each other so much

in overall appearance that one might consider them

"mimics" if there were not significant morphological

differences and color differences. These morphological

differences are difficult to see with a stereoscope during

the sorting process, but individuals mounted in a depression slide may be rapidly identified by noting the

absence of peduncular setae on uropods 1-2 of Ventojassa, the sparsity of setae on pereopod 3, the thin

article 2 of pereopods 1-2, and the distinctive setae (in

adults) on the outer ramus of uropod 3. Lateral cephalic lobes, gnathopods, accessory flagella, primary

flagella, and pleonal epimeron 3 also differ between the

two species; but pereopods and antennae are often

broken.

Hawaiian specimens of this species reach about 3.2

mm in length; adults from California as large as 4

mm have been reported; those adults have 3 pairs of

telsonic hooks. No gross differences from Californian

material occur in Hawaiian specimens, but careful

comparison of microscopic details may show consistent

differences in the apex of the outer ramus of uropod 3,

the notch of pleonal epimeron 3, and the cuspidorial

symmetry on the palm of gnathopod 2.

DISTRIBUTION.—Hawaiian Islands, and warm-temperate California.

Family LEUCOTHOIDAE

Leucothoe hyhelia J. L. Barnard

FIGURE 135

Leucothoe hyhelia J. L. Barnard, 1965a, pp. 489-490, fig. 5.

DIAGNOSIS.—Article 3 of antenna 1 about one-third

as long as article 1, article 1 with obsolescent terminal
206

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 133.—Ventojassa ventosa (J. L. Barnard), ovigerous female, 2.8 mm, JLB Hawaii 2:

a, left gnathopod 1; b, dactyl of gnathopod 1 enlarged and palmar serrations enlarged; c, body;

d, medial pereopod 3; e,f, dactyls of pereopods 3, 5; g, apex of gnathopod 2, enlarged, medial,

and extra dactyl showing lateral view. Ovigerous female, 2.6 mm: h, urosome. Specimen, 2.5

mm: i, inner ramus of uropod 3.
NUMBER 3 4 207

FIGURE 134.—Ventojassa ventosa (J. L. Barnard), ovigerous female, 2.8 mm, JLB Hawaii 2: a,

pereopod 1; b, mandibular molar showing flake; c, lateral cephalic lobe and eye; e, telson; /,

telsonic edges enlarged; g, right hook of telson; h, part of pereopod 5; i, mandibular palp. Female,

3.2 mm: d, outer ramus of uropod 3. Another specimen: ;, telson and uropod 3; I, outer ramus of

uropod 3. Ovigerous female, 2.6 mm: k, epistome and upper lip, left side.
208

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 135.—Leucothoe hyhelia J. L. Barnard, male, 2.5 mm, JLB Hawaii 3: a, lateral view of

body; b, article 1 of antenna 1, ventral; c, medial gnathopod 2; d, head; e, medial gnathopod 1;

/, telson. Male, 2.6 mm, JLB Hawaii 5: g, h, pleonal epimera 1,3.
NUMBER 3 4 209

cusps; rostrum of medium development, midanterior

cephalic keel rounded ventrally, lateral cephalic lobe

broadly rounded; eyes of medium size, round; coxa 1

with beveled anteroventral corner, coxae 2-4 longer

than wide, anteroventral corners decreasingly beveled,

posterior margin of coxa 4 long and deeply excavate

dorsally, ventral margin sloping obliquely downward;

gnathopod 1 of typical form, with long dactyl, posterior

margin of article 6 with wavelike serrations 3 times

as large as small round beads of apposing margin on

chela of article 5, article 6 otherwise bearing 3-6

marginal setules, article 5 otherwise smooth, gnathopod

2 of typical form, palmar margin of nearly smooth, untoothed variety, weakly undulate near finger hinge, anteromedial face with long row of setae; pereopods 1-2

simple; article 2 of pereopods 3-5 broadened, proximally commencing to be ovate but posteroventral margins strongly beveled and oblique; pleonal epimeron

1 with rounded-quadrate posteroventral corner bearing

2 weak notches in young individuals or 2 large serrations in terminal adults, epimeron 2 with strong sharp

tooth, epimeron 3 of two kinds, posterior margin either

weakly convex and posteroventral corner roundedquadrate (as in L. tridens Stebbing, herein) or with

straight-convex posterior margin and slight posteroventral rounded protrusion; uropod 2 inner ramus extending to end of peduncle on uropod 3; telson elongate, triangular, apex with obsolescent lateral wings

of a trilobation.

MATERIAL.—JLB Hawaii 2 (1), 3 (4), 5 (2), 8 (2),

10 (2), 11 (1), 13 (20), 14 (1), 15 (1). Devaney 1

(2).

COLOR IN 2-DAY FORMALDEHYDE.—Either albinid

with red eye, or bearing narrow vertical stripes at segmental junctions from head-pereonite 1 to pleonites

2-3, these stripes fading rapidly in alcohol, and eyes

fading to a rusty pink central core surrounded by clear

ommatidia.

REMARKS.—The Hawaiian populations have at least

2 phenotypes, those with a protuberant posteroventral

corner of epimeron 3 or perfectly rounded-quadrate.

Three differences of possible subspecific value occur in

the Hawaiian material in comparison with the unique

male type known from Micronesia but until Micronesian populations are more thoroughly studied I reserve

any subspecific appellation. The Micronesian holotype

has a shorter dactyl on gnathopod 1, a slightly stronger

cusp on epimeron 1, and faint setules on the chela of

gnathopod 1. Gnathopods of this species and the otherwise very different L. tridens are virtually identical.

The diagnoses demonstrate the divergencies in head,

anterior keel, coxae, pereopods 3-5, and epimeron.

DISTRIBUTION.—Micronesia and the Hawaiian

Islands.

Leucothoe lihue, new species

FIGURE 136

DIAGNOSIS.—Article 3 of antenna 1 about one-third

as long as article 1, article 1 with obsolescent terminal

cusps; rostrum small, midanterior keel with small,

sharp cusp at ventral end, lateral cephalic lobe obtusely

angular; eyes medium to large, round, red in formaldehyde; coxa 1 scarcely beveled anteriorly, coxae

2-4 longer than wide, anteroventral corners not beveled, posterior margin of coxa 4 long and deeply

excavate dorsally, ventral margin sloping obliquely

downward; gnathopod 1 of stout form, short, article 6

only about 130 percent as long as anterior margin of

article 5, dactyl very short, apposing margins of articles

5 and 6 macroscopically smooth; gnathopod 2 with

widely cusp-shaped article 6 tapering strongly toward

distal end, palm with several sinuations and bicastellate

distal process, dactyl apically blunt and slightly bent

distally, lobe on article 5 with villus-like apical extension, article 2 with mammilliform anteroventral hump,

anteromedial face of article 6 with long row of setae;

pereopods 1-2 simple; article 2 of pereopods 3-5

broadened, proximally commencing to be ovate but

posteroventral margins, especially of pereopods 4-5,

strongly beveled and oblique, margin of pereopod 5

concave; pleonal epimeron 1 with rounded-quadrate

posteroventral edge bearing 2 widely spaced weak serrations; epimeron 2 with strong, sharp tooth, epimeron

3 rounded-quadrate; (uropod 3 missing, thus*relationships of uropod 2 unknown) ; telson elongate, triangular, apex minutely trilobate.

HOLOTYPE.—Bishop Museum collections, catalog

number 7289, male, 3.3 mm.

TYPE-LOCALITY.—Devaney 1, off MokuManu,

Oahu, 33 m, stem of black coral, 13 May 1967.

MATERIAL.—Five specimens from the type-locality.

RELATIONSHIP.—In Hawaii, this species is clearly

distinguished from the other 2 known species in the

form of gnathopod 1 with short articles 6 and 7.

Leucothoe lihue is very close to L. alata J. L. Barnard (1959) from California and may exist as a sub-
210

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 136.—Leucothoe lihue, new species, holotype, male, 3.3 mm, Devaney 1: a, head; b,

coxa 4; c, apex of telson; d, telson; e, lateral cephalic lobe (hatched) and epistome-labrum; /, g,

gnathopods 1, 2; h, maxilliped; i, j , k, pereopods 5, 3, 4; /, left mandibular incisor; m, maxilla

2; n, article 1 of antenna 1; o, apex of gnathopod 1, enlarged; p, pleonal epimera 2—3.
NUMBER 34 211

species of that entity. There are several differences in

the Hawaiian population of a quantitative extent in

character expression and one character probably of

qualitative value. The article bearing the outer plate

on the maxillipeds is not alate as it is in L. alata. Other

character differences of minor extent in L. lihue are:

the cephalic lobe is slightly more angular; pleonal

epimeron 2 is slightly more protuberant at the posteroventral corner; coxa 1 is less strongly beveled; article 6

of gnathopod 1 lacks the minute beads on the posterior

margin; and article 2 of pereopod 5 is broader and

distinctly concave on the posteroventral oblique

margin.

Leucothoe minuscula Schellenberg (1938) has a

grossly ornamented palm on gnathopod 2, coxa 1 is

obtusely pointed anteroventrally, and article 2 of

pereopod 5 is not excavate.

Leucothoe brevidigitata Miers (see L. flindersi Stebbing, 1888) has a very large mandibular palp and

narrow incisor, and artid** 3 of antenna 1 is half as

long as article 1.

DISTRIBUTION.—Hawaiian Islands.

Leucothoe tridens Stcbbing

FIGURE 137

Leucothoe tridens Stebbing, 1888, pp. 777-779, pi. 47; Schellenberg, 1938, pp. 21-23, fig. 11.

DIAGNOSIS.—Article 3 of antenna 1 about 35 percent as long as article 1, article 1 with strong terminal

cusps; rostrum obsolescent, midanterior cephalic keel

sharply pointed at ventral end, lateral cephalic lobe

with sinuous anterior margin and weak to strong anterior cusp, becoming better developed in terminal

adults; eyes of medium size, round; coxa 1 with

rounded anteroventral corner, scarcely beveled, coxae

2-4 wider than long, short, anteroventral corners of

coxae 2 and 4 almost quadrate and slightly attenuate,

corner of coxa 2 rounded, posterior margin of coxa 4

short and nonexcavate, ventral margin sloping

obliquely upward; gnathopod 1 of typical form, with

long dactyl, posterior margin of article 6 with wavelike

serrations 3 times as large as small round beads of apposing margin on chela of article 5, article 6 otherwise

bearing 3-6 marginal setules, article 5 otherwise

smooth; gnathopod 2 of typical form, palmar margin

of nearly smooth, untoothed variety, weakly undulate

near finger hinge, anteromedial face with long row of

setae; pereopods 1-2 simple; article 2 of pereopods 3-5

evenly ovate; pleonal epimeron 1 with rounded-quadrate posteroventral corner, epimeron 2 with roundedquadrate or slightly protuberant corner (variable),

epimeron 3 with weakly convex posterior margin and

rounded-quadrate posteroventral corner; uropod 2,

inner ramus extending only two-thirds to three-fourths

along peduncle of uropod 3; telson of medium length,

ovatoconiform, with weak terminal trilobation.

MATERIAL.—JLB Hawaii 6 (9), 10 (1), 12 (1),

13(3).

COLOR IN 2-DAY FORMALDEHYDE.—Body a diffuse

burnt orange or golden (fading), eye red to bright

garnet.

REMARKS.—The Hawaiian populations of this

species differ from those of the Gilbert Islands and

Philippine Islands (Schellenberg, 1938) and the typematerial (Stebbing, locality presumably erroneous) by

the strongly projecting middle member of the trilobation on the telson.

DISTRIBUTION.—Tropical Pacific.

Leucothoides ? pott si Shoemaker

FIGURE 138

?Leucothoides pottsi Shoemaker, 1933a, pp. 249-250, fig. 3.—

Schellenberg 1938, pp. 26-28, fig. 13.

MATERIAL.—JLB Hawaii 2 (4), 3 (3), 6 (1), 8 (1),

10 (6), 13 (6), 14(1).

REMARKS.—Described from Tortugas Islands in the

Caribbean Sea, this species was reported by Schellenberg from the Gilbert and Marshall Islands in Micronesia. Schellenberg's figures and description correspond

to those of Shoemaker in head, coxae 1, 2, 4, article 6

of gnathopod 2, and vaguely in gnathopod 1. But the

anterior margin of article 5 on gnathopod 2 is longer

and the posterior lobe of article 5 stouter in Micronesian material than in the type-material. Antennae of

Micronesian specimens are subequal in length. Schellenberg did not clearly illustrate the details of the

carpal process of gnathopod 1 and Shoemaker may not

have clearly delineated the submergence of the distal

spines. In material at hand from Hawaii, the apex of

that carpal process has 2 partially immersed spines, one

long and one medium in length; the long spine has a

distal ornament and both are minutely serrate on one

edge. Shoemaker illustrated a distal ornament on a

short spine, but a declivity on the carpal process in his

drawing indicates the possible boundary of the immersed spine. Schellenberg's drawing was not sum-
212

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

d

FIGURE 137.—Leucothoe tridens Stebbing, male, 4.1 mm, JLB Hawaii 6: a, head and 5 pereonites; b, c, d, e, left coxae, 1, 2, 3, 4. Another specimen: /, head; g, maxilla 2; h, ij, pereopods 3,

4, 5; k, telson; /, articles 3-4 of antenna 2, medial; m, article 1 of antenna 1, ventral; n,

flattened view of fig. 1; o, p, pleonal epimera 2, 3. Male, 4.0 mm, JLB Hawaii 12: q, head.

Adult of JLB Hawaii 10: r, pleonal epimeron 2.
NUMBER 34

213

FIGURE 138.—Leucothoides Ipottsi Shoemaker, 1.3 nun, JLB Hawaii 3: a, apex of pereopod 1;

b, uropod 3; c, apex of gnathopod 1, articles 6 and 7; d, apex of gnathopod 1, article 5; e, f,

gnathopods 1,2.

ciently enlarged to show any of these details. Hawaiian

specimens have the fifth article of gnathopod 2 elongated like that of Micronesian material.

Comparison of minute details among the three populations, Caribbean, Micronesian, and Hawaiian probably would demonstrate differences of subspecific

magnitude at least. Hawaiian specimens, unlike those

from the Caribbean and Micronesian areas, have the

distally attenuate portion of coxa 1 broadened.

DISTRIBUTION.—Caribbean Sea, Micronesia, Hawaiian Islands.

Family LILJEBORGIIDAE

Liljeborgia heeia, new species

FIGURES 139, 140

DIAGNOSIS.—Eyes large, black; mandibular palp

article 3 shorter than article 1 and half as long as article 2, article 1 shorter than article 2; lateral cephalic

lobe long, apically truncate; coxa 1 with small posteroventral notch, coxa 4 projecting ventrally more than

in other species of genus, bearing 2—3 posterior serrations; palms of gnathopods evenly convex; dactyl of

gnathopod 1 with 3-5 basal teeth, dactyl of gnathopod

2 with 8-14 teeth, gnathopod 1 of medium size for

genus; pereopods 1-2 with subprehensile article 6 bearing small palm with weak cusp, posterodistal ends of

sixth articles with 2—4 spines; article 2 of pereopods

3-5 weakly serrate posteriorly, posteroventral corners

smooth and rounded; telson split nearly to base, each

apex with short notch, subapices asymmetrically extended, medial longer than lateral; medial -edge of

peduncle of uropod 1 with only 1 distal spine; only

pleonites 1-2 each with 3 small dorsal cusps; pleonal

epimera 1-3 each with small posteroventral tooth, no

distinct accessory sinuses.

DESCRIPTION.—Outline of eye truncate anteriorly,

nearly quadrate below, tapering dorsally by postero-
214

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 139.—Liljeborgia heeia, new specie*, female, 5.3 mm, JLB Hawaii 6: a, apex of left

mandible; b, apex of one telsonic lobe. Holotype, male, 4.2 mm: c, spines of gnathopodal palms,

enlarged; d, dorsal margin of plconites 1-2; e, f, outer and inner plates of maxilliped; g, telson;

h, right medial articles 1-3 of antenna 1, right to left; i, outer plate of maxilla 1; ;, maxilla 2; k,

lateral view of body; /, portions of dactyl and palm on gnathopod 2, enlarged; m, apex of pereopod

1; o, lateral cephalic lobe (shaded) and prebuccal complex.
NUMBER 3 4 215

FIGURE 140.—Liljeborgia heeia, new species, holotype, male, 4.2 mm, JLB Hawaii 6: a, article 6

of gnathopod 2, enlarged; b, e, d, pleonal epimera 1, 2, 3, right outlines; e, articles 6-7 of pereopod

5; f, g> h, uropods 1, 2, 3. Ovigerous female, 3.8 mm, JLB Hawaii 5: i, j , gnathopods 1, 2.

Ovigerous female, 5.3 mm, JLB Hawaii 2: k, lower lip; /, mandible; m, head.
216

dorsal curve; rostrum evenly acute; maxillipedal palp

and maxilla 1 like those of L. brevicornis (Bruzelius)

(Sars, 1895, pi. 187) except for the outer plate of

maxilla 1 figured herein; maxillipedal palp article 4

with weak distal nail; dactyl of gnathopod 2 with weak

lateral comb-row; midfacial palmar defining area of

gnathopod 1 with 1-2 spines, of gnathopod 2 with 2

spines; pereopods 3-5 with numerous setae on article

6 in large individuals (5.0 mm), but setae sparse or

absent on individuals 4 mm or smaller.

HOLOTYPE.—Bishop Museum collections, catalog

number 7290, male, 4.2 mm.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, coralline footballs, 29 January 1967.

MATERIAL.—JLB Hawaii 2(7), 3(10), 5(6), 6(9),

8(2),10(1),11(4),13(1).

RELATIONSHIP.—This is the only species so far

described in the genus with 3 dorsal denticules each

on pleonites 1-2 and no denticles on other pleonites.

Apart from dentations this species has affinities with

L. proxima Chevreux (1908, and see Schellenberg,

1938) from Gambier Archipelago and the Marshall

Islands. That species has one denticle each on pleonites

1-2 and 4, a distinct sinus above the tooth of epimeron

3, no tooth on epimeron 1, and equally extending subapices of the telson.

DISTRIBUTION.—Hawaiian Islands.

Liljeborgia laniloa, new species

FIGURE 141

DIAGNOSIS.—Eyes large, black; rnandibular palp

article 3 very slightly longer than article 1; lateral

cephalic lobe long, apically truncate; coxa 1 with small

posteroventral notch, coxa 4 projecting ventrally more

than in other species of genus, bearing 3 posteroventral

serrations; palms of gnathopods evenly convex, dactyl

of gnathopod 1 with 3-4 basal teeth, dactyl of gnathopod 2 with 6-7 teeth, gnathopod 1 large, dactyls of

gnathopods apically flagellate; pereopods 1 and 2

simple, posteroventral end of article 6 armed with 7-9

thin short setae; article 2 of pereopods 3-5 weakly serrate posteriorly, posteroventral corners smooth and

rounded; telson slit nearly to base, each apex with short

notch, subapices asymmetrically extended, lateral longer than medial; medial edge of peduncle of uropod 1

with only 1 distal spine; only pleonites 1-2 each armed

with 3 small dorsal cusps; pleonal epimera 1-3 each

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with 1 small posteroventral tooth, that tooth on epirneron 3 forming part of deep sinus.

DESCRIPTION.—Outline of eye circular; rostrum

evenly acute; mandible with spines reduced to 3 serrations, molar apparently absent; maxillipedal palp article 4 with very tiny distal nail; outer plate of maxilla 1

with 7 spines like L. heeia, new species, but all spines

of equal length; comb-row on dactyl of gnathopod 2

very weak; midfacial palmar defining area of gnathopod 1 with no spines, of gnathopod 2 with 3 spines;

articles 2 and 4 of pereopods 3-5 slightly broader than

those of L. heeia; outer rami of uropods 1-2 with only

1 spine, outer ramus of uropod 2 slightly shorter than

inner ramus, rami of uropod 3 very slender.

HOLOTYPE.—Bishop Museum collections, catalog

number 7291, male 2.3 mm.

TYPE-LOCALITY.—JLB Hawaii 15, 1 mile north of

Kualoa Point, Oahu, intertidal, block of ascidians, 23

May 1967.

MATERIAL.—Two specimens from the type-locality.

RELATIONSHIP.—This species has sculpture similar

to that of Hawaiian L. heeia but differs in the nonpalmate pereopods 1-2, in the deeper sinus of pleonal epimeron 3, in the terminally flagellate dactyls of the

gnathopods, the slender rami of uropod 3, and the circular eyes. Like L. heeia, this species also resembles

L. proxima Chevreux (1908), especially in epimeron

3, the slender rami of uropod 3, and in pereopods 1-2

but differs from L. proxima by the longer article 6 of

gnathopod 1, the flagellate gnathopodal dactyls, and

the presence of teeth on the dactyl of gnathopod 1.

The mandible, with its very weak spine row and

absence of molar, differs from any other species of

Liljeborgia so far described.

DISTRIBUTION.—Hawaiian Islands.

Family LYSIANASSIDAE

Lysianassa ewa, new species

FIGURES 142, 143

DIAGNOSIS OF FEMALE.—Lateral cephalic lobe

evenly conical; eyes medium in size; antennae short;

upper lip forming medium-sized, apically rounded

plate projecting in front of epistome and separated

from it by short slit; mandibular molar composed

mainly of clusters of sharp blades, very spinose, but

remainder of surface around molar not setulose; inner

plate of maxilla 1 long, ovovermiform, naked, outer
NUMBER 34 217

k

FIGURE 141.—Liljeborgia laniloa, new species, holotype, male, 2.3 mm, JLB Hawaii 15: a, b,

gnathopods 2, 1, medial views; c, d, parts of pereopods 3, 5; e, f, g, pleonal epimera 1, 2, 3; h,

head with outline of eye; i, right and left mandibular incisors and spine rows, left to right; ;,

uropod 3; k, apex of pereopod 1, enlarged.

plate with set of 3-4 small curved bifid spines and set

of 6 or 7 large serrate spines; outer plate of maxilla

2 narrower than inner, but inner not grossly expanded,

both lobes with very stout spine-setae; outer plate of

maxilliped with weak rugae and marginal beads, with

one spine-seta on medial edge, dactyl on palp of normal dimensions; posterior margin of article 6 on gnathopod 1 bearing 4 stout spines in line; dactyl of gnathopod 2 covering half of distal end of article 6; locking

spines of pereopods simple; posterior margins of article

2 on pereopods 5-7 straight, unexcavate; pleonal epimera 1 and 3 with rounded-quadrate posteroventral

corners, epimeron 2 with slight protrusion; constriction on inner ramus of uropod 2 of medium size; uropod 3 short, rami and peduncle equal to each other,

peduncle platelike, rami uniarticulate; telson truncate,

largest plumose setal pair near midlateral margins, a

pair of weak subterminal spines at lateral margins,
218

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 142.—Lysianassa ewa, new species, holotype, female, 3.6 nun, JLB Hawaii 13: a, body;

b, inner plate of maxilla 1; c, pleonal epimera 1, 2, 3; rf, palp of maxilla 1; e, uropod 3; f, inner

and outer plates of maxilliped; g, dactyl of pereopod 5; h, maxilla 2; i, medial view of gnathopod

1. Male, 3.3 mm: j , uropod 3.
NUMBER 34 219

FIGURE 143.—Lysianassa ewa, new species, holotype, female 3.6 mm: a, urosome; b, outer plate

of maxilla 1; c, d, right and left mandibles; e, telson; /, head; g, palp of maxilliped; h, dactyl of

pereopod 1; i, apex of gnathopod 2. Male, 3.3 mm: ;*, rami of uropod 3, flattened.

