Littoral Gammaridean Amphipoda

from the Gulf of California

and the Galapagos Islands

J. LAURENS· BARNARD

mut.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY · NUMBER 271
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y · NUMBE R 27 1

Littoral Gammaridean Amphipoda

from the Gulf of California

and the Galapagos Islands

J. Lanrens Barnard

SMITHSONIAN INSTITUTION PRESS

City of Washington

1979
ABSTRAC T

Barnard, J. Laurens. Littoral Gammaridean Amphipoda from the Gulf of

California and the Galapagos Islands. Smithsonian Contributions to Zoology,

number 271, 149 pages, 74 figures, 11 tables, 1979.—Less than 40 percent of the

Galapagan littoral Gammaridea are endemic and almost all affinity lies with

the eastern Pacific continental fauna. The strictly littoral Gammaridea of the

Galapagos Islands and the Gulf of California are impoverished of specific diversity

in comparison with the richer shores of California and Australia. Evolutionary

events at high level are few in the entire eastern Padfic Ocean. Only one family,

Bateidae, is endemic to American waters and the majority of species belong to

"invasive" cosmopolitan genera. Newly described genera are Anchialella, Posophotis, Varohios, and Zoedeutopus, and new species are described in the following

genera: Ampithoe, Anchialella, Dulzura, Elasmopus, Gitanopsis, Heterophlias,

Hyale, Lembos, Maera, Microjassa, Najna, Pontogeneia, Posophotis, Varohios,

Zoedeutopus.

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

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavemosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Barnard, Jerry Laurens, 1928-

Littoral gammaridean Amphipoda from the Gulf of California and the Galapagos Islands.

(Smithsonian contributions to zoology ; no. 271)

Bibliography: p.

Supt. of Docs, no.: SI 157:271

1. Amphipoda—California, Gulf of. 2. Amphipoda—Galapagos Islands. 3. Crustacea—California, Gulf of. 4. Crustacea—Galapagos Islands. I. Title. II. Series: Smithsonian Institution.

Smithsonian contributions to zoology ; no. 271.

QL1.S54 no. 271 [QL444.M315] 59l'.O8s [595'.37l'O9l64l] 78-606012
Contents

Page

Introduction 1

Acknowledgments 1

Methods 2

Figure Abbreviations 3

Exploration and Environment 3

The Fauna of Gammaridea 5

Biogeography 8

AMPELISCIDAE 13

Ampelisca lobata Holmes 13

Ampelisca schellenbergi Shoemaker 14

AMPHILOCHIDAE 14

Amphilochns }neapolitœnus Delia Valle 14

Amphilochus picadurus J. L. Barnard 14

Gitanopsis baciroa, new species 14

AMPITHOIDAE 16

Ampithoe guaspare, new species 16

Ampithoe plumulosa Shoemaker 18

Ampithoe plumulosa tepahue, new subspecies 18

Ampithoe pollex Kunkel 18

Ampithoe ramondi Audouin 20

Ampithoe tahue, new species 20

Ampithoe vacoregue, new species 21

Ampithoe kulafi J. L. Barnard 21

ANAMIXIDAE 21

Anamixis linsleyi J. L. Barnard 21

BATEIDAE 21

Batea rectangulata Shoemaker 21

Batea susurrator J. L. Barnard : 21

Batea transversa Shoemaker 23

COLOMASTIGIDAE 24

Colomastix species 24

COROPHIIDAE 24

Acuminodeutopus periculosus J. L. Barnard 24

Cheiriphotis megacheles (Giles) 24

Chevalia aviculae Walker 24

Corophium baconi Shoemaker 24

Ericthonius brasiliensis (Dana) 24

Gammaropsis tonichi (J. L. Barnard) 25

Lembos Bate 25

Key to the Males of the Species of Lembos from the Eastern Pacific

Ocean 25

Lembos macromanus (Shoemaker) 25

Lembos achire, new species 25

iii
IV

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page

Lembos tehuecos, new species 27

Microdeutopus Costa 30

Microdeutopus hancocki Myers 30

Microdeutopus schmitti Shoemaker 30

Photis Krøyer 30

Photis elephantis J. L. Barnard 30

Posophotis, new genus 30

Posophotis seri, new species 31

Rudilemboides J. L. Barnard 34

Rudilemboides stenopropodus J. L. Barnard 34

Varohios, new genus 34

Varohios topianus, new species 35

Zoedeutopus, new genus 37

Zoedeutopus cinaloanus, new species 38

DEXAMINIDAE 38

Polycheria osborni Caiman 38

EUSIRIDAE 38

Pontogeneia Boeck 38

Pontogeneia inermis (Krøyer) 43

Pontogeneia intermedia Gurjanova 43

Pontogeneia opata, new species 43

Pontogeneia rostrata Gurjanova 49

Tethygeneia J. L. Barnard 49

Tethygeneia nasa (J. L. Barnard), new combination 49

Tethygeneia? quinsana (J. L. Barnard), new combination 53

GAMMARIDAE 53

Anchialella, new genus 53

Anchialella vulcanella, new species 54

Ceradocus Costa 54

Ceradocus paucidentatus J. L. Barnard 54

Dulzura J. L. Barnard 54

Dulzura gal, new species 54

Elasmopus Costa 57

Specific Characters in Elasmopus 58

Key to the Males of the Species of Eastern Pacific Elasmopus and the

E. rapax Complex of Hawaii 60

Elasmopus antennatus (Stout) 61

Elasmopus bampo, new species 61

Elasmopus ?ecuadorensis Schellenberg 64

Elasmopus hau>aiensis Schellenberg, new status 67

Elasmopus mayo, new species 67

Elasmopus ocoroni, new species 68

Elasmopus ?rapax Costa from Eastern Pacific 69

Elasmopus serricatus J. L. Barnard, new status 73

Elasmopus temori, new species 75

Elasmopus tiburoni, new species 77

Elasmopus tubar, new species 79

Elasmopus zoanthidea, new species 79

Galapsiellus J. L. Barnard 83
NUMBER 271

Page

Galapsiellus leleuporum (Monod) 83

Maera Leach 83

Maera reishi, new species 83

Maera chinarra, new species 86

Maera simile Stout 88

Melita Leach 88

Melita sulca (Stout) 88

Meximaera J. L. Barnard 88

Meximaera diffidentia J. L. Barnard 88

HYALIDAE 90

A llorchestes Dana 90

Allorchestes angusta Dana 91

Allorchestes bellabella J. L. Barnard 94

Allorchestes carinata Iwasa, new status 96

Hyale Rathke 98

Key to the Males of the Species of Hyale from Mexico, California, and

the Galapagos Islands 98

Hyale darwini, new species 99

Hyale rubra (Thomson) 101

Hyale canalina, new species 102

Hyale yaqui, new species 104

Hyale zuaque, new species 108

Hyale guasave, new species Ill

Hyale frequens (Stout) 114

Hyale anceps (J. L. Barnard) 114

Hyale plumulosa (Stimpson) 114

The Hyale grandicornis Complex 114

Key to the Males of the Species in the Hyale grandicornis Complex

from the Pacific Ocean 115

Hyale species 115

Hyale californica J. L. Barnard, new status 116

Hyale humboldti, new species 116

Najna Derzhavin 118

Najna kitamati, new species 118

Parallorchestes Shoemaker and Parhyale Stebbing 119

Key to the Species of Parhyale and Parallorchestes 120

Parallorchestes ochotensis (Brandt) 120

Parhyale plumicomis (Heller) 121

Parhyale? zibellina Derzhavin 121

Parhyale? iwasai Shoemaker 121

Parhyale hawaiensis (Dana) 122

Parhyale eburnea Krapp-Schickel 122

Parhyale penicillata Shoemaker, new status 123

Parhyale fascigera Stebbing 123

Parhyale aquilina (Costa) 126

Parhyale species 126

Parhyale species of Bulycheva 127

Parhyale inyacka (K. H. Barnard) 127
VI

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page

ISCH YROCERIDAE 127

Microjassa chinipa, new species 127

Microjassa macrocoxa Shoemaker 128

LEUCOTHOIDAE 128

Leucothoe alata J. L. Barnard 128

Leucothoe spinicarpa (Abildgaard) 129

Leucothoides pacifica J. L. Barnard 130

Leucothoides pottsi Shoemaker 130

Leucothoides ?yarrega J. L. Barnard 130

LYSIANASSIDAE 130

Lysianassa dissimiïis (Stout) 130

Lysianassa holmesi (J. L. Barnard) 130

Lysianassa macromera (Shoemaker) 130

PHLIANTIDAE 131

Heterophlias seclusus escabrosa J. L. Barnard 131

Heterophlias galapagoanus, new species 131

PHOXOCEPHALIDAE 133

Metaphoxus frequens J. L. Barnard 133

"Paraphoxus" spinosus Holmes 133

PODOCERIDAE 134

Podocerus fidanus J. L. Barnard 134

Podocerus spp 137

Appendix: Station Data 138

Literature Cited 141

Index 147
Littoral Gammaridean Amphipoda

from the Gulf of California

and the Galapagos Islands

J. Laurens Barnard

Introduction

The intertidal faunas of crustaceous Gammaridea

in the Gulf of California and the Galapagos

Islands share many similar taxa. In neither area is

the intertidal zone richly populated with diverse

taxa though on occasion high densities of the particular species prevail. This study treats the hitherto

poorly known Gammaridea as based on widespread

but sparse sampling.

ACKNOWLEDGMENTS

I am indebted to Dr. John R. Hendrickson and

Dr. Newell Younggren of the University of Arizona

for their many kindnesses during my protracted

stay in their laboratories. I am grateful for the use

of the facilities of the marine station at Puerto

Peñasco operated jointly by the University of

Sonora and the University of Arizona. The University of Arizona and the Smithsonian Institution

provided funds for several field expeditions and

Sigma Xi generously contributed to these expenses

through a subvention to me and Dr. Richard C.

Brusca of the University of Arizona. I am indebted

to Dr. Brusca for many aids and fine fellowship

during our trips to Mexico. Mr. and Mrs. James

Wood and Dr. Hendrickson dived for me with

scuba gear at Bahía Kino. Dr. P. E. Pickens of the

J. Laurens Barnard, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560.

University of Arizona provided many collections for

examination. Many of the drawings were made by

Mrs. Wilma A. Findley; she, Carolyn L. Cox, Biruta

Akersbergs, Debra Homer and Katherine M. Hepburn inked the plates.

At the Smithsonian Institution I must thank Dr.

David L. Pawson, former Chairman of the Department of Invertebrate Zoology, Mrs. Montague

Smith, Mr. Roland H. Brown and Mrs. Dessie M.

Best for their extensive aid to me in the execution

of this project; Dr, I. E. Wallen kindly arranged

travel funds for moving my laboratory to Tucson

for a period of 4 years.

My collections in Panama in 1955 were greatly

assisted by Mr. and Mrs. R. L. Miller, then of

Balboa, Canal Zone, now of Sun City, California.

The late Dr. E. Yale Dawson gave me many fine

materials from El Salvador and the Galapagos

Islands. Over 60 colleagues on the 1964 Galapagos

International Scientific Expedition under the leadership of Dr. Robert I. Bowman, made possible my

9 week exploration of those islands, among them

deserving special mention are Dr. Donald P. Abbott,

Dr. J. Wyatt Durham, Dr. Richard N. Mariscal, the

late Allyn G. Smith and Dr. Victor A. Zullo, and

among them were the previously mentioned Dr.

Hendrickson and Dr. Dawson. My thanks go to the

Angermeyer family of Academy Bay who safely

conducted us by sea through the islands and provided our every need and good fellowship. The

Ecuadorian Navy also provided excellent transportation to Tower Island (Genovesa). The Charles
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Darwin Foundation (Contribution 230), University of California, California Academy of Sciences,

and the National Science Foundation made this

expedition possible with the superb cooperation of

the California Maritime Academy.

Mr. and Mrs. R. F. Dwyer of Corona del Mar,

California, kindly transported Dr. Brusca and me

to several islands in the Gulf of California aboard

their research vessel Sea Quest II.

Dr. A. A. Myers of University College, Cork,

Ireland, kindly offered valuable suggestions on this

manuscript. We differ, however, in the interpretation of certain relationships in the new corophiid

genera.

METHODS

Amphipoda were collected largely in "formalinwash" samples, for which intertidal flora and substrates are placed in a container of seawater laced

with formaldehyde. After death, the organisms are

strained from the residue through fine mesh screens,

preserved in a 5 percent solution of formaldehyde

and seawater, returned to the laboratory, sorted to

taxa in clean water and represerved for microscopic

study in 70 percent ethanol. Where seaside laboratories were available certain samples were sorted

after collection as rapidly as possible to facilitate

initial division into taxa by color pattern (J. L.

Barnard, 1970).

Aliquots of each sample were analyzed, and the

taxa and their individuals identified and counted.

The samples were then arranged into 6 groups: (1)

Puerto Peñasco, (2) Bahía Kino, (3) Topolobampo,

(4) Inner Baja California from La Paz to Bahía Concepcíon, (5) Cabo San Lucas region, and (6) the

Galapagos Islands. The average number of specimens per sample in each group was multiplied by

10 to provide whole numbers so as to compose

tables showing general rankings of abundance of

the species in the several areas.

Counts of juveniles in samples with more than

two species of a genus are divided among the

species in proportion to the ratio of adults. This

is clearly a poor practice as juveniles of different

species may be in quite different proportions than

their adults, but the identification of juveniles to

the species level in such difficult genera as Amphithoe, Hyale, and Photis could not be accomplished

in any event given the poor knowledge about juvenile morphology.

After aliquots had been sorted into taxa the

remainder of each sample was examined for any

species not found in the aliquot. In most cases these

were rare species at the locality and are reported

as the final members of the minor species for each

sample. In the grouping of samples for the comparative tables many of these species also are appended

as minor species; in other cases a minor species

in one sample may dominate another sample from

the same area and therefore is advanced to a higher

rank.

Unnamed species include a variety of taxa too

poorly represented in the collections to be described

herein and include two new species of Melita, new

species of Batea, Leucothoides, acanthonotozomatid,

Elasmopus, Hyale, Synchelidium from the Gulf of

California, and new species of Ericthonìus, Photis,

Lembos, Hyale, Maera, Melita, liljeborgiid, oedicerotid, from the Galapagos Islands and Cocos Island.

The few stenothoids and cyproidein found in the

collections are reserved for future studies when a

wider variety of tropical material facilitates their

identification. Many unanalyzed Galapagan and

Panamanian samples remain for subsequent study

in Smithsonian collections. Islands in the Galapagos are cited by their familiar names, rather than

strictly adhering to the Spanish appellations.

Types are registered either in the National Museum of Natural History, Smithsonian Institution

(USNM collection) or in the Allan Hancock Foundation, University of Southern California, Los Angeles (AHF). All remaining material collected and

noted in the station data was apportioned among

the Smithsonian Institution, the Allan Hancock

Foundation, and the University of Arizona. All new

names commemorate American Indians except those

specified.

Voucher material refers to specimens used, in

addition to the holotype, for illustration and verification of the specific identification. These materials are labeled and deposited under the taxonomic name in the collections of the Smithsonian

Institution or in the museum from which they

were borrowed. Other materials, cited elsewhere,

are not necessarily segregated into specific vials,

but may be returned, along with their compatriots

of other species, into the original sample jars, which

are in the museums as unsorted samples and labeled

according to collector and locality.

Diagnoses and descriptions are not internally
NUMBER 271

parallel in this study; rather they follow conventions in the literature according to each genus.

The term "Mark" (M.) followed by a number

0-100 refers to a point on an appendage, article,

or ramus, the distance from which point to the base

of the structure is expressed as the percentage of

the total length of that structure. For example, a

spine at Mark 70 (or M. 70) on the outer ramus of

uropod 3 lies 0.70 (or 70 percent) of the distance

from base to apex of the stated ramus.

FIGURE ABBREVIATIONS

Capital letters indicate structures; lower case

letters on the left side of capital letters denote specimens cited in the legends and voucher material of

the text; lower case letters on the right side of

capital letters (or within a drawing, as is often the

case with "e" and "s") indicate characteristics of

the structures. Other lower case letters not in the

list below are defined in the legends. Where space

does not allow a horizontal alinement of the label,

the resulting vertical arrangement has the same

order of elements top to bottom as found in the

horizontal from left to right.

A

B

C

D

E

F

G

H

I

J

K

L

M

N

O

P

Q

R

S

T

U

V

antenna

prebuccal from lateral

head

dactyl of pereopod

coxa

accessory flagellum

gnathopod

peduncle

identified in legend

lacinia mobilis

spine, seta, or denticle

lower lip

mandible

molar

sternum

pereopod

cuticle

uropod

maxilliped

telson

upper lip

palp

W

X

Y

Z

a

d

e

i

I

m

0

P

s

V

pleon (pleonites 1-3 bearing epimera; pleonites

4-6, also referred to as

urosomites 1-3, comprising urosome; often

shown with attached

structures: adjoining

pereonites, telson, and

uropods 1-3)

maxilla

pleopod

calceolus

outer (ramus or plate)

dorsal

broken

inner (ramus or plate)

lateral

medial

opposite or right

apex

setae removed

ven tral

EXPLORATION AND ENVIRONMENT

Impetus for this study came from the Symposium

on Biogeography in the Gulf of California (Garth

and Savage, editors, 1960) and the low degree of

knowledge about Amphipoda in the east Pacific

Ocean outside of California and to a degree the

Oregonian Province. However small in body size,

Amphipoda are generally a diverse and, in the aggregate, a conspicuous element of intertidal regions,

though they usually are far more dominant on level

mud bottoms in relation to other groups of organisms. Prior to 1960 the known Amphipoda of the

tropical eastern Pacific Ocean totaled fewer than

35 species and most of these had been described

by Shoemaker (1925, 1942) from sublittoral trawls

made by the Albatross and Houston in the Gulf of

California. Actually, most of these species were

collected in Bahía Magdalena, in Baja California,

just outside the Gulf of California. Literature on

amphipods in the waters of the Gulf of California

and Galapagos Islands was contained in these few

papers (Shoemaker, 1925, 1926, 1942, Schellenberg,

1936, J. L. Barnard, 1954b, 1960, 1969a).

Collections of Amphipoda from both areas deposited with the Allan Hancock Foundation, University of Southern California, were examined; these

are mainly composed of occasional specimens or

lots from scattered dredge hauls; they do not comprise a comprehensive or consistent series of material. The first extensive Galapagan collection was

provided by the late Dr. E. Y. Dawson, who, during

his exploration of Galapagan algae in 1962 collected 40 special samples of phycophilic amphipods.

I supplemented these with my collections made

during the Galapagos International Scientific Expedition of 1964 (Figure 1).

Massive collecting in the Gulf of California (Figure 1) commenced in 1962 (J. L. Barnard, 1969a)

when opportunity was afforded through the

Beaudette Foundation to make several journeys to

Bahía de Los Angeles. More widespread collecting

was accomplished between 1970 and 1974 when

intensive study of the intertidal was undertaken

near the marine laboratory at Puerto Peñasco operated by the University of Arizona and the University of Sonora. This was capped by an expedition

to Cabo San Lucas, Bahía Concepcíon, and other

places in Baja California.

Several colleagues at the University of Arizona

also provided materials they collected during sojourns to the Gulf of California and these are

reported upon where they yield new information.

Despite a large amount of field effort, the results

of this study barely scratch the surface of ignorance

about amphipods in these two regions. The study
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

San Francisco

^Partida

V-Esp¡r¡tu Santo

WESTERN MEXICO

FIGURE 1.—Gulf of California and the Galapagos Islands.
NUMBER 271

had to be confined to intertidal or very shallow

water zones because of the lack of any other collecting means. This biotic zone in both regions is

very improverished either of suitable substrates or

of amphipod taxa.

The intertidal zone of the Galapagos Islands, in

the two places intensively studied, is very barren

of the ordinary substrates in which amphipods are

usually found in midlatitudes (Hedgpeth, 1969).

Apparently most of the intertidal algae are consumed by marine iguanas except in the most fierce

of the breaking seas. Multitudes of Sally Lightfoot

crabs appear to scrape away most of the microscopic

algal mats remaining on the hot barren rocks. Intertidal amplitudes are moderate in any event. E. Y.

Dawson braved the treacherous breaking seas in his

search for algae and acquired the better set of phytic

samples in the Galapagos. My colleagues and I,

however, were able to pry apart slabs of intertidal

strata to find rich encrustations secreted in crevices

out of reach of reptiles and crabs. Two scuba divers

were able to acquire sublittoral substrates in quiet

waters but for the most part these yielded few small

crustaceans.

Tidal amplitudes in the Gulf of California range

from low to high. At Puerto Peñasco, the main

focus of this study, the amplitude on occasion

reaches 5 or more meters, revealing during certain

times of the year a wide belt of intertidal flora,

which, at other times, is depleted by dieoff. Individuals of Amphipoda here are very numerous but

diversity of taxa is low compared to such numbers

as are found at Cayucos, California (J. L. Barnard,

1969b: 10). Surface sea temperatures commonly range

between 10° and 32° C (Brusca, 1974:7) so that

amphipods living in such shallow waters all year

must have wide thermal tolerance. Others may

oscillate from deep to shallow waters to remain in

suitable thermal regimes.

Somewhat richer samples were found at Topolobampo, but these came from an area of low tidal

amplitude in sublittoral depths constantly underwater. The rocks here were covered with a recently

settled spatfall of tunicates.

Samples from Cabo San Lucas, the tip of Baja

California exposed to upwelling and open Pacific

swells, were not as rich as predicted, but collecting

was hampered by precipitous cliffs, sharply sloping

reefs, moderately heavy seas, and severe restrictions

against trespass.

In the area between La Paz and Concepción our

expedition, arriving in November and December,

met a heavy dieoff of intertidal algae along a shore

of low tidal amplitude. Many of these intertidal

samples were destitute of amphipods yet one might

predict that amphipods would thrive as scavengers

during such dieoffs. The richest locality in that

inner apex of Baja California appeared to be Bahía

San Evaristo where the intertidal region not only

supported a rich flora but a heavy concentration of

influent mud among the rocks.

THE FAUNA OF GAMMARIDEA

TABLES 1-10

Despite the vagaries confronted in an attempt to

compare intertidal localities, a certain resemblance

occurs in the fauna of amphipods from place to

place; but where differences occur, partial explanation can be suggested.

Intensive sampling during two seasons, October

(peak summer) and February (cool winter), in

Puerto Peñasco (Table 1) showed relatively similar

dominance ranking, Ericthonius brasiliensis being

an order of magnitude more abundant than the

next dominants, Hyale yaqui, Ampithoe plumulosa,

Lembos macromanus, and Posophotis seri. In Table

1 all of the major species except Zoedeutopus

cinaloanus and the species of Elasmopus were well

represented in both seasons, Zoedeutopus appearing

TABLE 1.—Rank in abundance of intertidal Gammaridea

from Puerto Peñasco, Mexico (based on al¡quots from 21

samples, total specimens averaged per sample, multiplied by

10; asterisks denote domicolous species)

*Ericthonius

brasiliensis

Hyale yaqui

*Ampithoe plumulosa

*Lembos macromanus

*Posophotis seri

Podocerus fulanus

*Gammaropsis tonichi

Tethygeneia nasa

Elasmopus rapax

1851

326

235

81

76

72

39

34

29

• Zoedeutopus cinaloanus

Elasmopus tìburoni

Melita sulca

*Photis elephantis

*A mpithoe pollex

Heterophlias seclusus

Podocerus brasiliensis

*Photis sp.

Leucothoe alata

26

26

19

16

15

13

11

7

5

Minor species: Elasmopus bampo, *Rudilemboides stenopropodus, *Corophium baconi, Gitanopsis baciroa, Podocerus brasiliensis, *Acuminodexitopus periculosus, Elasmopus serricatus, Paraphoxus spinosus, Synchelidium sp.,

Lysianassa dissimilis, *Lembos tehuecos, Leucothoides

pacifica.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

only in the February samples. The minor species

fluctuated greatly from season to season. Many of

these, except perhaps the domicolous forms, may

actually fluctuate hourly as tides change.

The strong dominance of Ericthonius in Puerto

Peñasco, almost at the head of the Gulf of California, where maximum thermal ranges occur in the

gulf, suggests a permanent stressful situation.

Ericthonius brasiliensis (or a series of close siblings),

though widely distributed throughout the world

seas, is best known to dominate shallow water habitats in or near harbors where human influences

have altered the natural environment. To a slight

degree the species may show a pattern of distribution similar to that of the polychaete, Capitella

capitata, which generally abhors the presence of

other organisms and thus signifies by its presence

a great tolerance to physical variables but virtually

no tolerance to the presence of other Metazoa. Several genera of amphipods, Corophium, Podocerus,

and Jassa also become dominant in weakly polluted

or artificial environments.

The dominance of Ericthonius over Hyale by an

order of magnitude in frequency is a precise reversal of the usual situation found in California

(J. L. Barnard, 1969b: 19), the only other place

where intertidal amphipods have been counted

and ranked extensively. The dominance of Hyale

also occurs in the Galapagos Islands but not elsewhere in the Gulf of California samples analyzed

herein. Specific samples often have a dominance of

Hyale but the aggregate from a locality does not.

Hyale dominated in about a third of the samples

from New Zealand intertidal Q. L. Barnard,

1972b:201-204), but only once by an order of magnitude. Because Hyale is associated with algae in

greatest degree, any sample strictly limited to preferential algae would be dominated by that genus.

At Puerto Peñasco almost half of the top 18

dominant species are domicolous or belong to families in which domicoly is thought to be universal.

This includes the premier dominant Ericthonius.

Domicolous amphipods usually spin a tube or loose

nest of weblike material emitted from apical meati

on pereopods 1 and 2. The nest is usually attached

to a substrate, or inside the hole or tube of another

animal, though occasional domicolous amphipods,

such as Cerapus, may drag the tube about in the

fashion of a hermit crab. Inquilinous genera at

Peñasco include only Leucothoe and Leucothoides

as far as known. Inquilinous species usually live in

close association with sessile invertebrates, often

inside sponges or tunicates. The remaining species

are loosely classified as nestlers, those simply hiding

in anastomoses but often swimming freely from

place to place. However, this category is a simplification because such genera as Podocerus may be

highly sedentary; they are often found crawling

slowly about on hydroids in the fashion of caprellids. Species of Hyale, Elasmopus, Pontogeneia, and

Melita are presumed to be errant nestlers, actively

moving about amongst the flora and encrustations,

but at low tide finding a moist hiding place to

withstand desiccation.

Rarely in my studies of intertidal amphipods in

California, Hawaii, New Zealand, and Australia

have more than one species of Hyale been collected

in a small sample; but the species of Elasmopus appear to occupy small territories and perhaps are

separated from each other by much less obvious

niche criteria than volume of space. One species in

California was found by Barnard (1969b: 117) to be

associated with sponges in contrast to algae. The seasonal fluctuation of Elasmopus at Peñasco suggests

that in the intertidal the species might replace each

other from sublittoral sources.

As in California, the species of Ampithoe and

Photis are largely composed of juveniles. Adults

generally form much less than 5 percent of the total

individuals found in a sample, whereas significantly

higher percentages of the other species occur in

adult morphology adequate for identification. To

further complicate the taxonomic procedure, adult

males of Ampithoe and Photis are usually extremely

rare; identification of the several species is most

easily accomplished by examining males.

The fauna of Peñasco (Table 1) may be compared

with the richest locality in California, Cayucos

(Table 7). Peñasco has 30 species, Cayucos 52. Ten

of the Peñasco species do not occur in California

as far as known and more than 20 of the Cayucos

species apparently do not occur, at least commonly,

in the intertidal of the Gulf of California, though

several are known in sublittoral depths. Several

species occur at Cayucos that are predominantly

tropical and will be expected to occur very abundantly in sublittoral zones of the Gulf of California,

for example Chevalia aviculae. But such species

have not, as yet, been found intertidially in the Gulf

of California.
NUMBER 271

One is therefore left with the impression that the

major factors constraining the composition and diversity of the Peñasco fauna are simply the wide

ranging sea temperatures, and the aridity and summer heat from the sun impinging on the intertidal

zone during low tides. As a result, the dominance

of the main nestling genus, Hyale, is suppressed

slightly, whereas the tube dweller Ericthonius brasiliensis, able to retain moisture within its tube during

dry periods, and highly tolerant of wide ranging

sea temperatures, and released to some extent from

pressures of interordinal diversity, comes to prominence in the Peñasco fauna.

The high consistency of occurrence of Heterophlias seclusus is also of interest. This armoured,

dinosaurian-like amphipod, indeed all members of

its family Philantidae, are rarely collected by marine

explorers. Species of the family, but not specimens,

are numerous in museums. J. L. Barnard (1969b: 16)

suggested that Heterophlias lives mainly in or on

sands between rocks in the intertidal, and that the

rod-like translucent but red or ochre stained posterior gut contents might be coagulates from a lign¡n diet during scavenging of dead algal rhizomes

or other parts of woody algae.

The isopod-like body shape of Heterophlias suggests that it may be a functional replacement in the

impoverished intertidal of Puerto Peñasco for some

other crustacean group unable to survive the wide

yearly extremes of climate.

In any event the taxonomist is likely to see more

specimens of this family in one sample from Puerto

Peñasco than during a lifetime of other exploration.

The fauna in the five samples from Bahía Kino,

330 km south of Peñasco, though sparse and poorly

diverse (perhaps from low sampling density), has a

distinctive composition, the only place in this study

where Parhyale is prominent and where a species

of Elasmopus is the premier dominant (Table 2).

At Topolobampo, near the mouth of the gulf,

three underwater samples also show an occurrence

of Parhyale and Elasmopus bampo and species of

Podocerus, with Lembos macromanus being the

premier dominant (Table 3). Owing to the high

densities of the tunicate, Polycheria osborni, a

species usually found buried in colonies of Amaroucium sp., was also prominent. The samples at

Kino and Topolobampo may actually be typical of

sublittoral rocks in the Peñasco region, but that

zone has not been investigated.

TABLE 2.—Rank in abundance of intertidal Gammaridea

from Bahía Kino, Gulf of California (based on aliquots

from 5 samples, total specimens averaged per sample, multiplied by 10; asterisks denote dom¡colous species)

Elasmopus bampo

Parhyale penicillata

Hyale yaqui

Podocerus fulanus

Tethygeneia nasa

*Ericthonius brasiliensis

752

400

380

368

166

158

*Amphithoe pollex

Elasmopus tiburoni

amphilochids

Hyale californica

*Ampithoe plumulosa

Leucothoe alata

144

140

66

38

16

14

Minor species: *Corophium baconi, Lysianassa dissimilis,

*Lembos macromanus, * Gammaropsis sp. 2, lysianassid,

*Posophotis seri, Heterophlias seclusus.

TABLE 3.—Rank in abundance of subintertidal Gammaridea

from Topolobambo, Mexico (based on aliquots from 3 samples, total specimens averaged per sample, multiplied by

10; asterisks denote domicolous species)

*Lembos macromanus

Elasmopus bampo

Podocerus fulanus

Polycheria osborni

Podocerus brasiliensis

* Microjassa macrocoxa

Parhyale penici¡lata

Gitanopsis baciroa

*Gammaropsis tonichi

509

450

256

243

190

176

150

133

127

Tethygeneia nasa

* Corophium baconi

* Ericthonius brasiliensis

*Ampelisca lobata

Lysianassa dissimilis

*Ampithoe plumulosa

Hyale yaqui

*Lembos tehuecos

Maera simile

90

87

77

33

33

23

23

20

17

Minor species: stenothoid sp. 1, acanthonotozomatid, Colomastix sp., Leucothoe alata, stenothoid sp. 2, Leucothoe

spinicarpa, Melita sulca, *Photis sp., *Ampelisca schellenbergi.

Of the 16 samples from several localities between

La Paz and Bahía Concepción (about 350 km apart)

14 are grouped together in Table 4. Ten of these

samples contained fewer than 9 species each, reflecting the November red algal dieoff noticed throughout the region. The dieoff was especially intense at

certain subareas of Pichilinque Bay, Isla Espiritu

Santo, Isla Partida, and Bahía Concepción. Two of

these 10 samples lacked any amphipods and are removed from Table 4.

The eight impoverished samples (subtracting the

two without amphipods) were dominated by Ampithoe plumulosa (in 7 out of 8 samples), and

Lembos macromanus (3 out of 8), whereas other

species with numerous aggregate specimens never

occurred in more than two samples out of eight:

Pontogenia opata, Elasmopus rapax, E. bampo, and

Ericthonius brasiliensis.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 4.—Rank in abundance of intertidal Gammaridea

along Inner Baja California, from La Paz to Bahía Concepción (based on aliquots from 14 samples, parentheses =

number of samples in which species is present, total specimens averaged per sample, multiplied by 10; asterisks denote

domicolous species)

TABLE 5.—Rank in abundance of intertidal Gammaridea

from region of Cabo San Lucas, Mexico (based on aliquots

from 10 samples, total specimens averaged per sample, multiplied by 10; asterisks denote domicolous species)

*Ericthonius brasiliensis

(7)

*Zoedeutopus cinaloanus

(5)

* Ampithoe plumulosa

< " )

Hyaleyaqui (6)

* Ampithoe pollex (6)

•'Varohios topianus (5)

Elasmopus..rapax (4)

*Lembos macromanus

( " )

Elasmopus bampo (2)

Podocerus brasiliensis (1)

Pontogeneia opata (3)

421

248

215

194

172

169

159

129

123

97

59

Lysianassa holmesi (9)

•Ampithoe juveniles

Elasmopus tiburoni (1)

Podocerus fulcanus (2)

A mphilochus picadurus

Elasmopus juveniles

•Ampithoe ramondi (1)

Melita sulca (6)

Leucothoe alata (4)

*Rudilemboides

stenopropodus (1)

*Lembos tehuecos (2)

Batea rectangulata (3)

Paraphoxus spinosus (4)

51

50

34

25

24

22

21

19

19

15

14

11

11

Minor species: Maera reishi, Lysianassa macromera, Pontogeneia quinsana, *Microdeutopus schmitti, Batea susurrator,

*Microdeutopus sp., Tethygeneia nasa, Podocerus juveniles,

*Acuminodeutopus sp., Anamixis linsleyi, Leucothoides

pacifica, Podocerus cristatus, Synchelidium sp., Leucothoe

juveniles, *Corophium baconi, Pontogeneia juveniles, Ceradocus paucidentatus, Heterophlias seclusus, Metaphoxus

frequens, Polycheria osborni, Batea transversa, Lysianassa

dissimilis, Leucothoe spinicarpa, ?Gitanopsis neapolitanus,

Leucothoides ?yarrega.

The six other samples were collected at enriched

areas of Bahía Concepción and Isla Espiritu Santo

or at Bahía San Evaristo and Isla San Francisco;

they comprise the bulk of species and specimens

in Table 4. The maximum number of species in an

enriched sample is 20. This aggregation of samples

over 350 km of coast approximately matches the

diversity found at Cayucos, California (Table 7).

The most abundant species, Ericthonius brasiliensis, was found in greatest numbers in only a few

samples (predominantly the otherwise impoverished

samples). The next dominant Zoedeutopus cinaloanus was found in only five samples. However, the

more characteristic dominant, Ampithoe plumulosa

was found consistently abundant or present in 11

of the 14 samples. Lembos macromanus was also

found in 11 of the samples and would also be considered more characteristic than other species with

higher aggregate densities. Five of the enriched

samples are aggregated in Table 8 to show the preElasmopus serricatus

* Ampithoe pollex

Hyale yaqui

Hyale zuaque

Elasmopus antennatus

Elasmopus tubar

Elasmopus juveniles

*Photis elephantis

Elasmopus rapax

278

264

240

179

126

94

88

77

75

*Cheiriphotis megacheles

Hyale guasave

* Ampithoe plumulosa

Maera chinarra

Heterophlias seclusus

Tethygeneia nasa

• Varohios topianus

Hyale juveniles

Lysianassa macromera

29

28

27

19

15

12

7

4

4

Minor species: *Lembos macromanus, Amphilochus picadurus, *Ampelisca schellenbergi, Leucothoe alata, *Rudilemboides stenopropodus,

dominance of Varohios topianus and the sparsity of

Ericthonius brasiliensis.

The dominance of nestlers in the 10 samples

taken from the region of Cabo San Lucas (Table 5)

may reflect the exposure to open seas of the Pacific

Ocean. Only to a slightly lesser extent is this reflected in the samples from the Galapagos Islands

where many species of Elasmopus also came to

prominence (Table 6). The fauna from 54 samples

is presented in Table 6 in comparison with 54

samples from Cayucos, California, presented in

Table 7. Although the Galapagan samples represent a much greater diversity to sea exposure, rock

angles, depth of crevices explored and thermal

regimes from several different islands, fewer species

(42) were found than at Cayucos (52). Needless to

say the 42 days of arduous Galapagan exploration

(travel time subtracted) was much less productive

than the three days consumed in the Cayucos survey.

BlOGEOGRAPHY

The Galapagan fauna now comprises 51 species,

of which 17 are provisionally endemic (Tables 6, 9).

The two anchialine taxa, Galapsiellus and Anchialella, represent endemic genera probably descended

from Eriopisa and/or Paraniphargus. Neither of the

latter genera has yet been found along Galapagan

shores but the exploration of those islands has

scarcely commenced, even with the present work.

Except for the deepwater Proharpinia tropicana,

with subantarctic affinities, all of the remaining 49

species appear to have affinities with eastern Pacific
NUMBER 271

TABLE 6.—Rank in abundance of intertidal Gammaridea from

the Galapagos Islands (based on aliquots from 54 samples,

total specimens averaged per sample, multiplied by 10; asterisks denote domicolous species)

Hyale darwini

*Ericthonius brasiliensis

*Ampithoe pollex

complex

(includes A. tahue

and vacoregue)

Elasmopus tubar

Elasmopus mayo

Elasmopus serricatus

*Microjassa chinipa

*Posophotis seri

Elasmopus zoanthidea

*Lembos achire

*Ampithoe plumulosa

1176

384

234

212

197

130

93

82

79

75

63

Hyale zuaque

Elasmopus ocoroni

*Lembos macromanus

Amphilochus

neapolitanus

*Photis elephantis

*Chevalia aviculae

Maera chinarra

Meximaera diffidentia

Hyale humboldti

*Microdeutopus hancocki

Elasmopus ecuadorensis

*Ampelisca lobata

*Ampithoe guaspare

41

37

36

36

27

16

16

15

14

11

7

6

5

Minor species: Maera reishi, Hyale guasave, *Varohios

topianus, Heterophlias galapagoanus, Gitanopsis baciroa, Leucothoe spinicarpa, Podocerus spp., *Corophium baconi, Parh<yale hawaiensis, Polycheria osborni, Anamixis linsleyi, Melita

sp., Elasmopus temori, Colomastix sp., Dulzura gal, Leucothoides pottsi.

Other known species in shallow-water benthos from Galapagos Islands: Barnard (1954): Ampelisca milleri Barnard,

Ampelisca pugetica Stimpson; Barnard (1960): Paraphoxus

calcaratus (Gurjanova) (46-73 m); Proharpinia tropicana Barnard (27-46 m); Barnard (1967b): Hyachelia tortugae J. L.

Barnard, from turtle mouth; Myers (1968): *Microdeutopus

trichopus Myers (58-110 m); *Neomegamphopus roosevelti

Shoemaker; Monod (1970) and Barnard (1977): Galapsiellus

leleuporum (Monod); this paper: Anchialella vulcanella.

taxa (or with complexes of taxa known to have

representatives) in eastern Pacific waters. Many of

the endemic species are scarcely distinct from partners on the continent; and once subspeciation in

eastern Pacific Amphipoda has been studied, they

may be relegated to infraspecific level. Endemicity

is, therefore, not very strong in the Galapagan

fauna except in the anchialine element. The Panamic province, sensu stricto, has not been studied

for Amphipoda and may contain many of the

provisional Galapagan endemic species.

The remaining 29 species found in Galapagos

(Table 10) can be divided into those 12 species as

yet found elsewhere only in the Panamic and Gulf

of California provinces, the 6 species that extend

through those provinces as far as the warm-temperate of southern California, and 11 more that

are even more widespread, ranging northward into

TABLE 7.—Amphipoda from a rich intertidal locality, Cayucos,

California (based on 53 samples, specimens per 0.1 m2, recomposed from J. L. Barnard, 1969b; asterisks denote domicolous species)

Hyale frequens

*Aoroides columbiae

*Photis brevipes

*Photis bifurcata

Calliopiella pratti

*Microjassa litotes

*Photis conchicola

Parallorchestes

ochotensis

Elasmopus antennatus

*Jassa falcata

Oligochinus lighti

*Ampithoe pollex

Paraphoxus spinosus

750

262

167

136

130

96

81

73

70

63

57

50

33

*lschyrocerus sp. A

*Cheiriphotis megacheles

*Gammaropsis spinosus

*Gammaropsis

thompsoni

Pontogene¡a rostrata

*Ericthonius brasiliensis

Elasmopus mutatus

*A mpithoe simulans

*Megamphopus effrenus

*Chevalia aviculae

*Ampelisca schellenbergi

Parapleustes pugettensis

arr.philochids

31

24

21

17

14

13

12

10

9

6

6

6

5

Minor species: Maera vigota, *lschyrocerus sp. B, Elasmopus

holgurus, *Ampelisca lobata, Leucothoe alata, Pleustes de·

pressus, Parapleustes nautilus, Atylus levidensus, *Photis

californica, *Ampithoe lacertosa, Podocerus cristatus, *Cera·

pus tubularis, *Ericthonius hunteri, *Corophium baconi,

Leucothoides pacifica, Lysianassa macromera, Maera simile,

Fresnillo fimbriatus, Melita sulca, Podocerus brasiliensis,

Lysianassa pariter, *Ampithoe humeralis, *Ampithoe lind·

bergi, Allorchestes anceps, Pleusirus secorrus, Maera reishi.

TABLE 8.—Dominant species in 5 enriched samples from

Inner Baja California, PAZ 3, 10, 12, 13, 24, with 19 other

minor species not cited (based on aliquots, total specimens

averaged per sample, multiplied by 10; asterisks denote

domicolous species)

*Varohios topianus

Hyale yaqui

*Ampithoe pollex

*A mpithoe plumulosa

Podocerus brasiliensis

*Zoedeutopus cinaloanus

*Lembos macromanus

Amphilochus picadurus

Elasmopus tiburoni

948

510

444

344

272

252

112

96

92

Lysianassa holmesi

Podocerus fulanus

*Ampithoe ramondi

*Ericthonius

brasiliensis

Leucothoe alata

Lembos tehuecos

Batea rectangulata

90

78

58

54

46

42

36

cold American waters. Four of the species, Par hyale

hawaiensis, Colomastix "pusilla", Ericthonius brasiliensis, and Chevalia aviculae are in the category

"circumtropical" though intensive study on speciation in such complexes has not been carried out.

Polycheria osborni and Podocerus brasiliensis require better taxonomic investigations also.
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 9.—Geographic affinities of endemic Galapagan

Amphipoda (Gammar¡dea)

TABLE 10.—Geographic distribution of non-endemic

Galapagan Amphipoda (Gammaridea)

Species Affinity

1. Panamic and Gulf of California Provinces

Anchiallea vulcanella

Ampithoe guaspare

Ampithoe tahue

Ampithoe vacoregue

Elasmopus ecuadorensis

Dulzura gal

Elasmopus ocoroni

Elasmopus temori

Elasmopus zoanthidea

Galapsiellus leleuporum

Heterophlias galapagoanus

Hyale darwini

Hyale humboldti

Lembos achire

Microdeutopus trichopus

Microjassa chinipa

Proharpinia tropicana

Indo-Pacific Eriopisa

Circumtropical Ampithoe

ramondi

E. Pacific A. pollex complex

E. Pacific A. pollex complex

Tropical Pacific E. hawaiensis-antennatus complex

E. Pacific D. sal

Californian E. mutatus

E. Pacific Elasmopus rapax

complex

E. Pacific Elasmopus rapax

complex

Indo-Pacifica Eriopisa

E. Pacific Heterophlias

escabrosa

Pan-Pacific H. rubrafreq uens complex

Pacific H. grandicornis

complex, especially Japan

Gulf of California

L. tehuecos

Not elucidated

California Aí. claustris

Subantarctic

Recorded species from the Gulf of California and

close proximity in depths shallower than 200 meters

total 121 (Table 11). The treatment is very uneven

because in depths of 1-200 m only the Ampeliscidae

and Phoxocephalidae have been studied, to any

extent. By narrowing the treatment to species collected in the very shallow waters the species are

reduced to 86. Some of these have not yet been

collected in the gulf but their distribution pattern

in other parts of the Pacific suggests they will be

found in the gulf. Of these only 40 extend northward to southern California and very few occur

north of Point Conception, California.

One may conclude that about half of the Gammaridea so far found in the Gulf of California are

species ranging northward to the warm-temperature

region of California and which, in many cases, probably occur much to the south of the Gulf in the

Panam¡c province. The other half of the species

probably belong with the greater eastern Pacific

tropical fauna, which would include the Panamic

province and to a more limited extent the Gulf of

California.

Elasmopus tubar

Elasmopus mayo

Gitanopsis baciroa

Hyale guasave

Hyale zuaque

Lembos macromanus

Maera chinarra

Microdeutopus hancocki

Meximaera diffidentia

Neomegamphopus roosevelti

Parhyale hawaiensis

Posophotis seri

Varohios topianus

2. Item 1 plus Southern California

Anamixis linsleyi Maera reishi

Chevalia aviculae Photis elephantis

Elasmopus serricatus Polycheria osborni

3. Items 1 and 2 plus Extrinsic Areas

Ampelisca lobata

Ampelisca milleri

Ampelisca pugetica

Amphilochus neapolitanus

Ampithoe pollex

Ampithoe plumulosa

Colomastix sp.

Corophium baconi

Ericthonius brasiliensis

Hyachelia tortugae

Leucothoe spinicarpa

Leucothoides pottsi

Paraphoxus calcaratus

Podocerus spp.

4. Unknown Affinity

Melita sp.

Isolation from the great pantropical fauna is evident in the cluster of species in Elasmopus (though

several of those are very close to Hawaiian counterparts) and in the occurrence of endemic taxa (such

as Leucothoides pacifica, Anamixis linsleyi, Parhyale

penicillata, species of Photis, and in the endemic

genera of Corophiidae, some related to Hawaii,

and the endemic Bateidae). The Caribbean fauna

is too poorly known to suggest the degree of its influence on the eastern Pacific though J. L. Barnard

(1958) has presented a small amount of evidence

that Caribbean and western Atlantic warm-temperate faunas have Pacific imports. Exceptionally,

brackish water taxa, such as Grandidierella, may

have had easier ingress to the eastern Pacific from

the Caribbean Sea than from Hawaii.

Except in the Corophiidae, containing such

endemic genera as Varohios, Zoedeutopus, Rudilemboides, Acìtminodeutopus, and Neomegamphopus,

the east tropical Pacific is barren of important taxa

denoting evolutionary events at high levels. However, the family Bateidae is endemic to the greater
NUMBER 271 11

Table 11.—Gammaridea of the Gulf of California

Species Reference Depth in Gulf (m)

Extrinsic distribution

and depth (m)

Acuminodeutopus heteruropus

Acuminodeutopus periculosus .

Ampelisca agassizi (= vera,

= compressa)

Amþelisca brevisimulata

Ampelisca cristata

Ampelisca cristoides

Ampelisca hancocki

Ampelisca indentata

Ampelisca lobata

Ampelisca mexicana

Ampelisca milleri

Ampelisca pacißca

Ampelisca pugetica

Ampelisca romigi

Ampelisca schellenbergi .

Ampelisca venetiensis ...

Amphideutopus oculatus

Amphilochns hieapolitanus

Amphilochus picadurus ...

Ampithoe plumulosa

Ampithoe polle×

Ampithoe ramondi

Ampithoe tea

Anamixis linsleyi

Argissa hamatipes

Batea catharinensis

Batea rectangulata

Batea sussurator

Batea transversa

Byblis veleronis

Carinobatea conductor

Ceradocus paucidentatus ..

Cerapus tubularis

Cheiriphotis megacheles ..

Chevalia aviculae

Colomastix sp

Corophium baconi ...

Corophium uenoi

Elasmopus antennatus

Eìasmopus bampo

Elasmopus mayo

Elasmopus rrapax

Elasmopus serricatus ...

Elasmopus tiburoni ....

Elasmopus tubar

Ericthonius brasiliensis

Myers, 1968

Barnard, 1964a

here

Barnard, 1969a

Barnard, 1954a,

Barnard, 1969a,

Barnard, 1954a,

Barnard, 1954a,

Barnard, 1954a,

here

Barnard, 1969a

Barnard, 1969a

Barnard, 1954a,

Barnard, 1954a,

1971a

Barnard, 1954a,

here

Barnard, 1954a,

Myers, 1968

Barnard, 1969a

here

here

here

here

here

Barnard, 1969a

here

Barnard, 1969a

Shoemaker, 1942

here

here

here

Barnard, 1954a,

Barnard 1969a

here

Shoemaker, 1942

here

here

here

here

Barnard, 1969a

here

here

here

here

here

here

here

here

1971a

1971a

1967a

1971a

1967a

1966a

1969a,

1967a

1969b

1971a

unknown

unknown

0-38

15-137

38

15-58

0-46

59-110

38

0-137

25-46

18-101

unknown

0-119

15-128

0-128

37-274

0-41

1

0

0

0

0

0-25

0

0

18-27

0-40

0-37

0-37

38

0

0

11-27

probable

probable

0

0-9

0

probable

0

0

0

0

0

probable

0-42

California to Costa Rica, 1-82

Pan-American, 1—450

Pan-American, 9-456

Pan-American, 6-310

Pan-American, 9-73

Oregon to Costa Rica, 9-210

to California, 33-98

Pan-American, 0-234

Pan-American, 9-73

California to Ecuador, 0-187

Pan-American, 24-496

Washington to Peru, 0-765

Pan-American, 3-504

Pan-American, 0-128

Pan-American, 0-274

California to Costa Rica, 0-162

subcosmopolitan, 0-80

to California, 0-41

Canada to Ecuador, shallow

Pan-American, shallow

tropicopolitan, shallow

to California, 0-67

to California and Galapagos, 0

X. Hemisphere, 0-1096

Pan-American, 0-46

to California, 0-60

to Oregon, 27-422

to Outer Baja California, 0

questionable

California, Galapagos, and Cocos,

0-16

California and Galapagos, 0-35

questionable

Bering Sea to Peru, 0-55

Japan, California, 0-2

California to Cabo San Lucas,

0-11 m

to Galapagos and Ecuador, 0

to California and Ecuador, 0

to Galapagos, 0

cosmopolitan in lower latitudes,

0-171
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 11.—Continued

Species Reference Depth in Gulf (m)

Extrinsic distribution

and depth (m)

Gammaropsis spinosus

Gammaropsis thompsoni

Gammaropsis tonichi

Garosyrrhoe disjuncta

Gitanopsis baciroa

Gitanopsis pusilloides

Grandidierella notton¡

Heterophlias $edusus escabrosa

Heterophoxus oculatus

H¡ppomedon ?propinquns

Hyale californica

Hyale guasave

Hyale yaqui

Hyale zuaque

Jassa falcata

Lembos macromanus

Lembos tehuecos

Leucothoe alata .

Leucothoe spinicarpa ...

Leucothoides pacifica .. .

Leucothoides Tyarrega ..

Liljeborgia marcinabrio .

Listriella melanica

Lysianassa dissimilis ....

Lysianassa holmesi

Lysianassa macromera ..

Maera chinarra

Maera reishi

Maera simile

Megaluropus falciformis

Mega¡uropus visendus ..

Melita sulca

Melita sp. ("nitida")

Metapho×us frequens

Meximaera diffidentia

Microdeutopus schmitti

Micro jassa macrocoxa

Monoculodes hartmanae .

Neomegamphopus roosevelti

Orchomene magdalenensis ..

Pachynus barnardi

Parajassa angularis

Paraphoxus cognatus .

Paraphoxus epistomus

Paraphoxus floridanus

Paraphoxus gemmatus

Barnard, 1969b

Barnard, 1969b

here

Barnard, 1969a

here

Barnard, 1969a

Shoemaker, 1935b

here

Barnard, 1969a

Barnard, 1969a

here

here

here

here

Barnard, 1969a, b

here

here

here

here

here

here

Barnard, 1969a

Barnard, 1969a

here

here

here

here

here

here

Barnard, 1969a

Barnard, 1969a

here

Shoemaker, 1935b

here

here

here

here

Barnard, 1969a

Barnard, 1969a

Myers, 1968

Barnard, 1969a

Barnard, 1969a

Shoemaker, 1942

Barnard, 1969a

Barnard, 1969a

Barnard, 1969a

Barnard, 1960

Barnard, 1969a

probable

0

0-38

0-24

0-1

0-9

probable

0

0-46

15-30

0

probable

0-7

0

7-18

0-9

0-1

0

0-1

0

1

46

7-44

0-1

0

0

probable

0

1

2

2-17

0-24

probable

0

0-24

0-44

1

0-41

0-24

6-46

30-46

18-27

6

9

unknown

2-9

California to Bahía Magdalena,

0-27

to Washington, 0-218

Galapagos, 0

to California, 0-20

only Mazatlán, 0

to California, 0-16

Washington to Panama, 0-1941

to Subarctic, 0-183

to California, 0

to Galapagos, 0-6

to bays of outer Baja California,

0-7

to Galapagos and Ecuador, 0

cosmopolitan, 0-18

to California, Hawaii, and

Galapagos, 0-9

to California, Galapagos, Japan,

0-24

cosmopolitan, 0-1505

to California, 0-8

Australia, shallow

to California, 2-97

to California and Galapagos, 0-73

California to Ecuador, 0-183

to California, 0

to Galapagos, 0

to California and Galapagos, 0-6

to Puget Sound, 0-221

to California, 2-108

to Puget Sound, 0-101

only Mazatlán, 0

Costa Rica to Ecuador and Cocos

Oregon to Isla Isabel, 0-496

to Galapagos, 0-24

California to Costa Rica, 0-221

to outer Baja California, 0-27

to California, 2-146

Pan-American, 0-60, Galapagos

to outer Baja California, 0-46

to California, 12-373

California to Bahía Magdalena,

0-27

to California, 0-325

Pan-American, 0-507

Pan-American, 4-44
NUMBER 271 13

TABLE 11.—Continued

Species Reference Depth in Gulf (m)

Extrinsic distribution

and depth {m)

Paraphoxus obtusidens

"Paraphoxus" spinosus

Paraphoxus tridentatus

Parapleustes commensalis

Parhyale fasc¡gera

Parhyale hawaiensis

Parhyale penicillata

Photis bifurcata

Photis brevipes

Photis californica

Photis elephant¡s

Photis spinicarpa

Platyischnopus metagracilis

Platyischnopus viscana

Podocerus spp? ("brasiliensis", and

"cristatus")

Podocerus fulanus

Polycheria osborni

Pontogeneia opata

Posophotis seri

Rhachotropis luculenta

Rildardanus tros

Rudilemboides stenopropodus ...

Synchelidium rectipa¡mum

Tethygeneia nasa

Tethygeneia quinsana

Vristes entalladurus

Varohios topianus

Westwoodilla cornuta

Zoedeutopus cinaloanus

Barnard, 1960

here

Barnard, 1969a

Barnard, 1969a

here

here

here

Barnard, 1969a

Barnard, 1969a

Barnard, 1969a

here

Shoemaker, 1942

Barnard, 1969a

Barnard, 1969a

here

here

here

here

here

Barnard, 1969a

Barnard, 1969a

here

Barnard, 1969a

here

here

Barnard, 1969a

here

Barnard,1969a

here

unknown

0-24

19-38

9

probable

probable

0

38-46

9-36

34-46

0

probable

13-46

0

0

0-42

0-1

0

0

38-46

9-16

0-38

0-24

0-1

0

38

0

19-16

0

Kuriles to Columbia, 0-459

to Puget Sound and Atlantic, 0-519

to Washington, 0-89

to California, 9

Pan-American trc JÌCS, shallow

pantropical, shallow

to California, 0-93

to Oregon, 0-266

to California, 0-139

to California, Galapagos, 0-6

Bahía Magdalena, 18-27

to outer Baja California, 13-73

to California, 0-27

California, Mexico, 0-171

to California bays, 0-42

to California, Galapagos, 0-1

to California and Cocos, 0-7

to Panama, Galapagos, 0-6

to California, 0-68

Canada to Costa Rica, 0-100+

to outer Baja California, 0-6

to California, 2-38

Pan-American region and may be said to be the

only event of familial magnitude yet discovered in

the evolutionary pattern of eastern Pacific Gammaridea. Once the so-called Pontogeneias, Tethygeneias, and similar taxa are well studied, other

endemic genera may be discovered. The Panamic

province remains unworked although I have examined more than 100 samples from Pacific Panama

and El Salvador and find little of potential importance in the evolutionary picture. For the most part

the east tropical Pacific fauna is composed of widespread tropical or warm-temperate genera, many

with endemic species and evidence of adaptive

radiation (Elasmopus), but which one might call

"invasive" in the botanical vein. The dominance

by Ericthonius, a cosmopolitan genus occurring in

many localities in the Gulf of California, supports

this attribution of the appellation "weedy" to the

fauna.

AMPEUSCIDAE

Atnpelisca lobata Holmes

Ampelisca lobata Holmes, 1908:517.—J. L. Barnard, 1954b:

11-14, pis. 5, 6; 1969a: 188.

MATERIAL.—TOP 3; PAZ 24; GAL 108, 113, 119;

DAW 15, 16, 23, 32.

DISTRIBUTION.—Gulf of California, 0-137 m; Galapagos Islands, 0-128 m; generally Caribbean Sea and

Eastern Pacific Ocean from Puget Sound, Washington, to Ecuador, 0-234 m.
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ampelisca schellenbergi Shoemaker

Ampelisca schellenbergi Shoemaker, 1933a:3-5, fig. 2.—J. L.

Barnard, 1954b:14-16, pis. 7, 8; l969a:l88.

MATERIAL.—TOP 2; PAZ 19, 20.

DISTRIBUTION.—Gulf of California, 0-128 m; generally Caribbean Sea, Gulf of Mexico and Eastern

Pacific Ocean from Cayucos, California to Peru,

0-128 m, probably rare below 20 m.

AMPHILOCHIDAE

Many of the specimens of this family were lost

in transit between Mexico and Washington, D.C.;

much of the data were reconstituted by sorting out

fresh specimens from new aliquots in the original

samples but the following samples could not be

reconstituted because all specimens had been sorted

out in the original examination and were therefore

lost: SCO 1, 18; KNO 1; TOP 1; PAZ 3, 21, 22.

Taxonomy of this family in the eastern Pacific

Ocean remains unsatisfactory. Close comparison of

specimens with European taxa will be needed to

determine if the type-species of Amphilochus and

Gitanopsis are distinct generically from eastern

Pacific species. The eastern Pacific taxa bear an accessory flagellum, lack setation on article 3 of the

mandibular palp, have more poorly developed

mandibular molars (even in Gitanopsis), have strong

setation on the palps of maxilla 1 and usually bear

much better developed gnathopods with distinctive

shapes.

Amphilochus ìneapolitanus Delia Valle

\}]Amphilochus neapolitanus Delia Valle, 1893:595 [probably

distinct from Pacific material].

Amphilochus neapolitanus.—J. L. Barnard, I962C:I26, fig.

3; 1969b: 83. [?Not Delia Valle.]

REMARKS.—Eastern Pacific specimens remain to

be compared with European types.

MATERIAL.—PAZ ?2; GAL 106, 107, 108, 113, 115,

116, 118, 121; DAW 4, 5, 6, 9, 16, 17, 18, 20, 26, 27,

31, 32, 33, 35, 37, 38; COCOS 3; ECU 2; PAN 14.

DISTRIBUTION.—Possibly pantropical and warmtemperate, 0-80 m; eastern Pacific Ocean specific

distribution; Cayucos California south to Ecuador

and the Galapagos Islands, in the Gulf of California

as yet recorded only from Bahía La Paz, 1 m; generally shallow water.

Amphilochus picadurus J. L. Barnard

Amphilochus picadurus J. L. Barnard, 1962c: 126-129, fig. 4;

l969b:82.

REMARKS.—Distinguished from Gitanopsis baciroa, new species, to which it bears great similarity

except in mandible, by the wavy serrations on the

palm of gnathopod 2 as contrasted to the extremely

fine comb of G. baciroa. The anterolateral spines on

the hand of gnathopod 2 in G. baciroa are difficult

to see except under highest magnification. The eyes

of A. picadurus in Gulf of California material have

distinct cores like those of G. baciroa.

MATERIAL.—PAZ 12, 13, 24.

DISTRIBUTOR—Goleta, California, southward into

Gulf of California at Bahía San Evaristo and Bahía

Concepción, 0-41 m.

Gitanopsis baciroa, new species

FIGURES 2, 3

DIAGNOSIS OF FEMALE.—Pleonites dorsally smooth.

Eyes large, formed of dark core surrounded by

unpigmented ommatidia (in alcohol). Gnathopod

1 large, similar in structure to gnathopod 2, subchelate, process of article 5 reaching about threefourths along posterior margin of article 6; gnathopod 2 larger than gnathopod 1, posterodistal end

of article 2 with slender seta, article 6 with 2

anterior thin spines laterally, process of article 5

reaching fully along posterior margin of article 6.

Telson much shorter than peduncle of uropod 3.

Coxa 1 more or less quadrate, coxa 2 rounded

ventrally.

MALE.—Body size much smaller than female,

ranging between 1.50 and 1.75 mm in length,

females of similar size with small eye, males with

eye relatively as large as that of ovigerous females

longer than 2.50 mm, males with juvenile-like

conditions of sparser spination and less expanded

hand of gnathopod 2.

ILLUSTRATIONS.—Following mouthparts magnified

identically: mandible, lower lip, maxillae 1-2,

maxilliped; coxae 3-4 reduced in magnification in

comparison with coxae 1-2; pereopod 2 similar to

pereopod 1; apices of pereopods 3-5 missing on

material; detached telson and uropod 3 magnified

to same degree.

HOLOTYPE.—USNM 169028, female "n," 2.77 mm

(illus.).
NUMBER 271 15

FIGURE 2.—Gitanopsis baciroa, new species, holotype, female "n," 2.77 mm (v = female "v"

2.89 mm).
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Gitanopsis baciroa, new species, holotype, female "n," 2.77 mm.

TYPE-LOCALITY.—DAW 8, Galapagos Islands, Isla

Santa Cruz, off Academy Bay on Isla Coamaño,

13 February 1962, wash of Cystophora (depth unknown).

VOUCHER MATERIAL.—Type-locality, female "v,"

2.89 mm (illus.).

RELATIONSHIP.—This species scarcely differs from

Gitanopsis vilordes J. L. Barnard (1962c) but consistently bears a thin posterodistal seta on article 2

of gnathopod 2, instead of a stout spine, and has a

much different shaped coxa 2, being narrow and

apically rounded rather than subquadrate.

MATERIAL.—SCO 19; TOP 3; GAL 116, ?I2O;

DAW 3, 8, 19.

DISTRIBUTION.—Gulf of California at Puerto

Peñasco and Topolobampo, 0-1 m; Galapagos

Islands, intertidal.

AMPITHOIDAE

Ampithoe guaspare, new species

FIGURE 4 (part)

DIAGNOSIS OF MALE.—Epimera 2-3 flat posteriorly,

posteroventral corners rounded-quadrate. Article 5

of gnathopod 1 shorter than article 6, posterior lobe

truncate, extended as sharp tooth distally, article 6

subrectangular, palm oblique, poorly developed, defined by strong spine, dactyl overlapping palm.