360-034 O - 15
220

those margins forming lateral rim, telson thus cupshaped.

MALE.—Eyes slightly larger than in female, flagellum of antenna 2 elongate, rami of uropod 3 thickened

and bearing long setae.

COLOR IN 2-DAY FORMALDEHYDE.—Body white, eyes

brown.

HOLOTYPE.—Bishop Museum collections, catalog

number 7292, female, ovigerous, 3.6 mm.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

Oahu, 3-4 m, corals, algae, corallines, 8 April 1967.

MATERIAL.—JLB Hawaii 2 (1), 3 (7), 12 (2), 13

RELATIONSHIP.—This species has close affinities

with Indonesian Lysianassa (=Arugella) heterodonta

(Pirlot, 1936), but differs by the slightly tuberculate

and unragged outer plate of the maxilliped, apparently

by the longer dactyl of the maxillipedal palp, the

slightly stronger spines in the small set on the outer

plate of maxilla 1, the truncate (not rounded) telson,

the presence of a slight protrusion on pleonal epimeron

2, and the unexcavate posterior margin on article 2

of pereopod 4.

The Caribbean L. cubensis (Stebbing, 1897; see

Shoemaker, 1935a and Hurley, 1963) is another closely

related species but L. ewa differs from it in the slightly

larger dactyl of gnathopod 2, the more ovate inner

plate of maxilla 1, the slightly narrower inner plate

of maxilla 2, the unexcavate posterior margin on article 2 of pereopod 4, the more weakly developed inner

ramus on uropod 2, the more strongly spinose mandibular molar, the absence of tiny spinules on the outer

plate of the maxilliped, the slight protrusion on pleonal

epimeron 2, and the presence of stout spines on article

6 of gnathopod 1. Lysianassa cubensis differs from

L. heterodonta mainly in the truncate telson. These

three species form a cohesive triad differing from

other members of the genus in gross characters. For

instance, the other tropical members of the genus have

an elongated peduncle on uropod 3 and L. holmesi

(J. L. Barnard, 1955b), the next nearest allopatriot to

L. cwa, has equally wide plates on maxilla 2, a biarticulate outer ramus on uropod 2, and a slightly excavate

telson with its armament all terminal. The lower lip

of L. ewa resembles that of L. holmesi.

DISTRIBUTION.—Hawaiian Islands.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Family OGHLESIDAE

Ochlesis alii, new species

FIGURES 144, 145

> Ochlesis innocens.—Pirlot, 1936, pp. 298-300, fig. 124 [not

Stebbing, 1910]

DESCRIPTION.—Body strongly compressed, very thin

and carinate dorsally, pereonite 7 with small but sharp

posterodorsal tooth, pleonites 1-2 dorsally elevated

posteriorly but not sharply, pleonite 3 with obtuse

dorsal hump, pleonite 4 elongate, pleonites 4-6 all

separate; head short, telescoped into pereonite 1, latter

extended forward strongly over head but closely appressed to it, rostrum and lateral cephalic lobe equal

in extent and size, coniform, eye moruliform, partially

covered by thin chitin of pereonite 1; article 1 of

antenna 1 longer than 2, 1-2 each with short ventrodistal cusp, cusp of article 2 extending only halfway

along short article 3; accessory flagellum absent;

primary flagellum 4-articulate, only article 1 elongate,

flagellum armed with large aesthetascs; antenna 2

shorter and thinner than antenna \, article 4 with

mediodistal cusp extending more than halfway along

article 5, flagellum 3-articulate, only article 1 elongate;

gland cone small; upper lip and epistome amalgamated, but epistomal portion with small anterior keel;

upper lip almost perfectly triangular; styliform mandibles with simple, pointed incisor and small, 1-serrate

lacinia mobilis on left member, molar small, weakly

cup-shaped and scarcely triturative, palp articles 1-3

subequal to each other in length, palp 3 with 3 short

terminal setae and linear comb-row; lower lip slender,

apices extended and bidigitate; maxilla 1 with small

inner plate, outer plate styliform, apex with about 7

tooth-ridges, 2-3 reverted cusps and about 4 articulated

hooks, palp tiny, uniarticulate, bearing one long seta;

maxilla 2 with slender, unequal but strongly setose

lobes; inner plates of maxillipeds each with pair of

distolateral spine-setae on lateral notch beside acute

apex, one more small distal setule, several marginal

setae; outer plates of maxillipeds slightly curled orally,

distal ends with a few deeply immersed setules; coxae

1-2 with slight proximal excavation, coxa 3 strongly

produced forward at base, but remainder of anterior

margin straight, posterior margin with midproximal

hump fitting excavation in coxa 4, latter broad, with

rounded posteroventral process below proximal excavation, ventral margin oblique and concave (called

posterior margin by Stebbing, 1910), coxa 5 with ante-
NUMBER 34 221

FIGURE 144.—Ochlesis alii, new species, holotype, female, 2.3 mm, JLB Hawaii 3: a, lateral view

of body; b, half of lower lip; c, d, right and left mandibular molars; e, palp of maxilla 1; f,

urosome; g, outer plate of maxilliped; h, gill (G) and broad lamella (L) of pereopod 2; i, coxa

(G), gill (G), and brood lamella (L) of gnathopod 2; ;, left obverse mandibular apex; k, apex of

mandibular palp; /, left mandibular apex; m, right mandibular apex, flattened; n, right mandibular apex, unflattened.

360 -034 O - 16
222 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 145.—Ochlesis alii, new species, holotype, female, 2.3 mm, JLB Hawaii 3: a, right pleopod

1; b, maxilla 2; c, maxilla 1; d, telson and uropod 3; e, upper lip; /, maxilliped; g, apex of antenna

1; h, mandible; i, maxilla 1 apex; j , k, I, m, gnathopod 1; n, o, gnathopod 2.
NUMBER 3 4 223

riorly directed lobe and posterior lobe longer than

anterior; coxa 6 narrower than 5 and not anteriorly

produced; posterior margin of coxa 7 with deep,

oblique concavity; gnathopod 1 with article 2 basally

inflated, articles 3 and 4 subequally elongate, articles

5-6 elongate, but 5 longer than 6, latter with weak,

oblique palm armed with sharp medial cusp, one

immersed feather-seta at denning corner, one feathered

strap-seta near hinge; dactyl very complex, clawlike

basic portion nearly enveloped by its own 4 feathered

strap-setae; gnathopod 2 with subrectangular article

2, short article 3, elongate article 4 obliquely jointed

to article 5 and distoventrally extended and distally

truncate, article 5 ventrodistally produced (thus carpochelate) along article 6, but only weakly forming

a guard, setose, article 6 shorter than 5, simple, dactyl

stout, rigid, coniform; pereopods 1-2 with long anterodistal process on article 4 extending more than halfway along article 5 and arising from anterior lobe,

article 4 with narrow neck proximal to lobe, dactyls

simple but bearing cornified distal nail, article 6 with

a few posterior spinules; pereopods 3-5 similar in

structure of articles 3-7 as to pereopods 1-2 except

article 4 without proximal neck, second articles broadly

expanded, posterior margins increasingly excavate and

roughly serrate; pleonal epimera 1-2 with midposterior

extensions above slightly concave margins, posteroventral corners rounded but protruding, epimeron 3 with

very short posterior straight margin and medium-sized

posteroventral tooth with accessory ventral serration;

outer rami of uropods 1-3 slightly to strongly

shortened; telson linguitriangular, with ventral keel.

COLOR IN I-DAY FORMALDEHYDE.—Body and appendages with dull gray background and texture of

concrete, pereonites 1-7 with anterodorsal spots of

burgandy color, all other color spots marked in Figure

144 light orange in color, those spots (chromatophores) or diffuse stripes occurring on rostrum,

lateral cephalic lobes, antennae, anterior lobe only of

epistome, coxae 1, 2, 3, article 2 of pereopod 5, pleonites 1-3 laterally, stripes on articles 5-6 of pereopods

1-5, stripe on article 4 of pereopod 2, spot on article

6 of pereopod 5, eyes orange; antennal stripes occurring on distal halves of article 1 of flagella; other shading in figure represents shadows.

HOLOTYPE.—Bishop Museum collections, catalog

number 7293, female, 2.3 mm. Unique.

TYPE-LOCALITY.—JLB Hawaii 3, off Ewa Beach,

Oahu, 18 m, Pocillopora, bryozoans, 29 January 1967.

RELATIONSHIP.—If Pirlot's identification of O.

innocens from lies Aru, 13 m, proves to be erroneous,

and if he had in reality another species, perhaps the

one at hand, then O. alii differs from O. innocens not

only in the following external characters but also in

mouthparts. Ochlesis alii has a shorter article 2 on

antenna 1 and shorter cusps on articles 1 and 2 than

does O. innocens; the flagellum is longer; article 4 of

antenna 2 has a long cusp; coxa 3 is much more

strongly geniculate, coxa 4 is broader, shorter, deeply

excavate anteriorly, and the ventral margin less deeply

excavate than in O. innocens; coxa 7 has a deep posterior excavation; pleonite 3 is less strongly acute dorsally; the pereopodal dactyls are much shorter, the

telson lacks a ventral tooth but has a keel. Mouthparts

of O. alii differ from those of O. innocens as described

by Stebbing (1910) in the triangular, not lobulate

upper lip, the much broader extent over which the

ornaments of the first maxilla extend, and the shorter

but digitate processes of the lower lip. Stebbing may

have overlooked the lacinia mobilis and molar, and

there may be differences in the maxilliped not evident

from his small figure. The dactyl of gnathopod 2 in

O. alii seems much stouter and more rigid than that of

O. innocens.

Ochlesis lenticulosus K. H. Barnard (1940) from the

littoral of Simonstown, South Africa, has larger and

sharper dorsal teeth on pleonites 1-3 and a smaller

epimeron 3 than does O. alii. Coxae 1-6 of O. lenticulosus resemble those of O. alii, but coxa 4 is less strongly excavate anteriorly. Coxa 7 is not posteroventrally

excavate in O. lenticulosus; the rostrum is thinner, the

cephalic lobe blunter, and gnathopod 1 has a simpler

morphology than in O. alii. Article 5 of gnathopod 1

is longer than 6 in O. lenticulosus, the lower lip is not

apically bidigitate on either lobe, pereopod 1 has a

proximal anterior lobe on article 4 apart from the

normal lobe, and article 2 of pereopod 5 has a distal

mammilliform lobe projecting posteriorly. Apparently,

articles 1-3 of antenna 1 are equal in length, articles

1-2 are scarcely dentiform, the flagellum is very short,

and article 4 of antenna 2 is not dentiform in O.

lenticulosus. No palp occurs on maxilla 1 and apparently neither mandible has a lacinia mobilis in

O. lenticulosus.

Ochlesis alii differs from O. levetzowi Schellenberg

(1953) by the strong antennal cusps and the tooth of

pleonal epimeron 3.

DISTRIBUTION.—Hawaiian Islands.
224

Family OEDICEROTIDAE

Kanaloa, new genus

DIAGNOSIS.—Head very massive, with short, stout

rostrum, eyes large, yellowish-orange in life, contiguous,

occupying dorsal part of head and separated from each

other by minute raphus; antennae short, antenna 1

reaching about halfway along flagellum of antenna 2,

article 3 of antenna 1 more than half as long as article

1; mandibular incisor slightly projecting and with 1 or

2 teeth, molar large, mostly smooth but edges triturative, palp of medium size, articles 2-3 subequal in

length, article 3 falcate; inner lobes of lower lip free;

inner plate of maxilla 1 medially setose; maxillipedal

palp very stout, scarcely exceeding outer plate, article 4

short, tumid, apically acute, bearing small subapical

spine(s) ; coxa 1 of different shape than coxa 2, expanded midanteriorly, coxa 4 strongly excavate posterodorsally but midposterior lobe not strongly or acutely

extended, coxa 5 much shorter than coxa 4; gnathopods

1-2 small, similar to each other in size and general

morphology, but gnathopod 2 distinctly larger than 1,

palms transverse, article 5 of gnathopod 1 broadly but

shallowly lobate, lobe setose, article 6 narrower and

shorter than 5, dactyl strongly overlapping simple palm,

article 5 of gnathopod 2 truncate and asetose posteriorly, article 6 relatively longer than in gnathopod 1,

dactyl fitting palm armed with large tooth-spines each

bearing elephantine accessory setule; pereopods 1-2

simple, slender, weakly setose, pereopods 3-5 notably

weakly setose, dactyl of pereopod 5 very stout and

apically setose; uropods all extending equally, rami on

uropods 1-2 bearing long terminal spines; telson

square, truncate.

TYPE-SPECIES.—Kanaloa manoa, new species.

RELATIONSHIP.—This remarkable oedicerotid appears morphologically as if it had filled both an oedicerotid and a synopiid (tironid) niche in Hawaiian

waters. Superficially it has the aspect of a tironid like

Garosyrrhoe J. L. Barnard in its head and a reduction

of pereopodal setae but, of course, the vestigial accessory flagellum, elongate peduncle of uropod 3, and

short entire telson mark Kanaloa manoa as an oedicerotid. The family is confined to cold waters.

Kanaloa has affinities with Antarctic genera armed

with medial setae on the inner plate of maxilla 1 and

bearing weak gnathopods. It resembles Parhalimedon

Chevreux (1906) and Methalimedon Schellenberg

(1931) in its weak gnathopods, head, mandible, lower

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lip, maxilla 1, maxillipeds, and spines of uropods 1-2

but differs from those genera by: (1) the transverse

palms of the gnathopods and the relative distinctions

(not strong) between gnathopods 1 and 2 not occurring

in the other genera; (2) the more strongly excavate

coxa 4; (3) the equally extending uropods. Kanaloa

differs more strongly from Methalimedon than from

Parhalimedon in its falcate mandibular palp article 3

and slightly elongate article 3 of antenna 1. But the

eyes of the type-species of Parhalimedon are supposed

to be bilaterally paired and those of Methalimedon

not, although J. L. Barnard (1961) has described a

tropical Parhalimedon (Coral Sea, 230 m) with unpaired eyes. Article 2 on pereopod 5 of Parhalimedon

is constricted ventrally, whereas in Methalimedon and

Kanaloa it has a ventrally extended lobe.

Kanaloa resembles antiboreal Bathyporeiapus Schellenberg (1931), but the latter genus has a nearly simple,

elongate gnathopod 2 with vestigial dactyl, a short

mandibular palp article 3, and no setae on the inner

plate of maxilla 1.

Carolobatea Stebbing (1899b) (see Halimedon

schneideri Stebbing, 1888, pi. 59) from Kerguelen Island also is related to this group of genera and has a

transverse palm on gnathopod 2, but the gnathopods

are otherwise morphologically equal to each other and

have small pendant posterior lobes on the fifth articles,

and elongate coxa 5, weakly excavate coxa 4, elongate

maxillipedal palp with thin article 4, generally stronger

setation on the pereopods, and a mandibular palp typical of Westwoodilla Bate.

Exoediceros Stebbing (1899b), with 2 species in

southeast Australia, has uropod 2 shortened, no dactyls

on pereopods 1-2, and lobed fifth articles on both

gnathopods 1-2.

No resemblance of Kanaloa exists toward oedicerotid

genera occurring in the Northern Hemisphere or toward the one other tropically oriented genus, Perioculopsis Schellenberg.

Kanaloa manoa, new species

FIGURES 146, 147

DIAGNOSIS.—With the characters of the genus.

DESCRIPTION.—Pleonal epimera 1-2 each with

slightly convex posterior margin and small or large

posteroventral point; pleonal epimeron 3 with 3 posteroventral serrations; lateral cephalic lobe weakly and

obtusely attenuate; spine row of mandible with 1 coni-
NUMBER 34

225

FIGURE 146.—Kanaloa manoa, new genus, new species, holotype, female, 1.8 mm, JLB Hawaii 6:

a, body; b, mandible; c, telson; d, e, gnathopods 1,2;/, uropod 3; g, urosome; h, maxilla 2.
226

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 147.—Kanaloa manoa, new genus, new species, holotype, female, 1.8 mm, JLB Hawaii 6:

a, medial gnathopod 1; b, c, medial and lateral gnathopod 2; d, lower lip; e, inner and outer

plates of maxilla 1; /, medial pereopod 4; g, article 7 of pereopod 5; h, maxilliped; i, pleonal

epimera 1-3, posteroventral corners.

cal, unarmed spine and 2 apically frilled spines; majority of spines on outer plate of maxilliped clavate; dactyl

of gnathopod 2 simple, but that of gnathopod 1 with

subapical clavate setae; margins of palm and lobe on

article 5 of gnathopod 1 serrate but smooth on gnathopod 2. Male unkown.

COLOR IN 2-DAY FORMALDEHYDE.—Body white, eye

yellow-orange, fading completely in alcohol.

HOLOTYPE.—Bishop Museum collections, catalog

number 7294, female, 1.8 mm.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Oahu, 30 m, coralline footballs, 29 January 1967.

MATERIAL.—JLB Hawaii 6 (15), 10 (?1 damaged).

DISTRIBUTION.—Hawaiian Islands.

Family PHLIANTIDAE

Palinnotus Stebbing

DIAGNOSIS.—Rostrum a small point; coxa 1 as long

as or extending nearly as far ventrally as coxa 2, coxa
NUMBER 3 4 227

4 with posterodistal excavation; maxilla 1 with variable

palp, either represented by a spine, possibly absent or

formed of a short flat plate, inner plate either absent

or unobserved; maxilla 2 with lobes distinct from one

another apically but coalesced basally; palp of maxilliped 3-articulate, article 3 blunt and apically setose;

gnathopods 1-2 simple; article 2 of pereopod 5

smaller than article 2 of pereopods 3-4; pleopods with

subequal rami, only peduncle of pleopod 2 extending

medially more or less; uropod 1 biramous; uropod 2

uniramous; uropod 3 represented by a pair of small

anal flaps ventral to telson and lacking rami; telson

simple and entire.

Palinnotus alaniphlias, new species

FIGURES 148, 149, 150

DIAGNOSIS OF FEMALE.—Article 2 of pereopod 5

ovatoquadrate, very small and scarcely broader than

article 3, lacking posterior keel, coxa 7 coalesced with

pereonite 7; article 4 of pereopod 3 subquadrate or

trapezoidal, article 4 of pereopod 5 subtriangular, thus

distally expanded; outer plates of maxilliped large and

extending equally with palp; maxilla 1 with small, flat,

coniform palp.

DESCRIPTION.—Eyes black in alcohol; pereonites

1-7 with lateral bulges just above coxae, pleonites 1

and 2 with subdorsal bulge, pleonal epimera all rounded-quadrate at posteroventral corners, urosome with 3

segments, but segment 3 not visible dorsally; gnathopods 1-2 nearly equal in size, article 6 on gnathopod

1 medially furnished with asparagoid spines and numerous short prickles, gnathopod 2 lacking these; brood

lamellae of female furnished with complex curl-tipped

setae.

HOLOTYPE.—Bishop Museum collections, catalog

number 7295, female, 3.4 mm.

TYPE-LOCALITY.—Fee 1, Waikiki Beach, Oahu, intertidal, Viva lactuca, 25 April 1967.

MATERIAL.—The type-specimen and a female, 4.2

mm, from JLB Hawaii 17.

REMARKS.—Both specimens are females, the largest with 5 hatched juveniles fully occupying the brood

pouch.

HATCHED JUVENILES.—These have the aspect of

eophliantids more than of phliantids; body subcylindrical, lacking dorsal and lateral sculpture, head large,

tall, broad, long, lateral lobes unprotruding and lateral

margins long and weakly convex, eyes small, black; antenna 1 like that of adult, antenna 2 stouter; coxae

1-4 (and coxae 5-7 to some extent) not contiguous,

much less broadened than in adult and scarcely

splayed, anterior coxae forming elongate rectangles,

coxa 4 unexcavate, coxae 5 and 6 unlobed, coxa 7

coalesced with pereonite 7; pereonite 1 a simple cube

not encroaching on head; gnathopod 1 lacking medial

armature of adult, otherwise gnathopods and pereopods 1-2 with form of adult; article 2 of pereopods 3

and 4 smaller than that of pereopod 5, thus in adult

article 2 of pereopods 3-4 becoming enlarged whereas

that of pereopod 5 remaining proportionally similar to

its condition in juvenile; article 2 of pereopod 3 more

nasiform than in adult; pleopods without setae; pleonal

epimera scarcely developed; urosome unflattened but

broad; uropod 3 relatively much larger than in adult,

flaps reaching or exceeding telsonic apex.

RELATIONSHIP.—Palinnotus alaniphlias differs from

the 3 other known species of the genus in the presence

of a flat coniform palp on maxilla 1 and the coalescence

of coxa 7 with the body. These characters do not seem

to be of generic value if one considers the remarkable

conformity of the pleopods, urosome, head, and gnathopods with other members of the genus, plus the

general congruence of the other mouthparts except for

the outer plates of the maxillipeds, which in P. holmesi

Gurjanova (1938, 1951) are much smaller than in

the other known species. That Japanese species also

has a larger article 2 on pereopod 5 than does P.

alaniphlias.

The type-species, P. thomsoni (Stebbing, 1899a),

from New South Wales also has a larger article 2 of

pereopod 5 and a less quadrate article 2 of pereopod

3.

Palinnotus natalensis K. H. Barnard (1940) from

Natal, has a much larger article 2 of pereopod 5 and

more strongly nasiform article 4 of pereopod 5 than

does P. alaniphlias.

DISTRIBUTION.—Hawaiian Islands.