Gnathopod 2 with oblique and weakly excavate

palm defined by medium sized subacute projection,

anterodistal margin of hand tumid, weakly produced, heavily setose, dactyl slender, fitting palm,

article 5 elongate, with moderately developed,

broad, blunt posterior lobe, article 2 with large
NUMBER 271 17

fGl mG2

FIGURE 4.—Ampithoe guaspare, new species (m = holotype, male "m," 6.24 mm; ƒ = female "f,"

6.32 mm; / » remainder of antenna). Ampithoe ramondi Audouin (c = male "c," 5-23 mm).

Ampithoe plumulosa tepahue, new subspecies (n = holotype male "n" 10.49 mm).
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

blunt lobe at anterodistal corner laterally. Antenna

1 longer than antenna 2, both pairs slender,

peduncle of antenna 1 reaching apex of article 4

on peduncle of antenna 2, antenna 2 moderately

setose, flagellum slender, about three-fourths as long

as article 4-5 of peduncle together. Lateral apical

lobule of lower lip projecting strongly, medial

lobule very broad, blunt, scarcely projecting. Coxa 1

produced forward. Peduncular process of uropod 1

absent. Body poorly pigmented, eyes red in alcohol.

FEMALE.—Gnathopod 1 with article 5 shorter

than in male and less strongly produced apicoposteriorly; gnathopod 2 small, article 5 shorter than

in male, palm oblique, slightly concave.

DESCRIPTION.—Lower lip as shown for Ampithoe

ramondi by J. L. Barnard (1965a, fig. l5e), pereopods generally similar but articles 2, 6, and 7 of

pereopod 7 shown herein; outer ramus of uropod 3

as in figure l6j of Barnard (1965a), but inner ramus

with only 3 spines and 6 setae.

HOLOTYPE.—USNM 169014, male "m," 6.24 mm

(illus.).

TYPE-LOCALITY.—DAW 27, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 16 February 1962,

intertidal, rock wash.

VOUCHER MATERIAL.—Type-locality, female "f,"

6.32 mm (illus.).

RELATIONSHIP.—This species is very similar to

Ampithoe ramondi and might appear to be a youthful stage of that species; but in the Galapagos

Islands A. ramondi is much smaller than this

species when the hand of male gnathopod 2 is fully

developed with deep incision and blunt tooth. The

palm of male A. guaspare remains scarcely excavate,

the hand is broadened and the anterodistal tumidity

remains weaker than in A. ramondi.

MATERIAL.—GAL 114, 116; DAW 27, 31, 35, 40.

DISTRIBUTION.—Galapagos Islands, Academy Bay,

Isla Santa Cruz, Darwin Bay, Tower Island; intertidal.

Ampithoe plumulosa Shoemaker

Ampithoe plumulosa Shoemaker, 1938:16-19, fig. 1; 1942:39.—

J. L. Barnard, 1959:37; l965b:2O, figs. 11, 12; l969a:l9O;

1969b: 84.

MATERIAL.—SCO 1, 5, 7, 10, 12, 14, 16, 18, 19,

21, 22; KNO 1; TOP 3; PAZ 2, 3, 6, 7, 8, 9, 10, 11,

13, 18, 20, 24, 25; GAL 101, 107, 108, 115, 120;

DAW 3, 4, 5, 9, 16, 20, 27, 30, 33, 35, 38, 40; ECU

2, 3; PAN 14.

DISTRIBUTION.—British Columbia to Ecuador, including Gulf of California and Galapagos Islands,

intertidal and shallow sublittoral.

Ampithoe plumulosa tepahue, new subspecies

FIGURE 4 (part)

DIAGNOSIS.—Differing from the nominate subspecies in the terminal male by the development of

a sinus and square tooth on the palm of gnathopod

2.

DESCRIPTION.—Subterminal males as in typical

subspecies.

HOLOTYPE. USNM 169015, male "n," 10.49 mm

(illus.).

TYPE-LOCALITY.—DAW 31, Galapagos Islands,

Isla Santa Cruz, halfway between Academy Bay and

Tortuga Bay, 22 February 1962, 6-9 m, algae-rock

wash.

MATERIAL.—DAW 31 (7 specimens).

DISTRIBUTION.—Galapagos Islands: Isla Santa

Cruz, 6—9 m.

Ampithoe pollex Kunkel

FICURE 5 (part)

Ampithoe pollex Kunkel, 1910:92-94, fig. 36.—J. L. Barnard,

1954a:29-31, pis. 27, 28; l969a:l9O, fig. 8; l969b:84.

VOUCHER MATERIAL.—GAL 102, male "k," 3.44

mm (illus.).

RELATIONSHIP.—Galapagan specimens differ from

continental specimens in the broader article 2 of

pereopods 4-5, which resembles that seen in Ampithoe tahue. The rami of uropod 3 are generally

smaller as in A. tahue. Tracing of the ancestry of

A. tahue and A. vacoregue appears possible through

these features, as well as conformity in lower lips

and gnathopod 1. Ampithoe pollex often occurs in

samples with A. tahue and A. vacoregue. The latter

2 species are known only in the female phase. Until

their reproductive biology is studied, they are segregated at specific level rather than being considered

phases of A. pollex.

The samples in which occur A. tahue and A.

vacoregue also contain occasional juvenile males

and very rarely terminal males of A. pollex; the
NUMBER 271 l!>

cG2o

FIGURE 5.—Ampithoe pollex Kunkel (A = male "k," 3.44 mm). Ampithoe tahue, new species

{c — female "c," 4.01 mm; / = remainder of antenna).
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

juveniles have stout antenna 2 like juvenile females

of the other two species, but whether or not these

represent males of those species is unknown. The

samples have been retained in mixed condition to

facilitate future disentanglement of the problem.

MATERIAL.—SCO 1, 12, 13, 14, 19; KNO 2, 3, 5,

6, 10; PAZ 3, 7, 10, 12, 13, 14, 15, 16, 17, 19, 21, 22,

23, 24; GAL 102, 108, 109, 113, 114, 115, 116; DAW

1, 2, 3, 5, 9, 10, 13, 16, 19, 21, 22, 26, 27, 33, 37;

BRU 1.

DISTRIBUTION.—Bermuda and eastern Pacific

Ocean from Coos Bay, Oregon into the Gulf of

California and across to the Galapagos Islands,

littoral and shallow sublittoral.

Ampithoe ramondi Audouin

FIGURE 4 (part)

Ampithoe ramondi Audouin, 1826:93.—Shoemaker, 1942:40.

—J. L. Barnard, 1955a:28-29; 1965b:25-27, figs. 15, 16;

1969a: 190, fig. 7o,p.

VOUCHER MATERIAL.—GAL 105, male "c" 5.23

mm (illus.).

REMARKS.—Males of this species in the Galapagos

Islands reach only 5.5 mm in length, about 1 mm

shorter than A. guaspare, new species, and much

smaller than in other parts of its range. The palm

of gnathopod 1 is definitely excavate. This may be

a distinction marking the specificity of this taxon

in the Galapagos Islands.

The inner ramus of uropod 3 has 4 spines and

5 setae apically. The antennae generally resemble

those of A. guaspare in contrast to specimens illustrated from the Hawaiian Islands by J. L. Barnard

(1970; fig. 18a). Article 2 of pereopod 7 has 1

spinule at the posteroventral corner.

MATERIAL.—PAZ 24; GAL 105, 109, 114, 115,

118, ?I2O; DAW 6, 17, 18, 23, 28, 31, 35.

DISTRIBUTION.—In the Gulf of California only

recorded from Bahía de Los Angeles and Bahía Concepción; Galapagos Islands; eastern Pacific northern

record at Bahía de San Ramon, Baja California;

said to be tropicopolitan; phycophilous, intertidal

or shallow sublittoral.

Ampithoe tahue, new species

FIGURE 5 (part)

DIAGNOSIS OF FEMALE—Pleonal epimera 2-3

with small setule notch at posteroventral corner,

no lateral ridges. Article 5 of gnathopod 1 shorter

than article 6, posterior edge of article 5 with

rounded-truncate lobe, palm oblique, gnathopod 2

especially stout, posterior lobe of wrist thin and

elongate, hand tumid; article 2 of both gnathopods

with anterodistal lobe laterally, not prominent on

gnathopod 2. Article 2 of pereopods 1-2 about 2.6-

2.8 times as long as broad. Ventral edge of article

1 on antenna 1 not spiniferous; peduncles of antennae 1-2 short, peduncle of antenna 2 stout,

article 5 only 0.7 times as long as article 4, medial

surfaces of articles 4-5 with weak conical protuberances, 2 or 3 cones on article 4, 2 on article 5, first

article of flagellum elongate, with apicomedial cone

and apicoposterior tooth, article 2 of flagellum very

short but also with cone and tooth, flagellum only

7-articulate and about as long as article 5 of peduncle. Apical lobules of lower lip about equally short,

bulbous, appressed. Coxa 1 produced forward.

Peduncular process of uropod 1 vestigial. Rami of

uropod 3 especially small. Article 2 of pereopods

4 and 5 weakly expanded, with protruding posteroventral corner. Telson ordinary.

MALE.—Unknown.

HOLOTYPE— USNM 169016, female "c," 4.01 mm

(illus.).

TYPE-LOCALITY.—GAL 102, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 23 January, 1964,

intertidal, from rocks in tidepool.

RELATIONSHIP.—This species is an apparent sibling of A. pollex Kunkel. The female is very similar to the female of A. pollex, especially in the

critical features of lower lip and gnathopods but

differs from that species in the shortened antennae,

especially antenna 2, which resembles in rudimentary form the very stout antennae of the A. lindbergi group. In addition, the medial surfaces of

articles 4-5 on the peduncle bear weak cone-teeth

and article 1 of the flagellum on antenna 2 is elongate and also bears teeth. Uropod 3 is like that of

male pollex with shortened rami. Gnathopod 2 is

very tumid for a female of the genus.

DAW 27 has most males and females of the

pollex form with thin bodies, thin antennae, but

one stout female of A. tahue is present and one

large stout male of this species with juvenoid

gnathopod 2 like that of A. pollex is present. At

4.8 mm this stout male is significantly larger than

the normal terminal males (about 3.5 mm) but

lacks any of the modifications on the antennae of
NUMBER 271 21

females of A. tahue and A. vacoregue except for the

stoutness of antenna 2 and the relatively short flagellum. One might suggest that A. tahue and A.

vacoregue are therefore simply aberrant superfemales, though they bear normally setose brood

plates and carry eggs.

MATERIAL.—GAL 102, 103; DAW 27, 40.

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz, Academy Bay, intertidal.

Ampithoe vacoregue, new species

FIGURE 6 (part)

DIAGNOSIS OF FEMALE.—Pleonal epimera 2-3

with weak setule notch at posteroventral corner, no

lateral ridges. Article 5 of gnathopod 1 shorter

than article 6, posterior edge of article 5 with

rounded posterior lobe, palm oblique, gnathopod 2

of female form, article 5 short and lobate, article 2

of both gnathopods with anterodistal lobe laterally.

Article 2 of pereopods 1-2 about twice as long as

wide. Ventral edges on articles 1-3 of antenna 1

not spiniferous, furnished with elongate setae, especially dense on articles 1-2; antenna 2 much shorter

than antenna 1, flagellum of antenna 1 slightly longer

than peduncle, flagellum of antenna 2 shorter

than article 5 of peduncle, article 4 of peduncle

with sharp posterodistal extension medially. Apical

lobules of lower lip obsolescent, apices of outer

lobes weakly sinuate. Coxa 1 produced forward.

Peduncular process of uropod 1 absent; rami of

uropod 3 especially small. Article 2 of pereopods

3-5 weakly expanded, with protruding posteroventral corner. Knobs on telson slightly enlarged.

MALE.—Unknown.

HOLOTYPE.—USNM 169017, female "g," 5.65 mm

(illus.).

TYPE-LOCALITY.—DAW 31, Galapagos Islands,

Isla Santa Cruz, halfway between Academy Bay and

Tortuga Bay, 22 February 1962, 6-9 m, algae and

rock wash.

RELATIONSHIP.—The only known species in the

eastern Pacific to which this taxon, known only for

the female, is related, appear to be A. pollex and

A. tahue. The shapes of gnathopod 1 and the lower

lip are the clues to this relationship. Ampithoe

vacoregue appears on first sight to be identifiable

with A. tahue but differs from A. tahue in the duplication on pereopod 3 of the shapes on pereopods

4-5, in the lack of medial cones on antenna 2, the

sharpness of the posterodistal and medial extensions

on article 4 of the peduncle on antenna 2, the much

more setose peduncle of antenna 1 and the more

slender gnathopod 2.

MATERIAL.—DAW 31 (4 females only).

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz, vicinity of Academy Bay, intertidal.

Ampithoe kulafi J. L. Barnard

Pleoncxes(T) species.—J. L. Barnard, 1965a:542-543, fig. 34.

Ampithoe kulafi J. L. Barnard, 1970:30.

HOLOTYPE.—USNM 106907, male, 4.0 mm.

TYPE- LOCALITY.—Abbott Station 141-D 3, Ifaluk

Island, west reef between Elangalap Islet and north

tip of Falarik Islet, 20 October 1953, 1 fathom,

wash from algae, lagoonward reef margin.

REMARKS. Heretofore a holotype had not been

designated.

ANAMIXIDAE

Anamixis linsleyi J. L. Barnard

Anamixis linsleyi J. L. Barnard, 1955c:28-30, figs. 2a-d, £-m,

o-w; l969b:89.

MATERIAL.—PAZ 3, 12; DAW 31.

DISTRIBUTION.—Gulf of California: Bahía San

Evaristo and Isla Espiritu Santo, intertidal; generally, from Carmel, California, to Bahía San Evaristo, Gulf of California and across to the Galapagos

Islands, intertidal, probably associated with tunicates and sponges.

BATEIDAE

Batea rectangulata Shoemaker

Batea rectangulata Shoemaker, 1925:31, figs. 7-9; 1926:9-13,

figs. 5-7.—J. L. Barnard, 1969a: 193.

MATERIAL.—PAZ 10, 12, 13.

DISTRIBUTION.—Gulf of California at Bahía de

San Francisquito, at Isla San Francisco, and Bahía

San Evaristo, 0-40 m.

Batea susurrator J. L. Barnard

Batea susurrator J. L. Barnard, 1969a: 193-195, fig. 10.

MATERIAL.—PAZ 1, 11.

DISTRIBUTION.—Gulf of California; Bahía de Los

Angeles, Isla San Francisco, and La Paz, 0-37 m.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 6.—Amptthoe vacoregue, new species (g = holotype, female "g," 5.65 mm). Ceradocus

paucidentatus J .L. Barnard (j = juvenile "j, " 3.74 mm).
NUMBER 271 23

Batea transversa Shoemaker

FIGURE 7

Batea transversa Shoemaker, 1926:13-18, figs. 8-11.—Hewatt,

1946:204.—J. L. Barnard, l962b:8O, fig. 6.—Reish and

Barnard, 1967:15-16.—J. L. Barnard, 1969b:94-95.

Batea transversa coyoa J. L. Barnard, 1969a: 195, fig. 7d,e,f,g.

VOUCHER MATERIAL.—AHF 4818, male "a," 5.7

mm (illus.).

REMARKS.—Intertidal specimens (3^1 mm long)

are half to two-thirds the size of sublittoral specimens (5-6 mm) but are sexually mature. They have

been identified by the key in J. L. Barnard (1969a)

according to coxa 2, pereopod 3, epimeron 3, and

the mandibular palp; but the small size of the specimens complicates identification because of the

poorly toothed dactyls on gnathopod 2, the poor

development and setation of article 2 on pereopods

3-5, and the generally more stout gnathopod 2 than

in sublittoral adults.

DESCRIPTION OF FEMALE.—Apart from presence of

brood plates no apparent differences from male

except that all definitely identified females bearing

deep, triangular posterior excavation on coxa 4.

Largest specimen a female 8.1 mm long (brood

plates vestigial), others exceeding 4.0 mm in length;

generally recognizable females between 4.8 and 6.2

mm in length; poorly excavate coxa 4 confined to

males and juveniles between 3.7 and 4.5 mm long

(smallest specimen, 3.7).

VARIATIONS.—Coxa 4, see discussion under

"Female," above; dorsal process of pleonite 3 highly

variable, not apparently associated with size (above

3.7 mm), varying from high rhombic (Figure 7) to

quadrato-rhombic to transverse-rhombic, pleonite 2

often bearing similar but smaller rhombic process

or rounded lobe; pereopod 5 occasionally with exXI

FIGURE 7.—Batea transversa Shoemaker, male "a," 5.7 mm.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

treme rounded protrusion on article 2 at posterodorsal corner, protrusion weak in most specimens,

absent in small specimens and males.

ADDITIONAL RECORD.—Allan Hancock Station

4818, southern California, 34°3O'2O"N, l2O°3O'3O"W,

19.8 m, 16 January 1957.

MATERIAL.—PAZ 8.

DISTRIBUTION.—California, Morro Bay, Channel

Islands (abundant) off southern California, and

coastal shelf of southern California (rare) southward to Gulf of California at Isla Partida, 0-60 m,

with subspecies coyoa at Bahía de Los Angeles,

9-37 m.

COLOMASTIGIDAE

Colomastix species

Colomastix pus¡lla.—Shoemaker, 1942:12.—J. L. Barnard

1969b: 100-101 [not Grube].

REMARKS.—Specialists studying this group have

advised me that the so-called "Colomastix pusilla"

of Hawaii and California may represent a new

species. Until their works can be completed, the

specimens of this genus in the collections at hand

are held for future study.

MATERIAL.—TOP 2, 3; GAL 115.

COROPHIIDAE

Acutninodeutopus periculosus J. L. Barnard

Acum¡nodeutopus periculosus J. L. Barnard, 1969a:190-192,

fig. 9.

MATERIAL.—SCO 13; unidentified specimens of

genus from PAZ 8, DAW 31.

DISTRIBUTION.—Gulf of California: Puerto Peñasco

and Bahía de Los Angeles, 0-38 m.

Cheiriphotis megacheles (Giles)

FIGURE 34 (part)

Melita megacheles Giles, 1885:70-71, pi. 3.

Cheir¡phot¡s megacheles (Giles).—J. L. Barnard, 1962a: 17,

fig. 4; 1969b: 141-142.

VOUCHER MATERIAL.—DAW 9, male "h," 2.26 mm

(illus.).

REMARKS.—This species is presumably tropicopolitan but more investigation is required on the

taxonomy of the complex.

VARIATION.—The Galapagos material has the

middle tooth on the palm of male gnathopod 2

shifted more proximally than in continental specimens (Figure 34: hG2).

MATERIAL.—PAZ 22; GAL 102, 103, 116; DAW

3, 8, 23, 26, 27, 35, 40; COCOS 1, 9.

DISTRIBUTION.—Eastern Pacific Ocean: Cayucos,

California southward to Cabo San Lucas but not

yet recorded in the Gulf of California; Galapagos

Islands; Cocos Island; intertidal to 16 m.

Chevalia aviculae Walker

Chevalia aviculae Walker, 1904:288-290, pi. 7: fig. 50, pi. 8:

fig. 50.—J. L. Barnard, 1962a: 17-20, fig. 5; 1969b: 142.

VARIATION.—The Galapagos specimens are like

those of California (J. L. Barnard, 1962a) in contrast to those of Hawaii (J. L. Barnard, 1970) in the

shape of article 2 on pereopod 5 and shape of palm

on gnathopod 2.

MATERIAL.—DAW 7, 9, 35, 40.

DISTRIBUTION.—Eastern Pacific form, from Cayucos, California southward to shelf off Bahía de San

Quintín, outer Baja California, 0-35 m; Galapagos

Islands, Isla Santa Cruz, intertidal, but not recorded

in the Gulf of California.

Corophium baconi Shoemaker

Corophium baconi Shoemaker, 1934:356-359, fig. 1.—J. L.

Barnard, l969h:lOl.

MATERIAL.—SCO 14, 16, 19, 21, 22; KNO 1; TOP

2, 3; PAZ 3, 13; DAW 31, 36.

DISTRIBUTION.—Bering Sea to Peru, and into the

Gulf of California and across to the Galapagos

Islands, 0-55 m.

Ericthonius brasiliensis (Dana)

Pyctilus brasiliensis Dana, 1853-1855:976-977, pi. 67: fig.

5a-h.

Ericthonius brasiliensis (Dana).—J. L. Barnard, 1955a:37-38;

1959:39; 1961:183; l964a:H2; l964b:2l9; l969a:!97; 1969b:

102, fig.24f-i.

MATERIAL.—SCO 1, 5, 7, 8, 9, 10, 11, 12, 13, 14,

15, 16, 17, 18, 19, 21, 22; KNO 1, 10; TOP 2, 3;

PAZ 2, 3, 10, 11, 24, 25; GAL 106, 107, 108, 113,

114, 115, 116, 118, 119, 120; DAW 1, 3, 4, 5, 6, 8,

9, 16, 17, 18, 19, 21, 26, 27, 28, 32, 33, 35, 36, 37,

40; COCOS 1, 3; ECU 2; PAN 14.
NUMBER 271 25

DISTRIBUTION.—Eastern Pacific Ocean from Puget

Sound, Washington, southward to Punta Centinela,

Ecuador; into the Gulf of California as far as Puerto

Peñasco, 0-42 m; and at the Galapagos Islands and

Cocos Island; generally cosmopolitan in tropical and

boreal seas, 0-171 m.

Gamtnaropsis tonichi (J. L. Barnard)

Eurystheus tonichi J. L. Barnard, l969b:2l2, fig. 23.

MATERIAL.—SCO 9, ?I5, 16, ?I8, 19; KNO ?l;

TOP 2, 3.

DISTRIBUTION.—Gulf of California, Puerto Peñasco, Bahía de Los Angeles, Bahía Kino, Topolobampo, 0-38 m.

L·embos Bate

Only adult males are identified to species.

The following key contains known species from

the eastern Pacific Ocean. A key to Hawaiian species

(Barnard, 1971b) may also be used to sort out

probable new species that will be encountered in

the eastern Pacific Ocean. L·embos concavus and

L·. audbettius are treated by J. L. Barnard (1962b).

Key to the Males of the Species of L·embos from the Eastern Pacific Ocean

1. Thorax lacking sternal teeth L·. concavus

Thorax bearing sternal teeth 2

2. Some sternal teeth doubled 3

Each sternite with only one tooth 4

3. Coxa 1 strongly setose, setae elongate L·. tehuecos, new species

Coxa 1 poorly setose, setae minute L·. achire, new species

4. Thorax with only 2 sternal teeth, eyes small, anterior margin of article 6 on male gnathopod

1 deeply convex L·. macromanus

Thorax with 6 sternal teeth, eyes enlarged, anterior margin of article 6 on male gnathopod

1 almost straight L·. audbettius

L·embos macromanus (Shoemaker)

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

Lembos macromanus.—J. L. Barnard, l962a:9, fig. 3; 1964a:

110; 1969a: 192; 1969b:90-91 pmisidentification].

MATERIAL.—SCO 1, 12, 13, 14, 15, 16, 17, 18, 19;

KNO 1; TOP 3; PAZ 3, 5, 6, 7, 8, 10, 12, 13, 18, 19,

24; GAL ?l 16, 117, 120; DAW 3, 4, 16, 32, 36.

DISTRIBUTION.—Gulf of California, 0-9 m; outer

Baja California northward definitely to Estero de

Punta Banda and provisionally nortlrward to Cayucos, California, 0 m; Galapagos Islands, 0 m; Hawaii, 0 m; depth range, 0-9 m.

L·embos achire, new species

FIGURE 8

DIAGNOSIS OF MALE.—Lateral cephalic lobes

strongly extended forward, truncate, anteroventral

margin of head excavate, nearly flat-bottomed, then

produced to weak, obtuse anteroventral cusp. Eyes

subcircular, brownish black cores in alcohol surrounded by 1 or 2 layers of clear ommatidia. Coxa 1

short and broad, rectotrapezoidal, scarcely attenuate

and subsharp anteroventral corner; anterior coxae

poorly setose, setae sparse and short. 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, with anteromedial setal brush curving

freely distad, palm oblique, bearing deep incision

separating 2 processes, unarmed defining process

of medium length or short and projecting strongly

posteriad, dactyl slender and scarcely overlapping

palm, dactyl overriding face of hand; article 2 of

gnathopod 2 with large anterodistal lobe similar

to that of gnathopod 1, article 5 slightly broader

and longer than article 6, palm weakly oblique,

defined by spine, setation dense on anterior margins

of articles 5-6. Thoracic sternites usually with 2

teeth each, one tooth occasionally obsolescent,

sternite 5 with 2 tiny nipples, sternite 6 smooth,

sternite 1 with 1 thick ventral tooth, teeth not

arranged in bifid fashion as in L·. intermedius, teeth
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Aim

FIGURE 8.—Lembos achire, new species, holotype, male "a," 4.1 mm (u = female "u," 5.4 mm).

XI

of sternite 3 closely contiguous. Pleonal epimera

1-3 with bulging posterior margins, weak lateral

ridges and slight notch and tooth at posteroventral

corners, ventral setules present on epimera 2-3.

Outer ramus of uropod 3 uniarticulate.

FEMALE.—Coxa 1 longer and slightly less pointed

anteroventrally than in male, bearing only short,

dispersed setae ventrally; gnathopod 1 of normal

kind in Lembos, anterodistal corner of article 2

lacking process, palm oblique, defined by 1 spine,

terminal females with slightly sinuous palm and

weak defining tooth; gnathopod 2 like that of male

but lacking protuberance on article 2; epimeron 2

with ventral setae generally shown for L. tehuecos

but lacking posterior setae, epimera 1 and 3 lacking

all but 1 small seta at posteroventral corner.

DESCRIPTION.—Length of antennae unknown,

antennae broken distally; epistome rounded anteriorly; article 1 of antenna 1 with 3 or 4 ventral

spines as in L. tehuecos, rarely apical spine with

partner; mouthparts generally as shown by Shoemaker (1925, figs. lOd, llb-e) for L. macromanus,

left mandible similar and bearing molarial seta

about half as long as right, with large shark-tooth

flake, right mandible as shown for L. tehuecos, palp

as shown by Shoemaker, see Figure 8:í/ and L of

upper lip and lower lip herein, see Figure 8: XI of

maxilla 1, similar for L. tehuecos and L. achire,

article 1 of palp with 1 long seta, palp with 4 facial

setae; gnathopod 1 of female illustrated, note sinuous palm and weak defining tooth, this condition

typical of L. tehuecos and L. achire, see Figure 9:

nGlo of female gnathopod 1 in L. tehuecos for

subterminal condition of female gnathopod 2 in

both species; see Figure 8:G2 of female gnathopod

2 in L. achire for condition typical of both L.

tehuecos and L. achire; pereopods 3-5 unknown,

missing on all specimens; pigment unknown, specimens long aged in alcohol.

ILLUSTRATION.—Setal densities of gnathopods 1-2
NUMBER 271 27

have been reduced in Figures 8 and 9.

HOLOTYPE.—USNM 142294, male "a," 4.1 mm

(illus.).

TYPE-LOCALITY.—DAW 27, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 16 February 1962,

intertidal, rock wash.

VOUCHER MATERIAL.—Type-locality, female "u,"

5.4 mm (illus.).

RELATIONSHIP.—This species is almost identical

to L·. tehuecos from Gulf of California and may

represent a subspecies of L. tehuecos, but this

taxonomic refinement is difficult to ascertain because of the close relationship of L. tehuecos and

L. intermedius Schellenberg from Hawaii. Both

L·. tehuecos and L·. achire differ from L·. intermedius

primarily in the male sternal teeth, which in L.

intermedius are usually paired and thrust outward

on a pedicle, whereas in the other two species the

teeth sit directly on the sternites. Lembos achire

resembles L·. intermedius in the poorly developed

setae of the male coxae and thus lacks the second

good character distinguishing L·. intermedius and

L·. tehuecos. The only other character suggesting

that L·. achire and L·. tehuecos are distinct species

is the odd attachment region of the dactyl on the

hand of male gnathopod 1, which is somewhat extended, truncate, and appears to force the dactyl

to override the medial face of the hand. All males

in the collections are preserved with the dactyl

folded in this fashion. The upper lip of L. achire

is not excavate as it is in L·. tehuecos. Epimeron 2

of the male lacks a facial seta, but this character

is typical only of a few of the oldest males of L·.

tehuecos.

Lembos achire, L·. intermedius, and L·. tehuecos

form a triad of species showing strong interrelationships between the Galapagos, Hawaii, and America

continents.

MATERIAL.—GAL 105, 107, 109, 113, 114, 116;

DAW 1, 5, 9, 13, 19, 23, 26, 27, 31, 35, 36, 40;

COCOS 3.

DISTRIBUTION.—Galapagos Islands, 0-6 m; Cocos

Island, intertidal.

Lembos tehuecos, new species

FIGURES 9, 10

DIAGNOSIS OF MALE.—Lateral cephalic lobes

strongly extended forward, truncate, anteroventral

margin of head excavate, nearly flat-bottomed, then

produced to weak, obtuse anteroventral cusp. Eyes

subcircular, brownish black cores in alcohol surrounded by 1 or 2 layers of clear ommatidia. Coxa 1

short and broad, rectotrapezoidal, with slightly

attenuate but rounded anteroventral corner; anterior coxae heavily setose below, setae dense and

long and parallel to flat plane of coxa. 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, with anteromedial setal brush curving

freely distad, palm oblique, bearing deep incision

separating 2 processes, unarmed defining process

of medium length or short, dactyl slender and

scarcely overlapping palm; article 2 of gnathopod 2

with large anterodistal lobe similar to that of

gnathopod 1, article 5 slightly broader and longer

than article 6, palm weakly oblique, defined by

spine, setation dense on anterior margins of articles

5-6. Thoracic sternites 2, 3, 4 usually with 2 teeth

each, one tooth often obsolescent or absent, sternite

5 often with 2 tiny teeth (nipples), sternite 6 often

smooth, occasionally with rudiments of nipples,

sternite 1 with 1 thick ventral tooth, teeth not

arranged in bifid fashion as in L·. intermedius.

Pleonal epimera 1-3 with bulging posterior margins, weak lateral ridges and slight notch and tooth

at posteroventral corners, ventral setules present on

epimera 2-3, epimeron 2 with long subventral seta.

Outer ramus of uropod 3 uniarticulate.

FEMALE.—Coxa 1 longer and slightly less pointed

anteroventrally than in male, bearing only short,

dispersed setae ventrally; gnathopod 1 of normal

kind in Lembos, anterodistal corner of article 2

lacking process, palm oblique, defined by 1 (aberrantly by 2) spine, terminal females with slightly

sinuous palm and weak defining tooth; gnathopod 2 like that of male but lacking protuberance on

article 2; epimera heavily setose, especially posterior

margins of epimera of 1-2 and ventral margin of

epimeron 2.

DESCRIPTION.—Antenna 1 as long as head,

pereon and first 2 pleonites together, antenna 2

scarcely shorter than antenna 1; epistome rounded

anteriorly; mouthparts generally as shown by Shoemaker (1925, figs. lOd, llb-e) for L·. macromanus,

left mandible similar and bearing molarial seta
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gl

FIGURE 9.—Lembos tehuecos, new species, holotype, male "a," 4.4 mm (m »» male "m," 3.6 mm;

n — female "n," 4.8 mm; u «* male "u," 4.4 mm; y •= male "y" 4.3 mm; ƒj «= remainder of

antenna; l¿ — molarial flake).
NUMBER 271 29

mD3

FIGURE 10.—Lembos tehuecos, new species, holotype, male "a," 4.4 mm (m = male "m," 3.6 mm;

n = female "n," 4.8 mm).

about half as long as right, with large shark-tooth

flake, right mandible figured, palp as shown by

Shoemaker, palp of maxilla 1 with 4-6 facial setae,

see Figures 9: U and Si of upper lip and inner plate

of maxilliped, see Figures S:Xl and U of L. achire

for similar maxilla 1 and lower lip but mandibular

lobes uncurled apically; pereopods as illustrated;

uropods 1-2 each with long interramal tooth;

telson with weak apicolateral slit: female gnathopod

1 terminal condition as for L. achire (Figure 8:

uGlo).