Family PLEUSTIDAE

Parapleustes derzhavini makiki, new subspecies

FIGURES 151, 152

DESCRIPTION.—Head with small rostrum, lateral

cephalic lobes mammilliform, anteroventral corner of

head sharply angular; eyes large, subcircular, formed

of dark-purplish brown core surrounded by clear
228

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 148.—Palinnotus alaniphlias, new species, holotype, female, 3.4 mm, Fee 1: a, body; b,

urosome; c, maxilla 1; d, antenna 2; e, ventral urosome showing uropod 3; /, dorsal view of

head and anterior thorax.

ommatidia; antenna 1 equal to head and 8 body segments, antenna 2 equaling head and 4 body segments,

peduncle of antenna 1 reaching to middle of peduncular article 5 on antenna 2; upper lip obtusely projecting anteriorly from lateral view, its ventral plate

from anterior view deeply incised; left mandible with

large lacinia mobilis more deeply serrate than incisor,

spine row with about 8 spines, molar nontriturative

but lacking spines, slightly fuzzy on one rim, palp

large, article 3 slightly falcate and armed distally and

unimarginally with stout spine setae; lower lip normal;

maxillae 1-2 generally like those of Neopleustes pulchella (Kroyer) (Sars, 1895, pi. 122, fig. 1), but inner

plate of maxilla 1 much smaller and bearing only 1

seta; maxillipeds normal structurally but palp broad,

4-articulate; coxa 1 with 1 or 2 posteroventral notches,

coxa 2 with 1; gnathopods 1-2 of medium size, tumid,

strongly subchelate, article 2 strongly setose anteriorly
NUMBER 34 229

FIGURE 149.— Palinnotus alaniphlias, new species, female, 4.2 mm, JLB Hawaii 17: a, mandible.

Holotype, female, 3.4 mm, Fee 1: b, outer plate of maxilla 2, opposite side of 150<f; c, inner plate

of maxilliped; d, view of gnathopod 1; e, right aspect of pereopod 5 articles 2-3 and coxa; /,

lateral gnathopod 1: g, lateral pereopod 5; h, lateral pereopod 2.

on gnathopod 1 only, article 4 produced sharply only

on gnathopod 2, article 5 long, with thick lobe on

gnathopod 1, thinner on gnathopod 2, article 5 with

1 anterior spine, palm as long as posterior margin of

article 6, defined by spines and bearing 1 set of palmar

spines and obtuse cusp in middle, dactyl fitting palm;

pereopods 1-5 of normal size for family, article 2 of

pereopods 3-5 broadly ovatorectangular, articles 4-6

slender, article 4 produced nearly halfway along article

5; uropods 1-3 with increasingly shortened outer ramus, rami of uropod 3 of medium length for family, telson long, about 70 percent as broad as long, linguiform,

apically rounded, with small, obtuse lateral notch

bearing setule on each side; body smooth dorsally;

pleonal epimera 1-3 with small posteroventral cusp, 2

with larger, sharply extended tooth, ventral margins of

1 and 3 spinose, 2 with ventrolateral spines, all spines

small.

HOLOTYPE.—Bishop Museum collections, catalog

number 7297, female, 4.2 mm.

TYPE-LOCALITY.—JLB Hawaii 13, Kaneohe Bay,

3-4 m, corals, corallines, fleshy algae, 8 April 1967.

MATERIAL.—Nine specimens from the type-locality.

RELATIONSHIP.—Parapleustes d. derzhavini (Gurjanova, see 1951) from the Japan Sea has the blunt

article 4 of gnathopod 1 seen in P. d. makiki, and

gnathopod 1 has the broad lobe on article 5, but the

lobe of article 5 on gnathopod 2 is slightly narrower

and article 5 slightly shorter, while article 4 of gnathopod 2 is also blunt; both pairs of gnathopods have
230

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 150.—Palinnotus alaniphlias, new species, female, 4.2 nun, JLB Hawaii 17: a, e, f,

pleopods 3, 1, 2; g, maxilla 1. Holotype, female, 3.4 mm, Fee 1: b, apices of curl-tipped setae on

brood plates: c, mandible, right obverse; d, maxilla 2; h, half of lower lip; i, maxilliped; ;,

pleon, left lateral.

long setae on article 2. These small differences do not

warrant full specific nomenclature for the Hawaiian

population at present.

This subspecies from Hawaii has affinities with P.

pugettensis (Dana) from the cold-warm temperate

of Pacific North America, but differs in the longer

article 5 of the gnathopods, the stouter posterior lobes,

the blunter article 4 of gnathopod 1, and the more

strongly convex posterior margin of pleonal epimeron

2.

Parapleustes d. makiki is also closely reated to P.

gracilis (Buchholz) from boreal waters but differs in

the same ways it does from P. pugettensis, in addition

to the more rounded lateral cephalic lobes and spinier

pereopods and uropods.

DISTRIBUTION.—Hawaiian Islands.
NUMBER 3 4 231

FIGURE 151.—Parapleustes derzhavini makiki, new subspecies, holotype, female, 4.2 mm, JLB

Hawaii 13: a, telson; b, head; c, articles 3—5 of lateral gnathopod 1; d, e, f, g, pereopods 2, 3,

4, 5; h, i, coxae 3, 1; ;, articles 3-5 of gnathopod 2, right lateral aspect; k, gnathopod 2.
232

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

0

FIGURE 152.—Parapleustes derzhavini makiki, new subspecies, holotype, female, 4.2 mm, JLB

Hawaii 13: a, head; b, c, uropods 1, 3; d, left mandibular body; e, mandibular palp; /, upper

lip; g, upper lip protrusion from left lateral view, arrow pointing dorsally; h, lower lip; i, pleonal

epimera 1—3; j , inner plate of maxilla 1; k, maxilliped; /, dimensions of uropod 2; m, dactyl of

pereopod $;n,o, medial gnathopods 2, 1.
NUMBER 3 4 233

Parapleustes (?) honomu, new species

FIGURES 153, 154

DESCRIPTION.—Head with medium-sized rostrum,

lateral lobe vertically truncate but forming softly

rounded right angle marking boundary of ventral

quadrate excavation for attachment of antenna 2;

eyes very large, ochraceous in alcohol, red in formaldehyde, ovatocircular, composed of tightly packed ommatidia on slight lateral bulge; antennae 1-2 missing;

upper lip deeply incised; mandibles with large serrate

lacinia mobilis matching incisors, spine-row with 2-3

spines, molar narrow and nontriturative, conically

geniculate, bearing apical spinules, palp very large,

article 3 clavate and armed distally and unimarginally

with stout spine-setae; lower lip, maxillae 1-2 generally

like those of Neopleustes pulchella (Kroyer) (Sars,

1895, pi. 122, fig. 1), except inner plate of maxilla 1

slightly smaller and with only 1 seta; maxillipedal

inner plate armed with 3 small nobs, one of them

articulate and 2 apicolateral setae, outer plate slightly

hooked, apex with 2 setules, inner margin with 4 spines,

palp 3-articulate, article 3 longest and strongly spinose

and possibly bearing vestige of article 4; coxae short,

1-4 about as broad as long, 4 weakly excavate posteriorly, posterior lobes of 5 and 6 large; gnathopods 1-2

small, slender, scarcely subchelate, articles 5-6 subrectangular or sublinear, palms with several long and

short spines, dactyls long; pereopods 1-5 very large

and stout for family, dactyls long, article 6 of pereopods

3-5 elongate, article 2 strongly lobed posteroventrally;

uropods 1-3 with shortened outer rami, rami on uropod 3 of medium length for family; telson short, as

broad as long, softly triangular; pereonites 6-7 and

pleonites 1-2 with posterodorsal hump-keel, pereonite

7 with small posterolateral bulge, pleonites 1-2 with

lateral keel continuing to posterior quadrate corner,

connected with epimeron directly, pleonite 3 with

partial anterolateral keel; pleonal epimera 1-2 with

lateral ridge, posteroventral corner formed of small

cusp, posterior edge slightly convex and with 1-2

serrations, epimeron 3 rounded and serrate posteriorly.

HOLOTYPE.—Bishop Museum collections, catalog

number 7296, male, 2.3 mm. Unique.

TYPE-LOCALITY.—Devaney 1, off Moku Manu Island, Oahu, 33 m, stem of black coral, 13 May 1967.

RELATIONSHIP.—This species obviously will require

the erection of a new genus for its reception, but this

event should await a firmer foundation in unbroken

material of both sexes. Parapleustes honomu is apparently derived from boreal pleustids of either the genera

Parapleustes or Neopleustes, but the absence of palp

article 4 on the maxilliped obscures the relationship,

for the two boreal genera differ from each other in

the presence of a process on article 3 that cannot be

detected in P. honomu because article 4 is absent and

the relative relationships between the 2 articles no

longer occur.

Parapleustes (?) honomu differs from other pleustids generally in the weak lateral cephalic lobe, large

eye on slight lateral bulge, the absence of maxillipedal

palp article 4, presence of lateral ridges on pleonites

1-3, serrate epimeron 3 (recalling a similar situation

in warm-water members of Pontogeneia), elongate

article 6 of pereopods 3-5, short coxae, stenothoid-like

mandibular incisors and lacinia mobili, and the extraordinarily heavy setae of the mandibular palp.

This species resembles Neopleustes boeckii (Hansen)

(Sars, 1895, pi. 122, fig. 2) in the presence of 3 dorsal

teeth and to some extent in the head, but N. boeckii

has a dorsal crest on the head among the usual

differences.

Gnathopods 1—2 of P. honomu are very thin and

poorly developed, but Parapleustes oculatus (Holmes,

see Barnard and Given, 1960) also has weak gnathopods and large apical setae on the mandibular palp,

but the outer lobes of the lower lip of P. honomu are

normally broad and P. oculatus has no dorsal teeth.

Neopleustes rasmyslovi Gurjanova (1951) also has

a thin but elongate gnathopod 1 and P. johanseni Gurjanova (1951) has thin but strongly subchelate gnathopods and the lateral cephalic lobes are poorly

developed.

DISTRIBUTION.—Hawaiian Islands.

Family PODOCERIDAE

Laetmatophilus hala, new species

FIGURE 155

DIAGNOSIS OF FEMALES AND JUVENILES.—Head

with 3 dorsal cusps in transverse row, anteroventral

corner with small cusp; pereonite 1 with 2 axial dorsomedial cusps and 2 lateral cusps on each side, pereonites 2, 3, 4, 5, 7 with 1 dorsal and 1 lateral cusp on

each side, pereonite 6 with 1 dorsal cusp and 1 veryweak ridge on lateral surface, pereonites 5-7 coalesced;

lateral margins of pereonites 1-7 with small ridge-
234

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

9

FIGURE 153.—Parapleustes (?) honomu, new species, holotype, male, 2.3 mm, Devaney 1: a, b,

c, d, pereopods 3, 4, 5, 1; e, f, apices of medial gnathopods 2, 1; g, pereonites 6—7 and pleonites

1-3, left lateral.
NUMBER 3 4 235

FIGURE 154.—Parapleustes (?) honomu, new species, holotype, male, 2.3 mm, Devaney 1: a,

head; b, maxillipedal palp; c, d, gnathopod 1; e, article 1 of mandibular palp; /, articles 2—3 of

mandibular palp; g, mandibular body; h, outer plate of maxilliped; i, coxa 4; j , inner plate of

maxilliped; k, telson; I, upper lip; m, n, o, uropods 1, 2, 3.
236

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 155.—Laetmatophilus hala, new species, holotype, female, 1.8 mm, JLB Hawaii 6: a,

body; b, medial gnathopod 2; c, head; d, lateral gnathopod 2; e, urosome, dorsal; /, pleon, lateral;

g, lateral gnathopod 2; h, dactyl of gnathopod 1. Juvenile, 0.9 mm, JLB Hawaii 3: i, head,

pereon, metasome from right side.
NUMBER 34 237

cusp above coxae; pleonites 1-2 each with dorsal cusp

and pleonite 1 with small dorsolateral ridge on each

side; young specimens with lateral supracoxal ridges

and dorsolateral teeth absent, middorsal cusps represented by humps, anteroventral cephalic cusp absent;

only coxae 3-4 in adult female with ventral points;

article 2 of gnathopod 2 with sharp anterodistal cusp,

article 4 with medium-sized posterodistal process;

gnathopod 1 palm longer than posterior margin of

article 6 and lined with long setae, defined by sharp,

slender, short, feathered spine, dactyl slightly longer

than palm and distally multifid, lobe of article 5

evenly bulging; outer ramus of uropod 1 half as long

as inner.

DESCRIPTION.—Pereopods 2-7 and antennae 1-2

missing from all material except pereopods 3-4 of juvenile in JLB Hawaii 3, generally like those of L. tuberculatus Bruzelius (Sars, 1895, pi. 226) ; mouth parts

also like that species but inner plate of maxilla 1 obsolescent, right mandibular molar with large flake and

seta, left molar with small flake and no seta; epistome

strongly conical; uropod 3 a small flap with medial

setule. Specimen of JLB Hawaii 5 lacks lateral cusps

on pleonite 1, and the adult of JLB Hawaii 3 has all

cusps less produced than on the holotype, and pereonite 1 and pleonite 1 have no lateral cusps.

HOLOTYPE.—Bishop Museum collections, catalog

number 7298, female, 1.8 mm.

TYPE-LOCALITY.—JLB Hawaii 6, off Barbers Point,

Hawaii, 30 m, coralline footballs, 29 January 1967.

MATERIALS.—JLB Hawaii 3 (2), 5 (1), 6 (1).

RELATIONSHIP.—This species probably has its closest relationships with L. hystrix (Haswell, 1880b), but

that species has at least one more dorsolateral cusp on

each side of pereonites 1-7 and pleonite 1 has 2 dorsal

medials, the anteroventral cephalic cusp is larger and

coxae 5-6 have ventral cusps. Considerable developmental change occurs in L. hala, and perhaps terminal

adults develop the characters of L. hystrix.

Laetmatophilus armatus (Norman) (Sars, 1895, pi.

227, fig. 1) has fewer dorsal cusps than L. hala, the

former having primarily a double dorsal spiniform set

along the midline of pereonites 2-7 and pleonites 1-2

and lacking an anteroventral cephalic cusp.

Laetmatophilus leptocheir K. H. Barnard (1937)

has the outer ramus of uropod 1 about two-thirds as

long as the inner, the head and pereonite 1 lack dorsal

cusps or keels and the hand of gnathopod 1 is very

slender.

DISTRIBUTION.—Hawaiian Islands.

Podocerus Leach

Identifications of the tropical species in this genus

have been subject to confusion after 1906 when Stebbing adequately summarized the known species which

at that time had limited distributions. Since then, two

species, Brazilian P. brasiliensis (Dana) and New Zealandian P. cristatus (Thomson), have been reported

from localities far from tropical waters and P. brasiliensis has come to be considered circumtropical.

One of those records identified by J. L. Barnard

(1962a, fig. 30) from open-sea southern California is

undoubtedly erroneous as his figure shows a head entirely different from that seen in P. brasiliensis from

Hawaii and vaguely seen in Dana's (1853) drawings

of the original establishment of the species. The ocular

lobes of P. brasiliensis fill the anteroventral corner of

the head, whereas in the Californian material they

occur behind a sharp corner. The record by J. L.

Barnard (1959) from Newport Bay has the correct

head although the dactyl of gnathopod 1 is apically

multispinose as in the open-sea populations and thus

differs from the dactyl in tropical individuals. The

Californian open-sea material should be accorded a

name different from P. brasiliensis but estuarine-lagoonal populations in California may represent a form

of P. brasiliensis imported by human agencies.

Gnathopod 1 of male P. brasiliensis from Hawaii

(J. L. Barnard, 1955a) and Ceylon (Walker, 1904, as

P. synaptochir) has a distal hemispherical lobe on a

truncate article 5, whereas Dana's (1855) and J. L.

Barnard's (1959) materials have the lobe on article 5

stretching the breadth of the article.

Podocerus brasiliensis (Dana)

FIGURES 156,157

Platophium brasiliense Dana, 1853, pp. 838-839, pi. 55, fig. 9.

Platophium synaptochir Walker, 1904, pp. 296-297, pi. 8,

fig. 52.

Podocerus brasiliensis.—Schellenberg, 1938, p. 94; J. L.

Barnard, 1953, p. 87 [with synonymy]; 1955a, p. 39.

DIAGNOSIS OF HAWAIIAN MALE.—Lateral cephalic

lobe rounded anteroventrally and filled with red or

clear eye in ocular bulge; accessory flagellum 1-articulate, long but thin, antenna 2 very long, antenna 1

reaching slightly beyond article 4 on peduncle of antenna 2; epistome conically produced; coxa 1 weakly

extended forward, coxa 2 with large medial stridulating flange; article 5 of gnathopod 1 shorter than article

360-034 O - 17
238

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 156.—Podocerus brasiliensis (Dana), male, 5.3 mm, Pearl Harbor: a, part of pereopod 3,

medial; b, mandibular palp minus setae; c, palm of gnathopod 2; d, dorsal telson, setae marked

with pits; e, body; /, maxilla 2; g, part of lateral pereopod 1; h, spines at defining corner of"

gnathopod 1; i, dorsal urosome. Female, 2.9 mm: ;, dorsal telson; k, aberrant uropod 1; /,

normal uropod 1.
NUMBER 3 4 239

FIGURE 157.—Podocerus brasiliensis (Dana), male, 5.3 mm, Pearl Harbor: a, lateral telson and

uropod 3 (shaded) ; b, coxa 2, medial, showing stridulation flange; c, gnathopod 1; d, lateral

pereopod 5, less coxa; e, lateral gnathopod 2. Female, 2.9 mm: f, lateral gnathopod 1; g, medial

gnathopod 2; h, dorsal head.
240 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6, slightly cup-shaped and bearing tumid posterior lobe

slightly marked off from main part of article, article 6

ovatotriangular, but oblique palm longer than posterior margin of article 6, defined by obtuse corner

bearing several short spines, dactyl with weak inner

serrations and fitting palm; article 2 of gnathopod 2

with obsolete anterolateral lobe bearing spine, article

4 short and stout, weakly protuberant posterodistally,

article 6 elongate, heavily setose, posterior margin bearing all of palm, undefined and with obsolescent distal

bulge hidden among setae, dactyl about half as long

as article 6, strongly bent basally; article 2 of pereopods

1-5 of medium stoutness, slightly bilobate on pereopod

3 and heavily spinose on pereopods 3-4; uropods 1-2

each with stout peduncular tooth between rami, outer

ramus of uropod 1 varying between 67 and 75 percent

length of inner ramus, of uropod 2 about 67 percent

length of inner ramus, uropod 3 small but with apical

and subapical spine; telson quadrotrapezoidal, dorsally

produced as large nob armed with 8 or more long setal

spines; pleonites 1-2 with obsolescent dorsal hump,

pereonite 7 slightly longer than others, all pereonites

articulate; pleonal epimera 1-3 evenly rounded below.

FEMALE.—Gnathopod 2 short and stout, article 4

elongate and grossly extended distally, article 5 small,

cup-shaped, article 6 subcircular, palm longer than

posterior margin of article 6, defined by 2 large spines,

dactyl fitting palm; telson with 6 spines; article 5 of

gnathopod 1 lobate but lobe not marked off from main

article, palmar defining corner without short spines.

MATERIAL.—Same as J. L. Barnard (1955a).

DISTRIBUTION.—Circumtropical.

Podocerus hanapepe, new species

FIGURES 158, 159

DIAGNOSIS OF MALE.—Lateral cephalic lobe quadrate

anteroventrally, head extended quadrately forward,

lateral ocular bulges of medium extent, divorced from

anteroventral corner, red in formaldehyde, ochraceous

in alcohol, with weak core of dark pigment; accessory

flagellum 1-articulate and very small, antennae 1-2

short; coxa 1 strongly extended forward, article 6 of

gnathopod 1 triangular, distally expanded, palm transverse, defined by small spine, dactyl with weak inner

serrations, article 6 with extremely stout semipinnate

setal spines, article 5 slightly longer than 6, posterior

lobe broadly bulging; article 2 of gnathopod 2 with

small, distal, anterolateral, obtusely pointed lobe, article 4 long and apically extended as blunt triangle, palm

oblique and defined by stout cusp and 2 spines, dactyl

almost fitting palm, palm weakly setose and bearing

broad, low, castellate distal process and smaller proximal quadrate cusp; article 2 of pereopods 1-5 very

short and stout, bilaterally lobate on pereopod 3, outer

ramus of uropod 1 about three-fourths as long as inner

ramus but appearing shorter from lateral view, uropod

2 very stout, outer ramus about 60 percent as long as

inner; uropod 3 small but with 2 apical-subapical

spines; telson subcircular-quadratiform, bearing dorsal

nob armed with 2 long stout spines; pereonites 1-3 with

weak transverse dorsal depressions, pereonites 4-5

shortest of all but pereonite 6 not extremely broadened

nor heavily muscularized and all segments clearly articulate; pereonite 7 and pleonites 1-2 scarcely elevated

posterodorsally; pleonal epimera 1-3 evenly rounded

below.

FEMALE.—Unknown.

HOLOTYPE.—Bishop Museum collections, catalog

number 7299, male, 2.4 mm. Unique.

TYPE-LOCALITY.—Fee 1, Waikiki Beach, Oahu, intertidal, Ulvalactuca, 21 April 1967.

RELATIONSHIP.—The overall appearance of this

species resembles that of P. mangarevae Chevreux

(1908), but P. hanapepe has a slightly more extended

and squared-off head, a fully transverse palm on gnathopod 1, no anterior taper on coxa 1, a short accessory

flagellum, a strong distal extension on article 4 of

gnathopod 2, a slightly stronger defining tooth on the

the palm of gnathopod 2, and only 2 telsonic spines

that are larger than on P. mangarevae. Antenna 2 of

P. mangarevae is very conspicuously longer than antenna 1, a condition not true of P. hanapepe.

Possibly P. mangarevae is a synonym of P. zeylanica

(Walker, 1904), and the same differences are approximately applicable between P. hanapepe and P. zeylanica as mentioned above except for coxa 1 and the defining tooth of gnathopod 2 palm. Antennal lengths for

P. zeylanica were not mentioned, and the telson has 1

large and 2 small spines.

Podocerus inconspicuus (Stebbing, 1888), from

southeast Australia, but apparently not that taxon

identified by Pirlot (1938), is very close to P. hanapepe,

even though the 2 species are based on opposite sexes

and various missing parts of P. inconspicuus preclude

their description. Gnathopod 1 of P. inconspicuus

(based on a female) has a very narrow article 5 in

comparison to that of P. hanapepe.

DISTRIBUTION.—Hawaiian Islands.
NUMBER 3 4 241

FIGURE 158.—Podocerus hanapepe, new species, holotype, male, 2.4 mm, Fee 1: a, body; b,

right mandibular molar; c, telson and uropod 3, dorsal; d, mandible; e, pereonite 7 and pleonal

epimera 1-3; /, apex of maxillipedal palp, some setae removed; g, lower lip; h, maxilla 2; i,

urosome.

Podocerus talegus lawai, new subspecies

FIGURES 160, 161

DIAGNOSIS OF MALE.—Lateral cephalic lobes

rounded anteroventrally, eyes on strong lateral bulges

divorced from anteroventral corners, red in formaldehyde, ochraceous in alcohol, with weak core of darker

ochre; accessory flagellum 1-articulate but thick and

elongate, antennae 1-2 long; coxa 1 strongly extended

forward; article 6 on gnathopod 1 of normal expansion,

slightly longer than article 5, palm about twice as long

as posterior margin of article 6, defined by 2 or more

spines, one of them thorny, dactyl with long inner

serrations near apex, article 6 with long, anteriorly

submarginal, semipinnate setal spines, article 5 with

evenly bulging posterior lobe; article 2 of gnathopod 2

with quadrate anterodistal lobe, article 4 with small

posterodistal cusp, palm undefined, occupying most of

posterior margin of article 6, dactyl about two-thirds

as long as article 6, palm fringed with long setae and

medial face of hand with many stout but long setae, no

palmar teeth showing laterally or marginally from
242

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 159.—Podocerus hanapepe, new species, holotype, male, 2.4 mm, Fee 1: a, uropod 2;

b, head; c, d, pereopods 3, 5, in whole or part; e, f, gnathopod 2, medial and lateral; g, uropod 1;

h, apex of lateral gnathopod 1; i, part of pereopod 4; ;, medial gnathopod 1.

medial view in young male but bearing mediosubmarginal, low, broad, crenulate distal process, next

proximally a rounded smooth process and finally a

small or large obtusely conical process, terminal male

with these processes much enlarged and medial edge

of article 2 heavily setose; article 2 of pereopods 1-5 of

medium stoutness; outer ram us of uropod 1 about

three-fourths as long as inner ramus, peduncle with

ventrodistal sharp tooth of medium length between

rami, outer ramus of uropod 2 about two-thirds as long

as inner ramus, often appearing shorter from lateral

view, uropod 3 small but with 3 apical-subapical spines;

telson subcircular, bearing extended dorsal nob armed

with 2 long stout spines; pereonites 1-5 with weak

transverse dorsal depressions, pereonites 4-5 shortest of

all, pereonites 6-7 exceptionally long, articulations be-
NUMBER 3 4 243

tween segments 5-7 and 6—7 incomplete, segments 6-7

heavily muscularized and rigid, pereonite 7 and pleonites 1-2 each with dorsal hump and occasionally weak

dorsolateral accessory hump often marked only with

setae; pleonal epimera almost evenly rounded below.

FEMALE.—Article 2 of gnathopod 2 lacking anteroventral process, distal cusp of article 4 small, article 6

stout, short, broadly ovate, palm long, strongly defined

by 2 stout spines and weak cusp, palm minutely castellate; coxae broader than in male.

HOLOTYPE.—Bishop Museum collections, catalog

number 7300, male, 2.6 mm.

TYPE-LOCALITY.—JLB Hawaii 5, off Ewa Beach,

Oahu, 30 m, Pocillopora, 29 January 1967.

MATERIAL.—JLB Hawaii 2 (2), 5 (1), 12 (3), 13

(3),17(1).Fee l (5).

RELATIONSHIP.—This subspecies differs from P. t.

talegus J. L. Barnard (1965a) from Micronesia only in

minor ways such as the rounder anteroventral cephalic

margin, the shorter and broader article 5 of gnathopod

1, and apparently the slightly longer serrations on the

dactyl of gnathopod 1. The terminal male of P. t.

talegus is unknown.

Podocerus talegus seems to have close affinities with

P. inconspicuus (Stebbing) and P. Plobatus (Haswell)

as identified by Pirlot (1938). There is considerable

evidence that Pirlot's materials should be given new or

different names than those he applied. Unlike Pirlot's

species, P. talegus has a strong dorsal hump or keel

only on 3 segments and the bulges are thick and not

lamellar. A few individuals of those species identified

by Pirlot also have only 3 segments keeled. Article 2

of gnathopod 2 on P. talegus lawai fits P. inconspicuus

better than P. Plobatus, but the palm of male gnathopod 2 fits P. Plobatus better than P. inconspicuus. P.

lobatus (Haswell, 1885) and P. Plobatus of Pirlot have,

however, a defining tooth on the palm, whereas, if

that tooth is present on P. talegus, it has moved distally. Podocerus talegus does not fit the original P.

inconspicuus (Stebbing, 1888) at all.

Poderus fulanus J. L. Barnard (1962a) from California {—Podocerus species of J. L. Barnard, 1959)

has a few resemblances to P. talegus, but also has 4

keeled segments and has only 1 process on male gnathopod 2 palm, weaker ocular bulges, shorter posterior

margin of pleonal epimeron 3, fewer spines on uropod

3, and a slightly shorter ventrodistal peduncular cusp

on uropod 1, larger segment 6 of the pleon, and longer

palm on gnathopod 1 than does P. talegus.

The so-called P. cristatus (Thomson) of California,

as identified by J. L. Barnard (1962a), has 5 dorsally

keeled segments and dorsolateral humps, but gnathopod 2 has only 2 palmar processes, 1 of them very thin,

and article 2 of gnathopod 2 has a weak, mammilliform anteroventral lobe unlike that of P. talegus and

the species of Pirlot; coxa 1 is less extended, epimeron

3 short posteriorly, the palm of gnathopod 1 is longer,

lacks an anterodistal process on article 2 of gnathopod

2, and the cephalic lobes are sharp.

The Hawaiian specimens of P. talegus resemble

Cyrtophium dentatum Haswell (1880a, p. 342) in

dorsal keel. That species has been synonymized with P.

cristatus. Chilton (1926) has drawings of an identification of P. cristatus from New Zealand (type-area)

and P. talegus differs from those in the absence of a

distinction between the thin palmar setae and 5-6 thick

spines radiating around the defining corner of gnathopod 1 palm plus slight differences in the shape and

arrangement of teeth on the palm of male gnathopod

2; Chilton does not show anterodistal lobes on article

2 of gnathopod 2.

Thomson's (1881) second paper on Cyrtophium

cristatum has a figure with a dorsal keel similar to

several specimens of P. talegus with 4 weak dorsal

humps, but the eyes of the New Zealand species appear

weak, and he mentions 3-4 spines defining the palm

of gnathopod 1.

Haswell's (1885) Dexiocerella dentata ( = Cyrtophium dentatum of 1880a) is shown to have uropods

1-2 similar to those of P. talegus, but both pairs have a

ventral peduncular tooth instead of one as in P. talegus.

Perhaps, that is an error.

The identity of Haswell's Dexiocerella laevis by

Walker (1904) (as Platophium laeve) seems to fit the

Hawaiian species very closely. The palm of gnathopod

2 has no processes but that of young males in P. talegus

also shows no processes from marginal view, and they

might have been overlooked in Walker's material.

The fringing palmar setae of Walker's material however, are very short whereas they are long in P. talegus.

Walker's material has similarities to P. talegus in dorsal

ornamentation, telson, ocular lobe, gnathopod 1, the

small proximal palmar acclivity on gnathopod 2, but

coxa 1 is not as strongly extended forward, uropod 3

is less spinose, and the rami of uropods 1-2 are not

clearly described as to length.

DISTRIBUTION.—Hawaiian Islands.
244

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 160.—Podocerus talegus lawai, new subspecies, holotype, male, 2.6 mm, JLB Hawaii 5:

a, body; b, head, dorsal; c, d, pereopods 2, 4, in part. Male, 3.7 mm, Fee 1: e, f, uropods 1, 2; g,

h, antennae 1, 2; i, palm of gnathopod 1.
NUMBER 34 245

FIGURE 161.—Podocerus talegus lawai, new subspecies, male, 3.7 mm, Fee 1: a, gnathopod 2,

lateral, setae removed. Female, 3.0 mm: b, medial gnathopod 2. Holotype, male, 2.6 mm, JLB

Hawaii 5: c, medial gnathopod 2, setae removed; d, palm of gnathopod 2, enlarged; e, telson and

uropod 3, dorsal; /, lateral pereopod 5; g, gnathopod 1, with enlarged dactyl and spine offset to

left.
246

Family SEBIDAE

Seba ekepuu, new species

FIGURES 162, 163

DIAGNOSIS.—Anterior coxae evenly rectangular,

lacking sharp extensions; gnathopod 1 of the normal

chelate form lacking palmar ornaments, posterior margin of article 6 with setose declivity; article 6 of gnathopod 2 much longer than article 5; article 2 of pereopods 3-5 broadly expanded and increasingly so on

pereopod 5, latter with smooth posterior margin and

long, oblique, sparsely notched posteroventral margin;

fourth articles of pereopods 3-5 very slender and

weakly produced posterodistally; pleonal epimeron 1

with small sinuosity at posteroventral corner, epimeron

2 with sharp tooth, epimeron 3 with quadrate posteroventral corner; outer ramus of uropods 1 and 2 shorter

than inner rami, rami naked; uropod 3 with distal

spines on the single ramus; telson short and ovate.

DESCRIPTION.—Eyes apparently absent, but ocular

region with subcircular mass of dense granular tissue,

similar tissue occurring in other cephalic patches and

in article 2 of pereopods 3-5; accessory flagellum long

and 2-articulate; antenna 1 with 5 articles, ultimate

minute, antenna 2 with 3 flagellar articles; inner

ramus of pleopods 2-articulate, outer ramus 3-articulate.

HOLOTYPE.—Bishop Museum collections, catalog

number 7301, ?sex, 1.4 mm.

TYPE-LOCALITY.—J. L. Barnard Hawaii 17, Kawela

Bay, Oahu, intertidal, Sargassum, algae, rocks, corals,

24 May 1967.

MATERIAL.—Two specimens from the type-locality.

RELATIONSHIP.—Relationships among the 7 other

species of this unique genus are unclear owing to polymorphism in gnathopod 1 and variation in the extent

of flabellation on articles of the pereopods. Several

minute characters shown in drawings herein have not

been described for all of the species. Seba ekepuu appears to have closest affinities with S. typica (Chilton,

1884), from New Zealand, but differs in having a narrower article 4 of pereopods 3-5 and a broader and

characteristically shaped article 2 of pereopod 5. That

shape appears not to occur in any other species of the

genus. Seba antarctica Walker (1907) and S. subantarctica Schellenberg (1931) have, in adult stages,

broadly inflated fourth articles of pereopods 3-5, and

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

article 5 of gnathopod 2 is longer than article 6. Seba

saundersi Stebbing (1888) has an evenly ovate article

2 on pereopod 5, conspicuous marginal spines on the

ramus of uropod 3, and a sharp posteroventral extension on coxa 2. Seba innominata Bate (1862) has a

large tooth on the posteroventral corners of pleonal

epimera 1-3. Seba armata Chevreux (see Ghevreux

and Fage, 1925) has an elongate-oval article 2 on

pereopod 5 and apparently has urosomites 2 and 3

freely articulate.

Seba dubia Schellenberg (1926) has an ovate article

2 on pereopod 5.

DISTRIBUTION.—Hawaiian Islands.

Family STENOTHOIDAE

Stenothoe haleloke, new species

FIGURE 164

DIAGNOSIS OF FEMALE.—Lateral cephalic lobe

broadly truncate, eyes medium in size, red in formaldehyde, clear in alcohol; antennae 1-2 extending equally,

peduncles short; article 6 of gnathopod 1 strongly

dominating articles 4 and 5 in size, latter with narrow

posterior lobe, palm and posterior margin of article 6

equal to each other in length, article 2 thick; article 2

of gnathopod 2 with medial and lateral sharp distal

lobes, lobe of article 3 sharp, articles 4-6 of regular

form, palm oblique, simple, and equal to posterior

margin of article 6, denned by 2 pairs of large spines;

locking spines of pereopods 1-5 simple; posterodistal corner of article 2 on pereopod 3 angular,

pereopods otherwise of medium stoutness, fourth articles poorly expanded and weakly extended posterodistally; pleonal epimera 1-3 of regular form;

urosomite 2 strongly telescoped into urosomite 1;

uropod 1 with 2 spines on each ramus, uropod 2 with 1

spine on each ramus; peduncle of uropod 3 shorter

than article 1 of ramus, bearing 2 sets of spine (s),

article 1 of ramus with 2 sets of spine (s), article 2 of

ramus simple, equal in length to article 1; telson with

2 spines on each side.

HOLOTYPE.—Bishop Museum collections, catalog

number 7302, female, 2.0 mm. Unique.

TYPE-LOCALITY.—Fee 1, Waikiki Beach, Oahu, intertidal, Ulva lactuca, 25 April 1967.

RELATIONSHIP.—The female of this species has its

closest known affinities with, but differs from S. mono-
NUMBER 3 4 247

FIGURE 162.—Seba ekepuu, new species, holotype, ?sex, 1.4 mm, JLB Hawaii 17: a, body; b, c,

gnathopods 1, 2; d, telson; e, uropod 3, dorsal view, unflattened; /, maxilla 2.
248

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 163.—Seba ekepuu, new species, holotype, ?sex, 1.4 mm, JLB Hawaii 17: a, head, with

patches of glandular tissue, no eyes; b, mandible; c, maxilla 1; d, pleonal epimera 1-2; e,

maxilliped; /, mandibular palp; g, pereopod 5, with gland; h, pereopod 3; i, half of lower lip;

j , uropod 3, dorsal view, unflattened; k, accessory flagellum on antenna 1; /, lateral view of

urosome.
NUMBER 3 4 249

FIGURE 164.—Stenothoe haleloke, new species, holotype, female, 2.0 mm, Fee 1: a, body; b,

palmar defining area of gnathopod 1; c, medial gnathopod 2; d, uropod 3; e, medial gnathopod 1;

/, upper lip and epistome; g, telson; h, head; i, maxilliped.

culoides (Bate) (Sars, 1895, pi. 82, fig. 1), in the truncate cephalic lobe, and the presence of a few spines

on the inner ramus of uropod 1, both rami of uropod

2 and on article 1 of the ramus on uropod 3 (the middle segment), and on the telson.

The sympatriot S. gallensis Walker (J. L. Barnard,

1955a) differs in its female from S. haleloke in the

longer peduncle of uropod 3, the weak gnathopod 1

with thin article 2, and the relatively large size of

article 4 compared with article 6 and in the smallness

of the lobe on article 5. Male S. gallensis has a distinctive gnathopod 2 and uropod 3, but male S. haleloke

is unknown.

DISTRIBUTION.—Hawaiian Islands.
250 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Stenothoe ?valida Dana

FIGURE 165

PStenothoe validus Dana, 1853, pp. 924-925, pi. 63, figs.

1 a-o.

?Stenothoe valida.—Stebbing, 1906, p. 194.—J. L. Barnard,

1953,pp. 83-87 [with synonymy].

MATERIAL.—Devaney 1 (84).

REMARKS.—These specimens differ from J. L. Barnard's (1953) representation of the species from California in the much more slender article 4 of pereopods

3-5 in both sexes. Female palms of gnathopod 2 are

short and have denning spines like those of S. gallensis,

but a terminal female may not be present in these

collections.

In Hawaii, S. gallensis Walker (J. L. Barnard,

1955a) differs from S. valida and extrinsic populations

of S. gallensis (like S. crenulata Chevreux, 1908) in the

nonexcavate posteroventral margin of coxa 2 in both

sexes, a condition occurring in S. valida of Hawaii

and California. It thus appears that neither species is

FIGURE 165.—Stenothoe valida Dana, male, 4.2 mm, Devaney 1: a, medial gnathopod 2; b, c,

pereopods 3, 5; d, e, coxae 2, 3. Female, 2.8 mm: /, g, lateral gnathopods 1, 2; h, i, coxae 2

(sideways), 3.
NUMBER 3 4 251

of typical condition in Hawaiian waters, S. gallensis

differing in coxa 2 and S. valida differing in article 4

of pereopods. Males of S. gallensis continue to exhibit

the rugose apex of uropod 3 and even projectings of

the bifid tooth on gnathopod 2 and males of S. valida

continue to exhibit a simple apex of uropod 3 and

uneven projections of the bifid tooth on gnathopod 2.

DISTRIBUTION.—Cosmopolitan in low latitudes.

Stenothoe species A

FIGURE 166<J-«

DESCRIPTION.—Lateral cephalic lobe rounded, eyes

large; antennae extending equally, article 3 of antenna

1 as long as article 2; mouthparts and pereopods of

form in genus Stenothoe; article 5 of gnathopod 1

scarcely lobate, palm oblique, longer than posterior

margin of article 6, minutely crenulate, defined by long

spine and short spine, dactyl fitting palm and with

accessory tooth on inner margin, gnathopod 2 of juvenile condition, palm with small cusp in middle; dactyls

of pereopods 1-5 with accessory claw, sixth articles

stout and almost prehensile; rami of uropods 1-2

naked, outer ramus of uropod 2 slightly shorter than

inner ramus, inner ramus exceeding uropod 3; peduncle of uropod 3 with 1 spine, ramus slightly longer

than peduncle, articles 1 and 2 equal in length, naked;

telson naked.

MATERIAL.—JLB Hawaii 3 (1 juvenile, less than

1.0 mm long).

REMARKS.—The form of articles 6 and 7 on pereopods 1-5 distinguish this species from other Hawaiian

stenothoids.

Stenothoe species B

FIGURE 166;

DESCRIPTION.—Lateral cephalic margins broadly

rounded, scarcely extended forward, eyes large;

mouthparts and pereopods in form of genus Stenothoe;

gnathopod 1 like Stenothoe species A, but palm slightly

shorter and less oblique and defined by 1 spine, palm

bearing 1 other lateral spine adjacent to defining spine

and 3 medial spines, all evenly spaced for about half

of palm length (proximal end), dactylar accessory

tooth smaller than in Stenothoe species A; gnathopod

2 also like that of species A, thus juveniliform, defining

spine of palm short, middle cusp obsolescent; pereopods 1-5 lacking spines, of normal slendemess, dactyls

simple, fourth articles slender and produced one-fourth

to one-third along article 5 on pereopods 3-5; expansion of article 2 on pereopods 4-5 narrow for genus

(see figure), of equal form on both pairs of pereopods,

posteroventral lobe obsolescent; rami of uropod 1 extending equally, outer ramus with 2 spines, inner with

1; outer ramus of uropod 2 about two-thirds as long

as inner ramus, latter reaching end of uropod 3, inner

ramus with 2 spines, outer with 1; uropod 3 long and

slender, lacking spines, peduncle, articles 1 and 2 of

ramus all equal in length to each other; telson lacking

lateral spines.

MATERIAL.—JLB Hawaii 10 (3 specimens, largest

1.4 mm).

REMARKS.—The spineless pereopods with simple

dactyls and relatively narrow article 2 of pereopods 4-5

distinguish this species in Hawaii. Juveniles of S. gallensis and S. valida have a few spines on the pereopods.

Superfamily TALITROIDEA

Hyale affinis Chevreux

FIGURE I67e-h

Hyale affinis Chevreux, 1908, pp. 503-506, figs. 21-22.—

Schellenberg, 1938, pp. 67-68.—J. L. Barnard, 1955a,

p. 14 (in part, not fig. 6).

DESCRIPTION.—Following characters and parts as

illustrated for H. laie, new species: female gnathopods

1 and 2, male and female head, eye shape, pereopods 1,

2, 3,4, uropods 1, 2, 3, urosome, telson.

Characters differing from H. laie slightly: eyes paler

purple, aesthetascs on antenna 1 shorter, flagellum of

antenna 2 slightly longer, coxae 1—4 with posterior acclivity sharper and nearer ventral margin; articles 4-6

of pereopod 5 slightly stouter; pleonal epimera 2-3

slightly sharper.

DIFFERENTIAL CHARACTERS (male).—Hand and

palm of gnathopod 1 slightly expanded, dactyl not

reaching full length of palm but reaching spine near

softly rounded defining corner armed with several

short setae, not bulging or cuspidate, palm otherwise

setose; gnathopod 2 with palm more than twice as

long as free posterior margin of hand, oblique and

slightly convex, defined by 2 spines, overall palm recessed deeply from posterior defining bulge, densely

spinose.

MATERIAL.—JLB Hawaii 17 (5). Poipu Beach,

Kauai, 17 September 1959, collector H. Caspers

(50+).
252

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 166.—Stenothoe species A and B. Figures a-i, Stenothoe species A, juvenile, 1.0 nun,

JLB Hawaii 3: a, apex of gnathopod 1; b, coxa 3; c, d, e, f, pereopods 2, 5, 4, 3; g, h, apices of

dactyls on pereopods 2, 1; i, gnathopod 2. Stenothoe species B, female, 1.4 mm, JLB Hawaii 10:

;, pereopod 5.
NUMBER 3 4 253

FIGURE 167.—Hyale honoluluensis Schellenberg and Hyale affinis Chevreux. H.h., male, 6.4 mm,

Fee 1: a, articles 1—4 of lateral gnathopod 2 ; b, gnathopod 1; c, pleonal epimera 1—3; d, palm of

gnathopod 1. H. a., female, 4.9 mm, Jarvis Island: e, maxilla 1. Male, 6.6 mm, JLB Hawaii 17:

f> S> gnathopod 1; h, lateral gnathopod 2.
254 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—The Oahuan population of this species

resembles that described by Schellenberg from Laysan

Island, but the rami of uropod 1 extend equally, the

ramus of uropod 3 is equal to the peduncle, and the

palm of male gnathopod 1 is without a defining bulge.

The Hawaiian population (of the high islands) thus

differs from that of the Gambier Archipelago in the

softly rounded, unbulging defining corner of male

gnathopod 1 palm, the strongly defined acclivities on

the anterior coxae, the presence of a pair of posterior

setae on the hands of the female gnathopods, and the

longer palm of male gnathopod 2.

Male H. affinis in Hawaii differs from young males

of H. honoluluensis Schellenberg in the dactyl of

gnathopod 1 overlapping the palmar spine and the

presence of a small spine on the posterior margin of the

hand of gnathopod 2, proximal to the palm. Females

differ in gross characters only in the coxal acclivities of

H. affinis being nearer the ventral margins than in H.

honolulensis.

Figure 6 of Barnard (1955a) is erroneous, representing actually female gnathopods of H. bishopae J. L.

Barnard.

DISTRIBUTION.—Tropical Pacific.