VARIATION.—One terminal male of PAZ 24 with

coxa 1 like that of L. achire, long setae absent;

smaller males in same sample with long setae of

coxa 1 moderately well developed.

REMARKS.—In alcohol, on male pigmentary

blotches composed of trabeculae covering posterodorsal part of head, patches on epimera 1-3 and

medial to epimera outlining bases of pleopods, vertical bands, 1 each on pereonites 1, 7, pleonites 1,

2; 2 bands on pereonites 4, 5, 6; 1.5 bands on pereonites 2, 3; coxae 2 and 5 clear, other coxae pig-
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mented; occasionally pereonite 5 clear (this formula

represents specimens with densest pigmentation,

numerous other specimens with less pigmentation);

female with 1 anterior and 1.5 posterior bands on

pereonite 1 (or no posterior bands), 2 bands on

pereonites 2, 3, 4, 6, and 1 band on pereonite 5

(or clear), and on pleonites 1, 2, 3; pereonite 7 clear.

Note that L. macromanus (Shoemaker) is characterized by pigmentary trabeculae organized into

stellate figures, whereas trabeculae of L. tehuecos

meander.

ILLUSTRATIONS.—Setal densities of gnathopods 1-2

have been reduced in Figure 9.

HOLOTYPE.—USNM 142310, male "a," 4.4 mm

(illus.).

TYPE-LOCALITY.—TOP 3, Gulf of California,

Topolobampo, 25 November 1971, 1 m, rock wash,

with tunicates and sponges.

VOUCHER MATERIAL.—Type-locality, female "n,"

4.8 mm (illus); male "m," 3.6 mm (illus); male

"u," 4.4 mm (illus.); male "y," 4.3 mm (illus.).

RELATIONSHIP.—This species bears close resemblance to L. intermedius Schellenberg from the

Hawaiian Islands. This relationship is marked by

the doubling of ventral teeth on certain sternites

of the male, but in L. intermedius the outline

forms a single bifid process on the several segments

whereas in L. tehuecos the teeth are separate basally

from each other. The heavy brush of medial setae

on male gnathopod 1 is also congruent between the

two species; but L. tehuecos has a setal brush on

male coxa 1, and dense, long setae on coxae 2-4,

untrue of L. intermedius. The two species show a

relationship in the absence of article 2 on the outer

ramus of uropod 3, and in the mammilliform lobe

on article 2 of the male gnathopods. Perhaps these

2 species should be ranked at subspecific level.

Lembos tehuecos is easily distinguished from

L. macromanus (Shoemaker), its sympatriot, in the

setal brush of male gnathopod 1 and in the diffuse

pigment, which in L. macromanus forms trabecular

stellate figures.

Lembos concavus Stout (J. L. Barnard, 1962a)

may form a subspecies of this complex or be an

extreme clinal member of the L. tehuecos epigenotype, in which case L. concavus would have nomenclatural priority. Lembos concavus, however, lacks

elongate setae on the anterior male coxae and has

a very narrow article 6 on male gnathopod 1; sternal

teeth on the male are obsolescent.

MATERIAL.—SCO 14; TOP 2, 3; PAZ 13, 24.

DISTRIBUTION.—Gulf of California at Puerto

Peñasco, Topolobampo, Bahía Concepción, and

Bahía San Evaristo, 0-1 m.

Microdeutopus Costa

Microdeutopus hancocki Myers

Microdeutopus hancocki Myers, 1968:497-501, figs, lb,c,e-l,k,

6a; 1969:124, figs. 12a, l3a,c, l4d,h,m, 15, l6c,e, 20k, pi. la.

MATERIAL.—GAL 108; DAW 32.

DISTRIBUTION.—Galapagos Islands, 0-18 m; generally from Costa Rica to Ecuador, 3-15 m.

Microdeutopus schmitti Shoemaker

Microdeutopus schmitti Shoemaker, 1942:18-21, fig. 6.—J. L.

Barnard, 1959:32-33, pi. 9; 1961:180; l964a:llO; l964b:2l8;

1966a: 17; l966b:6O.—Reish and Barnard, 1967:15.—Myers,

1968:497, fig. la,d,j,l; 1969:120-122, figs. 11, l3b,d, l4a-<:,f,g4,

15, l6a,b,d, 20m, pi. 16.—J. L. Barnard, 1969a: 192;l969b:9l.

MATERIAL.—PAZ 11, ?I3.

DISTRIBUTION.—Gulf of California from Isla San

Francisco to Bahía de Los Angeles as yet on Baja

California side only, 0-44 m; generally from

Monterey Bay, California, south to Costa Rica,

0-221 m, rare in depth greater than 65 m.

Photis Krøyer

Photis elephantis J. L. Barnard

Photis elephantis J. L. Barnard, 1962a:39-42, figs. 16, 17;

1969b: 151.

MATERIAL.—SCO ?6, 8, 10, 14, ?I5, 18; PAZ 18,

21; GAL ?IOI, 105, 108; DAW ?5, 16, 31.

DISTRIBUTION.—Gulf of California, Puerto Peñasco, Cabo San Lucas, 0 m; Galapagos Islands,

0-6 m; to Corona del Mar, California; depth range,

0-6 m.

Posophotis, new genus

ETYMOLOGY.—Posophotis is from the Greek

posos (how much) and photis (from the classic generic name).

DIAGNOSIS.—Article 3 of antenna 1 as long as

article 1; accessory flagellum 2-3 articulate (possibly

with tiny additional article). Head deeply recessed
NUMBER 271 31

for reception of antenna 2. Coxae 1-5 elongate,

large, and overlapping. Mandible with 3-articuIate

palp, article 3 shorter than article 2, clavate,

strongly setose. Dactyl of maxilliped short, stubby,

multisetose apically. Gnathopod 1 small, weakly

subchelate, article 5 about 1.1 times longer than

article 6. Gnathopod 2 of both sexes slightly larger

than gnathopod 1, wrist of moderate length, broadly

but weakly lobate, hand slightly longer than

wrist, broad, subchelate, palm oblique, sculptured.

Peduncle of uropod 3 elongate, rami styliform,

slightly shorter than peduncle, outer raraus 1-

articulate, inner ramus as long as outer or occasionally in male shortened, about 60 percent as

long as outer ramus. Telson ordinary, with small

posterior recessment.

TYPE-SPECIES.—Posophotis seri, new species (here

designated).

COMPOSITION.—Monotypic; possible member is

Gammaropsis photosimilis Ruffo, 1969.

RELATIONSHIP.—This genus has the external

facies of Photis and Cedrophotis but differs in the

absence of article 2 on the outer ramus of uropod 3,

the well-developed accessory flagellum and the

stubby dactyl on the maxillipedal palp. This genus

would key to the Gammaropsis complex (J. L. Barnard, 1973) but differs from all the genera of that

complex except Pseudeurystheus in the stubby

dactyl of the maxillipedal palp and from the vast

majority in the Photis-like appearance of the coxae.

Males of Pseudeurystheus have an elongate wrist

on gnathopod 2, much longer than the hand, and

the coxae are not elongate.

Posophotis seri is diagnosed and described below

in expanded form for purposes of comparison to

members of the Gammaropsis complex and Photis.

Posophotis seri, new species

FIGURES 11, 12

DIAGNOSIS.—Accessory flagellum composed of 2-3

long articles possibly tipped by minute fourth. Coxae

of elongate form as in Photis, coxae 1-2 longer

than broad but slightly shorter than coxae 3-5,

latter as long as coxa 4, coxae 3-4 of male only

expanded in middle; stridulation ridges absent

everywhere. Article 5 of gnathopod 1 longer than

article 6, palm oblique and weakly excavate, poorly

defined but with 2 widely spread defining spines,

dactyl overlapping defining spines. Male gnathopod

2 small, article 2 not lobate, article 5 large, weakly

lobate, article 6 broad, only 1.4 times as long as

article 5, palm oblique, bearing 1 sinus and weak

defining tooth, dactyl not reaching apex of palm;

female gnathopod 2 with deep palmar excavation

guarded by tooth bearing large spine, dactyl overreaching palm. Article 2 of pereopod 3 very broad

and grossly lobate posteroventrally, of pereopod 4

narrower and unlobate, of pereopod 5 scarcely expanded and unlobate; article 4 of pereopod 3 expanded and strongly spinose posteriorly, of pereopods 4-5 narrow and poorly spinose; locking spines

composed of 1 long and 1 short simple spines;

dactyls simple. Epimera 1-3 almost evenly rounded

posteriorly, posteroventral corners with weak notch

and setule, epimeron 3 with additional setule and

notch, these together not contiguous. Peduncle of

uropod 1, but not uropod 2, with long interramal

tooth, outer rami of both pairs shorter than inner

rami, spines not heavily thickened. Peduncle of

uropod 3 elongate, rami styliform, rami shorter

than peduncle, outer ramus bearing 3 thick marginal spines, apex lacking article 2, bearing 2 spinules, inner ramus of female as long as outer ramus,

inner ramus of male about 65 percent as long as

outer ramus, with 1 subapical spine in male, 3 marginal spines in female (mixed on opposite margins).

Telson trapezoidal, apex broad, with ventroapical

protrusion and apicodorsal excavation marked by

2 large dorsal spines, 1 apicolateral setule each side,

2 minute apicolateral knobs each side, with pair of

penicillate setules each side in middle.

DESCRIPTION.—Upper lip with ventral notch; incisors of mandibles with 5 teeth, right Iacinia mobilis

bifid, spine row with 3 spines, molar with large

flake and long seta, weakly triturative at cutting

edge, left mandible with 4 spines, no flake or seta

on molar; lower lip as shown for Photis reinhardi

by Sars (1895, pi. 202) but mandibular lobes sharper;

maxilla 1 as shown by Sars, outer plate with 11

spines but inner plate lacking seta, palp with 5

spines and 4 setae (both sides); maxilla 2 as shown

by Sars, inner plate with submarginal oblique setal

row medially; maxilliped as shown by Sars except

for palp article 4 illustrated herein, inner plate

with 3-5 stout apical spines, 1 medial locking spine

and about 7 terminal and 5 medial setae, outer

plate with about 6 medial and 4 apical spines and

setae.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gl

Fir.LRF 11.—Posoplìotis seri, new species, holotype, male "a," 3.12 mm (w = female "w," 2.77 mm).
NUMBER 271 33

FIGURE 12.—Posophotis seri, new species, holoiype, male "a," 3.12 mm (/ = female "f," 2.70 mm;

w = female "w," 2.77 mm).
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VARIATION.—Galapagan males have inner ramus

of uropod 3 as long as that of female.

ILLUSTRATIONS.—Gnathopod 1 enlarged greater

than gnathopod 2; female pereopod 4 like that

shown for male; only terminal male lacking pereopod 5 but presumed to resemble that of female

illustrated herein; two main setae of uropod 3 elongate but appearing short and stout in illustration.

HOLOTYPE.—USNM 149507, male "a," 3.12 mm

(illus.).

TYPE-LOCALITY.—SCO 10, Gulf of California,

Puerto Peñasco, 23 February 1971, intertidal, Sargassum sp.

VOUCHER MATERIAL.—Type-locality, female "w,"

2.77 mm (illus.): SCO 19, female "f," 2.70 mm

(illus.).

MATERIAL.—SCO 1, 10, 18, 19, 21; KNO 1; GAL

101, 105, 106, 107, 118, 120; DAW 1, ?3, 4, 5, 6, 9,

20, 21, 26, 28, 31, 33, 35, 36, 38, 40; PAN 14.

DISTRIBUTION.—Gulf of California at Puerto Peñasco, Bahía Kino, 0 m; Galapagos Islands, 0-6 m;

Panama, 0 m.

Rudilemboides J. L. Barnard

Rudilemboides stenopropodus J. L. Barnard

Rudilemboides stenopropodus J. L. Barnard, 1959:31-32, pi.

8; 1961:180; l964a:llO; 1969a:192-193.

MATERIAL.—SCO 18, 19; PAZ 11.

DISTRIBUTION.—Gulf of California at Puerto Peñasco, Bahía de Los Angeles and Isla San Francisco,

0 m; from Point Conception, California, to Bahía

de San Quintín, Baja, California, 1-68 m.

Varohios, new genus

DIAGNOSIS.—Antenna 2 inserted deeply below

head and behind eye, ocular lobes strongly protuberant; article 3 of antenna 1 longer than article

1, accessory flagellum 2-articulate; antennae 1-2

extending subequally, thin, of medium length.

Mandibular palp article 3 tumid, heavily setose.

Inner plate of maxilla 1 small, bearing 1 medial

seta. Maxillipedal palp article 4 short, stubby, with

stout apical spine and several subapical setae. Male

gnathopod 1 much larger than gnathopod 2, bearing only 6 articles (either article 5 or article 7

absent or resorbed in adjacent articles), article 6

forming dactyl bearing large inner tooth and closing on long chela of article 5 (forming hand),

gnathopod 2 small, moderately setose, article 6

longer than article 5, female gnathopods 1-2 small

like male gnathopod 2, subequal in size to each

other. Coxae overlapping serially, coxa 1 of male

broader than coxa 2, coxa 5 with anterior lobe as

long as coxa 4. Rami of uropod 3 extending equally

and subequal to peduncle, latter slightly elongate

(in familial context), outer ramus with small barrelshaped article 2.

TYPE-SPECIES.—Varohios topianus, new species.

RELATIONSHIP.—This genus keys to Aloiloi J. L.

Barnard or Pseudomegamphopus Myers because of

the enlarged male gnathopod 1 but differs so radically from those genera that discussion of similarities is pointless. Like those genera, Varohios appears

to be another example of axial reversal in gnathopods 1-2, which in a hypothetical ancestor would

have had gnathopod 2 enlarged but in evolutionary

descent has that gnathopodal form shifted to the

position of gnathopod 1. An alternative hypothesis

is that the ancestor of Varohios bore a male gnathopod 1 which conceivably might have lost an article

to assume the form seen in Varohios. The best

example of this possibility could be visualized in

gnathopod 1 of Neomegamphopus Shoemaker. That

gnathopod bears the normal 7 articles and the 6th

bears a large inner hump similar to the hump of

article 6 on Varohios. In this hypothesis one would

presume that article 7, the dactyl, was fused to

article 6, the latter now assuming the function of a

dactyl and closing on the chela of article 5, found

in both Varohios and Neomegamphopus. I find no

line of fusion separating two parts of the new dactyl

on Varohios except near its base, proximal to the

hump; this small segment does not qualify as a remnant of the fully articulate system in Neogamphopus since the hump of Neomegamphopus lies

on article 6, whereas the hump in Varohios, by

this definition, would lie on article 7. There are

no articular remnants visible in the region of articles

3-4 on Varohios, where one might expect fusion to

be more common than apically. Nevertheless, I find

the first hypothesis more attractive, that articles 6-7

of a Neomegamphopus-\ike gnathopod 1 have become fused to assume the condition found in

Varohios.

In the other proposal, that gnathopod 1 of Varohios is a direct result of axial reversal from a Gam-
NUMBER 271

maropsis-Podoceropsis-Megamphopus ancestor, few

concrete examples can be found, because no extant

species of those 3 genera is known to have lost an

article on male gnathopod 1, and only the following species have a chelate gnathopod 2 closely fitting the form seen in Varohios: Gammaropsis Una

(Kunkel, 1910), G. semichelatus (K. H. Barnard,

1957), and Audulla chelifera Chevreux (1901, see

Sivaprakasam, 1970:564).

In those species, however, the chela is found on

article 6, whereas the chela of Varohios is found on

article 5. In the transformation from one stage to

another, one would have to assume loss of article 3

or 4 in Varohios, thus increasing the number of

steps in evolutionary descent. The most parsimonious system is one involving descent of Varohios

from a Neomegamphopus-\ike ancestor.

Because female and juvenile gnathopod 1 bears

7 articles it follows that adu)t males must lose the

7th article during maturation; the material at hand

lacks any males in stages showing this supposed

transformation.

Varohios topianus, new species

FIGURES 13, 14

DESCRIPTION OF MALE.—Eyes subovate, with black

core surrounded by 3 or 4 layers of tiny ommatidia.

Upper lip broadly rounded below, prebuccal parts

rounded anteriorly, no epistomal spike. Lower lip,

maxilla 2 and dactyls of pereopods 1-2 as shown

FIGURE 13.—Varohios topianus, new species, holotype, male "a," 2.53 mm (m

2.61 mm).

male "m.
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Varohios topianus, new species, holotype, male "a," 2.53 mm (ƒ — juvenile "j, "

1.57 mm).
NUMBER 271 37

for Zoedeutopus cinaloanus; only right mandibular

molar with small plumose seta. Hand (article 5) of

gnathopod 1 with secondary palmar tooth inside

chela. Uropod 1 with immense interramal tooth,

uropod 2 lacking interramal tooth. Pleonites 4-5

each with 1 dorsolateral seta on each side marking

dorsal limit of pellucid chitinous sleeve overlapping

next posterior segment. Telson with short, deep

dorsoposterior excavation guarded by stout spine

each side, pair of lateral setae each side, 1 distal

setule each side, 1 small boss each side and lateral

to setule. Pereonal sternites lacking teeth.

Pigment in alcohol; following parts covered with

dark purple brown pigment: pereonites 4-5, posterior half of coxa 4, dorsoposterior parts of head or

scattered lightly over entire head; uropods 1-2 bearing peduncular bands of similar pigment, pereonites

5, 7, pleonites 1, 2 each with small dorsal spot, coxa

2 with medium density spot on posteroventral face,

coxa 5 with scattered pigment on anteroventral face,

pleonites 1-2 with scattered lateral pigment, bases

of pleopods behind epimera 1-3 outlined with

diffuse purple pigment.

FEMALE (largest, 2.88 mm).—Gnathopod 1 smaller

than gnathoped 2, gnathopods as illustrated herein

for juvenile. Pigment similar to male but pigment

dark on head, pereonites 1-2, coxae 1-2, no pigment

on pereonites 4—5 and dorsal spots present only on

pereonite 7 and pleonite 1.

HOLOTYPE.—USNM 142349, male "a," 2.53 mm

(illus.); pereopod 4 missing both sides.

TYPE-LOCALITY.—PAZ 19, Baja California, 11 km

E of Cabo San Lucas, 4 December 1971, intertidal,

from small polychaete-like (possibly amphipod tubes

of this species).

VOUCHER MATERIAL.—Type-locality, male "m,"

2.61 mm (illus.), juvenile "j, " 1.57 mm (illus.).

MATERIAL.—PAZ 3, 12, 13, 19; GAL 108; DAW

16.

DISTRIBUTION.—Gulf of California at Bahía San

Evaristo, Isla Espiritu Santo and Cabo San Lucas,

0 m; Galapagos Islands, 0 m.

Zoedeutopus, new genus

DIAGNOSIS.—Lateral cephalic lobes strongly produced, antenna 2 deeply inserted along ventral

cephalic margin. Mandibular palp articles 2 and 3

subequal in length to each other, article 3 linear,

setose apically, article 1 more than half as long as

articles 2-3. Mandibular lobes of lower lip subacute.

Inner plate of maxilla 1 broadly expanded, apically

rounded, bearing 2 facial setae. Gnathopod 1 larger

than gnathopod 2 in female, in male enlarged and

of Microdeutopus-íorm, carpochelate, article 6

shorter and narrower than article 5, gnathopod 2

in male also enlarged and larger than gnathopod

1, weakly carpochelate, article 6 shorter and narrower than article 5, chelate. Rami of uropod 3

subequally long but inner distinctly shorter than

outer, latter with distinct article 2. Telson ordinary.

TYPE-SPECIES.—Zoedeutopus cinaloanus^ new species.

RELATIONSHIP.—This genus superficially resembles

Microdeutopus Costa, which has been monographed

by Myers (1969). Zoedeutopus cinaloanus resembles

the armatus-chelifer group of that genus in which

male gnathopod 2 is enlarged and propodochelate

but Zoedeutopus differs from Microdeutopus generally in the deeply inserted antenna 2, the short,

linear article 3 of mandibular palp on which all

setae are terminal, in the facial and not apical

setae on the inner plate of maxilla 1, and in the

presence of a distinct article 2 on the outer ramus

of uropod 3. In these respects, especially the head

and uropod 3, Zoedeutopus appears to belong to

the group of genera including Neomegamphopus

Shoemaker, Amphideutopus J. L. Barnard, Acuminodeutopus J. L. Barnard, Konatopus J. L. Barnard, from the east tropical Pacific and Hawaii, all

of which have the Microdeutopus-ìike male gnathopod 1. Those four genera may be divided into two

groups based on the flabellate expansion of article 3

on the mandibular palp in Amphideutopus, Neomegamphopus, and Konatopus, and its simple,

linear condition in Acuminodeutopus and Zoedeutopus. The new genus appears to be the most primitive member of this generic group, because of the

simple article 3 of the mandibular palp, moderate

insertion of antenna 2 (deep but not as extreme

as in the other genera), and the slight shortness of

the inner ramus of uropod 3. The facial setae on

the inner plate of maxilla 1, however, may be an

advancement. Acuminodeutopus appears to be the

most closely related genus to Zoedeutopus but is

more specialized by virtue of the strong shortening

of the inner ramus on uropod 3 while retaining the

small male gnathopod 2 typical of Microdeutopus.

Zoedeutopus is peripheral to the triangle of
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Amphideutopus-Neomegamphopus-Ko?iatopus discussed by J. L. Barnard (1970:70), a group characterized by shortening of the outer ramus of uropod

3 or the precise equality of both rami, bears the

enlarged male gnathopod 2 characteristic of Amphideutopus and in Neomegamphopus and Konatopus has article 4 of the maxillipedal palp

reduced in size. In the four genera related to

Zoedeutopus the setae on the inner plate of maxilla

1 appear to have migrated to the medial margin.

In a phyletic sequence one would place Zoedeutopus in the center, with Acuminodeutopus forming

one line of descent and Amph¡deutopus forming

the first member of a second line of descent, followed by Neomegamphopus. The Hawaiian Kona·

topus would appear to be descendent from

Neomegamphopus.

These 5 genera, containing only 8 known species,

form a group so far discovered only in East Africa,

middle America, and Hawaii. The counterpart and

presumedly more primitive ancestors comprise the

genera Microdeutopus and Chevreuxius Bonnier

from both tropical and cooler latitudes. A second

counterpart, composed of Grandidierella Coutière

and Neomicrodeutopus Schellenberg ( = }Grandidierella) is found in the tropical eastern hemisphere

but appears to be descendent from Microdeutopus

because of the poorly inserted antenna 2.

Zoedeutopus cinaloanus, new species

FIGURES 15-17

DESCRIPTION.—Antenna 1 about as long as

pereonites 1-6 together, antenna 2 about threefourths as long as antenna 1, article 1 of antenna

1 with 1 midventral spine, 1 apicoventral spine,

article 3 approximately half as long as article 1,

accessory flagellum 2-articuIate, antenna 2 inserted

deeply into head because of protrusion of lateral

cephalic lobe but insertion reaching only posterior

margin of eye and not as deep as in Amphideutopus·

Neomegamphopus. Upper lip broadly rounded below. Molar of left mandible with seta only half as

long as shown for right mandible. Interramal tooth

of uropod 1 extending nearly 40 percent along outer

ramus, uropod 2 lacking interramal tooth. Male

lacking sternal teeth.

REMARKS.—Pigment in alcohol brownish purple

on posterior part of head and marked by vacuoles,

on pereonite 5 and parts of pereonites 4 and 6,

broad stripe on article 2 of pereopod 3, blotch on

anterior face of epimeron 1, pigment generally consistent on more than 80 percent of specimens.

ILLUSTRATIONS.—Maxilla 1 enlarged in comparison to maxilla 2.

HOLOTYPE.—USNM 142350, male "a," 2.37 mm

(illus.).

TYPE-LOCALITY.—SCO 14, Gulf of California,

Puerto Peñasco, 24 February 1971, intertidal, rock

wash.

VOUCHER MATERIAL.—Type-locality, female "w,"

3.14 mm (illus.); PAZ 12, male "b," 2.70 mm (illus).

MATERIAL.—SCO 1, 12, 14, 17, 18; PAZ 3, 5, 9,

12, 24.

DISTRIBUTION.—Gulf of California, at Puerto

Peñasco, Bahía Concepción, Bahía San Evaristo,

Isla Partida, and Isla Espiritu Santo, 0 m.

DEXAMINIDAE

Polycheria osborni Caiman

Polycheria osborni Caiman, 1898:268-269, pi. 32: fig. 2 —

Skogsberg and Vansell, 1928:268-282, figs. 1-26.—J. L.

Barnard, l969a:2OO, fig. 25g; 1969b: 103.

Polycheria antarctica—Alderman, 1936:63.—J. L. Barnard,

l954a:2l [not Stebbing, 1888].

MATERIAL.—PAZ 3; TOP 2, 3; DAW 16.

DISTRIBUTION.—Gulf of California, at Bahía de

Los Angeles, Topolobampo, and Isla Espiritu Santo,

0-1 m; Galapagos Islands, 0 m; California, northward to Monterey Bay, intertidal; usually burrowing into tests of Amaroucium spp.

EUSIRIDAE

Pontogeneia Boeck

J. L. Barnard (1972a: 186) redefined several genera

of the Eusiridae and excluded from Pontogeneia all

but 1 valid species and 2 questionable species. Part

of that decision rested on the erroneous basis that

the rostrum of the type-species P. inermis (Krøyer)

was short and blunt, according to figures in Sars

(1895, pi. 159c). I have now examined specimens

of P. inermis and find the rostrum larger than supposed, somewhat smaller but similar to that of

several mid-American species which should now be
NUMBER 271

39

FIGURE 15.—Zoedeutopus cinaloanus, new species, holotype, male 'a," 2.37 mm.
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

wG2

FIGURE 16.—Zoedeutopus ánaloanus, new species, holotype, male "a," 2.37 mm (b — male "b,'

2.70 mm; u> = female "w." 3.14 mm: note Gl bottom left with tilted article 6).
NUMBER 271 41

R3p

P5

FIGURE 17.—Zoedeutopus cinaloanus, new species, holotype, male "a," 2-37 mm.

reinstated in Pontogeneia and removed from Tethygeneia where they were placed in 1972. Maxilla 2

of P. inermis has also been clarified and shown to

have 4 enlarged or partially enlarged medial setae

similar to the mid-American species with 2 or 3

of these setae. The calceoli of several of those midAmerican species have now been studied and are

found to be similar to the calceoli of P. inermis as

shown by Sars (1895); therefore, these species can

now be considered to be congeneric. Pontogeneia

inermis has distinct but vestigial inner lobes on the

lower lip, whereas the mid-American species have

these lobes so evanescent as to be nearly invisible.

I believe that these lobes should be discounted in

this case, although this would require revision of

the arrangement in Figure 111 of J. L. Barnard

(1972a). Needless to say that arrangement is but a

strawman ready to be modified as various facts are

uncovered to regulate the confusion in the pontogeneiid genera. The correspondence in rostra, accessory flagella (their absence), maxillae 1-2 and calceoli among P. inermis, P. rostrata, P. intermedia,

P. opata, P. quinsana (calceoli unknown), and P.

bartschi seems sufficient to group these warmtemperate and cold-temperate species from the

northern hemisphere in Pontogeneia and to contrast

them with Tethygeneia and Gondogeneia from the

southern hemisphere. Tethygeneia includes the

northern P. nasa J. L. Barnard from the Gulf of

California. Aesthetascs of Pontogeneia longleyi Shoemaker (1933b) and P. quinsana J. L. Barnard

(1964a) are unknown. The main distinctions among

these genera involve the calceoli of the male antennae. A calceolus of Pontogeneia consists of 2

large subequal parts, that of Gondogeneia consists

of 4 parts arranged linearly and that of Tethygeneia

consists of 1 main part and a cluster of 3 small

basal lobes. Rostra of Pontogeneia and Tethygeneia

are long and thin (except P. bartschi with short

and thin rostrum), whereas the rostrum of Gondo·

geneia is very short and blunt, the head very massive, and the eyes greatly enlarged. The head of

Pontogeneia quinsana is somewhat similar to that

of Gondogeneia and that species cannot be firmly
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

s

FIGURE 18.—Above: Pontogeneia inermis (Krøyer), male "a," 4.4 mm. Below: Tethygeneiaî

quinsana (J. L. Barnard), new combination, female "a," 3.2 mm.
NUMBER 271 43

assigned to Pontogeneia because male calceoli have

never been observed. Most specimens of P. quinsana

lose their antennae in preservative; the few males

observed with antennal remains have lacked calceoli. Nevertheless, P. quinsana is very close to

T. nasa and must be presumed to be a sibling of

that species; quinsana is therefore tentatively placed

in Tethygeneia.

Pontogeneia inermis (Krøyer)

FIGURE 18 (part)

Amphithoe inermis Krøyer, 1838:275-278, pi. 3: fig. 11.

Pontogeneia inermis.—Sars, 1895:451-453, pi. 159.

VOUCHER MATERIAL.—Devon Island, Dundas Harbor, Canada, British Columbia, 23 July 1946, sta.

54, Duvall and Handley, U.S. Fish and Wildlife

Service, USNM 174174, young male "a," 4.4 mm

(illus.).

REMARKS.—Sars' fine portrayal of this species is

supplemented here with drawings of the rostrum,

accessory flagellum, setation of maxilla 2, and the

locking spines and dactyl of pereopods 1-5.

DISTRIBUTION.—Panboreal.

Pontogeneia intermedia Gurjanova

FIGURES 19-21

Pontogeneia intermedia Gurjanova, 1938:333-335, fig. 41;

1951:722, fig. 502.—J. L. Barnard, l969b:llO, figs. 16, 17.

VOUCHER MATERIAL.—Smith 13 (male "a," 5.8

mm, illus.), 14 (female, 7.12 mm). California, Bodega Bay, intertidal, August 1972, collected by Dr.

Ralph I. Smith.

VARIABLES.—The inner plate of maxilla 2 has

3-5 enlarged marginal setae; antenna 1 varies in

length, often on the same specimen, from 30 to 40

percent of the body length; the illustrated male has

the short form of flagellum on antenna 1 but generally the flagellum is 1.6 times as long as that

shown; antenna 2 generally half as long as body;

the cuticle is like that of P. rostrata (Figure 25:

QW)·, the deep spine sheath shown on the dactyl

of pereopod 3 is abnormal; all figures are from the

male; the female is similar except in the presence

of brood plates and absence of penes.

DISTRIBUTION.—Japan Sea; California: Bodega

Bay to Corona del Mar.

Pontogeneia opata, new species

FIGURES 22-24

Pontogeneia minuta.—J. L. Barnard, 1959: 23, pi. 3;

1964a: 106, fig. 2lb,c—Reish and Barnard, 1967:16-17.

[Not Chevreux.]

DIAGNOSIS.—Rostrum large, extending more than

halfway along article 1 of antenna 1, broad dorsoventrally, tapering apicad to rounded point. Accessory flagellum absent but article 3 of antenna 1

with large quadrate extension on mediodistal margin; male calceoli bipartite, weakly bell-shaped.

Article 3 of mandibular palp about 70 percent

length of article 2. Inner lobes of lower lip obsolescent, marked only by weak ridges and swellings.

Inner plate of maxilla 1 with 2 terminal and 1 medioterminal setae; inner plate of maxilla 2 with 2

enlarged medial setae, 1 of these weakly submarginal. Coxae 1-3 each with 1 large posterior spine.