Hyale ayeli J. L. Barnard

FIGURES 168,169

DIAGNOSIS OF MALE.—Lateral cephalic lobes broadly truncate, anteroventral cephalic excavation shallow

but asymmetrically indented, anteroventral corner of

head rounded; eyes large, subcircular, black cores in alcohol; antennae of medium length, antenna 1 reaching more than halfway along antenna 2, flagellum with

numerous aesthetascs, peduncle of antenna 2 of medium length, thick, and bearing a few medioposterior

setular brushes, flagellum of medium thickness and

bearing weak brushes; coxa 1 of normal breadth,

scarcely extended forward, coxae 1-4 with slightly attenuate but strong, rounded acclivities, coxa 4 with

simple excavation; article 2 of gnathopod 1 unproduced anterodistally, article 4 with medium distal extension, lobe on article 5 broad, article 6 evenly rectangular but not thin, palm oblique, defined by pair

of spines and about one-half of posterior margin of

hand armed with short setae, dactyl fitting palm; article 2 of gnathopod 2 with broad anterodistal lobe,

article 3 with broad anterior lobe, article 4 weakly

extended, article 5 immersed posteriorly, hand very

large, palm oblique, spinose, and simple in young males

but progressively marked with strong, distal, quadrate

protrusion, palm about as long as posterior margin of

hand, defined by weak hump and pair of small spines,

dactyl fitting palm; pereopods 1-2 with one large striate distal locking spine partially hidden by a few setae,

one of which strongly hooked, next proximal armament composed of 2 slender setae and 1 slightly thickened spine-seta scarcely one-fourth as wide as main

spine; pereopods 3-5 with distal locking spine small

and slightly striate spirally, next spine in tandem much

larger and more heavily striate, almost forming screw,

next proximal spines forming pair of small unstriate

members; dactyls of pereopods 1-5 slightly striate,

bearing weak distal setule and facial setule; article 6

of j>ereopods 3-5 lacking posterior spines or setae;

pleonal epimeron 1 rounded posteroventrally, epimera

2-3 slightly sinuous posteriorly and with strong blunt

posteroventral protrusion; uropod 1 with large distolateral spine on peduncle, outer rami of uropods 1-2

without dorsal spines, uropod 3 with peduncle and

ramus of medium breadth and spinose apex.

FEMALE.—Gnathopod 1 like that of male, but

gnathopod 2 with narrower lobe on article 5 and

slightly bulging posterior margin of hand.

JUVENILE.—Gnathopods 1-2 like adults of H. laie,

thus with only 2-3 posterior setae on hands.

MATERIAL.—The original material, especially the

lot from Hanauma Bay, Oahu (36).

REMARKS.—Ruffo (1956) considered Hyale ayeli

to be a synonym of H. media (Dana) in view of Hurley (1957) and Stephensen (1949) reporting specimens with setular brushes on antenna 2, but the following characters of the Hawaiian specimens differ from

those described by Hurley from New Zealand: (1) the

palm of gnathopod 2 has a strong protrusion in adult

and subadult males; (2) the posteroventral corner of

article 2 on pereopod 5 is rounded; (3) uropod 1 has a

strong distolateral spine on the peduncle (not present

in H. media) ; (41 uropods 1-2 lack dorsal spines on

the outer rami; (5) the posterior setae on gnathopod 1

of both sexes and gnathopod 2 of the female are more

broadly spread than in H. media.

Hyale grandicornis (Kreyer)

Hyale grandicornis Kreyer.—Hurley, 1957, pp. 904-907, figs.

1-23 [with references] [not forma thomsoni\.

Hyale novaezealandiae Thomson.—Iwasa, 1939, pp. 276-278,

fig. 16, pi. 16.
NUMBER 3 4 255

FIGURE 168.—Hyale ayeli J. L. Barnard, male, 8.0 mm, Hanauma Bay, 24 November 1951: a, b,

c, pereopods 3, 4, 5; d, e, medial gnathopod 2; /, uropod 3; g, pereopod 2; k, coxa 3. Female,

7.0 mm: i, lateral gnathopod 1;;', left side of telson from lateral view.

360-034 O - 18
256

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 169.—Hyde ayeli J. L. Barnard, male, 8.0 mm, Hanauma Bay, 24 November 1951: a,

head; b, c, gnathopods 1, 2; d, e, dactyls of pereopods 2, 5; /, palm of medial gnathopod 2; g,

pleon.
NUMBER 3 4 257

Hyale grandicornis bishopae J. L. Barnard, new

combination

FIGURE 170

Hyale bishopae J. L. Barnard, 1955a, pp. 16-17, fig. 8.

DIAGNOSIS OF MALE.—Lateral cephalic lobes broadly

truncate, anteroventral cephalic excavation shallow

and flat, anteroventral corner of head rounded-quadrate; eyes large, subovate, with black-brown-purple

cores; antennae short, antenna 1 reaching about halfway along antenna 2, flagellum with numerous

aesthetascs, peduncle of antenna 2 of medium length,

flagellum thick and bearing few setules; coxa 1 extraordinarily broad, slightly extended forward, coxae 1-4

with strong cuspidiform posterior acclivities, coxa 4

with simple excavation; article 2 of gnathopod 1 unproduced anterodistally, article 4 with weak posterodistal process, lobe of article 5 of medium breadth,

article 6 evenly rectangular, palm oblique, defined by

pair of spines and about one-fourth of posterior margin

of hand armed with short setae, dactyl fitting palm;

article 2 of gnathopod 2 with weak anterodistal lobe,

article 3 with distinct but small tumid lobe, article 4

weakly extended, article 5 immersed posteriorly, hand

very large, palm oblique and simple, spinose, slightly

longer than free posterior margin of hand, defined by

two enlarged spines, posterior margin with several

distal setules, dactyl weakly curved, fitting palm; pereopods 1-2 with pair of small, distal, straight, unstriate

spines in tandem; pereopods 3-5 with straight distal

locking spine enlarged and slightly striate axially, in

tandem with smaller spine; dactyls of pereopods 1-5

slightly striate distalward, bearing large, stiff, spirally

striate setule in middle, dactyl with one facial setule;

article 6 of pereopods 3-5 lacking posterior setae or

spines; pleonal epimeron 1 rounded-quadrate posteroventrally, bearing 2 weak notches, epimeron 2 concave

posteriorly and bearing rounded posteroventral protrusion, epimeron 3 convex posteriorly and bearing

smaller protrusion than epimeron 2; uropod 1 with

medium-sized mediodistal spine on peduncle, rami of

uropods 1-2 spinose, uropod 3 with broad peduncle

not especially short, ramus slender, with narrow

oblique spinose apex and one dorsal, subterminal, discontiguous spine seen from lateral view; telson with

quadrate lobes.

FEMALE.—Gnathopod 1 like that of male, but

gnathopod 2 like that of female H. iole, new species.

MATERIAL.—The type material: Hanauma Bay,

Oahu, 24 November 1951, algae, collector Donald

Bates (6 specimens, holotype not seen again).

REMARKS.—This species differs from H. iole, new

species, in the broader, shorter coxa 1, the shorter

article 5 of gnathopod 1, and larger hand with fewer

posterior setae, in the distinct tumid lobe of article 3

on male gnathopod 2, in the smallness of the distal

locking spine on pereopods 1-2 and the presence of a

second spine in tandem, in the spiral striations of the

dactylar seta, in the smallness of the mediodistal spine

on the peduncle of uropod 1, in the disjunct ramal

spine on uropod 3 (seen from lateral view), in the

more strongly rounded epimeron 1 with small notches

bearing setules, and in the absence of heavy brushes of

setules on antenna 2.

Hyale bishopae seems to fall into the H. grandicornis

complex represented by populations from New Zealand, South Africa, and Japan. The characters uniting

these widespread populations are especially: (1) a

large dactylar seta on pereopods 1-5; (2) a disjunct

subterminal spine on the ramus of uropod 3; (3) a

small or medium-sized straight distal locking spine on

pereopods 1-5 with another spine proximal to it; (4)

the dominant spine on the peduncle of uropod 1 is

mediodistal and scarcely enlarged; (5) coxae 1-4 have

a strongly attenuate posterior acclivity; (6) hands of

male and female gnathopod 1 and female gnathopod 2

have a narrowly spread clump of several setae on the

posterior margin.

Tiny pectinations seen on the dactyl of pereopods

1-5 in New Zealand and Japanese individuals are not

present in Hawaiian specimens; uropod 3 is thin like

that on New Zealand but not on Japanese specimens.

The latter population has a pyriform article 6 on male

gnathopod 2. The dactylar seta on pereopods 1—5 of

H. g. bishopae has spiral striations not mentioned for

the other members of the complex.

DISTRIBUTION.—Hawaiian Islands.

Hyale honoluluensis Schellenberg

FIGURES 167a-d, 171

Hyale honoluluensis Schellenberg, 1938, pp. 69-71, figs. 35b,

c—J. L. Barnard, 1955a, p. 18; 1965a, p. 520, figs. 22, 23.

DESCRIPTION.—Following characters and parts as

illustrated for H. laie, new species: Female gnathopods
258 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 170.—Hyale grandicornis bishopae J. L. Barnard, male, 6.9 mm, Hanauma Bay, 1951:

a, pleonal epimeron 1 (tilted); b, articles 2-3 of pereopod 1; c, uropod 3; d, coxa 4; e, uropod 1;

/, locking spines of pereopod 2; g, dactyl of pereopod 4; h, left lateral side of telson; i, j , medial

gnathopods 1, 2.

1 and 2, eye shape, pereopods 1-5, uropods 1, 2, 3,

urosome, telson, coxae 1-4, but coxa 1 not as strongly

extended forward.

Characters differing from H. laie slightly: articles

2-3 of antenna 1 equal in length to each other, female

coxa 2 slightly longer.

DIFFERENTIAL CHARACTERS.—Hand and palm of

gnathopod 1 slightly expanded, dactyl not reaching

full length of palm and not reaching spine near softly

rounded corner bearing 2 stout spines, margin near

spine slightly bulging, palm otherwise setose; gnathopod 2 of terminal male with palm occupying entire

posterior margin of hand, dactyl overlapping free

palmar margin, younger male with shorter palm defined by two spines but not deeply recessed and lacking

posterior spinule seen in H. affinis; presumed juvenile

male with short oblique palm defined by 2 spines, palm

shorter than posterior margin of hand, latter with 1
NUMBER 3 4 259

FIGURE 171.—Hyale honoluluensis Schellenberg, male, 6.4 mm, Fee 1: a, b, medial gnathopods

1, 2. Female, 4.8 mm: c, d, medial gnathopods 1, 2. Juvenile, 5.2 mm: e, palm of gnathopod 1;

/, lateral gnathopod 2. Juvenile, 3.5 mm: g, medial gnathopod 1; h, lateral gnathopod 2.

setule (thus juvenile male possibly that of H. affinis

and not H. honoluluensis).

Eyes ruby or garnet in formaldehyde but rapidlyturning dark purple in formaldehyde or alcohol.

MATERIAL.—JLB Hawaii 12 (2). Fee 1 (51). Canton Island, February 1958, collector Drs. Degener

(100 + ). French Frigate Shoals, Tern Island, 13 August 1959, collector H. Caspers (25).

REMARKS.—Probably H. honoluluensis should be

reduced to a subspecies of H. chevreuxi K. H. Barnard

(see J. L. Barnard, 1965a, for references). Specimens

from Canton Island, a member of the Phoenix Islands

and thus close to the Gilberts from which Schellenberg

reported H. chevreuxi, have the palm of male gnathopod 2 as in H. honoluluensis, and suggest possibly that

a cline exists between the original material of H. chevreuxi from the Seychelles and the eastern populations.

Barnard (1965a) also identified H. honoluluensis from

the Marshall Islands.

DISTRIBUTION.—Hawaiian, Phoenix, and Marshall

Islands.
260

Hyale iole, new species

FIGURE 172

DIAGNOSIS OF MALE.—Lateral cephalic lobes broadly

truncate, anteroventral cephalic excavation shallow

and flat, anteroventral corner of head rounded-quadrate; eyes large, subcircular, with black-brown-purple

cores; antennae of medium length, antenna 1 reaching

halfway along antenna 2, flagellum with few aesthetascs, peduncle of antenna 2 of medium length,

both peduncle and flagellum base with dense brushes

of setules; coxa 1 broad, slightly extended forward,

coxae 1-4 with strong posterior acclivity projecting as

tooth, coxa 4 with simple excavation; article 2 of

gnathopod 1 with slight, subterminal, quadrate extension on anterodistal margin, article 4 with strong posterodistal process, lobe of article 5 of medium breadth,

article 6 evenly rectangular, palm slightly oblique, defined by pair of spines and about half of posterior

margin of hand armed with short setae, dactyl fitting

palm; article 2 of gnathopod 2 with slightly bilobate,

strong anterodistal protuberance, article 3 unlobate,

article 4 weakly extended, article 5 immersed posteriorly, hand very large, palm oblique and simple,

spinose, slightly longer than free posterior margin of

hand, defined by two slightly enlarged spines, posterior margin with 2 distal setules, dactyl curved, fitting

palm; pereopods 1-2 with 1 small, nearly straight distal

locking spine, slightly striate axially; pereopods 3-5

with distal locking spine similar and slightly larger

than next proximal spine; dactyls of pereopods 1-5

slightly castellate in middle and bearing very large

stiff seta in middle, seta slightly striate axially, dactyl

with one facial setule; article 6 of pereopods 3-5 lacking posterior setae or spines; pleonal epimeron 1

rounded-quadrate posteroventrally, epimeron 2 concave posteriorly and bearing rounded posteroventral

protrusion, epimeron 3 convex posteriorly and bearing

smaller protrusion than epimeron 2; uropod 1 with

large mediodistal spine on peduncle, rami of uropods

1-2 spinose, uropod 3 with especially short broad

peduncle, ramus with oblique, spinose distal margin,

no discontiguous spine from lateral view; telson normal.

FEMALE.—Article 2 of gnathopods 1-2 with rounded

or subquadrate anterodistal lobe, gnathopod 1 otherwise like that of male; gnathopod 2 stouter than

gnathopod 1, article 4 like that on gnathopod 2 of

male, article 4 with narrow lobe, article 6 stouter than

in gnathopod 1, palm more oblique, and short posteSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rior setae extending only about one-third of hand

length.

HOLOTYPE.—Bishop Museum collections, catalog

number 7303, male, 11.3 mm.

TYPE-LOCALITY.—Kahalui Harbor, Maui, 28 May

1953.

MATERIAL.—Thirty-three specimens from the typelocality.

REMARKS.—This species has a superficial resemblance to H. ayeli, but differs in numerous characters,

including: (1) narrower lobes on article 5 of gnatho-

|x>d 1 in both sexes and gnathopod 2 in the female;

(2) the shorter, broader peduncle of uropod 3; (3)

the presence of spines on both rami of uropods 1-2;

(4) the shift of the major peduncular spine on uropod

1 from lateral to medial side; (5) the broader hand

and thinner dactyl of male gnathopod 2, plus the presence of distinct, though small defining spines on the

palm; (6) the presence of a large, dominant dactylar

seta on the pereopods; (7) the slightly stronger coxal

cusps.

Hyale iole has affinities with H. grandicornis bishopae from Hawaii, but the former differs from the latter in the larger mediodistal spine on the peduncle of

uropod 1, the absence of a disjunct spine on the ramus

of uropod 3, the slightly castellate dactyls of pereopods

1-5, the stout uropod 3, the antennal brushes and the

longer row of posterior setae on the hands of gnathopods 1 and 2.

The strong brushes of setules on antenna 2 resemble

those on Hyale plumulosus (Stimpson) from coolwater North America. Presumed specimens of that

poorly known species from Point Richmond, Contra

Costa County, California, differ from H. iole in the

presence of a distinct point and notch on pleonal epimeron 1, in a slightly more extended posteroventral corner on epimeron 2, the absence of black pigment in the

eyes, more attenuate and sharper coxal acclivities, and

the blunt, unextended article 4 on male gnathopod 1.

Fully adult male gnathopod 2 of H. plumulosus has not

been observed in this comparison.

DISTRIBUTION.—Hawaiian Islands.

Hyale laie, new species

FIGURES 173, 174

DIAGNOSIS OF MALE.—Lateral cephalic lobes broadly

truncate, anteroventral cephalic excavation shallow

and flat, anteroventral corner rounded-quadrate; eyes
NUMBER 3 4 261

n

FIGURE 172.—Hyale iole, new species, holotype, male, 11.3 mm, Kahului Harbor, Maui: a, coxa

4; b, locking spine of pereopod 1; c, half of telson, right side, lateral; d, uropod \; e,f, gnathopods

I* 2; g, pleonal epimera 1-3; h, uropod 3. Female, 10.4 mm: i, head and antennae; ;, uropod 3;

k, palp article 4 of maxilliped; /, articles 2-3 of gnathopod 1; m, dactyl of pereopod 5; n,

gnathopod 2.
262

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 173.—Hyale laie, new species, holotype, male, 4.2 mm, JLB Hawaii 14: a, b, gnathopods

1, 2; c, d, e, pereopods 3, 4, 5; /, part of pereopod 5, medial view. Female, 5.6 mm: g, coxa 3;

h, gnathopod 1; i, uropod 3.
NUMBER 3 4 263

d

h

FIGURE 174.—Hyde laie, new species, holotype, male, 4.2 mm, JLB Hawaii 14: a, head; b,

posterodistal end of article 5 on pereopod 2 (upside down); c, pleonal epimera 1-3. Female,

5.6 mm: d, right half of telson, arrow pointing posteriorly; e, pereopod 2; /, .dactyl of pereopod 5;

g, gnathopod 2; h, urosome.

large, subcircular, with black-brown-purple cores; antennae of medium length, antenna 1 reaching halfway

along flagellum of antenna 2, flagellum with numerous

aesthetascs, peduncle of antenna 2 short, flagellum

setose only; palp of maxilla 1 uniarticulate and reaching to apex of outer plate; coxa 1 very broad, slightly

extended forward, coxae 1-3 with strong, quadrate

posterior acclivity, coxa 4 with bisinuate posterodorsal

excavation; article 2 of gnathopod 1 unproduced anterodistally, article 4 weakly extended posterodistally,

lobe of article 5 narrow, article 6 broadly expanded,

anterodistal corner tumid, palm oblique, defined by

large, blunt cusp, palm convex and bearing large spine

in middle, dactyl fitting palm; article 2 of gnathopod

2 broadly lobate anterodistally, article 3 with broad

hemicircular lobe, article 4 weakly extended, article 5

immersed posteriorly but bearing slight lobe, hand very

large, bearing 2 anteroproximal spines, palm oblique,

much longer than free posterior margin of hand, bearing distal truncate process, remainder of palm excavate and bearing slight hump near rounded defining

corner, palm spinose throughout, dactyl fitting palm

and slightly S-shaped on inner margin; pereopods 1-5

with main locking spine and smaller spine heavily

striate in corkscrew pattern distally, dactyls bearing

small distal setule and very weak castellations and
264

striations on inner edge; article 2 of pereopods 3-5

broad, subcircular or slightly flattened posteriorly on

pereopod 4, weakly serrate, articles 4-6 moderately

slender and without special features, article 6 lacking

posterior setae; pleonal epimera rounded-quadrate posteriorly, epimeron 2 slightly sinuous and scarcely extended, epimeron 3 slightly smaller, less sinuous, with

slight posteroventral extension, subacute corner; uropod 1 with large distolateral spine on peduncle, rami

of uropods 1-2 dorsally spinose, uropod 3 with row of

peduncular setae, thin ramus slightly shorter than

peduncle, bearing only terminal spines; telson normal.

FEMALE.—Article 2 of gnathopod 1 slightly tumid

anterodistally, article 4 with small subacute spinose

protrusion posterodistally, article 5 longer than in

male, article 6 slender, subrectangular, with 2 posterior

setae, palm oblique, simple, slightly convex, defined by

spine, dactyl fitting palm; gnathopod 2 similar to

gnathopod 1 but larger, lobe of article 5 more slender.

HOLOTYPE.—Bishop Museum collections, catalog

number 7304, male, 4.2 mm.

TYPE-LOCALITY.—JLB Hawaii 14, 1 mile N of Kualoa Point, Oahu, intertidal, algae, dead coral heads,

23 May 1967.

MATERIAL.—JLB Hawaii 14 (17), 15 (14), 17 (7).

18 (3). Kauai, Poipu, 17 September 1959, Chir. haw.,

H. Caspers (1).

This species has strong affinities with H. affinis

Chevreux (1908), but male gnathopod 2 palm of H.

affinis is simple, the palm of gnathopod 1 lacks a midspine, and the defining hump is very weak.

Hyale pusilla Chevreux (1908) has the ornamented

palm seen in H. laie, but it has 2 distal humps instead

of 1 and otherwise lacks spines except on those humps.

Male gnathopod 1 of H. pusilla is slightly expanded

distally but the palm is simple and not defined by a

cusp.

Gnathopod 1 of H. laie resembles the male of H.

galateae Stebbing (1899a), but the dactyl of Stebbing's

species is bifid.

Hyale laie probably has its closest affinities with that

species identified by Iwasa (1939) from Japan as H.

dollfusi Chevreux, a species originally described from

the Mediterranean Sea. Perhaps the Japanese form

should become a subspecies of H. laie and not be identified with H. dollfusi. Iwasa's material differs from the

Mediterranean species in the distinct palmar protrusion of male gnathopod 2, though it is weaker than in

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

H. laie. The Japanese material also has relatively

longer antenna 2 than that from Hawaii.

DISTRIBUTION.—Hawaiian Islands.

Hyale Rathke

Lelehua, new subgenus

DIAGNOSIS.—Hyale with article 4 of male maxillipedal palp bearing long whiplike terminal seta.

TYPE-SPECIES.—Hyale (Lelehua) waimea, new

species.

Hyale (Lelehua) waimea, new species

FIGURES 175, 176, 177

DESCRIPTION.—Lateral cephalic lobes scarcely truncate, slightly convex, anteroventral cephalic excavation

weak: eyes large, subovatc, red or yellow in formaldehyde, clear in alcohol; antennae short, 1 slightly shorter

than 2, flagellum of 1 with aesthetascs; palp of maxilla

1 uniarticulate and reaching to apex of outer plate;

palp article 4 of female maxilliped with several long

stiff setae lining medial margin (when flattened) ; coxa

1 very broad, slightly extended forward, coxae 1-3

with obsolescent posterior acclivity, 4 with small sharp

acclivity in posterodorsal excavation; article 2 of

gnathopod 1 unproduced anterodistally, article 4 unextended posterodistally, lobe of article 5 of medium

breadth, article 6 narrow and rectangular, palm

oblique, defined by pair of spines, posterior margin

setose, medial face of hand with row of stout setae,

dactyl fitting palm; article 2 of gnathopod 2 with broad

but not deeply extended anterodistal lobe, article 3

with small mammilliform hump, article 4 distally triangular, article 5 hidden between articles 4 and 6 and

bearing short lobe extending over article 6, latter

normally broad, palm slightly oblique, bearing deep

narrow rounded excavation in middle, with medial

flanges narrowing the excavation on medial side, palm

defined by weak tooth adjacent to small sinus, bearing

2 medial spines, dactyl thick, fitting palm, bearing inner hurnp; pereopods 1-5 with small, striate. locking

spine adjacent to larger simpler spine, dactyl slightly

striate, bearing distal sharp flake, marginal setule and

facial setule; pereopods 3-5 with broadly expanded

article 2, article 4 on pereopods 3-4 very broad and

nasiform, posteriorly spinose, article 5 also broad on

pereopod 4, article 6 lacking posterior setae; pleonal
NUMBER 34 265

FIGURE 175.—Hyale (Lelehua) ivaimea, new subgenus, new species, holotype, male, 3.8 mm,

JLB Hawaii 5: a, telson; b, maxilla 2; c, inner plate of maxilliped, setae removed; d, lower lip;

e, uropod 3; f, mandible; g, maxilla 1; h, inner plate of maxilliped, with setae; i, maxilliped.