Gnathopods small, subequal in size to each other,

similar between the sexes, palms oblique, posterior

lobe of article 5 on gnathopod 1 short, tapering

conically, lobe on gnathopod 2 extended, blunt,

turned distad partially, extending about halfway

along hand towards palm, occasionally longer.

Pereopods elongate, with setae on both margins of

article 6, distal pair of locking spines equal or subequal to each other in length, smaller than adjacent

spines, smooth, distalmost simple, next proximal

with setular trigger, spines weakly curved, dactyl

smooth marginally, bearing subap¡cal slit and long,

thin setule proximal to slit. Pleonal epimera with

moderate posterior bulge sweeping on straight margin to weak posteroventral tooth on epimera 1-2,

rounded posteroventral corner on epimeron 1, ventral margins of epimera 1-3 with about 2, 5, and 5

spines in tandem respectively on ventral margins,

epimera 1-2 each with lateral ridge. Telson ordinary, apices blunt and naked. Cuticle covered with

conspicuous imbedded spicules resembling thin

fibrous highly elongate crystals, spicules often

nearly straight but especially parabolic on epimera.

DESCRIPTION.—Eyes, when originally preserved,

with dark purple core of pigment concealing all

but outer row of ommatidia, now pigment stronglybleached (23 years of preservation); dimensions of

rostrum variable (J. L. Barnard, 1959, pi. 3); male

antenna 2 much longer than antenna 1 and thus

longer than female antenna 2 (male antenna 2 of
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Pontogene¡a intermedia Gurjanova, male "a," 5.8 mm.
NUMBER 271 45

Gl

FIGURE 20.—Ponlogetieia intermedia Gurjano\a, male "a," 3.8 mm.

Figure 22: A2 drawn to same scale as antennae attached to female head, Figure 22: wC), articles of

flagella of antennae 1-2 on male especially swollen

alternately, weakly so on female antenna 1 but not

female antenna 2, calceoli of male antenna 1 flagellum also alternating in zigzag pattern of attachment

but this pattern weakly developed, calceoli poorly

fingerprinted with striae in comparison to calceoli

of P. rostrata; right lacinia mobilis bifid, with short

accessory cusp; male gnathopods slightly larger and

better developed than gnathopods of female, posterior margins of hands in female with one group

of elongate setae, in male with 2 short spines in

tandem; uropod 3 of female as shown for male but

lacking elongate setae, bearing only short spines;

posteroventral tooth of epimeron 2 occasionally

absent; spines of uropods 1-2 reduced in number

in young adults, outer ramus of uropod 1 with 0-1

dorsal spines, of uropod 2 with no dorsal spine,

inner rami more conservative, generally maintaining normal condition of 2 spines on inner ramus

of uropod 1, 1 spine on inner ramus of uropod 2

(hidden in Figure 24: W).

VARIATION.—Cocos Island male gnathopods 1-2

with posterior margin of hand scarcely notched and

armaments thin as in female.

HOLOTYPE.—AHF 4923, male "a," 4.20 mm

(illus.).

TYPE-LOCALITY.—Upper Newport Bay, California, inter tidal, 28 June 1949, collected by Dr. John

L. Mohr.

VOUCHER MATERIAL.—Type-locality, female "a,"
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Pontogene¡a intermedia Gurjanova, male "a," 5.8 mm.

4.4 mm; female "w," 3.5 mm (illus.); female "p,"

3.8 mm (illus.); male "k," 4.16 mm.

RELATIONSHIP.—Pontogeneia opata may be a sibling of P. longleyi Shoemaker (1933b) from the

Caribbean Sea, but P. opata has a much longer

lobe on article 5 of gnathopod 2, spines on coxae

1-3, a slightly longer article 3 of the mandibular

palp and does not have the weak dorsal protrusions

of pleonites 1-2.

Pontogeneia minuta Chevreux (1908) differs from

P. opata in the shorter lobe of article 5 on gnathopod 2 and the absence of coxal spines.

MATERIAL.—PAZ 2, 25; COCOS 9.

DISTRIBUTION.—California, in bays such as Morro,

Newport, 0 m; Baja California, in Bahía de San

Quintín, 7 m; Gulf of California, Pichilinque Bay

at Las Paz, Bahía Concepción, 0 m; Cocos Island,

Om.
NUMBER 271

47

FIGURE 22.—Pontogeneia opata, new species, holotype, male "a," 4.20 mm (»• = female "w,

3.5 mm).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

wG2o

FIGURE 23.—Pontogeneia opata, new species, holotype, male "a," 4.20 mm (n; — female "w,"

3.5 mm).
NUMBER 271 49

pW

FIGURE 24.—Pontogeneia opata, new species, holotype, male "a," 4.20 mm (p = female "p,"

3.8 mm).

Pontogeneia rostrata Gurjanova

FIGURES 25-27 (part)

Pontogeneia rostrata Gurjanova, 1938:330, 398, fig. 39;

1951:719, fig. 500.—J. L. Barnard, •l962b:8l; 1964a:114-116,

fig. 20; 1969b: 111-114.

VOUCHER MATERIAL.—Smith 16, female "a," 6.2

mm (illus.); male "v," 4.2 mm (illus.); female "w,"

4.6 mm.

REMARKS.—Small (4.0 mm) southern specimens

from Mexico were previously figured by Barnard

(1964a, figure 20); figures of the larger (6.0 mm)

specimens from middle California are here presented but no adult male with calceoli has been

found; therefore, an old collection of males from

southern California has been dissected to obtain a

male antenna 1 and enlargements of calceoli. The

latter conform in gross generalities to those found

on Pontogeneia bartschi Shoemaker by J. L. Barnard (1972a: 186).

The cuticle bears scattered setules, those on

pereonites and pleonites with conspicuous basal

bulbs; the epimera bear blunt scales.

Antennae 1-2 project almost equally, antenna 1

slightly the shorter, the antennae about 50-55 percent as long as the body and head together.

MATERIAL.—Smith 16 (8), 17 (9), 18 (15); California, Bodega Bay, intertidal, August 1972, collected

by Dr. Ralph I. Smith. Barnard 39-K-1 (male)

(see Barnard, 1969b:111).

DISTRIBUTION.—Japan Sea; California from

Bodega Bay to Bahía de San Quintín, Mexico,

0-100 m.

Tethygeneia J. L. Barnard

Tethygeneia nasa (J. L. Barnard),

new combination

FIGURE 27 (part)

Pontogeneia nasa J. L. Barnard, l969a:2OO, figs. 14, 15.

VOUCHER MATERIAL.—SCO 19, male "a," 4.6 mm

(illus.); female "w," 4.6 mm (illus.); TOP 1, male

"v," 3.7 mm (illus.).
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Pontogeneia rostrata Gurjanova, female "a," 6.2 mm (y = male "v," 4.2 mm;

/ = cuticle of pereonite 7).
NUMBER 271 51

Glo

FIGURE 26.—Pontogeneia rostrata Gurjanova, female "a," 6.2 mm {y = male "v," 4.2 mm).
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

w

FIGURE 27.—Above: Tethygeneia nasa (J. L. Barnard), new combination, male "a," 4.6 mm

(y = male "v," 3.7 mm; w = female "w," 4.6 mm). Below: Pontogeneia rostrata Gurjanova,

female "a," 6.2 mm.

KT
NUMBER 271

53

ILLUSTRATIONS.—The figures provided herein supplement and complete the original description of

P. nasa. Epimeron 2 bears the ordinary lateral ridge

of this genus; the cuticle bears scattered, ordinary

long projecting setules, no surficial patterns on

cuticle.

MATERIAL.—SCO 1, 7, 12, 14, 15, 18, 19, 22; KNO

l;TOP l;PAZ6, 18,20.

DISTRIBUTION.—Gulf of California, at Puerto

Peñasco, Bahía de Los Angeles, Bahía Kino, Topolobampo, Isla Espiritu Santo, and Cabo San Lucas,

0-1 m.

Tethygeneia? quinsana (J. L. Barnard),

new combination

FIGURE 18 (part)

Pontogeneia quinsana J. L. Barnard, 1964a: 106-108, fig. 19.

VOUCHER MATERIAL.—SQ 53 (Barnard, 1964:57)

female "a" 3.2 mm (illus.).

REMARKS.—Specimens from an embayment near

La Paz, Baja California, are assigned to this species

despite one strong distinction from T. quinsana as

formerly known only from Bahía San Quintín,

hundreds of kilometers north of La Paz. Specimens

from La Paz have the telson as deeply cleft as in T.

nasa. In other respects the La Paz specimens differ

from T. nasa in the shorter rostrum, poorly protruding anteroventral cephalic corner, broader article 2 of pereopod 5 and straight posterior lobe on

article 5 of gnathopod 2, all characters distinguishing T. quinsana from T. nasa. This second record

of T. quinsana suggests that the species is a sibling

of T. nasa confined to embayments whereas T. nasa

is confined to the open sea.

I believe that the heads in Figure I9j,n in Barnard (1964a) are erroneously attributed to T. quinsana and should be deleted hereafter from the

visual image of the species. The heads as shown in

Figure l9a,b (Barnard, 1964a) are valid.

One specimen of this series from PAZ 3 bears

apices on pereopods 1 and 4, confirming that the

locking spines and dactyls of these pereopods

resemble the same appendages of T. nasa.

MATERIAL.—PAZ 3.

DISTRUBUTION.—Gulf of California, Isla Espiritu

Santo, 0 m; outer Baja California, Bahía de San

Quintín, 0-6 m.

GAMMARIDAE

Anchialella, new genus

TYPE-SPECIES.—Anchialella vulcanella, new species.

ETYMOLOGY.—Anchialella (Greek), little inhabitant of seaside pool.

DIAGNOSIS.—Coxal gills 2-6, ovate, scarcely pedunculate, not 2-articulate. Sternal gills absent. Only

female known, sexual dimorphism unknown, oostegites elongate, evenly slender, with 0-2 apical

setae. Body subvermiform. Head unnotched. All

coxae very short, of similar length. Uropod 3 greatly

exceeding uropod 1, parviramous, outer ram us

1-articulate, peduncle weakly elongate, longer than

urosomite 3 but shorter than rami of uropod 1.

Telson cleft, lobes triangular, pointed, each bearing apicomedial spine. Female gnathopod 1 of

ceradocin form, wrist weakly elongate, palm oblique,

articles 4-6 pubescent. Female gnathopod 2 of

melitin form, enlarged, palm well defined, oblique,

armed with hooked setules, unsculptured, wrist

weakly elongate and lobate, posterior margin of

hand with hadziid-form, setal groups weakly developed, pubescence absent. Mandibular palp article 3

linear, shorter than article 2, bearing 1 D seta, 2 E

setae (Stock, 1974:77). Lower lip with weak inner

lobes marked as creases on outer lobes. Inner plate

of maxilla 1 with 4 apical setae, of maxilla 2

weakly setose medially, lacking oblique facial row

of setae. Outer plate of maxilla 1 with 7 spines.

Dactyl of maxillipedal palp with well-developed

nail. Pleopods biramous. Urosomites free, only urosomite 2 with dorsolateral spine on each side, no

other ornamentation.

RELATIONSHIP.—This genus forms a transitional

grade of morphology between Eriopisa Stebbing

and Galapsiellus J. L. Barnard (1976:422). It differs

from Eriopisa in the absence of article 2 on the

outer ramus of uropod 3, in the elongation of the

peduncle on uropod 3, in the extremely weakened

inner lobes on the lower lip, and in the generally

greater reduction in medial setosity on the maxillae.

Galapsiellus is an apomorph bearing severely

enfeebled female gnathopod 2 with greatly elongate

wrist, an even more elongate peduncle on uropod 3,

turgid telsonic lobes, only 6 spines on the outer

plate of maxilla 1, the absence of a nail on the

dactyl of the maxilliped and uneven spination on
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the outer plate of the maxilliped.

Anchialella appears to be very close to Paraniphargns Tattersall, a genus with 2 freshwater,

probably hypogean, species in Java and the Andaman Islands, but the anterior coxae are short as in

Eriopisa, maxilla 2 retains a few medial setae, the

mandibular lobes of the lower lip are not strongly

extended, and mandibular palp article 2 retains at

least 3 medial setae. Anchialella, therefore, cannot

be descendent from Paraniphargus but instead

appears to be an apomorph of Eriopisa.

Psammoniphargus Ruffo, a monotypic genus

found in hypogean waters of Madagasgar, has no

direct relationship to Anchialella because of several

apomorphic characters, such as weakly fused telesonic lobes, loss of articles on the mandibular palp,

medially unsetose maxilla 2, complete loss of inner

lobes on lower lip, and smaller female gnathopod 2.

The shape and setation of the mandibular palp

are good indications of the relationship of Anchialella to Eriopisa and not to Paraniphargus even

though one may hypothesize an evolutionary step

from Paraniphargus directly to Anchialella in other

attributes.

Anchialella vulcanella, new species

FIGURES 28, 29 (part)

DESCRIPTION.—As in diagnosis of the genus. See

illustrations.

NOTES.—Pereopod 4 slightly smaller than pereopod 3. Telson of holotype aberrant, one side apparently damaged, shortened and missing main

spine, only other known specimen, juvenile, 2.10

mm, with somewhat more normal telson, though

one side larger than other side. Juvenile with only

1 basofacial spine on uropod 1 at Mark 44.

HOLOTYPE.—USNM 169018, female "u," 4.06 mm

(illus.).

TYPE-LOCALITY.—GAL 103, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 23 January 1964,

intertidal, mangrove tidepool 305 m from sea near

lower bodega, anchialine.

VOUCHER MATERIAL.—Type-locality, juvenile "j, "

2.10 mm (illus.).

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz, anchialine.

ETYMOLOGY.—The epithet vulcanella is from

Latin meaning "fire."

Ceradocus Costa

Ceradocus paucidentatus J. L. Barnard

FIGURE 6 (part)

Ceradocus paucidentatus J. L. Barnard, 1952b:5.õ-58, pis.

11-13.

REMARKS.—This is the second known collection

of this elusive species originally described from the

region of Punta Eugenia, Baja California.

A figure of the juvenile gnathopod 2 is shown to

represent juvenile or female-like conditions. Heretofore only males were known. Ep¡meron 2 bears 1

spine, epimeron 3 bears 3 ventral spines, in each

case the spines being much enlarged in comparison

with adult males. Possibly these spines do not increase in size with age, as they are almost as large

as those of males near 8 mm in length.

MATERIAL.—PAZ 13, juvenile, 3.74 mm (illus).

DISTRIBUTION.—Near Punta Eugenia, outer Baja

California and Bahía San Evaristo, Gulf of California, intertidal.

Didzura J. L. Barnard

Dulzura gal, new species

FIGURE 34 (part)

DIAGNOSIS.—Article 2 of pereopods 5-7 with only

1-2 serrations at posteroventral corners. Apicolateral spine on peduncle of uropod 1 extending to

Mark 60 of outer ramus. At least 2 spines on each

lobe of telson as long as telson.

DESCRIPTION.—Other appendages and mouthparts

generally similar to the type-species of the genus,

Dulzura gal J. L. Barnard (1969b) except peduncle

of uropod 1 with 5 small dorsolateral spines, of

uropod 2 with 4 dorsolateral spines, apicalmost

spine unpaired; urosomite 1 with 1 spine-seta dorsolaterally on each side, urosomite 2 with 1 spine and

1 setule on each side, urosomite 3 with 1 spine on

each side at base of telson; rami of uropods 1-2,

dorsal spines consist of uropod 1 outer 1, inner 3,

uropod 2 outer 3, inner 4; epimeron 1 lacking

ventral spines, epimeron 2 with 2 ventral spines,

epimeron 3 with 4 ventral spines; mandibular palp

article 2 with only 1 apical seta, 1 inner basad

setule; palm of gnathopod 1 slightly oblique.
NUMBER 271

55

FIGURE 28.—Anchialella vulcanella, new species, holotypc, female "u," 4.06 mm (ƒ = juvenile "j ,

2.10 mm).
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VM

FIGURE 29.—Above: Anchialella irulcanella, new species, holotype, female "u," 4.06 mm. Below:

Maera chinarra, new species, holotype, male "a," 3.7 mm (ƒ = female "f," 3.4 mm; j = juvenile

"j, " 2.9 mm; p = male "p," 3.6 mm).
NUMBER 271

57

HOLOTYPE.—USNM 169019, male "w," 3.00 mm

(illus.). Unique.

TYPE-LOCALITY.—GAL 108, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 24 January 1964,

offshore of lower bodega in shallow water, wash

from 3 rocks.

REMARKS.—This species differs from the typespecies in the greater length of both the telsonic

spines and the main spine of uropod 1 and in the

fewer posteroventral serrations on article 2 of

pereopods 3-5.

Dulzura gal is so similar to Dulzura hamakua

(J. L. Barnard, 1970, formerly in Eriopisa) that the

two species may necessarily be reduced to subspecific

level when more can be learned of speciation in the

genus. During the interim, the two species can be

distinguished by the great length of spines on uropod 1 and the telson of D. gal.

MATERIAL.—GAL 108.

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz, 1 m.

Elasmopus Costa

Several subspecies and phenotypes of Elasmopus

rapax Costa from California and Mexico have been

described by J. L. Barnard (1962b, 1969b). A larger

quantity of widely distributed material is now

available so that replication of identifications is

possible and certain limits to variability can be

seen. I conclude that all forms, phenotypes and

subspecies defined in the key on pages 118-119 of

J. L. Barnard (1969b) should be given full specific

names. The so-called formae are now shown to be

widely distributed sympatric populations minutely

distinct from one another. Characters of recognition are confined mainly to terminal males, though

subadult males and females can usually be distinguished from one another by minute characters

of epimeron 3 or telson. Juveniles of the several

species appear very similar to each other but I have

not studied the life history of each species intensively and thus have not discriminated juveniles

adequately.

Because of the consistency of certain conservative

characters, these species of Elasmopus from California and Pacific Mexico appear to have a common ancestry. For the purposes of this discussion

the common ancestor is stated to be E. rapax Costa,

a species widely distributed throughout the world

and apparently living also in the California-Mexico

area. There is room for doubt, however, in this

contention, as the so-called Elasmopus rapax of

western America can be distinguished minutely

from specimens reported in the type-area of Europe

in the same way that American species can be distinguished from the local representative known as

E. rapax. These minute distinctions have generally

been ignored throughout the world in the various

identifications of E. rapax. Such minutiae must

now be recognized in light of the adaptive radiation found in eastern Pacific elasmopuses.

My approach is to consider as E. rapax those

specimens closest to the European type in terms of

male gnathopod 2 and to consider this founder

taxon as a race of E. rapax from which have developed the several descendent species.

These observations confound the taxonomy of

this genus throughout the world as it becomes apparent that several identifications in the earlier

literature, including papers by me, are erroneous.

For example, the species identified as E. rapax from

Hawaii by J. L. Barnard (1970:131) is probably

synonymous with E. bampo, new species, from

Mexico, so that true E. rapax appears not to have

been collected as yet in Hawaii, if indeed it exists

there at all. If the cluster of species in Mexico is

truly a tightly radiated group, in this Hawaiian

case, we may be observing one of the descendants of

E. rapax having already migrated to Hawaii.

One of the descendent species of E. rapax in

Mexico has gnathopod 2 of the male in form similar to that of E. pocillimanus (Bate), another European species identified throughout the Pacific

Ocean. This remarkable convergence suggests the

probability that the Pacific E. pocillimanus may

actually be distinguishable from the European E.

pocillimanus because the Pacific species has highly

characteristic locking spines on the pereopods in

contrast to the European species. The Mexican

species, E. tiburoni, appears closer to the European

E. pocillimanus than does the Hawaiian taxon

identified by J. L. Barnard (1970:130) as E. pocillimanus. Nevertheless, the Mexican species can be

distinguished from European E. pocillimanus by

other minute characteristics.

The European and eastern Pacific evolutes in

Elasmopus are probably secondary and are derived

from species endemic to the heart of the IndoPacifk fauna, as yet poorly studied. Virtually noth-
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ing is known of variability correlated with biogeographic increments. We therefore have few bases

on which to erect a taxonomic hierarchy in this

highly diverse genus.

Specific Characters in Elasmopus

The main taxonomic characters in Elasmopus

are confined to a few criteria in contrast to other

diverse amphipodan genera, such as Pamdexamine

(J. L. Barnard, 1972a) where virtually every part

of the external body structure bears significant

taxonomic characters. In Elasmopus these are limited primarily to male gnathopod 2, epimeron 3,

telson, pereopods, article 3 of the mandibular palp,

and the eyes. Few taxonomic characters have been

reported on mouthparts other than article 3 of the

mandibular palp, or on the coxae, uropods 1-2,

gnathopod 1, antennae, and epimera 1-2. Now

that most carinate species of Elasmopus have been

removed to Mallacoota J. L. Barnard (l972a:243)

the ornaments of the urosome are found in only

a few species in the western Pacific Ocean. The

antennae differ among species in setosity and

lengths of articles or flagella but these tenuosities

are poor taxonomic devices. Uropod 3 on occasion

has variables of taxonomic use but not as significantly as believed by J. L. Barnard (1969b: 118).

Gnathopod 2 of the terminal male is of highest

taxonomic value, but this usefulness is obviously

confined to a few specimens of any collection. The

palm and posterior margin of the hand are generally confluent but often are distinguished from

each other by a defining tooth. Near the dactylar

hinge most species of Elasmopus bear a spinose

hump and, slightly proximal to that, a naked tooth.

Setae are often sparse and evenly distributed along

the palm and posterior margin of the hand. This

description fits E. rapax of Europe. Other species

often differ in one of the following modifications:

the loss of spines on the main hump or its total

loss, the loss or reduction of the naked tooth or the

defining tooth, or the reduction of the defining

tooth to a ridge bearing a spine, or the anomalous

development of setal patterns, the palm naked and

the posterior hand margin heavily setose, or setae

developed so densely as to obscure the palm. In

several species a fourth palmar tooth forms a companion to the spinose hump, whereas the ordinary

tooth proximal to the hump is generally attached

more medially than the fourth companion tooth.

The apices of pereopods 1-5 provide identification characteristics among many species, but in the

Mexican rapax group these apices are similar

among the several species and afford no help in

identifying juveniles. In the simplest condition the

pair of apical locking spines is short, thin, almost

straight and poorly striate; in many species one or

both of these are enlarged or heavily striate, flabellate or curved, occasionally truncate or claviform.

The pereopodal dactyls in the simplest cases have

smooth inner margins and a thin or tiny subapical

setule. In other species the setule may be grossly

enlarged, even spine-like, the inner margin may

have minute notches or grossly extended castellat¡ons and an internal sclerotic slit is found near

the subapical setule. The Mexican rapax group

has the simplest case of locking spines and dactyl.

The Mexican rapax group is characterized by

long posterior setae on article 2 of pereopods 3-5

but E. rapax from Europe and many other species

of Elasmopus lack those setae.

Pleonal epimeron 3 varies among the species but

often so insignificantly as to defy identification.

Elasmopus rapax from Europe has a straight posterior margin bearing several small denticles or

notches evenly projecting, the dorsal notches often

bearing setules. The denticle-notches are highly reduced in the northern European populations of

E. rapax. Asymmetry occurs in the denticles of

many other species of Elasmopus, the ventralmost

denticle being enlarged as a tooth, the posterior

margin becoming bulbous to form a weak S-shaped

curve, the denticles dorsal to the main tooth often

being lost, or the main tooth often being marked

by a small sinuosity just dorsad. Juveniles of the

Mexican E. rapax group generally have the latter

condition. The denticle pattern is generally similar between fully terminal males and females but

subadults usually have fewer denticles, smaller

denticles, or smooth margins in varying degrees in

the several species.

The telson of species in the E. rapax group everywhere in the world appears to be characteristic.

The apices of each lobe are protuberant and naked

but are defined laterally by a subtendant notch or

incision, itself defined laterally by a sharp protrusion and largely covered with one or more spines

inserted dorsally just proximal to the invagination.
NUMBER 271 59

No spines are fully apical nor generally medial.

Spine number varies in terminal adults among the

species, and spine length may vary between the

sexes of a single species; for example, the females of

the Mexican E. rapax group have highly elongate

telsonic spines whereas males have extraordinarily

short spines. Other species groups of Elasmopus

have apically truncate telsonic lobes bearing fully

apical spines often extending fully mediad.

Mandibular palp article 3 in Elasmopus is falcate

and medially pectonosetiferous. The degrees of

falcateness and stoutness of the article vary among

the species, but this shape is useful to the taxonomist mainly as a clue to differentiating subadults

of otherwise similar morphology and then only if

the adult palp is grossly distinct. Many species have

such similar palps that discriminations are difficult.

Ocular structure has been useful in rectifying

identification errors where gross ocular distinctions

can be made but these differences are not widespread. For example, J. L. Barnard (1970:115) was

able to distinguish E. calliactis from other similar

species in Hawaii mainly on the presence of red

ocular pigment in contrast to black and purple in

other species. The so-called E. pocillimanus of

Hawaii is easily recognized by the slight enlargement but extreme blackness of the eyes. This kind

of character loses its usefulness in all but the most

freshly preserved of materials because of the slow

deterioration of ocular pigment in various preservatives.

A distinction in the length of and degree of

spinosity between the rami of uropod 3 has limited

usefulness; J. L. Barnard (1969b: 118-119) used this

group of characters for distinction among several

members of eastern Pacific Elasmopus and suggested

that equality in rami marked the E. rapax group,

but this is now seen to be fallacious. Females and

juveniles of the Mexican E. rapax have a shortened

inner ramus of uropod 3, but in terminal males

and many terminal females this ramus enlarges to

match the outer ramus.

Clearly, the genus Elasmopus will be a major

problem for systematists until a monographer can

study a wide diversity of materials and until some

understanding of phenotypy can be obtained. My

treatment here is far from satisfactory but follows

the reasoning that the operable taxonomic units,

recognized by their replicability in several samples,

should be named as species. Because these units are,

as yet, recognizable only in a terminal male stage,

the material available is reduced to a small amount

of the total specimens in the genus, at least in the

E. rapax complex which dominates the eastern

Pacific Ocean. The literature contains other examples of this complex not yet integrated into the

system, such as Elasmopus pseudaffinis Schellenberg

(J. L. Barnard, l965a:5Ol), the so-called E. rapax

of Micronesia (J. L. Barnard, 1965a: 503), perhaps

E. hooheno J. L. Barnard and E. piikoi (J. L. Barnard, 1970:121, 127) and others. To make the system somewhat more understandable I have erected

new names for various taxa encountered in this

study and have raised to specific level other names

I had created earlier. Such OTUs as the Elasmopus

rapax of Bahía de Los Angeles (J. L. Barnard,

1962b: fig. 16 and J. L. Barnard, 1969b: 119, key)

are left nameless until more replicability can be

demonstrated. I cannot find distinctions between the

E. rapax of J. L. Barnard (1970:131, Hawaii) and

the new species, E. bampo, created herein from the

Gulf of California, except in telsonic spine lengths,

which themselves are considered to be variable and

were found to be so in the E. rapax of Alamitos

Bay, California by J. L. Barnard (l962b:94). One

might presume that the Alamitos Bay form of

E. bampo, residing in a bay and probably at the

northern margin of the species range, would exhibit

abnormalities.

Presumably the typical E. rapax Costa is represented in Chevreux and Fage (1925:244) because the

type-locality resides in Taranto, Italy, near Mediterranean France. Chevreux and Fage depict the

species as lacking long setae on article 2 of pereopods 3-5. The telson has on each lobe only 2 spines

strongly constrained to a small lateral notch. Male

gnathopod 2 has 3 palmar teeth, the hinge tooth

being simple, the middle tooth near but distinctly

separate from the hinge tooth, the defining tooth

small but distinct, the dactyl reaching beyond the

halfway mark between teeth 2 and 3; the palm is

poorly setose and the posterior margin of the hand

moderately setose. Mandibular palp article 3 is

elongate. The rami of male uropod 3 extend

equally. Epimeron 3 is multiserrate densely, with

a distinct tooth at the posteroventral corner.

Sars (1895, pi. 183) depicts this species from

Norway with a much broader spread of and more

numerous spines on the telson, especially in the

male. Epimeron 3 is poorly serrate. The inner
fífl SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ramus of uropod 3 is slightly shortened. The hinge in the Pacific Ocean many species are based on this

tooth on the palm of male gnathopod 2 is rounded kind of small difference. The problem to be faced

and not sharp as in the Mediterranean form. These is whether or not such variation in the eastern

minor character alternatives would appear to be Atlantic is phenotypy within a single species and

variations normally attributed to phenotypy but in the Pacific is a mark of speciation.

Key to the Males of the Species of Eastern Pacific Elasmopus and the E. rapax

Complex of Hawaii

1. Telson with medial part of each lobe protruding, rounded or subtruncate and free of spines,

spinose laterally from Mark 63 to Mark 100, rarely from Mark 40 2

Telson with truncate or slightly concave apices spinose laterally at least from Mark 50 and

usually from Mark 25 or Mark 0 9

2. Epimeron 3 lacking tooth at posteroventral corner 3

Epimeron 3 bearing tooth at posteroventral corner 4

3. Proximal palmar tooth on male gnathopod 2 well developed, palm heavily setose

E. ocor<mi, new species

Proximal palmar tooth on male gnathopod 2 vestigial, palm almost devoid of setae

E. mutattu

4. Male gnathopod 2 palm with only 2 processes, one near dactylar base, one defining palm,

hollow formed on medial face of hand E. tiburoni, new species

Male gnathopod 2 palm with 3-4 processes, no hollow on medial face of hand 5

5. Male gnathopod 2 palm with 4 distinct processes, epimeron 3 posterodistally serrate, teeth

sharp (includes part of E. rapax of Alamitos Bay) 6

Male gnathopod 2 palm with 3 processes (fourth rudimentary), epimeron 3 posterodistally

smooth or with minute notches 7

6. One or more telsonic spines half as long as telson (inner ramus of uropod 3 in male

shortened) (includes part of E. rapax of Alamitos Bay) E. bampo, new species

No telsonic spine more than one-fourth as long as telson (inner ramus of uropod 3 in male

not shortened) (includes part of E. rapax of Alamitos Bay and through E. rapax of Hawaii)

E. bampo, new species

7. Male gnathopod 2 palmar process near dactylar base bifid or trifid (includes E. rapax form I

of Bahía de Los Angeles, J. L. Barnard, l969b:ll9) £. ?rapax from eastern Pacific

Male gnathopod 2 palmar process near dactylar base simple 8

8. Middle palmar tooth on male gnathopod 2 immediately adjacent to hinge process, dactyl

of gnathopod 2 strongly shortened, article 2 of pereopod 5 strongly setose

E. mayo, new species

Middle palmar tooth on male gnathopod 2 slightly removed from hinge process, dactyl of

gnathopod 2 scarcely shortened, article 2 of pereopod 5 naked

typical E. rapax from Mediterranean

9. Male gnathopod 2 hand with large medial hollow E. tiburoni, new species

Male gnathopod 2 hand lacking medial hollow 10

10. Male pereopod 5 article 2 with deep castellations 11

Male pereopod 5 article 2 with weak serrations IS

11. Male gnathopod 2 article 6 with anterodistal stridulation hump, palm lacking middle tooth,

(epimeron 3 with small posteroventral tooth) E. tubar, new species

Male gnathopod 2 article 6 lacking stridulation hump, palm bearing middle tooth (epimeron

3 variable) 12

12. Pleonal epimeron 3 with slightly convex posterior margin, rounded-quadrate at posteroventral corner, sparsely serrate posteriorly; pereopodal dactyls strongly constricted; middle

process on palm of male gnathopod 2 short, broad, truncate E. serricatus, new status

Pleonal epimeron 3 of the "antennatus" morphology, posterior margin slightly convex,

smooth, with medium-sized posteroventral tooth; pereopodal dactyls moderately constricted; middle palmar process on palm of male gnathopod 2 very long and pointed

E. rapax form II of Bahía de Los Angeles
NUMBER 271 61

13. Midpalmar process of male gnathopod 2 absent, defining tooth absent 14

Midpalmar process(es) of male gnathopod 2 present, defining tooth present 15

14. Telson cleft more than halfway, epimeron 3 with posterior margin oblique and bearing

distal tooth £. antennatus

Telson cleft halfway or less, epimeron 3 with vertical posterior margin lacking tooth

E. ìecuadorensis

15. Eyes red in life, bleaching clear in alcohol in 2-3 weeks E. calliactis

Eyes with dark pigment not bleaching in alcohol in 2-3 weeks 16

16. Palm of male gnathopod 2 with 2 midprocesses, each conical, plus facial ridge and spine,

epimeron 3 with small sharp posteroventral tooth E. tetnori, new species

Palm of male gnathopod 2 with 1 midprocess, variable, epimeron 3 variable 17

17. Midprocess of male gnathopod 2 an asymmetrical cone, epimeron 3 with extended and

upturned posteroventral corner E. holgurus

Midprocess of male gnathopod 2 broad, truncate, ep¡meron 3 with small sharp posteroventral tooth E. zoanthidea, new species

Elasmopus antennatus (Stout)

Neogammaropsis antennatus Stout, 1913:645-646.