Female, 3.3 mm: ;, palp articles 3—4 of maxilliped.

epimeron 1 rounded posteroventrally, epimera 2-3

with slightly sinuous posterior margin and rounded

posteroventral protuberance; uropod 1 with large

distolateral spine on peduncle, rami of uropods 1-2

spinose, uropod 3 with 2 basal peduncular setae, broad

short ramus much shorter than peduncle, bearing only

terminal spines; telson sharply triangular.

FEMALE.—Gnathopod 1 like that of male, gnathopod

2 similar in size and shape to gnathopod 1, but article

4 with strong posterodistal extension, article 6 poorly

setose posteriorly and medially compared with gnathopod 1; articles 4—5 of pereopod 4 like those of pereopod

5 of male and not like pereopod 4 of male.

JUVENILE MALE.—Palm of gnathopod 2 with weak

excavation, dactyl without inner hump.

HOLOTYPE.—Bishop Museum collections, catalog

number 7305, male, 3.8 mm.

TYPE-LOCALITY.—JLB Hawaii 5, off Ewa Beach,

Oahu, 30 m, Pocillopora, 29 January 1967.

MATERIAL.—Ten specimens from the type-locality.
266 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 176.—Hyale (Lelehua) waimea, new subgenus, new species, holotype, male, 3.8 mm,

JLB Hawaii 5: a, coxa 4; b, c, d, e, pereopods 1, 3, 4, 5; /, dactyl of pereopod 1. Juvenile male,

2.6 mm: g, palm of medial gnathopod 2.
NUMBER 34 267

d

FIGURE 177.—Hyale (Lelehua) waimea, new subgenus, new species, male, 3.8 mm, JLB Hawaii

5: a, gnathopod 1; b, pleonal epimera 1—3; c, d, lateral and medial gnathopod 2; e, urosome.

Female, 3.3 mm: f, g, gnathopods 1, 2, setae removed from latter; h, head.
268 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—The long flagellate seta of the male

maxillipedal palp should be used as a subgeneric character to separate this species from the many other members of Hyale lacking it. Maxillipedal palps of few of

the 45 known species have been specifically mentioned

and others may have this character. Hyale carinata

(Bate) of the Mediterranean Sea (Chevreux and Fage,

1925) also has this seta but it is shorter. That species

probably also should be assigned to Lelehua.

DISTRIBUTION.—Hawaiian Islands.

Hyale species (cf. rubra [Thomson])

FIGURE 178

DIAGNOSIS OF MALE.—Lateral cephalic lobes broadly

truncate, anteroventral cephalic excavation shallow

and flat, anteroventral corner of head roundedquadrate; eyes dark in alcohol (shape destroyed by

preservative) ; antennae of medium length, antenna 1

reaching about halfway along antenna 2, flagellum

with a few aesthetascs, peduncle of antenna 2 of medium length, flagellum normal and bearing a few

setules; coxa 1 of regular expansion, scarcely extended

forward, coxa 1 weakly lobate posteriorly but with no

sharp acclivity, coxae 2-3 with strong, quadrate, softly

rounded acclivity; coxa 4 with slightly sinuous excavation; article 2 of gnathopod 1 with spinose, quadrate

anterodistal extension proximal to corner, article 4 almost unproduced distally, lobe on article 5 thin, article 6 narrowly rectangular and tapering distally, palm

very oblique and short, defined by 2 spines, dactyl

short, fitting palm, half of posterior margin of hand

FIGURE 178.—Hyale species (cf. H. rubra), male, 10.5 mm, Kahului Harbor, Maui: a, largest

spine on article 6 of pereopod 2; b, uropod 3; c, apex of pereopod 5; d, medial gnathopod 2;

e, f, coxae 3, 4 (in different magnifications) ; g, article 4 of pereopod 1; h, left side of telson,

lateral view; i, articles 2-4 of pereopod 2;;, gnathopod 1.
NUMBER 34 269

FIGURE 179.—Parhyalella pietschmanni Schellenberg, male, 10.3 mm, Lanikai: a, body; b, apex

of dactyl on gnathopod 1; c, d, medial gnathopods 1, 2; e, uropod 3; f, lateral palm of gnathopod

2; g, urosome. Female, 9.2 mm: h, apex of dactyl on gnathopod 2.
270

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 180.—Parhyalella pietschmanni Schellenberg, male, 10.3 mm, Lanikai: a, dactyl of

pereopod 2; b, maxilla 1; c, mandible; d, head; e, lower lip; /, maxilliped; g, apex of uropod 3;

h, pleonal epimera 1-3. Female, 9.2 mm: i, telson; ;, head; k, I, medial gnathopods 1, 2.
NUMBER 34 271

lined with short setae; article 2 of gnathopod 2 with

medium anterodistal lobe, article 3 with large tumid

lobe, article 4 weakly extended, article 5 immersed

posteriorly, hand very large, with 2 anteroproximal

spines, palm oblique and simple, spinose, about equal

to free posterior margin of hand, spines at defining

corner not enlarged, posterior margin of hand with 3

groups of tiny setules, dactyl curved, fitting palm;

pereopods 1-5 with small, weakly striate distal spine,

next proximal spine large and striate like other marginal spines, dactyl with small distal seta and facial

setule, with weak proximal castellations; pleonal

epimera and uropods 1-2 as figured for H. laie, new

species, thus largest peduncular spine of uropod 1 on

lateral side, outer ramus of uropod 2 much thinner

and shorter than outer ramus, rami of uropods 1-2

marginally spinose; uropod 3 slender, all spines of

ramus apical; telson with triangular lobes.

Female unknown.

MATERIAL.—Kahalui Harbor, Maui, 28 May 1953

(3 males).

REMARKS.—This species belongs with H. laie-honoluluensis-affinis complex in its uropod 1, but differs

from those species and H. g. bishopae and H. ayeli in

the thin hand of gnathopod 1, from the laie complex

by the numerous posterior setae on that hand, and

from H. g. bishopae and H. ayeli in the obsolescent

posterior acclivities on coxae 1—3 and the srnallness and

distal placement of the main dactylar seta on pereopods 1-5.

This population of 3 males appears to be related to

H. rubra (Thomson) (Hurley, 1957), but gnathopod

1 tapers distally proximal to the palm, and the lobe

on article 5 is thin; article 2 of pereopods 1-2 has

relatively long setae on the anterodistal corner. The

specimens appear similar to those identified by Iwasa

(1939) from Japan as Hyale schmidti.

Parhyalella pietschmanni Schellenberg

FIGURES 179, 180

Parhyalella pietschmanni Schellenberg, 1938, pp. 71-74, figs.

36,37.

The long description and several figures published by

Schellenberg, plus the additional figures presented

herein, should adequately define this species for the

present. The eyes in formaldehyde are a mixture of

garnet below frosty pink ommatidial outlines, turning

to dark brownish purple in alcohol, the eyes completely

pigmented to their periphery, unlike most Hawaiian

marine amphipods with dark ocular pigment.

Parhyalella pietschmanni differs from P. whelpleyi

(Shoemaker, 1933b) in the broader coxa 4, the larger

pleonal epimeron 2, the slightly broader sixth article of

gnathopods 1-2, and the much longer ramus of uropod

3.

MATERIAL.—Lanikai, Oahu, 20 May 1967, collection

D. H. Devaney, beach wash associated with algae (10).

DISTRIBUTON.—Hawaiian Islands.

Literature cited

Alderman, A. L.

1936. Some New and Little Known Amphipods of

California. University of California Publications

in Zoology, 41:53-74, figures 1-51.

Audouin, V.

1826. Page 93, plate 11 in J. C. Savigny's "Description

de I'Egypte, Publiee par les Ordres de sa Majeste

I'Empereur Napoleon-le-Grand. Histoire Naturella. Animaux articules. Crustaces. Vol. 1 iv.

Barnard, J. L.

1952. A New Species of Amphipod from Lower California (Genus Eriopisa). Pacific Science, 6:

295-299, figures 1, 2.

1953. On Two New Amphipod Records from Los

Angeles Harbor. Bulletin Southern California

Academy of Sciences, 52:83-87, plate 15.

1954a. Marine Amphipoda of Oregon. Oregon State

Monographs, Studies in Zoology, 8:1-103, 1

figure, plates 1-33.

1954b. A New Species of Microjassa (Amphipoda) from

Los Angeles Harbor. Bulletin Southern California Academy of Sciences, 53:127-130, plates

35-36.

1955a. Gammaridean Amphipoda (Crustacea) in the

Collections of Bishop Museum. Bernice P.

Bishop Museum, Bulletin 215:1-46, figures 1-20.

1955b. Notes on the Amphipod Genus Aruga with the

Description of a New Species. Bulletin Southern

California Academy of Sciences, 54:97—103,

plates 27-29.

1956. Two Rare Amphipods from California with Notes

on the Genus Atylus. Bulletin Southern California Academy of Sciences, 55:34—43, plates

12-14.

1959. Estuarine Amphipoda. In Ecology of Amphipoda and Polychaeta of Newport Bay, California,

by J. L. Barnard and D. J. Reish. Allan Hancock

Foundation Publications, Occasional Paper 21:

13-69, figures 1-14.

1961. Gammaridean Amphipoda from Depths of 400

to 6000 meters. Galathea Report, 5:23-128, figures 1-83.

360-034 O - 19
272

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Barnard, J.

1962a.

1962b.

1962c.

1965a.

1965b.

1967.

1969a.

1969b.

in press.

Barnard, J.

1960.

Barnard,K

1916.

1932.

1937.

1940.

Bate, C. S.

1862.

L.—Cont.

Benthic Marine Amphipoda of Southern California: Families Aoridae, Photidae, Ischyroceridae, Corophiidae, Podoceridae. Pacific Naturalist 3:1-72, figures 1-32.

Benthic Marine Amphipoda of Southern California: Families Tironidae to Gammaridae.

Pacific Naturalist 3:73-115, figures 1-23.

Benthic Marine Amphipoda of Southern California: Families Amphilochidae, Leucothoidae,

Stenothoidae, Argissidae, Hyalidae. Pacific Naturalist 3:116-163, figures 1-23.

Marine Amphipoda of Atolls in Micronesia. Proceedings of the United States National Museum,

117:459-552, figures 1-35.

Marine Amphipoda of the Family Ampithoidae

from Southern California. Proceedings of the

United States National Museum, 118(3522):

1-46, figures 1-28.

Bathyal and Abyssal Gammaridean Amphipoda

of Cedros Trench, Baja California. United

States National Museum Bulletin 260:i-vi, 1-

204, figures 1-92.

Gammaridean Amphipoda of the Rocky Intertidal

of California: Monterey Bay to La Jolla. United

States National Museum Bulletin 258:1-230,

figures 1-65.

A Biological Survey of Bahia de Los Angeles,

Gulf of California, Mexico, IV. Benthic Amphipoda. Transactions of the San Diego Society of

Natural History, 15:175-228, figures 1-30.

Keys to the Hawaiian Marine Gammaridea, 0-30

meters. Smithsonian Contributions to Zoology.

L. and R. R. Given

Common Pleustid Amphipods of Southern California, with a Projected Revision of the Family.

Pacific Naturalist, 1(17):37-48, figures 1-6.

.H.

Contributions to the Crustacean Fauna of South

Africa, 5: The Amphipoda. Annals of the South

African Museum, 15:105-302, plates 26-28.

Amphipoda. Discovery Reports, 5:1-326, figures 1-174, plate 1.

Amphipoda. The John Murray Expedition

1933-34 Scientific Reports, 4:131-201, figures

1-21.

Contributions to the Crustacean Fauna of South

Africa. XII. Further Additions to the Tanaidacea, Isopoda, and Amphipoda, Together with

Keys for the Identification of Hitherto Recorded

Marine and Fresh-Water Species. Annals of

the South African Museum, 32:381-543, figures

1-35.

Catalogue of the Specimens of Amphipodous

Crustacea in the Collection of the British Museum,

London, iv + 399, plates 1, la, 2-58.

Chevreux, E.

1901. Crustaces Amphipodes. Mission Scientifique de

M. Ch. Alluaud aux lies Sechelles (Man, Avril,

Mai, 1892). Mimoires de la Sociiti Zoologique de France, 14:388-438, figures 1-65.

1906. Crustaces Amphipodes. Expedition Antarctique

Francaise (1903-1905) Commandie par le Dr.

Jean Charcot. Sciences Naturelles: Documents

Scientifiques, 1-100, figures 1-56.

1908. Amphipodes Recueillis dans les Possessions Franchises de l'Oceanie par M. le Dr. Seurat, Directeur du Laboratorie de Recherches Biologiques de

Rikitea (lies Gambier), 1902-1904. Mimoires

de la Sociiti Zoologique de Prance, 20:470-527,

figures 1-35.

Chevreux, E. and L. Fage

1925. Amphipodes. Faune de France, 9:1-488, figures

1-438.

Chilton, C.

1884. Additions to the Sessile-eyed Crustacea of New

Zealand. Transactions and Proceedings of the

New Zealand Institute, 16:249-265, plates 17-21.

1921. Fauna of the Chilka Lake. Amphipoda. Memoirs of the Indian Museum, 5:521-558, figures

1-12.

1926. New Zealand Amphipoda: No. 6. Transactions

and Proceedings of the New Zealand Institute,

56:512-518, figures 1-4.

Crawford, G. I.

1937. A Review of the Amphipod Genus Corophium,

with Notes on the British Species. Journal of

the Marine Biological Association of the United

Kingdom, 21:589-630, figures 1-4.

Dana, J. D.

1853-55. Crustacea. Part II. United States Exploring Expedition, 14:689-1618, atlas of 96 plates,

de Laubenfels, M. W.

1957. New Species and Records of Hawaiian Sponges.

Pacific Science, 11:236-251, figures 1-15.

Delia Valle, A.

1893. Garnmarina del Golfo di Napoli. Fauna und

Flora des Golfes von Neapel.. . Monographic 20,

i-xi+1-948, atlas of 61 plates.

Doty, M. S.

1961. Acanthophora, a Possible Invader of the Marine

Flora of Hawaii. Pacific Science, 15:547-552.

Edmondson, C. H.

1940. The Relation of the Marine Fauna of Hawaii to

that of Other Sections of the Pacific Area. Proceedings of the Sixth Pacific Science Congress,

7939,3:593-598.

1951. Some Central Pacific Crustaceans. Occasional

Papers of the Bernice P. Bishop Museum, 20:

182-243, figures 1-38.

Ekman, S.

1953. Zoogeography of the Sea. Sidgwick and Jackson Ltd. London.
NUMBER 34

Garth, J. S.

1946. Distribution Studies of Galapagos Brachyura.

Allan Hancock Pacific Expeditions, 5:603—638,

charts 1-10.

Gosline, W. A.

1955. The Inshore Fish Fauna of Johnston Island, a

Central Pacific Atoll. Pacific Science 9:442-

480, figures 1-4.

1958. The Nature and Evolution of the Hawaiian Inshore Fish Fauna. Proceedings of the Eighth

Pacific Science Congress, 3:347-356, 1 figure.

Gosline, W. A. and V. E. Brock

1960. Handbook of Hawaiian Fishes. University of

Hawaii Press, Honolulu: ix + 372, illustrations.

Gurjanova, E.

1938. Amphipoda, Gammaroidea of Siaukhu Bay and

Sudzukhe Bay (Japan Sea). Reports of the

Japan Sea Hydrobiological Expedition of the

Zoological Institute of the Academy of Sciences

of the Union of Soviet Socialistic Republics in

1934, part 1, 241-404, figures 1-59 (in Russian

with English summary).

1951. Bokoplavy Morej SSSR i Sopredel'nykh Vod

(Amphipoda-Gammaridea). Opredeliteli po

Faune SSSR Akademija Nauk SSSR, 41:1-1029,

figures 1-705 (in Russian).

Gurjanova, E. F.

1965. K Voprosu o Sistematike i Rodstvennyx Otnoshchenijax Rodov Eriopisa, Eriopisella i Niphargus

(Sem. Gammaridae, Crustacea—Amphipoda).

Akademija Nauk SSSR, Trudy Zoologicheskogo

Instituta, 35:216-231, figures 1-4.

Hartman, O.

1966. Polychaetous Annelids of the Hawaiian Islands.

Occasional Papers of the Bernice P. Bishop Museum, 23:163-252, figures 1-2.

Haswell, W. A.

1880a. On Some Additional New Genera and Species of

Amphipodous Crustaceans. Proceedings of the

Linnean Society of New South Wales, 4:319-350,

plates 18-24.

1880b. On Some New Amphipods from Australia and

Tasmania. Proceedings of the Linnean Society

of New South Wales, 5:97-105, plates 5-7.

1885. Notes on the Australian Amphipoda. Proceedings of the Linnean Society of New South Wales,

10:95-114, plates 10-18.

Hurley, D. E.

1954a. Studies on the New Zealand Amphipodan Fauna

No. 5. Pleonexes Lessoniae, a New Species of

the Family Amphithoidae. Transactions of the

Royal Society of New Zealand, 81:619-626,

figures 1-2.

1954b. Studies on the New Zealand Amphipodan Fauna

No. 6. Family Colomastigidae, with Descriptions of Two New Species of Colomastix. Transactions of the Royal Society of New Zealand,

82:419-429, figures 1-3.

1957. Studies on the New Zealand Amphipodan Fauna

No. 14. The Genera Hyale and Allorchestes

(Family Talitridae). Transactions of the Royal

Society of New Zealand, 84:903-933, figures 1-9

(with subfigures).

1963. Amphipoda of the Family Lysianassidae from the

West Coast of North and Central America. Allan Hancock Foundation Publications, Occasional Paper 25, 1-165, figures 1-49.

Iwasa, M.

1939. Japanese Talitridae. Journal of Faculty of Science, Hokkaido Imperial University, Series 6,

Zoology, 6:255-296, plates 1-22.

Knox, G. A.

1958. The Distribution of Polychaetes within the IndoPacific. Proceedings of the Eighth Pacific Science Congress 3:403-411.

Kohn, A. J.

1959. The Hawaiian Species of Conus (Mollusca: Gastropoda). Pacific Science 13:368-401. figures

1^*, plates 1-2.

Myers, G. S.

1940. The Fish Fauna of the Pacific Ocean, with

Especial Reference to Zoogeographical Regions

and Distribution as They Affect the International

Aspects of the Fisheries. Proceedings of the

Sixth Pacific Science Congress, 1939, 3:201-210.

Nagata, K.

1961 a. A New Atylid Amphipod from Japan. Annotationes Zoologicae Japonenses, 34:216—218, figures

1-2.

1961b. Two New Amphipods of the Genus Eurystheus

from Japan. Publications of the Seto Marine

Biological Laboratory, 9:31-36, figures 1-2.

Nicholls, G. E.

1939. The Prophliantidae. A Proposed New Family of

Amphipoda, with Description of a New Genus

and Four New Species. Records of the South

Australian Museum, 6:309-334, figures 1-10.

Okada, Y. and S. Mawatari

1958. Distributional Provinces of Marine Bryozoa in the

Indo-Pacific Region. Proceedings of the Eighth

Pacific Science Congress, 3:391-402.

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 :

. . . 1: Lysianassidae . . . Gammaridae. Siboga

Expeditie, Monographie 33e: 237-328, figures

102-146.

1938. Les Amphipodes de l'Expedition du Siboga.

Deuxieme Partie: .. . Ill : .. . 2: .. . Dexaminidae . . . Podoceridae. Siboga Expeditie,

Monographie 33f:329-359, figures 147-161.
274 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ruffo, S.

1956. Nota su Lacuni Anfipodi Raccolti Sulle Coste

dell' India dal Dr. K. Lindberg. Memorie del

Museo Civico di Storia Naturale di Verona,

5:211-216.

Sars, G. O.

1895. Amphipoda. An Account of the Crustacea of

Norway with Short Descriptions and Figures of

All the Species, 1: i-viii + 1-711, plates 1-240,

supplementary plates 1-8.

Say, T.

1818. An Account of the Crustacea of the United States.

Journal of the Academy of Natural Sciences of

Philadelphia, 1:374-401.

Schellenberg, A.

1926. Die Gammariden der Deutschen Siidpolar-Expedition 1901-1903. Deutsche Sudpolar-Expedition, 18:235-414, figures 1-68.

1928. Report on the Amphipoda. In Zoological Research Cambridge Expedition Suez Canal, 1924.

Transactions of the Zoological Society of London, 22:633-692, figures 198-209.

1931. Gammariden und Caprelliden des Magellangebietes, Siidgeorgiens und der Westantarktis.

Further Zoological Results of the Swedish Antarctic Expedition, 1901-1903, 2 (6): 1-290, figures 1-136, plate 1.

1938. Litorale Amphipoden des Tropischen Pazifiks.

Kungl. Svenska Vetenskapsakademiens Handlingar, 16:1-105, figures 1-48.

1953. Erganzungen zur Amphipodenfauna SiidwestAfrikas nebst Bemerkungen viber Brutraumbildung. Mittheilungen aus dem Zoologischen

Museum in Berlin, 29:107-126, figures 1-7.

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, 7.0-

ology, 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.

1931. A New Species of Amphipod Crustacean (Acanthonotozomatidae) from California, and Notes

on Eurystheus tenuicornis. Proceedings of the

United States National Museum, 78: (2861):

1-8, figures 1-4.

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.

1935a. The Amphipods of Puerto Rico and the Virgin

Islands. Scientific Survey of Puerto Rico and

the Virgin Islands, New York Academy of Sciences, 15:229-253, figures 1-5.

1935b. A New Species of Amphipod of the Genus Grandidierella and a New Record for Melita nitida from

Sinaloa, Mexico. Journal of the Washington

Academy of Sciences, 25:65-71, figures 1-2.

1942. Amphipod Crustaceans Collected on the Presidentia 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.

1955. Notes on the Amphipod Crustacean Maeroides

Thompsoni Walker. Journal of the Washington

Academy of Sciences, 45:59.

1956. A New Genus and Two New Species of Amphipods from Dry Tortugas, Florida. Journal of

the Washington Academy of Sciences, 46:61-64,

figures 1-2.

Squires, D. F.

1959. Results of the Puritan-American Museum of Natural History Expedition to Western Mexico 7.

Corals and Coral Reefs in the Gulf of California.

Bulletin of the American Museum of Natural

History, 118:367-432, figures 1-20, plates 28-34.

Stebbing, T. R. R.

1888. Report on the Amphipoda Collected by H.M.S.

Challenger During the Years 1873-76. In Great

Britain, Report on the Scientific Results of the

Voyage of H.M.S. Challenger during the Years

1873-76. Zoology, 29: i-xxiv, 1-1737, plates

1-210 (in 3 volumes).

1897. Amphipoda from the Copenhagen Museum and

Other Sources. Transactions of the Linnean

Society of London, Zoology, series 2, 7:23—45,

plates 6-14.

1899a. Amphipoda from the Copenhagen Museum and

Other Sources Part II. Transactions of the Linnean Society of London, series 7, Zoology, 2:395—

432, plates 30-35.

1899b. Revision of Amphipoda (continued). The

Annals and Magazine of Natural History, series

7,4:205-211.

1904. Gregarious Crustacea from Ceylon. Spolia Zeylanica, 2:1-29, plates 1-6.

1906. Amphipoda I. Gammaridea. Das Tierreich,

21:1-806, figures 1-127.

1910. Crustacea. Part 5. Amphipoda. Sci. Res. Trawling

Exped. H.M.C.S. "Thetis". Memoirs of the Australian Museum 4, 2:565-658, plates 47-60.
NUMBER 34 275

Stephensen, K.

1944. Some Japanese Amphipods. Videnskabelige

Meddelelser fra Dansk Naturhistorisk Forening,

Kebenhavn, 108:25-88, figures 1-33.

1949. The Amphipoda of Tristan da Cunha. Researches of the Norwegian Scientific Expedition

to Tristan de Cunha 1937-1938, 19:1-61, figures 1-23.

Tattersall, W. M.

1922. Amphipoda and Isopoda. The Percy Sladen

Trust Expedition to the Abrolhos Islands (Indian

Ocean). Journal of the Linnean Society of

London, 35:1-19, plates 1-3.

Thomson, G. M.

1881. Recent Additions to and Notes on New Zealand

Crustacea. Transactions and Proceedings of the

New Zealand Institute, 13:204-221, plate 8.

Thorsteinson, E. D.

1941. New or Noteworthy Amphipods from the North

Pacific Coast. University of Washington Publications In Oceanography, 4:50-96, plates 1-8.

Walker, A. O.

1898. Crustacea Collected by W. A. Herdman, F.R.S.,

in Puget Sound, Pacific Coast of North America,

September, 1897. Transactions of the Liverpool

Biological Society, 12:268-287, plates 15, 16.

1904. Report on the Amphipoda Collected by Professor

Herdman, at Ceylon, in 1902. Supplementary

Report to the Government of Ceylon on the

Pearl Oyster Fisheries of the Gulf of Manaar

1904, 17:229-300, plates 1-8.

1905. Marine Crustaceans, 16: Amphipoda. The

Fauna and Geography of the Maldive and Laccadive Archipelagoes, 2 (1): 923-932, plate 88.

1907. Crustacea, III: Amphipoda. National Antarctic

Expedition, British Museum (Natural History),

3:1-39, plates 1-13.

1909. Amphipoda Gammaridea from the Indian Ocean,

British East Africa, and the Red Sea. Transactions of the Linnean Society of London, Series

2, Zoology, 12: 323-344, plates 42, 43.

Zimmerman, E. C.

1948. Insects of Hawaii, vol. 1. Introduction. University of Hawaii Press: i-xx, 1-206, figures 1-52.
Appendix 1

(Groups of Gammaridea not expected from the Hawaiian Islands sublittoral)

COLD-WATER GROUPS, EITHER COLD-TEMPERATE OR DEEPSEA

Astyridae, most Calliopiidae, Cressidae, Dogielinotidae,

most Eusiridae, Haustoriidae, Hyperiopsidae, Ischyroceridae (but 5 species occur in Hawaii), Lafystiidae,

Laphystiopsidae, Lepechinellidae, Melphidippidae,

most Oedicerotidae, Pagetinidae, Paramphithoidae,

Pardaliscidae, Pleustidae (2 species in Hawaii), Sebidae (1 species in Hawaii), Stegocephalidae, most

Stenothoidae except Stenothoe, Stilipedidae, most

Synopiidae except Synopia and Tiron, Thaumatelsonidae, Vitjazianidae.

ABSENCE OF KELPS FOR LIONIVORES

Eophliantidae, Phliantidae (1 species of each family in Hawaii).

ENDEMIC ELSEWHERE

Bateidae (America).

BRACKISH WATER

Grandidierella.

POORLY KNOWN OR RARE GROUPS

Kuriidae, Ochlesidae (1 species in Hawaii), Prophliantidae.

276
Appendix 2

List of samples and species. Samples represent aliquots reduced

to low denominators.

JLB Hawaii 2, open sea off W end of Pearl Harbor,

4-5 m, sand bottom, wash of giant encrustation mass

from hard sand reef, 29 January 1967, King Yacht

Pele. Amphilochus kailua 2, Amphilochus menehune 3,

Ampithoe ramondi 1, Chevalia aviculae 5, Cymadusa

fdosa 3, Elasmopus piikoi 32, Eriopisa hamakua 1,

Gammaropsis alamoana 6, Gammaropsis atlantica 14,

Konatopus paao 13, Lembos macromafius 2, Leucothoe hyhelia 1, Leucothoides Ppottsi 4, Liljeborgia

heeia 7, Lysianassa ewa 1, Maera kaiulani 12, Parajassa angularis 16, Photis aina 34, Photis kapapa 13,

PPodocerus talegus lawai 2, Ventojassa ventosa 69.

JLB Hawaii 3, off Ewa beach, 18 m, wash of Pocillopora, other corals, bryozoans and encrustations,

29 January 1967, King Yacht Pele. Amphilochus

menehune 2, Ampithoe species 1, Aoroides columbiae

31, Chevalia aviculae 17, Elasmopus molokai 15, Elasmopus piikoi 1, Ericthonius brasiliensis 6, Gammaropsis alamoana 1, Gammaropsis atlantica 23, Ischyrocerus kapu 1, Konatopus paao 5, Laetmatophilus hala

2, Lembos waipio 10, Leucothoides Ppottsi 3, Liljeborgia heeia 10, Lysianassa ewa 7, Melita appendiculata 1,

Nuuanu amikai 1, Ochlesis alii 1, Parajassa angularis

1, Photis aina 88, Photis kapapa 38, Stenothoe species

A 1, Ventojassa ventosa 7.

JLB Hawaii 5, off Ewa Beach, Oahu, 30 m, wash of

Pocillopora heads from long coral reef, 29 January

1967, King Yacht Pele. Amphilochus likelike 2, Amphilochus menehune 8, Ampithoe waialua 36, Aoroides

columbiae 5, Chevalia aviculae 8, Colomastix kapiolani 1, Elasmopus molokai 25, Elasmopus pocillimanus

2, Eusiroides diplonyx 2, Gammaropsis afra 15, Gammaropsis alamoana 3, Gammaropsis altantica 6,

Gammaropsis pokipoki 4, hybrid forms of above 10,

Ischyrocerus species C 1, Laetmatophilus hala 1, Lelehua waimea 10, Lembos leapakahi 1, Lembos waipio 1,

Leucothoe hyhelia 2, Liljeborgia heeia 6, Maera quadrimana 3, Maera species A (cf. hamigera) 1, Melita

appendiculata 1, Parajassa angularis 1, Photis aina 11,

Photis kapapa 4, Podocerus talegus lawai 1, Ventojassa

ventosa 3.

JLB Hawaii 6, off Barbers Point, Oahu, about 30 m,

wash of numerous coralline footballs, 29 January 1967,

King Yacht Pele. Amphilochus menehune 8, Ampithoe

waialua 8, Anamixis stebbingi 1, Aoroides columbiae

2, Ceradocus hawaiensis 9, Chevalia aviculae 10, Colomastix kapiolani 1, Colomastix species 1, Elasmopus

molokai 22, Elasmopus pocillimanus 1, Ericthonius

brasiliensis 3, Eriopisella sechellensis upolu 1, JEUMroides diplonyx 2, Gammaropsis atlantica 10, Gammaropsis pokipoki 3, Ischyrocerus kapu 1, Kanaloa manoa

15, Konatopus paao 11, Laetmatophilus hala 1, Limbos waipio 8, Leucothoe tridens 9, Leucothoides Ppottsi

1, Liljeborgia heeia 9, Maera Ppacifica 2, Melita appendiculata 3, Mokuoloe ninole 1, Paragrubia vorax

1, Photis aina 40, Photis kapapa 6.

JLB Hawaii 8, off W end of Pearl Harbor, 25-30 m,

several giant Pocillopora heads and large masses of

short-tufted red algae, calcareous encrustations, 29

January 1967, King Yacht Pe/e. Amphilochus menehune 2, Ampithoe waialua 3, Chevalia aviculae 150 + ,

Elasmopus molokai 6, Gammaropsis alamoana 26,

Gammaropsis atlantica 1, Konatopus paao 1, Lembos

macromanus 1, Leucothoe hyhelia 2, Leucothoides

Ppottsi 1, Liljeborgia heeia 2, Maera insignis 1, Paragrubia vorax 8, Parajassa angularis 7, Photis aina

180+ , Photis kapapa 7, Ventojassa ventosa 14.

JLB Hawaii 10, Kaneohe Bay, Oahu, 2 m, heads

of Pocillopora meandrina, recovered by divers Kruschwitz and Bowers, Sampan Pass, 23 February 1967.

Amphilochus menehune 3, Ampithoe ramondi 2,

Anamixis stebbingi 2, Colomastix kapiolani 2, Elasmopus molokai 6, Gammaropsis alamoana 13, Gammaropsis atlantica 20, Gammaropsis haleiwa 1, Gammaropsis pali 11, Gammaropsis pokipoki 6, Gitanopsis

pele 50, Ischyrocerus kapu 1, Ischyrocerus oahu 4,

Kanaloa manoa 1, Konatopus paao 4, Leucothoe

hyhelia 2, Leucothoe tridens 1, Leucothoides Ppottsi 6,

Lembos intermedius 5, Lembos leapakahi 3, Liljeborgia

heeia 1, Photis kapapa 9, Stenothoe species B 3.

JLB Hawaii 11, Kaneohe Bay, Coconut Island Inlet,

alga Dictyospora and coral head, 2 m, 23 February

1967, recovered by diver Bowers. Amphilochus likelike

4, Corophium Pbaconi 4, Elasmopus molokai 1.

277
278

Ericthonius brasiliensis 18, Lembos leapakahi 7, Lembos macromanus 28, Liljeborgia heeia 4, Leucothoe

hyhelia 1.

JLB Hawaii 12, Kaneohe Bay, Oahu, inside reef,

3-4 m, dredge of small corals, fleshy algae, corallines,

aboard U. Hawaii R/V Salpa, Jack M. Levitz, 4

March 1967. Ampithoe kaneohe 2, Ampithoe ramondi

10, Ampithoe waialua 2, Anamixis stebbingi 1,

Aoroides nahili 2, Cymadusa filosa 1, Elasmopus molokai 2, Elasmopus piikoi 12, Ericthonius brasiliensis 5.

Eriopisa laakona 1, Eriopisella sechellensis upolu 7,

Gammaropsis alamoana 1, Gammaropsis atlantica 4,

?Gammaropsis pokipoki 1, /fya/c honoluluensis 2,

Jassa lilipuna 1, Konatopus paao 2, Lembos intermedius 4, Lembos macromanus 1, Lembos pualani 17,

Leucothoe tridens 1, Lysianassa ewa 2 Maera serrata

5, Maera insignis 23, Paragrubia vorax 3, Photis kapapa 4, Podocerus talegus lawai 3, leucothoid, possible

new genus, gnathopod 2 like gnathopod 1 of 5«fea (1).

JLB Hawaii 13, same as JLB Hawaii 12, 8 April

1967. Aloiloi nenue 1, Amphilochus menehune 7,

Aoroides nahili 7, Ampithoe kaneohe 6, Anamixis stebbingi 2, Colomastix lunalilo 3, Cymadusa filosa 8, E/OJmopus molokai 3, Elasmopus piikoi 14, Ericthonius

brasiliensis 26, Eusiroides diplonyx 2, Gammaropsis

alamoana 10, Gammaropsis atlantica 18, Gitana liliuokalaniae 1, //ma lilipuna 6, Konatopus paao 43, L^m-

&o$ intermedius 13, Lembos macromanus 6, Lembos

pualani 24, Leucothoe hyhelia 20, Leucothoe tridens 3,

Leucothoides Ppottsi 6, Liljeborgia heeia 1, Lysianassa

ewa 9 (all in sample), Maera insignis 7, Maera serrata

3, Maera pacific a 3, Melita appendiculata 42, Paragrubia vorax 3, Parapleustes derzhavini makiki 9,

Photis kapapa 3( 1 aber. female), Podocerus talegus

lawai 3.

JLB Hawaii 14, 1 mi N of Kualoa Point, Windward

Oahu, near old sugar mill, intertidal, general wash of

algae and dead coral heads, 23 May 1967. Sample not

evenly sorted. Amphilochus menehune 13, Ampithoe

waialua 13, Aoroides nahili 2, Cymadusa filosa 4, ZS/OJmopus molokai 1, Elasmopus piikoi 25, Eusiroides

diplonyx 6, Gammaropsis alamoana 5, Gammaropsis

pali 1, Hyale laie 17, Konatopus paao 1, Lembos

pualani 6, Lembos waipio 2, Leucothoe hyhelia 1, L^ucothiodes Ppottsi 1, Maera insignis 9+ + , Maera pacifica 6, Maera quadrimana 1, Maera serrata 3,

Paragrubia vorax 3, Pontogeneia pacifica 2.

JLB Hawaii 15, same locality as JLB Hawaii 14,

wash of coral block with globular ascidians. Not evenly

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sorted. Amphilochus menehune 1, Ampithoe kaneohe

2. Ampithoe ramondi 4, Ampithoe waialua 7, ^4namtxif stebbingi 1, Chevalia aviculae 3, Cymadusa filosa

5, Hyale laie 14, Ischyrocerus oahu 1, Konatopus paao

I, Lembos intermedius 9, Lembos leapakahi 7, Lembos

macromanus 7, Lembos pualani 17, Lembos kamanu 1,

Leucothoe hyhelia 1, Liljeborgia laniloa 2, Maera insignis 18+, Maera serrata 2, Paradexamine {Wailele)

maunaloa 5, Pontogeneia pacifica 17.

JLB Hawaii 17, Kawela Bay, Oahu, intertidal wash

of Sargassum, other algae, rocks and corals, 24 May

1967. Sample dominated by Maera insignis, Hyale

species, Pontogeneia and Lembos, examined primarily

for faunal additions. Amphilochus kailua 4, Amphilochus menehune 3, Ampithoe akuolaka 2, Ampithoe

orientalis 1, Ampithoe ramondi 1, Ampithoe species 1,

Aoroides nahili 16, Atylus (Kamehatylus) nani 1,

Biancolina mauihina 1, Chevalia aviculae 3, Elasmopus pocillimanus 1, Elasmopus hooheno 5, Elasmopus

piikoi 2, Ericthonius brasiliensis 1, Eriopisa laakona 3,

Eusiroides diplonyx 1, Gammaropsis atlantica, hybrid

5, Gammaropsis kaumaka 1, //yak /atV 7, Hyale affinis

5, Ischyrocerus oahu 9, /awa lilipuna 9, Lembos leapakahi 8, Lembos pualani 7, Lysianassa ewa 1, Maera

insignis 5++, Maera pacifica 4, Maera species B 1,

Melita pahuwai 3, Palinnotus alaniphlias 1, Podocerus

talegus lawai 1, Pontogeneia pacifica 2, &6<z ekepuu 2.

JLB Hawaii 18, Kawela Bay, Oahu, intertidal, wash

of rocks and £//i/a lactuca, 24 May 1967. Report only

of unusual species. Ampithoe akuolaka 2, Ampithoe

species 1, i/ya/<? Zat> 3, Melita pahuwai 2, Neomicrodeutopus (?) makena 2.

Devaney 1. Seaward of Moku Manu, windward

Oahu, 33 m, stem of black coral, Antipathes irregularis

Verrill, including bivalve community attached to coral,

13 May 1967, collector D. Devaney. Amphilochus

menehune 1, Elasmopus diplonyx 8, Gammaropsis

afra 19, Leucothoe hyhelia 2, Leucothoe lihue 5, Parapleustes (?) honomu 1, Stenothoe valida, var. 84,

Ventojassa ventosa 1.

Fee 1. Waikiki Beach, Oahu, intertidal, wash of Ulva

lactuca and substrate, 25 April 1967, collector James

Fee. Amphilochus kailua 38, Ampithoe kaneohe 24,

Ampithoe orientalis 6, Ampithoe ?ramondi 6, Biancolina mauihina 1, Cymadusa filosa 3, Cymadusa

hawaiensis 1, Elasmopus ?molokai 1, Ericthonius

brasiliensis 3, //ya/e honoluluensis 51, /awa lilipuna

II, Maera insignis 27, Maera pacifica 2, Palinnotus

alaniphlias 1, Paragrubia vorax 1, Podocerus talegus

lawai 5, Pontogeneia pacifica 32, Stenothoe haleloke 1.
Index

Principal Hawaiian species and genera and their main page references are entered in boldface. Subjects in tables,

figures, and appendices are omitted.

abbotti, Megamphopus, 177

abyssicola, Gitana, 38

Acanthophora sp., 16

acherusicum, Corophium, 12, 17

Acropora, acroporans, see corals

Acuminodeutopus, 27, 92

adaptive radiation, 22

I'arqiiiinanus, Lembos, 72, 82, 85, 87

afer, Eurystheus, 170

affinis, Hyale, 24, 251, 255, 258, 259, 264, 271

afra, Gammaropsis, 24,170,177, 178, 184, 185

aina, Photis, 185, 188, 192, 193

alamoana, Gammaropsis, 27, 170, 174

alaniphlias, Palinnotus, 16, 226

alata, Leucothoe, 209, 211

alga, 16

algicola, Biancolina, 103

alii, Ochlesis, 220, 223

alluaudi, Ampithoe, 1

Aloiloi, 27, 29, 63

Aloiloi nenue, 63, 65

akuolaka, Ampithoe, 23, 44

America, 13, 14, 25, 29

warm-temperate, 12, 13, 14, 15, 16

amikai, Nuuanu, 166

Ampelisca, 22

schellenbergi, 7, 16

Amphideutopus, 25, 27, 70, 72

Amphilochus, 10, 11, 14, 23, 25, 30

kailua, 30, 33

likelike, 33

marionis, 7

menehune, 33, 37

neapolitanus, 33

picadurus, 37

Ampithoe (see also Pleonexes), 1, 14, 23, 50, 57

akuolaka, 23, 44

alluaudi, 1

kaneohe, 23, 44, 50, 57

kulafi, 1, 50, 57

lacertosa, 54

orientalis, 50, 54, 57

poipu, 23, 50, 57

pollex, 23, 44

ramondi, 23, 24, 44, 50,54

simulans, 23, 44, 54

sp., 50, 54, 57

waialua, 23, 53, 57

Anamixis stebbingi, 61

Anatylus, 93

anguipes, Ischyrocerus, 198, 205

angularis, Parajassa, 12, 202,204, 205

anomalus, Microdeutopus, 90

Anonyx, 22

antarctica, Seba, 247

Antipathes irregularis, 17

Aoridae, 27

Aoroides, 14, 23, 25, 27

californica, 68

?columbiae, 23, 27, 68, 70

nahili, 70

Apherusa, 12

appendiculata, Melita, 161

appendiculatus, Gammarus, 161

Aquarium, Honolulu, 17

arctica, Gitanopsis, 38, 39

armata, Seba, 247

armatus, Laetmatophilus, 237

Arugella heterodonta, see Lysianassa

ascidians, 11

asper, Dardanus, 115

atiantica, Gammaropsis, 24, 72, 170, 174, 177 178, 184, 185

atlanticus, Eurystheus, 174

Atylidae, 26

Atylopsis, 12

Atylus, 14, 25, 26, 93

collingi, 93

japonicus, 93

levidensus, 93

nani, 14,26,93, 110

Australia, 12, 16, 26

australis, Biancolina, 103

aviculae, Chevalia, 166

ayeli, Hyale, 24, 255, 260, 271

?baconi, Corophium, 17, 100

bannwarthi, Leucothoella, 11

Bateidae, 20

Bathyporeiapus, 224

Beaudettia palmeri, 26, 93, 94

Bemlos macromanus, 82

Biancolina algicola, 103

australis, 103

mauihina, 16, 103

bishopae, Hyale grandicornis, 24, 235, 257, 260, 271

black-coral, see Antipathes irregularis

boeckii, Neopleustes, 223

279
28

0 INDE

X

Brachyura, see crabs

brasiliense, Platophium, 237

hrasiliensis, Ericthonius, 17

Podocerus, 12, 17,237

brazieri, Colomastix, 100

brevicauda, Nippochelura, 17

brevicornis, Liljeborgia, 216

brevidigitata, Leucothoe, 211

Bryozoa, bryozoans, 3

Byblis, 12

cabindae, Neomicrodeutopus, 90

Caesaris, Eusiroides, 109

California, amphipods, 14, 20

Californian warm-temperature, 2, 14, 16, 19

californica, Aoroides, 68

calitemplado, Gitana, 37

Calliactis, 115

Elasmopus, 115, 128, 131

Caribbean Sea, 11, 16

carinata, Hyale, 268

capensis, Eriopisa, 138, 143

capensis, Eriopisella, 143, 147

Carolobatea, 224

castellata, Colomastix, 96

Cedrophotis, 118

centralis, Paraphoxus, 9

Ceradocus, 11, 14

hawaiensis, 12, 14, 115

"rubromaculatus," 11

Cerapus tubularis, 11, 12

chamissoi, Ischyrocerus, 198

Cheiriphotis, 12

megacheles, 11, 12

Chelura terebrans, 17

Chcvalia aviculae, 166

chevreuxi, Hyale, 259

chilkensis, Eriopisa, 143

China Sea, 19

cognates, 11, 27

collingi, Atylus, 93

Colomastix, 1, 23

brazieri, 100

castellata, 96

fissilingua, 96

kapiolani, 96

lunalilo, 16,96, 100

magnirama, 100

pusilla, 96, 100

sp., 100

subcastellata, 96

color, 2

?columbiae, Aoroides, 23, 27, 68, 70

commensalis, Ischyrocerus, 198

concavus, Lembos, 85

Conus, 3

cool-water, 11, 13, 24, 25

corals, 2, 3,11,13,19

Corophium, 17, 23

acherusicum, 12, 17

Corophium—Continued

?baconi, 17,100

insidiosum, 12, 17, 101

uenoi, 102

cortada, Eriopisella (Netamelita), 147

crabs, 18

crenulata, Stenothoe, 250

cristatum, Cyrtophium, 243

cristatus, Podocerus, 237, 243

Crustacea, crustaceans, 3

Cryptic elements, 16

cubensis, Lysianassa, 220

cumbrensis, Microjassa, 198, 202

current, Humboldt-Peruvian, 18

North Equatorial, 19

Cymadusa, 1, 10,23,24

filosa, 23, 24, 25, 61

hawaiensis, 25, 61

oceanica, 25

Cyproidea ornata, 11

Cyproidcinae, 27

Cyrtophium cristatum, 243

dentatum, 243

Dardanus asper, 115

dentata, Dexiocerella, 243

dentatum, Cyrtophium, 243

derzhavini, Parapleustes, 229

Paraplcustes makiki, 227, 229

Dexaminidae, 26, 93

Dexamininae, 26

Dexaminoides orientalis, 11, 102

Dexamonica, 26

reduncans, 26

Dexiocerella dentata, 243

laevis, 243

dezhnevi, Ischyrocerus, 198

digitata, Gammaropsis, 170,178

digitatus, Eurystheus, 128

diplonyx, Elasmopus, 115, 119

Eusiroides, 109, 110

discreta, Iphimedia, 11

dispersal, 17, 18,23

sweepstakes, 19, 20, 22

distribution, by ships, 6, 12, 16, 17

diversity, 6, 10, 16

dolichommata, Photis, 188

Dolium, 115

dollfusi, Hyale, 264

dubia, Seba, 247

echinoderms, 2, 3

ecuadorensis, Elasmopus hawiensis, 16, 24,119, 121

ekepuu, Seba, 16, 247

Elasmopus, 1, 9, 23, 24, 27, 95

calliactis, 115, 128, 131

?diplonyx, 115, 119

ecuadorensis hawaensis, 16, 24, 119, 121

holgurus, 115, 128

hooheno, 24, 121

insignis, 147
INDE

X 28

1

Elasmopus—Continued

minimus, 24, 125

molokai, 24, 119, 121, 124, 125, 128

pectcnicrus, 24, 119, 121, 125

piikoi, 24, 115, 121, 125,127, 128

pocillimanus, 121, 124, 127, 128, 131, 136

pseudaffinis, 115

rapax,12, 17, 115,125, 128, 131

serrula, 125

spinidactylus, 121, 136

spinimanus, 11

El Nino, 18, 19

elongata, Neomicrodeutopus, 92

Eriopisa, 138, 143

Ericthonius, 17

brasiliensis, 17

Eriopisa, 16,23,27, 138, 166

capensis, 138 (see also Eriopisella)