Elasmopus antennatus.—J. L. Barnard, 1962b:88-91, figs. 12,

13; l964b:222; 1969b: 115-117.

MATERIAL.—PAZ 21.

DISTRIBUTION.—California and Baja California

from Carmel to Cabo San Lucas, 0-11 m.

Elasmopus bampo, new species

FIGURES 30, 31

Elasmopus rapax of Alamitos Bay.—J. L. Barnard, 1962b,

fig. 17; 1969b: 119. [In part.]

7Elasmopus rapax of Hawaii.—J. L. Barnard, 1970:131-135,

figs. 79, 80.

DIAGNOSIS.—Eyes ordinary. Mandibular palp article 3 deeply falcate. Palm of male gnathopod 2 with

ordinary sp¡nose hump near dactylar hinge, then

supernumerary (4th) sharp enlarged tooth on same

plane, naked apically, rounded mediofacial tooth,

palm defined by tapering medial tooth, palm and

posterior margin of hand evenly and sparsely setose.

Article 2 of pereopods 3-5 with long posterior setae

in male only; locking spines of pereopods 1-5

ordinary, inner margin of dactyl smooth but with

minute shagreen proximally, main subapical setule

thick but apically flexible. Epimera 2-3 with long

ventral setae, epimera 1-3 with short ventral spines;

epimeron 3 with nearly straight posterior margin

bearing 5 or more sharp denticles becoming blunter

dorsally, each sinus armed with setule except

ventralmost, posteroventral tooth not larger than

other denticles and its guarding sinus not enlarged

greater than other sinuses. Rami of uropod 3

equally long in male, inner ramus shortened in

females and juveniles; inner ramus with several

stout basomedial spines. Telson with naked protrusive apices placed mediad, laterally defined by

sinus and sharp tooth, spines submarginal from

sinuosity and forming row of about 3 laterally;

some spines very long in female, all spines short in

male.

DESCRIPTION.—Female lacking long setae of article 2 on pereopods 3-5, small females and large

juveniles lacking serrations of epimeron 3 except

for ventralmost sinus and tooth; juvenile males with

fourth tooth of palm on gnathopod 2 much smaller

than in adults, small juvenile males apparently with

gnathopod 2 as in E. rapax and therefore indistinguishable from that species.

Following parts as shown for E. rapax (Figures

35-37) head, antennae, accessory flagellum, eye,

mouthparts (except for parts of mandibles illustrated), pereopods 1-5 (except 1 slightly anomalous

pereopod 5 illustrated), dactyls, gnathopod 1 except

article 4 as illustrated, urosome, uropods 1-2, female

gnathopod 2 except proportions as outlined in

figures, female uropod 3, coxae, cuticle, medial view

of article 1 on antenna 1 except for several additional ventral setae.

VARIANT.—Subadult males from PAZ 9 with long

spines on telson.

HOLOTYPE.—USNM 149390, male "a," 6.88 mm

(illus.).

TYPE-LOCALITY.—TOP 3, Topolobampo, Mexico,

25 November 1971, 1.6 km seaward of town, 1 m,

rock wash.
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

w

FIGURE 30.—Elasmo¡>us bampo, new species, holotype, male "a," 6.88 mm (p = male "p," 6.2 mm;

u = female "u," 6.56 mm).
NUMBER 271

63

PW

FIGURE 31.—Elasmopus bampo, new species, male "p," 6.2 mm (w = female "w," 6.3 mm).
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VOUCHER MATERIAL.—Type-locality, female "u,"

6.56 mm (illus.); PAZ 9, male "p," 6.2 mm (illus.),

female "w," 6.3 mm (illus.).

RELATIONSHIP.—This species differs from Mexican

E. rapax in the presence of a fourth tooth on the

hand of male gnathopod 2 and in the even serrations of epimeron 3 in terminal adults.

The Elasmopxis rapax described from Alamitos

Bay, California, by Barnard (I962b:94, 1969b: 119)

appears to belong with E. bampo. However, the

telsonic spines of the juveniles are not as long as

those of typical E. bampo and adults were said to

have the spines reduced to the size one sees in the

E. rapax of Hawaii described by J. L. Barnard

(1970:131). One must place the Hawaiian taxon

provisionally in E. bampo until the nature of telsonic spines can be elucidated; they would form

an excellent subspecific or specific distinction between the two widely separated regions were it not

for the variability encountered in Alamitos Bay,

California.

MATERIAL.—SCO 16; KNO 1; TOP 2, 3; PAZ 5, 9.

DISTRIBUTION.—Gulf of California: Puerto Peñasco, Bahía Kino, Isla Espiritu Santo and Isla Partida, intertidal; California, bay form in Alamitos

Bay; ?Hawaii, as a subspecies.

Elasmopus ?ecuadorensis Schellenberg

FIGURES 32, 33 (part)

[?]Elasmopus ecuadorensis Schellenberg, 1936:153-154, figf 1

[see identification below].

IDENTIFICATION.—These are the only specimens

found in the Galapagan collections matching Schellenberg's description and illustrations in the condition of epimeron 3, uropod 3, mandibular palp,

pereopod 5, accessory flagellum, lateral aspect of

gnathopod 2 and generally in the telson. The cleft

on the telson in the male is very slightly greater

than shown by Schellenberg but approximates that

found in the female.

DIAGNOSIS.—Eyes ordinary. Mandibular palp

deeply falcate. Palm of male gnathopod 2 with

rounded and poorly spinose hump near dactylar

hinge, no middle tooth, weak mediofacial ridge on

hand near apex of dactyl, medial face with line of

dense setal rows axially, palm and posterior margin

of hand densely setose throughout. Article 2 of

pereopods 3-5 lacking long posterior setae, article 2

of pereopod 5 normally serrate; locking spines of

pereopods 3-5 ordinary, thin, straight, weakly striate

longitudinally, inner locking spine on pereopods

1-2 also normal, but striate, outer locking spine

enlarged, curved, striate, proximal face of dactyl

striate or with fine prickles, main subapical setule

very thick, short, scarcely flexible apically. Epimera

bearing only short ventral spines, epimeron 3

weakly convex to straight posteriorly, sparsely

notched, posteroventral corner rounded. Uropod 3

of male with shortened inner ramus bearing 1

medial spine. Telson with truncate apices each

armed with 3-4 spines, three spines always elongate,

lateral spine most elongate, fourth spine when

present medial and very short.

DESCRIPTION.—Generally like E. antennatus

(Stout) (J. L. Barnard, 1962b) but herein compared

to Figures 35-37 of E. rapax for expediency. Eyes

stained brownish magenta; antennae stout, flagellum

of antenna 1 especially stout (abnormally shortened

on head view, see Figure 52:Alm of normal length),

articles 2-3 with ventral setae weakly elongate,

accessory flagellum 2-articulate; mandibles like

rapax except palp like E. tubar, article 3 short, very

stout, each mandible with 4 rakers, article 2 of

palp with inner setal formula of 1-2-1, first seta

highly basal; upper lip, lower lip like E. rapax,

maxilla 1 with thin palp, spines on outer plate with

more numerous and longer cusps, outer plate of

maxilla 2 thin, dactyl of maxilliped shorter and

nail longer than in rapax. Gnathopod 1 like rapax

but article 4 evenly rounded apically (details of

hand on Figure 32: Glo). Anterior margin of coxa 1

straight, coxa 2 broader, coxa 3 slightly tapering

and coxa 4 with posterior excavation shallower and

tooth blunter than in rapax, long ventral setae of

coxae 1-4 in male "a" =5-6-6-6. Pereopods 1-2

slightly stouter than in rapax, article 4 with only 2

posterior bundles of setae, 3 anterior sets of spines,

article 5 with one set of midanterior setae and one

set apically, posterior margin with 5-7 spines, in

larger specimens 2 spines paired in basal set, article

6 with only 5 posterior spine positions besides locking set, some middle sets with 2-3 spines (pereopods

3-5, epimera, uropods and telson on Figures 32

and 33). Males with ventral spines on epimeron 3

largely absent in posterior third, in largest male

"v" 7.52 mm (not illustrated), epimeron 1 with

pair of ventral spines, epimeron 2 with 2-1 (anterior

to posterior), epimeron 3 with 1-1-1-1, male "a"
NUMBER 271 65

FIÍ.URE 32.—Elasinopus ?ecuadorensis Schellenberg, male "a," 6.35 mm (w = female "w," 7.29 mm).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 33.—Above: Eiasmopus ?ecuadorensis Schellenberg, male "a," 6.35 mm. Below: Eiasmopus

mayo, new species, holotype, male "a," 8.72 mm (w = female "w," 7.75 mm).
NUMBER 271 67

with formulae of 1, 2-1, and 1-1-2-1. Most specimens lacking abnormal gap in peduncular spines

of uropod 1 (in contrast to Figure 32: W); uropod 3

especially small.

FEMALE.—Like male but gnathopod 2 smaller,

medial face of hand immensely setose, bearing

mediofacial spine on hand near apex of dactyl;

spines on epimeron 3 proceeding fully posteriad,

formula for epimera 1-3 of female "w" = 2, 1-2-1

and 1-1-2-2-1-1.

ILLUSTRATIONS.—Flagellum on antenna 1 of male

head abnormal, see normal flagellum in Figure

$2:Alm; telsonic apices of normal adult males and

females are like Figure 32:wT, but adult males

have a slightly deeper cleft; note abnormal spination and apical shapes for male "a" telson (Figure

33: T, above).

TYPE-LOCALITY.—Galapagos Islands, Albemarle

(Schellenberg, 1936:153).

VOUCHER MATERIAL.—DAW 2, male "a," 6.35 mm

(illus.); female "w," 7.29 mm (illus.); male "v,"

7.52 mm.

RELATIONSHIP.—This species resembles E. antennatus (Stout) but differs in the quadrate epimeron 3

lacking a posteroventral tooth. Schellenberg (1938a:

54) was undoubtedly correct that E. ecuadorensis

hawaiensis from the Hawaiian Islands (J. L. Barnard, 1970:119) is closely similar to the Galapagan

taxon but the redescription of E. e. ecuadorensis

herein suggests that the differences between the

two are sufficient to elevate E. hawaiensis to full

specific level. Elasmopus hawaiensis has a deeper

telsonic cleft, fewer apical spines on the telson,

more strongly excavate telsonic apices with more

protrusive middle limbs. If terminal males were

indeed described by J. L. Barnard (1970:119) then

E. hawaiensis has a much more poorly setose male

gnathopod 2 than does E. ecuadorensis.

MATERIAL.—DAW 2.

DISTRIBUTION.—Galapagos Islands, intertidal.

Elasmopus hawaiensis Schellenberg, new status

Elasmopus ecuadorensis hawaiensis Schellenberg, 1938a:54-55,

fig. 27.—J. L. Barnard, 1970:119-121, fig. 69.

Schellenberg's subspecies is here elevated to full

specific status.

Elasmopus mayo, new species

FIGURE 33 (part)

Elasmopus rapax form I of Tiburon Island—J. L. Barnard,

1969b: 119 [part].

DIAGNOSIS.—Eyes ordinary. Mandibular palp article 3 deeply falcate and stout as in E. bampo.

Palm of male gnathopod 2 with ordinary but narrowed spinose hump near dactylar hinge, next tooth

naked, subcolumnar, closely proximal to main

hump, palm defined by strong conical tooth, palm

and posterior margin of hand poorly setose, dactyl

reaching only part way along palm. Article 2 of

pereopods 3-5 with posterior setae in male only,

locking spines of pereopods generally ordinary but

largest spine apically curved on pereopods 1-2,

inner margin of dactyl smooth, main subapical

setule thick but apically flexible. Epimera 1-3

similar to those of E. rapax but long setae on

epimera 1-2 absent, epimeron 3 with nearly straight

posterior margin armed with 1-2 notches bearing

setules, posteroventral tooth small in female, nearly

obsolescent in male, sinus above tooth naked. Rami

of uropod 3 equally long in male, inner ramus with

3 basomedial spines, inner ramus shortened in

females. Telson with naked protrusive apices placed

mediad, laterally defined by sinus and medium

sized tooth, with 1 or 2 spines on each lobe in male

(second spine developing for next instar seen on 1

male), female with 3 spines, 1 spine much longer

than spines of male.

DESCRIPTION.—Following parts as shown for E.

rapax (Figures 35-37): head, lateral antenna 1, accessory flagellum (3-articulate), antenna 2, upper lip,

right mandible (but lacinia mobilis illustrated), left

mandible, lower lip, maxilliped, gnathopod 1, general appearance and spination of pereopods 1-4 (but

posterior setae of article 2 on pereopods 3-4 fewer),

coxae 1-3, urosome, uropods 1-2 (but spines on

peduncle of uropod 2 fewer), uropod 3 of both sexes

(but outer ramus with only 2 lateral spine groups

and peduncle lacking basolateral spines); female

gnathopod 2 (but article 4 with sharp posterodistal

point). Following parts as described or in Figure 33:

medial aspect of antenna 1 but proximal spine

group with only 1 spine; inner plate of maxilla 1

with 2 large and 3 or 4 small setae, palp with 8

spines and 6 or 7 setae; inner plate of maxilla 2

with only 1 medial seta; coxa 4 illustrated; formula
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of spines on epimera 1-3, front to rear, respectively:

l_2; 1-4-2-3; 1-1-1-2-3-2 (with 1 seta in last 2

positions on epimeron 3); pereopod 5 of male with

much narrower article 2 than in E. rapax (illustrated), the figure composed on a medial view for

articles 1-5 and lateral view for articles 6-7.

HOLOTYPE.—USNM 142469, male "a," 8.72 mm

(illus.)

TYPE-LOCALITY.—Bahía San Carlos, Mexico (near

Guaymas), 24 January 1971, intertidal, collected by

R. C. Brusca.

VOUCHER MATERIAL.—Type-locality, female "w,"

7.75 mm (illus.).

RELATIONSHIP.—This species differs from E. rapax

and E. bampo in the presence of only 3 palmar teeth

on male gnathopod 2; the fourth tooth as seen in

E. bampo can be visualized rudimentarily on E.

mayo. The latter and E. tiburoni were mixed together in my key (Barnard, 1969b: 119), as "Elasmopus rapax form I of Tiburon Island, couplet IC"

as I assumed that juveniles of E. mayo belonged

with what is now named E. tiburoni.

MATERIAL.—GAL 102, 107, 108, 109, 113, 114,

115, 116; DAW 1, 3, 19, 23, 27, 31, 32, 33, 35, 40;

ECU 2; BRU 1.

DISTRIBUTION.—Gulf of California, Bahía San

Carlos; Galapagos Islands; Ecuador; intertidal.

The distribution of this species is peculiar. It has

been found abundantly in Galapagan samples but

in only one sample from the Gulf of California.

Elasmopus ocoroni, new species

FIGURE 34 (part)

DIAGNOSIS.—Eyes ordinary. Mandibular palp

deeply falcate. Palm of male gnathopod 2 with

sharp falcate spinose hump near dactylar hinge,

palm short, middle tooth stout, blunt, palm defined

by broad conical tooth bearing spine, palm and

posterior margin of hand strongly setose throughout, setae highly elongate, medial face of hand

poorly setose. Article 2 of pereopods 3-5 lacking

long posterior setae, article 2 of pereopods 4-5

normally serrate; locking spines of pereopods 3-5

ordinary, thin, straight, weakly striate longitudinally, inner locking spine on pereopods 1-2 enlarged, curved, blade-like, face of dactyls on pereopods 1-2 with faint shagreen, margins unnotched

on pereopods 1-5, main subapical setule thick but

apically flexible. Epimera bearing only short and

thin ventral spines, epimeron 3 weakly convex posteriorly, sparsely notched, postoventral corner

rounded-quadrate. Uropod 3 of male with shortened inner ramus bearing 1 medial spine. Telson

with apices rounded and protruding medially, each

lateral acclivity broad, shallow, bearing 3 spines

grading from short to medium from medial to

lateral.

DESCRIPTION.—Generally like E. mutatus (J. L.

Barnard, l962b:96, fig. 18); eyes slightly enlarged,

ommatidia all stained burgundy, no pigment core,

antennae very similar, including elongate article 3

of antenna 1, article 2 with 2 pairs of very short

ventral setae, article 3 with several pairs of longer

setae, accessory flagellum 2-articulate; mouthparts

as shown for E. rapax (Figure 35) but mandibular

palp article 3 slightly less attenuate, mandibular

lobes of lower lip blunter, palp of maxilla 1 only

as wide as outer plate, latter with medial tooth formula of spines 2-5 = 2, 3 small, 3 large, 2 small;

outer plate of maxilla 2 much thinner; nail of

maxillipedal dactyl slightly elongate; palm of

gnathopod 1 with 3 defining spines, medial comb

of article 4 shortened, articular outline distinctive

(Figure 34: aGlo); coxae 1-4 like those of E. rapax

but coxa 4 with posteroventral lobe blunter and

broader, long setae similar, formula of coxae 1-4 =

10-7-7-9; pereopods 1-2 as in E. rapax but spines

and setae fewer; pereopods 3-5 as in E. mutatus or

E. zoanthidea (Figure 44); uropod 2 with only 1

subapical dorsal spine on peduncle.

FEMALE.—Unknown; dominant females of sample

from type-locality are not of this species but deposited in male collections for future reference.

HOLOTYPE.—USNM 169020, male "a," 5.00 mm

(illus.).

TYPE-LOCALITY.—DAW 24, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 18 February 1962,

intertidal, rock wash in mangrove cove.

VOUCHER MATERIAL.—Six males from typelocality.

RELATIONSHIP.—This species is closely similar to

E. mutatus from California but differs in male

gnathopod 2 having the proximal palmar ridge

bearing a spine extended onto the palmar margin

and forming a projecting tooth that defines the

palm. The palm is also heavily setose in contrast

to E. mutatus.
NUMBER 271

69

FIGURE 34.—Elasmopus ocoron¡, new species (a = holotype, male "a," 5.00 mm). Dulzura gal,

new species (w = holotype, male "w," 3.00 mm). Cheiriphotis megacheles (Giles) (h = male "h,"

2.26 mm).

MATERIAL.—DAW 24; COCOS 9.

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz; Cocos Island; intertidal.

Elasmopus ìrapax Costa from Eastern Pacific

FIGURES 35-37

Elasmopus rapax Costa, 1853:175.

Elasmopus rapax form I of Bahía de Los Angeles.—J. L.

Barnard, 1962b, fig. 16; 1969b: 119.

DIAGNOSIS.—Eyes ordinary. Mandibular palp article 3 deeply falcate. Palm of male gnathopod 2

with ordinary spinose hump near dactylar hinge,

then truncate proximal tooth on plane mediad,

palm defined by tapering medial tooth, palm and

posterior margin of hand evenly and sparsely setose.

Article 2 of pereopods 3-5 with long posterior

setae in male only; locking spines of pereopods

1-5 ordinary, inner margin of dactyl smooth but

with minute proximal shagreen or fuzz, main sub-
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 35.—Elasmopus ìrapax Costa, male "a," 10.5 mm (w = female "w," 7.7 mm).

apical setule thick but apically flexible (with 2

subsidiary setules apparently normal to genus).

Epimera 2-3 with long ventral setae, epimera 1-3

with short ventral spines; epimeron 3 with weakly

S-shaped posterior margin bearing setular notches,

margin flat between notches, final notch lacking

setule, guarded by small sharp tooth, then gross

invagination defined ultimately by slightly enlarged

posteroventral tooth. Rami of uropod 3 equally

long in male, inner ramus with several stout basomedial spines, inner ramus of females and juveniles

shorter then outer ramus. Telson with naked protrusive apices placed mediad, laterally defined by

sinus and sharp tooth, spines submarginal from

sinus and forming row of 2-5 laterally, some spines

very long in female, all spines very short in male.

NOMENCLATURE.—No synonymy or references are

provided as my application of this name to the
NUMBER 271 71

G2o

FIGURE 36.—Elasmopus }rapax Costa, male "a," 10.5 mm (w — female "w," 7.7 mm).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

w

FIGURE 37.—Elasmopus }rapax Costa, male "a," 10.5 mm (μ = male "u," 6.39 mm; v — male "v,

9.7 mm; w = female "w," 7.7 mm).

material in question is fraught with doubt that

European and eastern Pacific populations of similar

morphology are identical. These Mexican specimens are the closest to E. rapax of any found in the

area and may represent a race of E. rapax from

which has been derived many of the other local

species of Elasmopus.

REMARKS.—The Mexican E. rapax differs from

European E. rapa× (using Sars, 1895, and Chevreux

and Fage, 1925, as models) in the presence of long

posterior setae on article 2 of pereopods 3-5 in the

male; the female lacks these setae as do males of

European rapax; in the stoutness of the subapical

seta on the dactyls of pereopods 1-5; in the stronger

falcation on mandibular palp article 3; in the

apical truncation of the naked tooth proximal to

the main hump on male gnathopod 2 palm; in the

slightly broader inner ramus of uropod 3 bearing

distinct spines basomedially (male inner ramus as

long as outer, female inner ramus shorter than

outer in Mexican rapax, rami apparently subequal

or equal to each other in European rapax); in the

full definition of a small posteroventral tooth on

epimeron 3 distinct from other serrations; in the

presence of only 1 other distinct but small denticle

dorsal to the sinuosity defining the main tooth and
NUMBER 271 73

in the truncation of the other denticles (in European E. rapax the epimeron bears several evenly

developed serrations and no enlargement of the

ultimate corner tooth and no distinct sinuosity); in

the greater disparity in size of telsonic spines between males and females (in European E. rapax

the telsonic spines are subequally long). Note that

Norwegian E. rapax has 2 or 3 long spines, occasionally the third short, whereas Mediterranean

rapax has 1 long, 1 short spine only on each lobe

of the telson.

The pleonal epimera of Mexican rapax have

long setae and short spines ventrally whereas Sars1

(1895) rapax apparently has naked epimera, unknown in the Mediterranean specimens of Chevreux and Fage (1925).

The other species of the Mexican rapax group

are described herein as if descendent from this

species; adjectives describing shapes are simplified

so that references to illustrations are often required

for precise definition.

VOUCHER MATERIAL.—SCO 18: male "a," 10.5

mm (illus.); female "w," 7.7 mm (illus.); male

"v," 9.7 mm (illus.); PAZ 7: male "u," 6.39 mm

(illus.).

MATERIAL.—SCO 1, 13, 18, 19; PAZ 6, 7, 11, ?I6,

24.

DISTRIBUTION.—Gulf of California at Bahía de

Los Angeles, Puerto Peñasco, Bahiá Concepción,

Isla Espiritu Santo-Isla Partida, Isla San Francisco;

intertidal.

Elasmopus serricatus J. L. Barnard, new status

FIGURES 38, 39

Elasmopus rapax serricatus J. L. Barnard, 1969b: 121, fig.

24j-m.

DIAGNOSIS.—Eyes ordinary. Mandibular palp

article 3 deeply falcate. Palm of male gnathopod 2

with single falcate hump near dactylar hinge, then

blunt middle tooth, palm defined by ridge bearing

spine, palm and posterior margin of hand setose

throughout. Article 2 of pereopods 3-5 lacking long

posterior setae, article 2 of pereopod 5 deeply serricate; locking spines of pereopods ordinary, though

enlarged, weakly curved and apically blunt, inner

margin of dactyl smooth but with minute shagreen

proximally, main subapical setule thick but apically

flexible. Epimera bearing only short ventral spines;

epimeron 3 weakly convex posteriorly, evenly and

sparsely notched, posteroventral corner bearing

similar notch and weak tooth. Uropod 3 of male

with shortened inner ramus as in females and

juveniles, inner ramus with cluster of 2 medial

spines. Telson with excavate apices armed with 4

spines, 2 long and 2 short in male, 2-3 long and 2

short in female, latter with spines closer to margin

and margin more truncate than in male.

DESCRIPTION.—Following parts as shown for E.

rapax (Figures 35-37): head (but eye slightly

larger), antenna 1 (medial aspects of article 1 on

antenna 1 and base of antenna 2 illustrated for

minor details); antenna 2 (but setae fewer and

stouter), upper lip, right mandible (3 spines), left

mandible (4 spines), mandibular palp, lower lip,

maxilla 1 (but inner plate on Figure 38: XIi) (palp

with 9 spines, 5 setae), maxilliped (except for longer

apical spine, Figure 38: VS4, on palp article 4);

gnathopod 1 (except for article 4, Figure 39: Gl),

shape of pereopods 1-2 (but article 6 with sets of 2

spines, Figure 39: PI), articles 4-7 of pereoped 3;

female uropod 3 like that of male E. serricatus but

outer ramus with 1 less set of lateral spines, inner

ramus with only 1 medial spine instead of 2; accessory flagellum of both sexes 3-4 articulate.

VARIATIONS.—Panama material. Middle tooth on

palm of male gnathopod 2 becoming obsolete in

adults and in juvenile specimens more proximally

located (away from hinge tooth) than in Californian

specimens. Galapagan material: medial face of

hand on male gnathopod 2 either bearing or lacking the small ridge, when present ridge armed only

with a seta.

VOUCHER MATERIAL.—SCO 14: male "d," 7.91

mm (illus.). PAZ 22: male "a," 6.91 mm; male

"b," 6.21 mm (illus.); female "k," 5.24 mm (illus.).

REMARKS.—This taxon is elevated to full specific

status; it lies outside the rapax morph in the shape

and spination of the telson and the shortened inner

ramus of male uropod 3.

MATERIAL.—SCO 14, 19; PAZ 20, 22; GAL 101,

118, ?I2O; DAW 3, 4, 6, 8, 10, 17, 21, 26, 27, 28, 31,

37, 38; ECU 3; PAN 14; BRU 1, 2.

DISTRIBUTION.—California, from Carmel to La

Jolla; Gulf of California at Puerto Peñasco, Bahía

San Carlos, Guaymas, Cabo San Lucas; Galapagos

Islands; Ecuador; Panama; intertidal.
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

G2o

bR3 KG 2

FIGURE 38.—EUumopus serricatus J. L. Barnard, male "d," 7.91 mm (b •* male "b," 6.21 mm;

k m female "k," 5.24 mm).
NUMBER 271 75

FIGURE 39.—Elasmopus serricatus J. L. Barnard, male "d," 7.91 mm (A = female "k" 5.24 mm).

Elasmopus temori, new species

FIGURE 40 (part)

DIAGNOSIS.—Eyes ordinary. Mandibular palp

deeply falcate. Palm of male gnathopod 2 with small

quadrate spinose hump near dactylar hinge, then

immediately adjacent short conical tooth, middle

of palm with weak, broadly conical hump, palm

defined by minute protrusion bearing small spine,

medial face with ridge bearing large spine, palm

highly elongate, posterior margin of hand scarcely

one-third as long as palm, moderately setose, palm

almost naked, medial face of hand poorly setose,

dactyl elongate and fitting palm. Article 2 of pereopods 5-5 lacking long posterior setae, article 2 of

pereopod 5 normally serrate; locking spines of pereopods 3-5 ordinary, straight, one slightly larger

than other, smooth, inner locking spine on pereopods 1-2 enlarged, weakly curved, blade-like, inner

margin on dactyl of pereopods 1-5 with faintest

shagreen marginally, main apical setule thick but

apically flexible. Epimera bearing mixtures of short

spines and medium setae, epimeron 3 weakly convex posteriorly, sparsely notched, posteroventral

corner bearing weak sinus and weak sharp tooth.

Rami of male uropod 3 elongate, reaching equally;

inner ramus with several sets of medial spines. Telson with narrow, excavate apices armed with 3-4

spines, at least 1 spine highly elongate.

DESCRIPTION OF MALE.—Generally like E. rapax

but eyes with enlarged ommatidia clear of pigmentary core, article 3 of peduncle on antenna 1

shortened (illustrated), articles 1-3 with ventral

setae elongate. Following parts like E. rapax (Figures 35-37): upper lip, lower lip, mandibles (except

palp, Figure 40: aM), maxilla 1 (but with thin inner

plate, spine 2 (from medial edge) with 2 medial

teeth, spines 3-4 with 3 medial teeth, palp of medium stoutness), plates of maxilla 2 (but thin), dactyl

of maxilliped (but of shortened variety with elongate nail as in E. seiricatus). Coxae 1-4 (but slightly

broadened), posteroventral lobe of coxa 4 (but

slightly weaker, poorly setose, elongate setae on

coxae 1-4 = 3-1-2-1); article 4 of gnathopod 1

(but rounded posterodistally, article 6 slightly more

elongate and rectangular, illustrated); pereopods
76

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pld

FIGURE 40.—Elasmopus temori, new species (a = holotype, male "a," 4.17 mm). Heterophlias

seclu¡us escabrosa \. I,. Barnard (p = sex unknown "p," 3.12 mm).
NUMBER 271 77

1-2 (poorly setose, article 6 with 4-6 posterior spine

positions besides locking spines, all positions with

short spine, some with seta or additional spine);

pereopods 3-5 (generally as in E. zoanthidea but

article 2 slightly narrower and article 6 elongate, on

pereopod 3 about 0.7 times as long as articles 4-5

combined, on pereopods 4-5 almost 0.8 times). Urosome and uropods 1-2 (but less spinose).

FEMALES.—Females and juveniles associated with

unique male apparently of distinct species owing to

significant differences in dactyls of pereopods, epimera, uropod 3, and telson; males similar to these

not yet discovered; collection in the Smithsonian

Division of Crustacea with provisional name of "E.

temori—associated females" for future reference.

HOLOTYPE.—USNM 169022, male "a," 4.17 mm

(illus.). Unique.

TYPE-LOCALITY.—GAL 113, Galapagos Islands,

Tower Island (Genovesa), Darwin Bay, 29 January

1964, intertidal, rock wash.

VOUCHER MATERIAL.—The holotype.

RELATIONSHIP.—The unique male of this species

may be an aberration of E. zoanthidea; it finds its

closest relationship to that species in telson, epimera, and article 2 and dactyls of pereopods. It

differs in the palmar configuration of gnathopod 2

on which are borne 2 weak middle processes instead

of 1 strong process and in the full development of a

mediofacial ridge bearing a spine. The main dactylar seta of pereopods 1-5 is enlarged, the medial

face of the hand on gnathopod 2 is poorly setose

and uropod 3 and the epimera have greater maturity in the sense that the epimera bear setae as well

as spines and the inner ramus of uropod 3 is fully

elongate. Both right and left gnathopod 2 are similar so that the aberration, if so to be interpreted, is

bilateral and not simply confined to 1 appendage.

MATERIAL.—GAL 113.

DISTRIBUTION.—Galapagos Islands, Tower Island,

intertidal.

Elasmopus tiburoni, new species

FIGURE 41

Elasmopus rapax form I of Tiburon Island.—J. L. Barnard,

1969b: 119 [part].