chilkensis, 143

elongata, 138, 143

garthi, 138, 143

hamakua. 138, 140

laakona, 140, 143

philippensis, 140, 143

seurati, 140, 143

Eriopisella, 23, 27, 138, 143, 166

capensis, 143, 147

nagatai, 143

pusilla, 147

sechellensis 143, 147

upolu, 143

(Netamelita) cortada, 147

erythrophthalmus, Gammaropsis, 181

Eurystheus, 170

afer, 170

atlanticus, 174

digitatus, 178

ventosa, 205

Eusiroides Caesaris, 109

diplonyx, 109, 110

ewa, Lysianassa, 216, 220

Exoediceros, 224

falcata, Jassa, 11, 12, 202

filosa, Cymadusa, 23, 24, 25, 61

fishes, 2, 3,4, 12,20

fissilingua, Colomastix, 96

flindersi, Leucothoe, 211

flotsam, 12, 14,24

fouling masses, 17

founder effect, 17

francanni, Lembos, 81

fresneli, Gammarus, 61

Melita, 161

fulanus, Podocerus, 243

furina, Leucothoe, 11

Galapagos Islands, 18, 19, 24

galateae, Hyale, 264

gallensis, Stenothoe, 12, 17, 249, 250, 251

Gammaridea, tropical species, 3, 4, 7

gammaroides, Pleonexes, 57

Gammaropsis, 1, 9, 14, 23, 27, 29, 63, 65, 102

afra, 24, 170, 177, 178, 184, 185

alamoana, 27, 170, 174

atlantica, 24, 72, 170, 174, 177, 178, 184, 185

digitata, 170, 178

erythrophthalmus, 181

Gardineri, 174

haleiwa, 24, 170, 178, 181, 185

japonica, 184

kaumaka, 24, 181

maculatus, 181

pacifica, 177

pali,65, 170, 181, 183, 184, 185

pokipoki, 24, 170, 181, 185

semidentata, 184

thompsoni, 27, 170, 174

zeylanicus, 174

Gammarus appendiculatus, 161

fresneli, 161

quadrimanus, 152

Gardineri, Gammaropsis, 174

Garosyrrhoe, 224

garthi, Eriopisa, 138, 143

gastropods, see mollusks

Gitana, 14, 38

abyssicola, 38

calitemplado, 37

liliuokalaniae, 37, 38, 39

rostrata, 37

sarsi, 38

Gitanogeiton sarsi, 38, 39, 40

Gitanopsis, 1, 10, 11, 14, 30, 38

arctica, 38, 39

pele, 16, 33, 37, 40

pusilla, 33,40

pusilloides, 40

simplex, 39

tortugae, 16, 40

vilordes, 33, 37,40

gracilis, Parapleustes, 230

grandicornis, Hyale, 255, 257

Hyale bishopae, 24, 255, 257, 260, 271

Grandidierella, 72, 92

Grubia hawaiensis, 61

Guernea, 26

nordenskioldi, 26

Gulf of California, 24, 25

hala, Laetmatophilus, 16, 233

haleiwa, Gammaropsis, 24, 170, 178, 181, 185

haleloke, Stenothoe, 247, 249

Halimedon schneideri, 224

hamakua, Eriopisa, 138, 140

hamigera, Maera, 156

hanapepe, Podocerus, 16, 240

harbors, 17

Harpinia, 22

Haustoriopsis, 26
282 INDEX

hawaiensis, Ccradocus, 12, 14, 115

Cymadusa, 25, 61

Elasmopus ccuadorcnsis, 16, 24, 119,121

Grubia, 61

Photis, 188, 191

heeia, Liljeborgia, 213, 216

heterodonta, Arugella, see Lysianassa

Lysianassa, 220

Hilo Harbor, 17

Hippomedon, 22

holgurus, Elasmopus, 115, 128

holmesi, Lysianassa, 220

Palinnotus, 227

Honolulu Aquarium, 17

honoluluensis, Hyalc, 24, 255, 257, 259, 271

honomu, Parapleustcs ( ?), 26, 233

hooheno. Elasmopus, 24,121

Hoplopheonoides, 40

Humboldt-Peruvian current system, 18

Hyale, 1,9,14,23,24,27, 141

affinis, 24, 251, 255, 258, 259, 264, 271

ayeli, 24, 255, 260,271

bishopae, see grandicornis b.

carinata, 268

chevreuxi, 259

dollfusi, 264

galateae, 264

grandicornis, 255, 257

grandicornis bishopae, 24, 255, 257, 260, 271

honoluluensi, 24, 255, 257, 259, 271

iole, 257, 260

Iaie, 251,255,260, 264, 271

media, 255

novaezealandiae, 255

plumulosus, 260

pusilla, 264

rubra (see also Hyale sp.), 268, 271

schmidti, 271

sp. (cf. rubra), 268

(Lelehua),264,268

waimea, 16, 264

hyhelia, Leucothoe, 205

hystrix, Laetmatophilus, 237

IduneUa, 12

impoverishment, 6, 10, 12, 18, 27

inaequipes, Maera, 11, 12, 155

Maera serrata, 155

inconspicuus, Podocerus, 240, 243

India, 1,14, 20

Indo-Pacific, 1, 11,13, 16, 20, 29

innocens, Ochlesis, 220, 223

innominata, Seba, 247

inquilines, 14

insidiosum, Corophium, 12, 17, 101

insignis, Elasmopus, 147

insignis, Maera, 147

insulae, Tropichelura, 17

in termed ius, Lembos, 76, 77, 78

Introduced elements, 7, 16, 17

iole, Hyale, 257, 260

Iphimedia discreta, 11

irregularis, Antipathes, 17

Isaea, 12

Isaeidae, 27

Ischyroceridae, ischyrocerids, 11, 14, 23, 25

Ischyrocerus anguipes, 198, 205

chamissoi, 198

commensalis, 198

dezhnevi, 198

(?) kapu,26, 195

kraschcninnikovi, 198

oahu, 195, 198

isolation, 12, 18,25,26,27

Japan, Japanese, 9, 17

japonica, Gammaropsis, 184

japonicus, Atylus, 93

Jassa, 17,23

falcata, 11,12,202

(?) lilipuna, 195, 198,202

johanseni, Parapleustes, 233

Johnston Island, 12, 13

kailua, Amphilochus, 30, 33

kaiulani, Maera, 24, 147, 150, 151

kamanu, Lembos, 72, 74, 77, 89, 90

Kamehatylus, 25, 93

Kanaloa, 27, 224

manoa, 224

kaneohc. Ampithoe, 23, 44, 50, 57

kapapa, Photis, 188, 192, 195

kapiolani, Colomastix, 96

kapu, Ischyrocerus (?), 26, 195

kaumaka, Gammaropsis, 24, 181

Konatopus, 14, 25, 27, 70

paao, 70, 72, 185

koreana, Melita, 163

krascheninnikovi, Ischyrocerus, 198

kulafi, Ampithoe, 1, 50, 57

laakona, Eriopisa, 140, 143

lacertosa, Ampithoe, 54

Laetmatophilus armatus, 237

hala, 16, 233

hystrix, 237

leptocheir, 237

tuberculatus, 237

laeve, Platophium, 243

laevidorsum, Melita, 163

laevis, Dexiocerella, 243

Iaie, Hyale, 251, 255, 260, 264, 271

lamellifera, Photis, 188

laniloa, Liljeborgia, 216

lawai, Podocerus talegus, 241, 243

leapakahi, Lembos, 72, 76, 79, 81, 82, 85, 87, 89, 90

(Lelehua), 264, 268

Hyale waimea, 16, 264

Lembopsis tridens, 13

Lembos, 1, 9, 23, 24, 27, 63, 74

?aequimanus, 72, 82, 85, 87

concavus, 85
INDEX 283

Lembos—Continued

francanni, 81

intermedius, 76, 77, 87

kamanu, 72, 74, 77,89, 90

leapakahi, 72, 76, 79, 81, 82, 85, 87, 89, 90

macromanus (also see Bemlos m.), 24, 74, 79, 82, 85, 87

podoceroides, 11

processifer, 76, 87

pualani, 72, 74, 76, 79, 82, 85, 89, 90

sp., 87

waipio, 76, 82, 85, 87

websteri, 79

lenticulosus, Ochlesis, 223

Lepechinella, 26

leptocheir, Laetmatophilus, 237

lessoniae, Pleonexes, 61

Leucothoe, 1, 14, 23

alata, 209, 211

brevidigitata, 211

fiindersi, 211

furina, 11

hyhelia, 205

lihue, 209, 211

minuscula, 211

spinicarpa, 11

tridens, 209, 211

Leucothoella bannwarthi, 11

Leucothoides pacific a, 16

?pottsi, 16, 211

levetzowi, Ochlesis, 223

levidensus, Atylus, 93

lihue, Leucothoe, 209, 211

likelike, Amphilochus, 33

lilipuna, Jassa (?), 195, 198, 202

liliuokalaniae, Ghana, 37, 38, 39

Liljeborgia, 23, 24

brevicornis, 216

heeia, 213, 216

laniloa, 216

proxima, 24, 216

Line Islands, 12, 13

Linguimaera, 150, 152, 155

litotes, Microjassa, 202

lobatus, Podocerus, 243

longicaudata, Photis, 188, 195

longicaudatus, Photis, 188

longimanus, Photis, 188

longleyi, Pontogeneia, 110

lunalilo, Colomastix, 16, 96, 100

Lysianassa cubensis, 220

ewa, 216, 220

heterodonta, 220

holmesi, 220

Lysianassidae, 14, 20

macrocoxa, Microjassa, 202

macromanus, Bemlos, 82

Lembos, 24, 74, 79,82, 85, 87

maculatus, Gammaropsis, 181

Maera (see also Moera), 1, 9, 23, 24, 26, 27

hamigera, 156

inaequipes, 11, 12, 155

serrata, 155

insignis, 147

kaiulani, 24, 147, 150, 151

othonides, 161

pacifica, 24, 150, 151, 154, 155

quadrimana, 150, 152, 154

serrata, 155

sp. A, 156

sp. B, 159

subcarinata, 26

magnirama, Colomastix, 100

makena, Neomicrodeutopus ( ?), 27, 90, 92

makiki, Parapleustes derzhavini, 227, 229

mangarevae, Podocerus, 240

manoa, Kanaloa, 224

marionis, Amphilochus, 7

Marquesas Islands, 13

Marshall Islands, 13

mauihina, Biancolina, 16, 103

maunaloa, Paradexamine, 102, 103

media, Hyale, 255

megacheles, Cheiriphotis, 11, 12

Megamphopus, 177

abbotti, 177

Melita, 23, 138, 147, 166

appendiculata. 161

fresneli, 161

koreana, 163

laevidorsum, 163

nitida, 166

pahuwai, 161, 163

palmata, 161, 166

zeylanica, 163

Melitoides, 166

menehune, Amphilochus, 33, 37

Methalimedon, 224

Microdeutopus, 13, 27, 72, 90, 92

anomalus, 90

Microjassa, 195

cumbrensis, 198, 202

litotes, 202

macrocoxa, 202

Micronesia, 1, 12, 13, 20, 26, 29

minimus, Elasmopus, 24, 125

Minor elements, 16

minuscula, Leucothoe, 211

minuta, Pontogeneia, 110

Moera pectenicrus, 125

pocillimanus, 128

Mokuoloe, 27, 40

ninole, 40

mollusks, 2, 3

molokai, Elasmopus, 24, 119, 121, 124, 125, 128

monoculoides, Stenothoe, 247

nagatai, Eriopisella, 143

nahili, Aoroides, 70
284 INDEX

nani,Atylus, 14,26,93,110

natalensis, Palinnotus, 227

neapolitanus, Amphilochus, 33

nenue, Aloiloi, 63, 65

Neomegamphopus, 25, 27, 70, 72

Neomicrodeutopus, 27, 72

cabindae, 90

elongata, 92

(?) makena, 27,90,92

Neopleustes boeckii, 233

pulchella, 228, 233

rasmyslovi, 233

nestlers, 14, 20

Netamelita, 138, 166

Eriopisella cortada, 147

New Zealand, 22

ninole, Mokuoloe, 40

Nippochelura brevicauda, 17

nitida, Melita, 166

nordenskioldi, Guernea, 26

novaezealandiae, Hyale, 255

Nuuanu, 23, 27, 166

amikai, 166

oahu, Ischyrocerus, 195, 198

oceanica, Cymadusa, 25

ochlesid, 12

Ochlesis, 11

alii, 220, 223

innocens, 220, 223

lenticulosus, 223

levetzowi, 223

oculatus, Parapleustes, 233

Orchomene, 22

oriental is. Amp it hoe, 50, 54, 57

Dexaminoides, 11, 102

ornata, Cyproidea, 11

othonides, Maera, 161

paao, Konatopus, 70, 72, 185

Pacific, Central, 12, 13

pacifica, Gammaropsis, 177

Leucothoides, 16

Maera, 24, 150, 151, 154, 155

Pontogeneia, 14, 110

pahuwai, Melita, 161, 163

pali, Gammaropsis, 65, 170, 181, 183, 184, 185

Palinnotus, 226

alaniphlias, 16,226

holmesi, 227

natalensis, 227

thomsoni, 227

palmata, Melita, 161, 166

palmeri, Beaudettia, 26, 93, 94

Panama, Gulf of, 18

Paradexamine, 26, 102

maunaloa, 102, 103

(Wailele),102

Paragrubia vorax, 61

Parajassa, 11,202

angularis, 12, 202, 204, 205

Paraphoxus, 16

centralis, 9

Parapleustes, 23

derzhavini, 229

makiki, 227, 229

gracilis, 230

(?) honomu, 26, 233

johanseni, 233

oculatus, 233

pugettensis, 230

Parawaldeckia, 22

Parelasmopus suluensis, 11

Parhalimedon, 224

Parhyalella pietschmanni, 271

whrlplryi, 271

pectenicrus, Elasmopus, 24 , 119, 121 , 12 5

Mocra, 125

pele, Gitanopsis, 16, 33, 37, 40

pclerypods, sec mollusks

Peltocoxa, 40

IVrioculopsis, 224

philippensis, Eriopisa, 140, 143

Photidae, 27

Photis, 1,23

aina, 185, 188, 192, 193

dolichommata, 188

hawaiensis, 188, 191

kapapa,188, 192, 195

lamellifera, 188

longicaudata, 188, 195

longicaudatus, 188

longimanus, 188

reinhardi, 188

sp. of Pirlot, 188, 195

picadurus, Amphilochus, 37

pietschmanni, Parhyalella, 271

piikoi, Elasmopus, 24, 115, 121, 125, 127, 128

Platophium brasiliense, 237

laeve, 243

synaptochir, 237

Pleonexes, 1,23,50,57

gammaroides, 55

lessoniae, 61

sp., 57

pleustid, 23

plumulosus, Hyale, 260

pocillimanus, Elasmopus, 121, 124, 127, 128, 131, 136

Moera, 128

pocilloporans, see corals

podoceroides, Lembos, 11

Podocerus, 1, 17,23,237

brasiliensis, 12, 17, 237

cristatus, 237, 243

fulanus, 243

hanapepe, 16, 240

inconspicuus, 240, 243

lobatus, 243

mangarevae, 240

sp., 243
INDE

X 285

Podocerus—Continued

synaptochir, 237

talegus, 243

lawai, 241,243

zeylanica, 240

poipu, Ampithoe, 23, 50, 57

pokipoki, Gammaropsis, 24, 170, 181, 185

pollex, Ampithoe, 23,44

polychaetes, 3

Polynesia, 12, 13,19,26,29

Pontogeneia, 14

longleyi, 110

minuta, 110

pacific a, 14,110

Porifera, see sponges

?pottsi, Lcucothoides, 16, 211

(Prinassus),26

processifer, Lembos, 76, 87

Prophliantinae, 26

Prophlias, 26

proxima, Liljeborgia, 24, 216

pseudaffinis, Elasmopus, 115

pualani, Lembos, 72, 74, 76, 79, 82, 85, 89, 90

pugettensis, Parapleustes, 230

pulchella, Neopleustes, 228, 233

pusilla, Colomastix, 96, 100

Eriopisella, 147

Gitanopsis, 33, 40

Hyale, 264

pusilloides, Gitanopsis, 40

quadrimana, Macra, 150, 152, 154

quadrimanus, Gammarus, 152

rafting, 17, 18

ramondi, Ampithoe, 23, 24, 44, 50, 54

rapax, Elasmopus, 12, 17, 115, 125, 128, 131

rasmyslovi, Neopleustes, 233

reduncans, Dexamonica, 26

reinhardi, Photis, 188

Ronco, 110

sosa, 110

rostrata, Gitana, 37

rubra, Hyale (see also Hyale sp.), 268, 271

"rubromaculatus", Ceradocus, 11

Sargassum sp., 14, 18, 27, 96

sarsi, Gitana, 38

Gitanogeiton, 38, 39, 40

saundersi, Seba, 247

schellenbergi, Ampelisca, 7, 16

schmidti, Hyale, 271

schneideri, Halimedon, 224

scleractinians, see corals

Seba antarctica, 247

armata, 247

dubia, 247

ekepuu, 16, 247

innominata, 247

saundersi, 247

subantarctica, 247

typica, 247

sechellensis, Eriopisella, 143, 147

semidentata, Gammaropsis, 184

serrata, Maera, 155

Maera inaequipes, 155

serrula, Elasmopus, 125

seurati, Eriopisa, 140, 143

simplex, Gitanopsis, 39

simulans, Ampithoe, 23, 44, 54

sosa, Ronco, 110

spinicarpa, Leucothoe, 11

spinidactylus, Elasmopus, 121, 136

spinimanus, Elasmopus, 11

sponges (Porifera), 3

stebbingi, Anamixis, 61

Stenothoe, 1, 14, 17, 23, 24

crenulata, 250

gallensis, 12, 17,249,250,251

haleloke, 247, 249

monoculoides, 247

sp. A, 251

sp. B, 251

Pvalida, 12, 17,250,251

validus, 250

stenothoids, 16

stepping stone, 11, 12, 13

subantarctica, Seba, 247

subcarinata, Maera, 26

subcastellata, Colomastix, 96

suluensis, Parelasmopus, 11

sweepstakes dispersal, 19, 20, 22

synaptochir, Platophium, 237

Podocerus, 237

Synopia, 11

talegus, Podocerus, 243

Podocerus lawai, 241, 243

terebrans, Chelura, 17

thompsoni, Gammaropsis, 27, 170,174

thomsoni, Palinnotus, 227

Tiron, 12

tortugae, Gitanopsis, 16, 40

tridens, Lembopsis, 13

Leucothoe, 209, 211

Tropichelura insulae, 17

tuberculatus, Laetmatophilus, 237

tubicolous species, 14, 20

tubularis, Cerapus, 11, 12

typica, Seba, 247

uenoi, Corophium, 102

upolu, Eriopisa sechellensis, 143

?valida, Stenothoe, 12, 17, 250, 251

validus, Stenothoe, 250

Ventojassa, 11, 205

ventosa, 12, 203, 204, 205

ventosa, Eurystheus, 205

Ventojassa, 12, 203, 204, 205

vilordes, Gitanopsis, 33, 37, 40

vorax, Paragrubia, 61

waialua, Ampithoe, 23, 53, 57
286

Wailele, 102

waimea, Hyale (Lelehua), 16, 264

waipio, Lembos, 76, 82, 85, 87

warm-temperate, 11

websteri, Lembos, 79

Westwoodilla, 224

INDEX

whelpleyi, Parhyalella, 271

wood-borer, 17

Xcnocheira, 13

zeylanica, Melita, 163

Podocerus, 240

zeylanicus, Gammaropsis, 174

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and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings

may be worked out with the editor.

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

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

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

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

of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent lines

indented about three spaces. Within a taxon, use a period-dash (.—) to separate each reference.

Enclose with square brackets any annotation in or at the end of the taxon. For references within

the text, use the author-date system: "(Jones, 1910)" or "Jones (1910)." If the reference is

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

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

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

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

For measurements and weights, use the metric system instead of (or in addition to) the

English system.

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

throughout the printed text. They will be termed Figures and should be numbered consecutively;

however, if a group of figures is treated as a single figure, the individual components should be

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

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

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

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

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

separate from the text and not attached to the artwork.

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

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore

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

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

used alone).

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

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

50 free copies; order forms for quantities above this amount with instructions for payment will be

supplied when page proof is forwarded.