DIAGNOSIS.—Eyes ordinary. Mandibular palp

article 3 deeply falcate. Palm of male gnathopod 2

with ordinary spinose hump near dactylar hinge but

hump weak and concealed laterally by flange-like

inflation of entire palm, with dactyl overriding face

of palm, medial face of palm deeply hollowed as in

E. pocillimanus (Bate), palm and hollow defined by

small tapering medial tooth but third tooth absent,

hollow defined anteriorly by sharp setose ridge,

palmar margin nearly naked, posterior margin of

hand densely setose but setae in rows of short to

long. Article 2 of pereopods 3-5 with long posterior

setae in male only; locking spines of pereopods 1-5

ordinary, inner margin of dactyl usually smooth but

occasionally with 1-2 notches (forming broad castellations) and minute proximal shagreen, main subapical setule thick but apically flexible; articles 4-5

of pereopods 4—5 especially broad in comparison to

E. rapax, increase in breadth confined primarily

to male. Epimera 1-3 with long ventral setae and

short spines; epimeron 3 with nearly straight posterior margin armed with notches bearing setules,

margins between notches flat except for ultimate

posteroventral tooth and one tooth dorsal to it,

these teeth small and of subequal size, sinus above

ultimate tooth lacking setule. Rami of uropod 3

equally long in male, inner ramus with several stout

basomedial spines, inner ramus in females and

juveniles shorter than outer ramus. Telson with

naked protrusive apices placed mediad, laterally

defined by sinus and sharp tooth, spines submarginal from sinus and forming row of 2-4 laterally,

but row extending mediad across axis of each lobe,

male with 2 short spines, female with 4 long spines

on each lobe.

DESCRIPTION.—Following parts not illustrated

and as shown for E. rapax (Figures 35-37): head,

antennae, accessory flagellum, eye (slightly larger

than in E. rapax), mouth parts, pereopods 1-2,

dactyls, urosome, uropods 1-2. Female gnathopod 2,

coxae, cuticle, medial view of article 1 on antenna

1; female lacking long setae of article 2 on pereopods 3-5, small females and juveniles lacking serrations of epimeron 3 except for ventralmost sinus

and tooth. Juvenile males with gnathopods like

juvenile E. rapax; female uropod 3 with 3 apicomedial peduncular spines instead of 2 as in Figure 37: wR3 of E. rapax.

HOLOTYPE.—USNM 149405, male "i," 8.76 mm

(illus.).

TYPE-LOCALITY.—SCO 12, Puerto Peñasco, Mexico, 23 February 1971, intertidal, wash of Colpomenia sp.
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

W3

FIGURE 41.—Elasmopus tiburoni, new species, holotype, male "f," 8.76 mm (w = female "w,

6.58 mm).
NUMBER 271 79

VOUCHER MATERIAL.—Type-locality, female "w,"

6.58 mm (llus.).

REMARKS.—This species differs from Mexican E.

rapax in the pocillimanus-like gnathopod 2 of the

terminal male bearing only 2 palmar teeth and a

large hollow, with uneven setal distribution, in the

even serrations of epimeron 3, and in the enlarged

articles 4-5 of pereopods 4-5, with larger spines in

tandem on article 4 of pereopod 3.

MATERIAL.—SCO 12, 14, 19; KNO 3, 6; PAZ 3,

12, 13; BRU 1, 2; SNY 1. Also, AHF Velero III

1042 near Isla Tiburon.

DISTRIBUTION.—Gulf of California; Puerto Peñasco, Bahía Kino, Turners Island south of Isla

Tiburon, Bahía San Carlos, Guaymas, Bahía San

Evaristo, Isla Espiritu Santo; found in shells of

hermit crab in SNY 1; intertidal.

Elasmopus tubar, new species

FIGURES 42, 43

DIAGNOSIS.—Eyes ordinary. Mandibular palp

article 3 deeply falcate and extraordinarily stout.

Palm of male gnathopod 2 with hump near dactylar

hinge, hump setose but not spinose, medial face of

hand with weak ridge-hump hidden by setae, no

defining tooth, palm and posterior margin of hand

heavily setose throughout, apex of hand anterodistally bearing stridulation-like ridges or rays. Article

2 of pereopods 3-5 lacking long posterior setae,

male pereopod 5 with posteroventral margin of

article 2 serricate or castellate; locking spines of

pereopods 3-5 ordinary, on pereopods 1-2 composed

of one short truncate spine, one long, subtruncate

spine, inner margin of dactyl with minute shagreen

proximally, main subapical setule thick but apically flexible. Epimera 1-3 bearing short ventral

spines only; epimeron 3 with weakly sinuous posterior margin bearing weak acclivities each with

setule, posteroventral tooth and sinus very small.

Inner ramus of uropod 3 in both sexes shorter than

outer ramus, inner ramus with several widely separated medial spines. Telson with narrow, weakly

excavate apices bearing 1 long spine and 1 medium

spine, or in female with 1 long, 1 medium to long

and occasional short third spine.

DESCRIPTION.—Following parts as shown for E.

rapax (Figures 35-37): head (eyes slightly enlarged),

antennae (but much less setose), accessory flagellum

(2-articulate), upper lip, lower lip (but mandibular

lobes blunter), maxilla 2, maxilliped (but apical

spine on palp article 4 slightly longer), urosome,

uropods 1-2, pereopod 3 (but spines fewer and long

setae of article 2 absent). Pereopods 1-2 like those

of E. serricatus, bearing extra spine at each posterior

position. Outer plate of maxilla 1 with 7 spines (1

hidden in Figure 43: Xló), inner plate with 2 long

and 2 short setae, palp with 6 spines and 5 setae on

both sides; coxae 1-3 like those of E. rapax but

many fewer long setae (about 3 per coxa), coxa 4 as

Figure 43: E4; female epimeron 3 like that of male,

occasionally posteroventral tooth blunter; gnathopod 1 distinct from adults of other species of Elasmopus presented herein, retaining juvenile appearance in adult but heavily setose, slightly less setose

in female.

HOLOTYPE.—USNM 142482, male "a," 5.1 mm

(illus.).

TYPE-LOCALITY.—PAZ 17, Baja California, 11

km E of Cabo San Lucas, 4 December 1971, intertidal wash of Padina.

VOUCHER MATERIAL.—Type-locality, female "w,"

4.3 mm (illus.).

RELATIONSHIP.—This species approximates E.

antennatus (Stout) in male gnathopods 1 and 2, uropod 3, and telson but the main protrusion on the

hand of male gnathopod 2 is larger in E. tubar,

male pereopod 5 bears castellations on article 2, the

telson has fewer spines per apex and the spines are

much longer than in E. antennatus, male gnathopod

2 bears a "stridulating" hump and female gnathopod 2 has a much stouter hand in E. tubar than in

E. antennatus. Perhaps the odd declivity of article

4 on male pereopod 4 is also characteristic (Figure

43: P4) but insufficient material is available to

confirm this characteristic.

MATERIAL.—PAZ 17, 18; GAL 102, 108, 109, 119,

120; DAW 5, 9, 13, 16, 17, 18, 19, 23, 27, 33, 40.

DISTRIBUTION.—Baja California, Cabo San Lucas;

Galapagos Islands; intertidal.

Elasmopus zoanthidea, new species

FIGURE 44

ETYMOLOGY.—The epithet "zoanthidea" is from

Greek, zoön (animal) and antheros (blooming).

DIAGNOSIS.—Eyes ordinary. Mandibular palp

deeply falcate. Palm of male gnathopod 2 with
80

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

G2

FIGURE 42.—Elnsmo¡ms tubar, new species, holoi\pe, male "a," 5.1 mm (w = female "w," 4.3 mm).
NUMBER 271 81

FIGURE 43.—Elasmopus tubar, new species, holotype, male "a," 5.1 mm (w = female "w," 4.3 mm).

ordinary spinose hump near dactylar hinge but

hump weak, then broadly quadrate, weakly bifid

or trifid, middle tooth separated from spinose hump

by distance equal to width of hump 2, palm defined by obsolescent tooth on margin, palm and

posterior margin of hand strongly setose throughout, medial face of hand with numerous bundles

of setae in row paralleling palm and posterior

margin. Article 2 of pereopods 3-5 lacking long

posterior setae, article 2 of pereopods 4-5 normally

serrate; locking spines of pereopods 3-5 ordinary,

thin, straight, weakly striate longitudinally, inner

locking spine on pereopods 1-2 enlarged, curved,

blade-like, inner margin of dactyl on pereopods

1-2 with faintest shagreen, on pereopods 3-5 with

occasional obsolescent serration, main subapical

setule thick but apically flexible. Epimera bearing

only short ventral spines, epimeron 3 weakly convex

posteriorly, sparsely notched, posteroventral corner

bearing weak sinus and weak, sharp tooth. Uropod

3 of male with shortened inner ramus bearing 2

pairs of medial spines. Telson with excavate apices

armed with 3 spines, grading from short to greatly

elongate from medial to lateral.

DESCRIPTION.—Generally like E. rapax (Figures

35-37) as follows: Eyes slightly smaller and with

far fewer ommatidia, all stained brownish magenta,

antennae 1-2 like E. rapax but article 1 of antenna

1 as in Figure 44: Aim, articles 2-3 with ventral

setae elongate, accessory flagellum 3-articulate;

mandibles like rapax but left mandible with first

raker spine very thick and denticulate; upper lip,

and lower lip like rapax; maxilla 1 with spines 2,

3, and 6 (from medial) each with 2 elongate and

sharp side cusps, palp scarcely broader than outer

plate; plates of maxilla 2 thinner than in E. rapax;

dactyl of maxilliped shortened, main nail elongate,

almost as long as dactyl; palm of gnathopod 1 with

only 1 defining spine, medial comb of article 4

shortened, elements enlarged, article 5 with or without midanterior pair of setae; shapes of coxae 1-4

like those of E. rapax but elongate ventral setae

fewer, formula of coxae 1-4 = 5-3-4-1, coxae 2-3

smaller relative to coxae 1 and 4 than in E. rapax;

pereopods 1-2 generally as in E. rapax but spines

and setae fewer; pereopods 3-5 and uropod 1 as

in Figure 44, uropod 2 as in E. rapax but peduncle

bearing only 1 dorsolateral spine set apically.

FEMALE.—Unknown.

HOLOTYPE.—USNM 169023, male "a," 4.35 mm

(illus.).

TYPE-LOCALITY.—GAL 113, Galapagos Islands,

Tower Island (Genovesa), Darwin Bay, 29 January

1964, intertidal, rock wash.

VOUCHER MATERIAL.—Type-locality, male "v,"

4.41 mm (illus.) and 4 younger males.
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

G2o

FIGURE 44.—Elasmopus zoanthidea, new species, holotype, male "a," 4.35 mm (v = male "v,"

4.41 mm).

RELATIONSHIP.—This species resembles E. holgurus in terms of telson and male gnathopod 2

but differs in the breadth and shape of the middle

palmar tooth on male gnathoped 2, in the shallowness of posterior notches on article 2 of pereopod

5, and in the unextended epimeron 3. Other differences in setosity and shape may be a function of

the small body size of E. zoanthidea and, therefore,

juveniles of E. holgurus may resemble adults of

E. zoanthidea in those characteristics.

Elasmopus zoanthidea resembles E. mayo and E.

rapax especially in the palm of male gnathopod 2,

but E. zoanthidea has many more setae on the

medial face of the hand, lacks the protrusive medial
NUMBER 271 83

limbs on the telsonic apices, and lacks long posterior

setae on article 2 of pereopods 3-5.

MATERIAL.—GAL 102, 109, 113, 119; DAW 9, 23.

DISTRIBUTION.—Galapagos Islands, intertidal.

Galapsiellus J. L. Barnard

Galapsiellus leleuporum (Monod)

Paraniphargus leleuporum Monod, 1970:13-25, figs. 6-45.

Galapsiellus lelouparum [sic].—J. L. Barnard, 1976:422-423.

MATERIAL.—GAL 103.

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz, Academy Bay, anchialine.

Maera Leach

Maera reishi, new species

FIGURES 45-47

Maera inaequipes.—J. L. Barnard, 1959:25-26, pi. 5 [not

Costa]; l969a:2O5; 1969b: 121-122.

ETYMOLOGY.—Dedicated to Dr. Donald James

Reish for his numerous contributions to our knowledge of eastern Pacific lagoons and estuaries.

DIAGNOSIS.—Lateral cephalic lobes weakly mammilliform, anteroventral corner of head produced

as narrow elongate tooth. Eyes ovate, brownish pink

in alcohol. Article 1 of antenna 1 with 2 basoventral

spines and 1 apicoventral spine. Article 3 of mandibular palp 1.2 times as long as article 2. Coxa 1

weakly extended forward, coxae 1-4 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 spine, article 5 with several medial

rows of diverse setal spines, article 4 lacking posterodistal cusp; gnathopod 2 very large, article 2

heavily spinose anteriorly, 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 weak sinus, palm with one

deep incision in males, occasionally in females, latter usually without main sinus, dactyl fitting palm,

bearing inner acclivity near position of sinus,

acclivity weak to strong, usually absent in females

and juveniles. Article 2 of pereopods 3-5 narrowly

ovatorectangular, with slightly extended, angular

posterodistal corners, posteroproximal margins

rounded; dactyls of pereopods 1-5 with strong distal

constriction bearing sharp defining corner and

armed with seta, sharp but stunted nail bearing

accessory tooth, facial setule, constriction also bearing stout setule appressed to margin of nail, locking

spines small, straight or slightly curved. Pleonal

epimera 1-3 with slightly convex posterior margins,

each with small sharp posteroventral tooth, ventrally spinose; pleonites dorsally smooth. Uropod 3

extending flush with other uropods, rami flat, narrow, apically truncate and spinose, inner extending

to Mark 80 along outer, tiny article 2 on outer

ramus. Telson deeply but not fully cleft, with

strongly fixed medial gape, lobes thin, apically

truncate or convex, each lobe with 4 long spines,

some longer than telson. Dorsoposterior margin

of urosomite 3 rounded.

JUVENILES.—Smallest available, 1.9 mm long; proportions of mandibular palp articles 2-3 similar

to adult; inner ramus scarcely exceeding Mark 50

on outer ramus; each lobe of telson with 2 long

spines; cephalic tooth small but distinct.

DESCRIPTION.—Upper lip and epistome fused together; outer plate of maxilla 1 with 5 bifid and 4

serrate spines; cuticle smooth extent for occasional

bulbar setule and occasional seta on coxae; palmar

sinus of gnathopod 2 developing before dactylar

acclivity and before article 2 heavily spinose.

VARIATIONS.—At the northern end of its range,

in suburban California, specimens of Maera reishi

reach a length just less than 7 mm whereas their

maximum length in more southerly waters of

Mexico is less than 5.5 mm. In these large specimens from southern California, the gland cone of

antenna 2 extends only two-thirds along article 3

of antenna 2 whereas in southerly specimens the

gland cone overreaches the end of article 3. No

females with a palmar sinus on gnathopod 2 have

been found in the northern end of the range. The

dactylar acclivity on gnathopod 2 of many, but not

all, males in the north becomes extremely developed

as it is in Maera chinarra.

HOLOTYPE— USNM 142568, male "a," 5.3 mm

(illus.).

TYPE-LOCALITY.—PAZ 3, Gulf of California, Isla

Espiritu Santo, 28 November 1971, 0.3-1.0 m, algae,

rock wash.
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

l· ̄ir.L'RF. 43.—Maeia re¡shi, new species, holotype, male "a," 5.3 mm (ƒ = juvenile "j, " 2.6 mm;

v — male "v," 6.5 mm; u· »= female "w," 5.4 mm).
NUMBER 271 85

PI

FIGURE 46.—Maera reL·hi, new species, holotype, male "a," 5.3 mm (v «= male "v," 6.5 mm).

VOUCHER MATERIAL.—Type-locality: female "w,"

5.4 mm (illus.); juvenile "j, " 2.6 mm (illus.). Corona

del Mar, California, 6 February 1951, Sabellaria

bed, male "v," 6.5 mm (illus.).

RELATIONSHIP.—This species is distantly related

to M. rathbunae Pearse (1908) from Florida, the

latter bearing a distinctly oblique palm on gnathopod 2, equally extending rami of uropod 3 and

sharply pointed, poorly spinose telsonic lobes. The

identification of Maera rathbunae by Kunkel (1910)

from Bermuda appears to be a distinct species much

more similar to M. reishi than is the Floridian M.

rathbunae. The Bermudan species has truncate telsonic apices with numerous "setae," the inner ramus

of uropod 3 is shortened and the palm of gnathopod

2 is transverse as in M. reishi. The absence of spines

on article 2 of gnathopod 2 and the absence of all

but 1 spine on the palm of gnathopod 2 appear to

be good characters differentiating the Bermudan

species from M. reishi. The mandibular palp and

the inner plate of maxilla 1 also appear distinctive

and there are possible differences in epimera and
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 47.—Maera reishi, new species, holotype, male "a," 5.3 mm.

second maxillae. I reserve the naming of the Bermudan species to a person with material available

who can describe this species minutely and determine whether the "setae" of the telson are thin and

flexible or stout and spine-like.

Maera reishi differs from European M. inaequipes

in the blunt telsonic apices each bearing 4 spines,

whereas Aí. inaequipes has bifid apices bearing 2

spines. Actually 1 spine on each lobe in both species

emerges from the ventrolateral surface of the lobe.

Maera inaequipes has a very sharp anteroventral

corner on coxa 1, the gnathopodal palm in both

sexes slopes obliquely, and article 2 of gnathopod

2 is poorly spinose or free of spines.

Among several species of the central Pacific

Ocean (J. L. Barnard, 1970:147-161), Maera reishi

appears to be closest to M. pacifica Schellenberg,

from which M. reishi differs in the presence of

numerous stout spines on article 2 gnathopod 2 in

both sexes and in the male in the shallower sinus

adjacent to the defining tooth of the palm.

J. L. Barnard (1972b: 107) gives a brief synopsis

of species in the M. quadrimana complex, to which

M. reishi belongs.

MATERIAL.—PAZ 3; GAL 108; DAW 3, 8, 27, 31.

DISTRIBUTION.—Gulf of California at Isla Espiritu

Santo and Bahía de Los Angeles, 0 m; Galapagos

Islands, 0-6 m; California north to Cayucos, 0-3 m.

Maera chinarra, new species

FIGURE 29 (part)

DIAGNOSIS.—Lateral cephalic lobes weakly mammilliform, anteroventral corner of head with obsolescent sharp cusp. Eyes ovate, brownish pink in

alcohol. Article 1 of antenna 1 with 1 basoventral

spine and often no thickened apicoventral spine, or

small spine occasionally present. Article 3 of mandibular palp shorter than article 2. Coxa 1 weakly extended forward, coxae 1-4 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 spine, article 5 with several medial rows of diverse setal spines, article 4 lacking posterodistal

cusp; gnathopod 2 very large, article 2 heavily

spinose anteriorly, with medium-sized anterodistal

lobe, article 3 with similar lobe, article 4 slightly
NUMBER 271 87

extended posterodistally, article 5 very short, with

long tumid posterior lobe, article 6 very large, subrectangular, not expanded distally, palm transverse,

defined by medium tooth, bearing spine, tooth adjacent to weak sinus, palm with 1 deep incision in

males, smooth in females, dactyl fitting palm, bearing large inner acclivity near position of sinus, acclivity absent in females and juveniles. Article 2 of

pereopods 3-4 narrowly ovatorectangular, broader

on pereopod 5, wih slightly extended, angular posterodistal corners, posteroproximal margins slightly

extended but rounded; dactyls of pereopods 1-5

with strong distal constriction bearing sharp defining corner and armed with seta, sharp but

stunted nail bearing accessory tooth, facial setule,

constriction also bearing stout setule appressed to

margin of nail, locking spines small, straight or

slightly curved. Pleonal epimera 1-3 with slightly

convex posterior margins, each with small sharp

posteroventral tooth, ventrally spinose; pleonites

dorsally smooth. Uropod 3 extending flush with

other uropods, rami flat, broader than in M. reishi,

apically truncate and spinose, inner extending to

Mark 80 along outer, tiny article 2 on outer ramus.

Telson deeply but not fully cleft, with strongly fixed

medial gape, lobes thin to medium broad, apically

incised but weakly and broadly, each lobe with 3

or 4 spines, some longer than telson. Dorsoposterior

margin of urosomite 3 rounded.

DESCRIPTION AND ILLUSTRATIONS.—Following parts

similar to M. reishi (Figures 45-47): lateral views of

antennae 1-2; epistome-upper lip; anterior view of

latter also; mandibles, except for palp, left molar

with 5 sharp projecting spines (2 shown for reishi,

Figure 45: M) lower lip; maxilla 1 (but palpar apex

with 6 spines and 4 setae only); maxilla 2 (but

medial margin of inner plate with 1 additional

seta); maxilliped; coxae 1-4 (but these slightly

broader anterior to posterior and coxa 1 with anterior margin slightly less concave); gnathopod 1;

pereopods 1-4 (except article 4 of pereopods 3-4

slightly broader); dactyls; locking spines; uropods

1-2; uropod 3 (with slightly broader rami); epimera

(but epimeron 3 with only 3 ventral spines); cuticle.

HOLOTYPE.—USNM 142556, male "a," 3.7 mm

(illus.).

TYPE-LOCALITY.—PAZ 21, Baja California, 11 km

E of Cabo San Lucas, 4 December 1971, intertidal,

algal wash.

VOUCHER MATERIAL (all illustrated).—PAZ 22:

female "f," 3.4 mm; juvenile "j, " 2.9 mm; male

"p," 3.6 mm.

RELATIONSHIP.—This species appears to be a twin

of M. reishi and differs from the latter mainly in the

obsolescent cephalic tooth, the shortened article 3

of the mandibular palp and in the weakly excavate

telsonic apices. The greatly thickened acclivity on

the dactyl of male gnathopod 2 resembles that

found only in the northern specimens of M. reishi;

where M. reishi and M. chinarra occur sympatrically

only M. chinarra bears the extremely thickened acclivity; M. chinarra has not been found in southern

California coastal shallows where M. reishi specimens bear this thickened acclivity. Maera chinarra

can be considered to be a dwarf sibling of M. reishi

because specimens of M. chinarra appear to reach

terminal morphology at body lengths between 3.5

and 4.0 mm, whereas specimens of M. reishi reach

terminal stages at body lengths in excess of 5.0 mm

in warm waters and 5.5 mm in cool waters. My material is insufficient to confirm these approximations.

Maera chinarra differs from the Bermudan species

of Kunkel (1910:46) misident¡fied as M. rathbunae

in the presence of numerous spines on gnathopod

2, both on the palm and article 2, but apparently

has a similar mandibular palp. The excavate telsonic apices of M. chinarra are distinctive.

The apicoventral spine on article 1 of antenna 1

appears to be confined to females, whereas that

spine in males is a slender seta.

Galapagan specimens correspond in almost all

attributes to specimens from Mexico but the gland

cone extends two-thirds along article 3 of antenna 2.

Because this character is variable in M. reishi its

significance subspecifically is discounted at present

in Galapagan specimens of M. chinarra.

The value of 2 characters used to distinguish continental Maera chinarra from M. reishi is reduced

in Galapagan specimens of M. chinarra. The mandibular palp of a specimen from GAL 114 has article 3 over 90 percent as long as article 2. In another

specimen, a male from GAL 113, antenna 1 has the

thickened and enlarged apical spine on article 1.

Both of these specimens otherwise have the typical

head, telson and uropod 3 of M. chinarra.

MATERIAL.—PAZ 14, 17, 18, 21, 22; GAL 113, 114,

116, 118, 120; DAW 1, 6, 7, 8, 9, 17, 18, 19, 26, 27,

35, 40; COCOS 1, 3.

DISTRIBUTION.—Baja California, Cabo San Lucas,

0 m; Galapagos Islands, 0 m; Cocos Island, 0 m.
88

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Maera simile Stout

Maera simile Stout, 1913:644-645.—Shoemaker, 1942:12 —

Hewatt, 1946:199—J. L. Barnard, 1959:24-25; 1961:179;

l964b:222; l966a:l9; l966b:62; 1969b: 122-123.

Maera inaequipes—]. L. Barnard, 1954a:16-18, pis. 16, 17

[not Costa].

MATERIAL.—TOP 3.

DISTRIBUTION.—Gulf of California, Topolobampo,

1 m; generally from Puget Sound, Washington to

Bahía Magdalena, 0-221 m (depths greater than 50

m usually in submarine canyons).

Melita Leach

Melita sulca (Stout)

Caliniphargus sulcus Stout, 1913:641-642.

Melita palmata.—Shoemaker, 1941b: 187.—Hewatt, 1946:199

[not Montagu].

Melita sulca.—]. L. Barnard, l969a:2O9; 1969b: 126-130, figs.

22, 23.

MATERIAL.—SCO 1, 5, 7, 8, 14; TOP 3; PAZ 1, 2,

7, 10, 24, 25.

DISTRIBUTION.—Gulf of California, at Puerto

Peñasco, Bahía de Los Angeles, Topolobampo,

Bahía Concepción, Isla San Francisco, Isla Espiritu

Santo and La Paz, 0-24 m; generally Puget Sound,

Washington, to Isla Cedros, Baja California,

0-101 m.

R3

FIGURE 48.—Me×imaera d¡ffidentia J. L. Barnard, male "a,"

4.46 mm.

Me×imaera J. L. Barnard

Me×imaera diffidentia J. L. Barnard

FIGURES 48, 49

Me×imaera diffidentia J. L. Barnard, 1969a:209-210, figs. 21,

22.

VOUCHER MATERIAL.—GAL 111, male "a," 4.46

mm (illus.), male "b," 4.37 mm, female "c," 4.40

mm, female "f," 4.88 mm (illus.).

• REMARKS.—The Galapagan specimens differ from

those described from the Gulf of California in the

equally projecting apical cusps on each lobe of the

telson. At present, this seems inadequate for subspecific distinction.

Each mandibular molar bears a long pluseta; the

inner plate of maxilla 1 bears 2 long apical, 1 long

apicolateral and none or 1 short apical seta. In

Figures 48 and 49 gnathopod 1 is magnified 5 percent more than gnathopod 2.

The female was unknown heretofore. Females in

the Galapagos Islands resemble the males in all

characters, including size of gnathopods, except that

the medial face on the hand of gnathopod 2 has

several doublets of spines (Figure 49: fG2O).

The validity of Meximaera and its possible senior

synonym Linguimaera Pirlot, remain in question.

This hinges on rediscovery of the so-called typespecies of Linguimaera and a global evaluation of

the taxonomic importance of cephalic slits, presence or absence of article 2 on the outer ramus of

uropod 3, the loss of prehensility on gnathopods and

probably on many other characters (J. L. Barnard,

I972a:224).

MATERIAL.—GAL 108, 110, 111.

DISTRIBUTION.—Gulf of California, Bahía de Los

Angeles, 0-24 m; Galapagos Islands, 0 m.
NUMBER 271 89

FIGURE 49.—Meximaera diffidentia J. L. Barnard, male "a," 4.46 mm (ƒ = female "í,`` 4.88 mm).
90

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HYALIDAE

Allorchestes Dana

Allorchestes Dana, 1849:136; 1852:205.—Stebbing, 1905:581-—

J. L. Barnard, 1974:41.

Asp¡dophoreia Haswell, 1880:101.

DIAGNOSIS.—Talitroidean and hyalid with palp of

maxilla 1 vestigial. Article 4 of maxillipedal palp

unguiform. Gnathopods of both sexes subchelate,

male gnathopod 2 larger than 1, female gnathopod

2 generally similar to female gnathopod 1. Uropod

3 with 1 ramus, apical armament weak (setae always

present, spines rare or absent). Telson broad, rectangular, partially cleft but lobes not divergent.

TYPE-SPECIES.—A. compressa Dana (1852).

REMARKS.—J. L. Barnard (1974) has revised this

genus to exclude many species now transferred to

Hyale; those species have a long palp on maxilla 1,

no apical setae on uropod 3, just spines and triangular telsonic lobes, the telson fully or deeply

cleft. The presence or absence of a lobe on article

5 of male gnathopod 2 is discounted as a generic

character; all known species of Allorchestes have

this lobe, most species of Hyale lack this lobe. This

reorganization restricts Allorchestes to species in

cool waters, 3 in the North Pacific Ocean and 2 in

the South Pacific Ocean. The genus, therefore, has

biboreal discontiguity.

Taxonomic characters are focused on male

gnathopod 1. A key to species is provided by J. L.

Barnard (1974:42).

Allorchestes compressa Dana from Australia appears to have the least specialized male gnathopod

1, the hand being rectangular or bearing a slight

posterior bulge near the palm, the dactyl fitting the

palm or just failing this on the bulged form. The

inner palmar spines are of medium to small size, are

situated near the palmar margin and in adults number 3-4, often up to 7.

Allorchestes novizealandiae Dana from New

Zealand has a highly specialized male gnathopod 1,

the palm bearing a strong axial protrusion against

which the dactyl closes but overlaps. The freely projecting part of the dactyl is about one-third of the

length of the dactyl. Four palmar spines form a

semicircular row partially divorced from the palmar

margin.

All species of northern Allorchestes appear to be

united by the reduction in number and enlargement of palmar spines on male gnathopod 1. In

Allorchestes angusta only 2 inflexible spines remain

on the inner face of the hand, 1 near the defining

corner of the palm, another strongly submarginal

and near the middle of the palm. The dactyl is

strongly shortened and apically bifid.

Allorchestes carinata Iwasa has an ordinary rectangular hand with the dactyl fully fitting the palm.

Two enlarged spines occur on the medial palmar

face, 1 near the defining angle, the other submarginal from the middle of the palm. Both spines are

much smaller than in A. angusta.

The third species of Allorchestes in the North

Pacific resembles A. novizealandiae in the protrusive chela on the palm of gnathopod I and in the

dactyl strongly overlapping the palm, but in A.

bellabella the dactyl is tumid whereas in A. novi·

zealandiae it is slender. In terminal males only 1

enlarged spine occurs on the medial palmar face but

juvenile males have 2 of these spines.

Subsidiary taxonomic characters necessary to support the thesis that species of Allorchestes are definable by characters of male gnathopod 1 are difficult to find. The identification problem is especially

tedious in females and juveniles. Because the life

histories of the several species have not been studied

in detail, owing to an absence of materials of juveniles, the few minor subsidiary distinctions among

species cannot be confirmed as invariable. Allorchestes carinata and A. bellabella have posterior

body carinations, presumably absent in juveniles.

Spines on articles 4-5 of pereopods 3-5 are very

stout and blunt in A. carinata and A. bellabella but

not in other species. Allorchestes bellabella has

straighter posterior margins on epimera than do the

other species from the North Pacific. Allorchestes

carinata never develops more than one spine each

on the inner rami of uropods 1-2 but this character

is scarcely needed for identification in terminal

adults since carinations are so obvious. The presence

or absence of a stout spine mixed among the apical

setae of uropod 3 is of little value because apparently it is a phenotypic variable in A. angusta. The

adult shape of male gnathopod 2 varies from species

to species but has no value in identifying juveniles

because the latter have a female-like gnathopod.

Differences in female gnathopod 2 remain to be

confirmed; these differences are mainly of proportion and setosity and probably vary phenotypically

and with instar.
NUMBER 271 91

Allorchestes angusta Dana

FIGURES 50-52 (part)

Allorchestes angustus Dana, 1856:177.—J. L. Barnard, 1952a:

20-23, pi. 5: figs. 2-6; 1959:28.—Nagata, 1965:308—Reish

and Barnard, 1967:17—J. L. Barnard, 1974:42.

Allorchestes malleolus Stebbing, 1899:409-410, pi. 33A.—

Iwasa, 1939:285-288, figs. 20-22, pi. 20.—Bulycheva, 1957:

115-118, fig. 43.—Tzvetkova, 1967:183-184.

Allorchestes oculatus Stout, 1913:651.

Allorchestes vladimiri Derzhavin, 1937:95-96, pi. 5: fig. 2.—

Gurjanova, 1951:822, fig. 575.

Not Allorchestes angustus.—]. L. Barnard, 1954a:21-23, pi. 21

[ = A. bellabella].

DIAGNOSIS.—Mature male gnathopod 1 with hammer-shaped hand protruding at palmar corner,

dactyl much shorter than palm, weakly striate, not

heavily swollen, with bifid apical nail (or nail actually with outer accessory tooth), medial face of

hand with submarginal falcate spine divorced from

palmar corner, second spine simple, blunt, elongate, highly submarginal, set inward from middle

of palm and usually pointing anteriorly. Posteroventral protrusions of epimera 2-3 of medium extent. Body not dorsally carinate. Cuticle with ordinary bulbar setules, no plaques. Spines of pereopods

3-5 sharp, elongate.

DESCRIPTION.—See Figures 50-52 for generalities.

Right lacinia mobilis with 1-3 subbasal teeth (one

tooth shown in Figure 50: Jo). Hand of female

gnathopod 1 especially elongate and slightly curved.

Cuticle between bulbar setules smooth. Largest

specimen at hand, male, 9.3 mm, Yakutat Bay.

VARIABLES.—Apex of ramus on uropod 3 with 2-

4 apical setules and occasionally with short stout

spine, this character variable, apparently not correlated with adult body size nor geography (occurring both in Japan and California); castellations on

dactyls of pereopods increasingly numerous with

enlargement of body; simple spine of male gnathopod 1 hand especially elongate in large bodied

adults from high latitudes.

IDENTIFICATION.—Stebbing's original material of

Allorchestes malleolus from the Universitets Zoologiske Museum, Copenhagen, has been examined

in order to clarify his illustration of male gnathopod

1. That appendage fits the configuration of A.

angusta from the eastern Pacific and his various

Japanese specimens exhibit similar variations found

in eastern Pacific specimens; no subspecific distinctions have been found between the specimens from

the eastern and western Pacific, nor in specimens

from high latitudes, so that A. malleolus and A.

vladimiri must fall to A. angusta. Stebbing's erroneous view of male gnathopod 1 must be attributed to

a poor discernment of the dactylar apex which fits

the falcate palmar spine so closely that they often

appear as a unit.

LECTOTYPE.—Allorchestes malleolus Stebbing,

male, 8.3 mm, here selected. Deposited in Universitetets Zoologiske Museum, Copenhagen.

TYPE-LOCALITY.—34° 14'N, l29°34'E, Korlo-sho,

Andrea, 1869 [Tsushima, Japan].

VOUCHER MATERIAL.—Newport Bay, California,

male "a," 6.9 mm (illus.), female "n," 6.3 mm

(illus.), female "w," 6.3 mm; Laguna Beach, California, male "m," 7.8 mm (illus.); Yakutat Bay,

Alaska, male "u," 9.3 mm; Port Alice, British Columbia, male "y," 8.5 mm.

REMARKS.—Females and juveniles of this species

are probably difficult to distinguish from subadult

specimens of A. carinata but the problem does not

arise in materials at hand because of a sparsity of

samples and lack of juveniles in samples from the

known sympatric areas of the two species in high

latitudes.

A weak cline in adult body size apparently is

correlated with latitude or temperature or with

degree of enclosure of environment within lagoons

or bays, the species reaching sexual maturity at small

body sizes in southern warm latitudes or within enclosed bays. In southern California sexual maturity

appears to occur at a body length of about 4.0-4.5

mm, whereas in colder latitudes sexual maturity

may be reached at a body length of 6.0-7.0 mm

(observations minimal). Little if any modification

of spine shape occurs in cold water specimens so

that the specimens of A. carinata at hand can be

readily distinguished from A. angusta by their

heavily blunted pereopodal spines.

MATERIAL.—Stebbing's original material of

A. malleolus, Universitetets Zoologiske Museum,

Copenhagen: Japansker Haw, 9/3 1872, H. Koch

(3); Wladiwostock?, 9/3 1872, H. Koch (6); 3l°4O'N,

l25°5O'E, Tong-hai, F. Tang, Andrea 1869 (70);

Andrea 1869 (10); 34°I4'N, l29°34'E, Korlo-sho,

F. Tang [Tsushima, Japan]; 34°4O'N, l29°5O'E,

Japan, 50', Andrea 1869 (18); 37°O'N, l3l°2O'E,

Andrea 1869 (1); Smithsonian collections, full data

cataloged in museum: California: Laguna Beach,

Newport Bay, Pt. Fermin, Portuguese Bend, Venice,
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 50.—Allorchestes angusta Dana, male "a," 6.9 mm.
NUMBER 271

93

\Y

FIGURE 51.—Allorchestes angusta Dana, male "a," 6.9 mm (n - female "n," 6.3 mm; u — male

"u," 9.3 mm; y = male "y," 8.5 mm).
94

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 52.—Above: Allorchestes angusta Dana, male "a," 6.9 mm (m = male "m," 7.8 mm; w =

female "w," 6.3 mm). Below: Allorchestes bellabella J. L. Barnard, holotype, male "a," 12.4 mm

(j — juvenile, male "j, " 8.5 mm).

Morro Bay, Hazard Canyon, Monterey Bay, Berkeley Beach, Pt. Richmond, Oakland Estuary, Alameda, Bodega Bay; Washington: Meadow Point at

Seattle, Golden Gardens at Seattle, San Juan Island;

Canada: Port Alice, Vancouver Island; Alaska:

Saginaw Bay, Yakutat Bay; Ostrov Mednyy, 13

June 1906 [Medne Island, Commander Islands].

DISTRIBUTION.—Japan, about 3l°N, northward

through Kuriles, apparently across Aleutian chain

to Alaska, southward to California, about 32°N,

generally intertidal, phycophilous, rarely subtidal.

Allorchestes bellabella J. L. Barnard

FIGURES 52 (part), 53

Allorchestes bellabella J. L. Barnard, 1974:43.

Allorchestes angustus.—J. L. Barnard, 1954a:21-23, pi. 21 [not

Dana].

DIAGNOSIS.—Mature male gnathopod 1 with

weakly hammer-shaped hand bearing long tooth

projecting distally from palmar corner, dactyl overlapping palm, moderately to heavily striate, strongly

swollen, with simple, short apical nail, medial
NUMBER 271 95

FIGURE 33.—Allorchesles bellabella J. L. Barnard, holotype, male "a," 124 mm (J = juvenile "j ,

8. 5 mm; u = female "u," 12.6 mm).
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

face of hand with 1-2 medium to large spines,

terminal male with 1 large spine located posteriorly

but highly proximal to protrusion, second spine of

juvenile male apparently lost in adult, spines simple

(weakly striate); posteroventral protrusions of epimera weak to obsolete. Body dorsally with weak

posterior crest. Cuticle with complex alae surrounding bulbar setules, chitinous surface granular. Spines

of pereopods 3-5 blunt, short.

DESCRIPTION.—See Figures 52 and 53 for generalities. Following parts generally like those of A.

angusta (Figures 50-52): upper lip, lower lip, maxilliped, coxae; following parts like A. angusta with

modifications noted: setae of inner plate on maxilla

1 very tumid and palp thin (see Figure 53: VX1,

drawn to same scale as in A. angusta, Figure 50:

XI); outer plate of maxilla 2 with additional 2

medial setae, thus setae extending farther proximally; mandibles, but right lacinia mobilis with 3

fully developed subbasal teeth (also in largest specimens of A. angusta), main seta on molar vestigial in

male, fully developed on female (possibly male

aberrant); following parts similar to those of A.

carinata (Figure 54): pereopods 1-2 and their dactyls, inner setule of latter small as in A. carinata, 6

sets of sharp spines on article 6, including locking

spines; stoutness of pereopods 3-5 similar to A.

carinata; raker edges of basal gnathopodal articles

present as in A. carinata; female gnathopodal dactyls

with striae weaker than in A. carinata.

The distinctive pattern of cuticular pit-slits is

drawn fully on pereopod 3; female head and antenna

1 like male but antenna 2 (Figure 53: uA2) shorter

and thinner than in male; cuticle with alate slit-pits

especially prominent on article 2 of pereopods 3-5,

and the coxae, sparse elsewhere, granules or punctations very fine, occasional slightly enlarged prickle

spine emerging between alae.

HOLOTYPE.—USNM 140995, male "a," 12.4 mm

(illus.).

TYPE-LOCALITY.—Kyska Harbor, Alaska, beach,

W. H. Dall (presumably from sponge in 1873).

VOUCHER MATERIAL.—Type-locality, juvenile male

"j, " 8.5 mm (illus.). Bering Island, Alaska, female

"u," 12.6 mm (illus.).

MATERIAL.—Smithsonian collections, full data

cataloged in museum: Bering Island, 4 collections

by L. Stejneger, 29-30 December 1882 (22) (one

collection from crop of Simorhynchus pusillus,

which is old name of Aethia pusilla (Pallas), Least

Auklet); Izemleck Bay, Unimak Island, Alaska, from

duck stomach, 29 June 1925 (3); Kyska Harbor,

Alaska, beach, W. H. Dall (7).

DISTRIBUTION.—Bering Island, Alaska, to Cape

Arago, Oregon, intertidal and neritic ("planktonic").

Allorchestes carinata Iwasa, new status

FIGURE 54

Allorchestes malleolus carinatus Iwasa, 1939:288-289, figs. 23,

24, pi. 21.—Bulycheva, 1957:118-120, fig. 44.

DIAGNOSIS.—Mature male gnathopod 1 with ordinary or weakly hammer-shaped hand scarcely protruding at palmar corner, dactyl fitting palm,

weakly striate, not heavily swollen, with simple

apical nail, medial face of hand with 2 medium to

small spines resembling each other, 1 spine near

palmar corner, other spine situated submarginally

from middle of palm and pointing distalwards,

neither spine elongate. Posteroventral protrusions

of epimera 2-3 of medium extent. Body dorsally

carinate posteriorly. Cuticle with abnormal immersed setules bearing small plaque apically,

chitinous surface granular. Spines of pereopods 3-5

blunt, short.

DESCRIPTION.—See Figure 54 for generalities; following parts not illustrated and like those shown

for A. angusta (Figures 50-52): head, antennae 1-2,

upper lip, generalities of mandibles, lower lip,

maxillae 1-2, maxilliped, coxae and their marginal

setules, epimera 2-3; following parts similar to A.

angusta with stated exceptions: pereopods 1-2 with

spines relatively smaller in relation to thick and inflated articles, dactyl with numerous castelloserrations (15 or more), marginal dactylar seta relatively

shortened; article 2 of pereopod 3 similar but articles 4-6 stouter, article 2 of pereopods 4-5 slightly

broadened but posterior notch-setae more numerous;

flagellar articles of antenna 1 slightly more tumid;

epimeron 1 lacking posteroventral setule; inner

ramus of uropod 1 with only 1 spine, peduncle with

3 proximal spines, 3 peduncular spines of uropod

2 evenly separated from each other; female antennae

1-2 similar to those of male, thus stouter than in

A. angusta; dactyls of female gnathopods 1-2 with

fine striae unlike A. angusta; lacinia mobilis with

2 large and rudimentary third subbasal teeth.

REMARKS.—The sudden transformation of male

gnathopod 1 from the juvenile condition similar to
NUMBER 271

97

Glo G2o

FIGURE 54.—Allorchestes carinata Iwasa, male '1a," 15.9 mm (j =» juvenile "j,"6.3 mm; p =· female

"p," 15.6 mm).
98

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

that of the female to the male condition is demonstrated in 1 male specimen at hand preserved just

before ecdysis. The next instar is seen within the

old cast, the first gnathopod bearing 1 large new

spine replacing the 2 spines at the palmar angle

and a new large spine appearing on the medial subpalmar face; male gnathopod 2 is well developed at

this stage of transformation. The late C. R. Shoemaker noted these 2 stages of male gnathopod 1

as forms when he identified the material at hand.

The smallest juvenile at hand is 6.3 mm long

(from St. Paul Island) and, like adults, has evidence

of a dorsal crest, stout gnathopod 1, and blunt

pereopodal spines.

VOUCHER MATERIAL (all illustrated).—St. Paul

Island, Alaska, male "a," 15.9 mm; juvenile "j, " 6.3

mm; female "p," 15.6 mm

MATERIAL.—Universitetets Zoologiske Museum,

Copenhagen: Melne Bay, Simushir Island, Kuril

Islands, 23 June 1906, Steamer "Albatross" (1);

Smithsonian collection: St. Paul Island, Alaska,

C. H. Townsend (50); Bering Island, L. Stejneger

(3).

DISTRIBUTION.—Kurile Islands and Bering Sea,

presumably intertidal or very shallow water.

Hyale Rathke

This genus and Elasmopus in the Gammaridae

contain the most numerous individuals of intertidal

amphipods in the Californian-Mexican-Galapagan

area. Ericthonius (Corophiidae) is dominant also in

Mexico and the Galapagos Islands. The main taxonomic problem in this genus in the eastern Pacific

region has been Barnard's (1962c, 1969b) attempts

to find the affinities of H. frequens. He relegated the

species either to H. nigra or to H. rubra, but Barnard (1974:66-67) redescribed those 2 species from

Australia thereby demonstrating that H. frequens

is a valid species. Hyale rubra indeed occurs in the

eastern Pacific Ocean from Peru southward to Chile

and westward to the Juan Fernandez Islands. The

morph is slightly distinct from the Australian

morph and may eventually deserve a subspecific

name. The Hyale rubra found by Barnard (1969b:

138) from Isla Cedros is a distinctive species found

also in the Galapagos Islands, herein described as

H. darwini. Another new species, H. yaqui has very

close affinities with H. nigra from Australia.

The Hyale grandicornis complex is reviewed

again and the decision made to segregate all Pacific

taxa at specific level. This requires elevating H.

californica J. L. Barnard (1969b: 133) and H. rupi·

cola (see J. L. Barnard, 1974:54) and requires the

segregation of Iwasa's (1939:276) identification of

H. novaezealandiae to the status of Hyale sp. (p.

115).

Key to the Males of the Species of Hyale from Mexico, California, and

the Galapagos Islands

1. Uropod 1 with apicolateral peduncular spine on uropod 1 enlarged 2

Uropod 1 with apicolateral peduncular spine on uropod 1 not enlarged 9

2. Article 6 of gnathopod 1 with posterior setae widely spread 3

Article 6 of gnathopod 1 with posterior setae confined narrowly 7

3. Dactyl of maxillipeds with long setae 4

Dactyl of maxillipeds with short setae 6

4. Wrist of gnathopod 1 with midanterior spine H. canalina, new species

Wrist of gnathopod 1 lacking midanterior spine 5

5. Male gnathopod 2 with small hump on palm near hinge of dactyl, male gnathopod 1 with

posterior setae on article 6 divided into groups H. darwini, new species

Male gnathopod 2 with smooth palm, male gnathopod 1 with posterior setae on article 6

continuously and evenly set (eastern Pacific) H. rubra

6. Outer raraus of uropod 1 with dorsal spines, pereopods with 2 locking spines in tandem

H. yaqui, new species

Outer ramus of uropod 1 dorsally naked, pereopods with only 1 locking spine surrounded

by setae (Galapagan) H. zuaqut, new species

7. Coxae 2-4 with sharply extended posterior acclivity, outer ramus of uropod 1 dorsally

naked (Mexican) H. zuaque, new species

Coxae 2-4 with blunt, unextended posterior acclivity, outer ramus of uropod 1 with dorsal

spines 8
NUMBER 271 99

8. Locking spine of pereopods 3-5 very small but other anterior spines of article 6 enlarged,

male gnathopod 1 palm with large medial spine, palm of male gnathopod 2 fully occupying posterior margin of article 6 and densely setose H. guasave, new species

Locking spine of pereopods very small, other anterior spines of article 6 also small, male

gnathopod 1 palm lacking large medial spine, palm of male gnathopod 2 distinct from

posterior margin of article 6, spiny but not densely setose H. frequens

9. Main inner dactylar setule on pereopods 1-5 very thin 10

Main inner dactylar setule on pereopods 1-5 very thick, often striate 11

10. Posterior acclivity on coxa 1 situated in middle, large, male antenna 2 article 5 lacking

marginal posterior setae H. anceps

Posterior acclivity on coxa 1 situated dorsally, small, male antenna 2 article 5 bearing

marginal posterior setae H. plumulosa

11. Hand of male gnathopod 2 not tapering distally, cuticle smooth ....H. calijomica, new status

Hand of male gnathopod 2 strongly tapering distally, cuticle bearing weak polygons

H. humboldti, new species

Hyale darwini, new species

FIGURE 55

ETYMOLOGY.—Named for Charles Darwin.

DESCRIPTION OF MALE.—Anterior margin of lateral cephalic lobe slightly oblique, weakly concave.

Eyes of medium size, mostly formed of dark core

surrounded by 1-2 layers of clear ommatidia (in

alcohol). Antenna 2 about as long as head and

pereon together, peduncle slender, almost fully

naked, flagellum slightly less than 4 times as long as

peduncle, almost fully naked; antenna 1 about 45

percent as long as antenna 2, reaching less than 20

percent along length of flagellum of antenna 2,

aesthetascs 1-2 per article of flagellum. Upper lip

evenly rounded below, setules long. Mandibular incisors toothed, right lacinia mobilis evenly bifid and

with accessory side tooth, spine row with 1 large, 1

small spine, molar moderately triturative, right and

left molars with long plumose seta, left mandible

with 3 spines in spine row. Maxilla 1 as figured,

outer plate with 8 spines, inner plate softly acclivitous, 2 terminal setae short. Each terminal segment

of maxilliped with 1 or more lateral spines (setae),

article 3 of palp simple, apical spine of article 4

articulate, short, with 1 accessory setule and numerous elongate terminal setae and shorter proximal setae on inner margin. Posterior acclivities of

coxae 1-3 obsolescent and soft, 2 medial setae on

coxa 1 (lower part of plate), dorsoposterior excavation of coxa 4 not bisinuate or scarcely so. Article

2 of gnathopod 1 with 3 posterior spinules, of

gnathopod 2 with 4 spinules, article 2 of gnathopod

1 broadly lobate and protuberant distolaterally, article 4 with soft posterodistal protrusion, article 5

with 2 anterodistal spines, posterior lobe of article

5 large and broadly rounded, scarcely scalloped and

strongly spinose (setose), article 6 softly subrectangular but weakly tapering distally, posterior margin

with 5 setae in disjunct sets of 3, 1, 1, occupying 30

percent of marginal length, palm short, rounded

and weakly oblique, defined by 1 large medial and

1 small lateral spine, dactyl overlapping defining

spines, dactyl with medium development of minute

ridges; article 2 of gnathopod 2 grossly lobate and

weakly spinulose anteriorly, article 3 with nasiform

lobe, article 4 protrusively rounded apically, article

5 fully cryptic, hand of ordinary hyalid proportions,

with 3 anteroproximal spines on margin, palm oblique, slightly longer than posterior margin of hand,

latter with 1 acclivity bearing 2 short setae, palm

defined by weak hump and pocket for apex of

dactyl but enlarged spine absent, palm lined with

small stout spines, bearing weak quadrate protrusion near base of dactyl, latter fitting palm, with

weak hump near inner base. Pereopods ordinary,

like those of H. yaqui, pair of locking spines not

larger than other marginal spines on article 6, distalmost locking spine slightly shorter than proximalmost, distalmost scarcely striate, slightly bent, several setae between locking spines, dactyls minutely

striate, inner margins smooth, main inner seta very

small, thin and highly term¡nad. Uropod 1 with

enlarged apicolateral spine, apicomedial spine not
100

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICLRE 35.—Hyale darwini, new species, holotype, male "a," 4.9 mm (p = male "p," 4.5 mm;

a

1

 •» female "w," 4.0 mm).
NUMBER 271 101

enlarged, peduncles and rami of uropods 1-2 ordinarily spinose; uropod 3 ordinary but peduncle

slightly elongate, ramus short, peduncle with 2

apicodorsal spines, ramus with about 5 apical spines,

none disjunct from others. Telson split nearly to

base, lobes otherwise ordinary. Epimeron 1 ordinary, epimeron 2 with anterofacial ridge, medium

posteroventral sharp tooth, epimeron 3 with tooth

similar to but slightly stouter than that of epimeron

2. Cuticle with ordinary bulbar setules and surficial

matrix of widely scattered punctations.

FEMALE.—Body about three-fourths length of

male, antenna 2 reaching only three-fourths as far

as in male, peduncle even thinner than in male;

coxae 1-2 narrower relative to coxa 4 than in male;

article 2 of gnathopod 1 lacking posterior spines, of

gnathopod 2 with only 1 setule, dactyl with only a

few long ridges, hand narrower and more elongate

than in male, scarcely tapering distally, posterior

margin with setae similar to those of male, article

4 with long setae on strong apical protrusion;

gnathopod 2 like gnathopod 1 but slightly stouter,

posterior setae on article 6 composed only of group

of 3.

ILLUSTRATIONS. Following parts similar to those

shown for H. yaqui (Figures 57-59): head, lateral

views of antennae, epistome from lateral view, mandibles (but molarial seta present on both sides and

elongate), lower lip, maxilla 2, coxae 2, 3, pereopods 1-5 (but article 4 of pereopods 3—5 slightly

stouter), dactyls and locking spines, urosome, uropods 1, 2.

HOLOTYPE.—USNM 142488, male "a," 4.9 mm

(illus.).

TYPE-LOCALITY.—GAL 118, Galapagos Islands,

Isla Santa Cruz, Academy Bay, 4 February 1964,

intertidal, wash of alga.

VOUCHER MATERIAL.—Type-locality, male "p,"

4.5 mm (illus.); female "w," 4.0 mm (illus.).

RELATIONSHIP.—This species belongs with the.

group composed of H. rubra (Thomson), H. frequens (Stout), and H. yaqui, new species. It differs

from Australian H. rubra in the short nail on palp

article 4 of the maxilliped, in the discontiguous

setal groups on the posterior margin of article 6

on male gnathopod 1, and in the quadrate protrusion on the palm of male gnathopod 2.

Hyale darwini differs from H. frequens and H.

yaqui in the elongate apical setae of palp article 4

on the maxilliped, from H. frequens in the wider

spread of posterior setae on article 6 of male

gnathopod 1 and from H. yaqui in the narrower

spread, fewer setae and clear grouping of the posterior setae on article 6 of male gnathopod 1. Coxa

4 of H. darwini is especially broad and posteriorly

extended in comparison to H. frequens and H.

yaqui. The deep telsonic cleft of H. darwini contrasts to the shorter cleft of H. yaqui, and the

longer palm of H. darwini contrasts to the short

palm of gnathopod 2 in male H. frequens.

MATERIAL.—GAL 101, 108, 113, 114, 115, 116,

118, 119, 120; DAW 1, 3, 4, 5, 6, 7, 8, 9, 12, 13, 16,

17, 18, 19, 20, 21, 22, 23, 26, 27, 28, 29, 30, 31, 32,

33, 34, 35, 36, 37, 38, 40; PAN 14.

DISTRIBUTION.—Galapagos Islands, 0—6 m; Pacific

Panama, wharf.

Hyale rubra (Thomson)

FIGURE 56 (part)

Nicea rubra Thomson, 1879:236, pi. 10B: fig. 3.

Hyale rubra.—Stebbing, 1906:572.—Hurley, 1957:918-913, figs.

30-50.—J. L. Barnard, 1974:67-72, figs. 43-45.

?Hyale schmidti.—Iwasa, 1939:278-280, fig. 17, pi. 17.

Hyale species [cf. rubra].—J. L. Barnard, 1970:268-271, fig.

178.

Not Hyale rubra rubra.—J. L. Barnard, l969b:l38, fig. 24a-e

[= H. canalina, new species].

Not Hyale rubra frequens.—]. L. Barnard, 1969b: 139-141

[= H. frequens].

VOUCHER MATERIAL.—Antofogasta Bay, Chile,

male "h," 6.72 mm (illus.).

REMARKS.—Specimens from Chile, Peru, and the

Juan Fernandez Islands identified by C. R. Shoemaker have been compared minutely with J. L.

Barnard (1974:67), a depiction of H. rubra from its

type area, Australia. These specimens compare favorably with Australian specimens except that the

first proximal spine on the spinose margin of article

6 on pereopods 1-5 is larger than the distalmost

tandem-pair of locking spines and the posterior

row of setae on article 6 of male gnathopod 1 flows

evenly from proximal to distal end, with a slight

apical increase in the length of the setae. In Australian H. rubra, the proximal spines on the spinose

margin of article 6 on pereopods 1-5 are not enlarged and in terminal males the distalmost few

setae on article 6 of male gnathopod 1 are shorter

than the other setae. Reexamination of Australian

H. rubra reveals that young males develop 1 or

more distal setae near the defining spine fully dis-
102

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mW

FICIÌRE 56.—Hyale rubra (Thomson) (h = male "h," 6.72 mm). Hyale canalina, new species

(m = holotype, male "m," 6.33 mm).

junct from the posterior row and that as growth

proceeds these setae increase in number and eventually become contiguous with the main row.

These minute differences may have future value

in splitting the rubra complex into subspecies. The

main purpose of this analysis is to demonstrate

that H. frequens (Stout) and H. darwini are distinct from H. rubra. The former species has short

setae on the maxillipedal dactyl and the latter

species has a hump on the palm of male gnathopod

MATERIAL.— }uan Fernandez Islands, coll. W. L.

Schmitt, samples 34, 38, 59, 8-22 December 1926,

intertidal, one of these samples labeled "from tufts

of seaweed in Plagusia collected on weed covered

rocks along shore" (250+ ); Chile, Chanarel, 16

November 1926, W. L. Schmitt (2); Chile, Antofogasta Bay, 15 November 1926, W. L. Schmitt (male

"h" 6.72 mm, Figure 56) (30 + ); Peru, Independencia Bay, Velero 380-35, 14 January 1935, shore east

side of bay, W. L. Schmitt (20 + ). All material identified by Shoemaker.

DISTRIBUTION.—Species complex: Australia, Japan,

Hawaii, Juan Fernandez Islands, Peru, Chile.

Hyale canalina, new species

FIGURE 56 (part)

Hyale rubra rubra.—J. L. Barnard; 1969b: 138 [not Thomson].

DESCRIPTION OF MALE.—Anterior margin of lateral

cephalic lobe slightly oblique, weakly concave. Eyes

of medium size, completely and darkly pigmented.

Antenna 2 slightly longer than head and pereon

together, peduncle slender, almost fully naked,

flagellum about 4 times as long as peduncle, almost

fully naked; antenna 1 about 45 percent as long as
NUMBER 271 103

antenna 2, reaching less than 20 percent along

length of flagellum of antenna 2, aesthetascs 1-2

per article of flagellum. Upper lip evenly rounded

below, setu!es long. Mandibular incisors toothed,

right lacinia mobilis evenly bifid and with accessory side tooth, spine row with 1 large, 1 small

spine, molar moderately triturative, right and left

molars with long plumose seta, left mandible with

3 spines in spine row. Maxilla 1 as in Figure 55 of

Hyale darwini, new species, setae on inner plate

slightly more elongate. Maxilliped as in Figure 55

of H. darwini but article 3 of palp lacking medioproximal seta on face, apical spine of article 4 on

palp articulate, short, with 1 accessory seta and

numerous elongate apical setae and shorter setae

proximally on inner margin. Posterior acclivities of

coxae 1-3 obsolescent and soft, 2 medial setae on

coxa 1 (lower posterior part of plate), dorsoposterior

excavation of coxa 4 weakly bisinuate. Article 2 of

gnathopod 1 with 3-4 posterior spinules, of gnathopod 2 with 4 spinules, article 2 of gnathopod 1

broadly lobate and protuberant distolaterally, article 4 with soft posterodistal protrusion, article 5

with 5 apicodorsal spines and 1 midanterior spine,

posterior lobe of article 5 large and broadly rounded,

scarcely scalloped and strongly spinose, article 6

softly subrectangular but weakly tapering distally,

posterior margin with 9 setae in contiguous sets

of 3-1-2-3 or 4-3-2 (proximal to distal), occupying

35+ percent of marginal length, palm short,

rounded and oblique, defined by 1 small medium

and 1 larger lateral spine, dactyl overlapping defining spines, dactyl with medium development of

minute ridges; article 2 of gnathopod 2 grossly

lobate and weakly spinulose anteroventrally, article

3 with nasiform lobe, article 4 protrusively rounded

apically, article 5 fully cryptic but with sharp

posterior protrusion, hand stout, with 1-2 anteroproximal spines on margin, palm oblique, slightly

longer than posterior margin of hand, latter with

1 setule, palm defined by weak hump and pocket

for apex of dactyl but enlarged spines absent, palm

lined with small stout spines, bearing weak quadrate

protrusion near base of dactyl, latter fitting palm,

with weak hump near inner base. Pereopods ordinary, like those of Hyale yaqui, pair of locking

spines not larger than other marginal spines on

article 6, locking spines of identical size, both

weakly striate, unbent, several setae between locking spines, dactyls minutely striate, inner margins

with few very weak castellations proximally, stronger

on pereopods 1-2, main inner seta very small, thin

and highly terminad. Uropod 1 with enlarged

apicolateral spine on peduncle, apicomedial spine

not enlarged, peduncles and rami of uropods 1-2

ordinarily spinose; uropod 3 ordinary but peduncle

slightly elongate, ramus short, peduncle with 2

apicodorsal spines, ramus with 5 apical spines, not

disjunct from others. Telson split nearly to base,

lobes otherwise ordinary. Epimeron 1 ordinary,

epimeron 2 with anterofacial ridge, weakly protuberant posteroventrally, epimeron 3 with similar

protuberance but slightly sharper than that of

epimeron 2. Cuticle with ordinary bulbar setules

and surficial matrix of widely scattered punctations.

FEMALE.—Unknown.

HOLOTYPE.—AHF 5928, male "m," 6.33 mm

(illus.).

TYPE-LOCALITY.—J. L. Barnard, Sta. 36, Isla

Cedros, Baja California, 3.2 km SE of north point,

21 March 1959, in intertidal wash of Phyllospadix

sp.

VOUCHER MATERIAL.—AHF Velero III 1370-41,

Santa Catalina Island, 3 males.

RELATIONSHIP.—This species may prove to be a

subspecies of Hyale darwini, or, that species, this

species and H. rubra may form an eastern Pacific

Rassenkreis. The three species are minimally distinct in the sparse material in hand from essentially

3 areas, Peru and Juan Fernandez (rubra), the

Galapagos Islands (darwini), and Cedros-Catalina

(canalina), the offshore islands of the Californias.

Hyale canalina forms an intergrade between H.

rubra and H. darwini. Hyale canalina bears 1 or 2

spines on the anteroproximal margin of the hand

on male gnathopod 2, whereas H. rubra lacks these

spines and H. darwini bears 3. Hyale canalina bears

about 9 posterior setae on the hand of male gnathopod 1 very slightly divided into groups of 1-4

whereas these setae are unifromly set in H. rubra

and divided into fully disjunct groups in H. danvini.

Hyale canalina differs from either H. rubra or

H. darwini in the presence of a midanterior spine

on the wrist of gnathopod 1.

MATERIAL.—See holotype and voucher material

(4 specimens).

DISTRIBUTION.—Santa Catalina Island and Isla

Cedros, intertidal.
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hyale yaqui, new species

FICURES 57-59 (part)

Hyale nigra, bay form.—J. L. Barnard, 1964a: 109, fig. 21A

[not Haswell, nor other references to H. nigra by J. L.

Barnard].

Hyale rubra frequens.—J. L. Barnard, l969a:2!2 [not Stout,

nor other references to H. rubra or H. frequens by J. L.

Barnard].

IDENTIFICATION.—Material noted in synonymy

from Bahía de Los Angeles, Bahía de San Quintín,

and Estero de Punta Banda is referred to this

species.

DESCRIPTION OF MALE.—Anterior margin of

lateral cephalic lobe slightly oblique, weakly concave. Eyes of medium size, mostly formed of deep

purple to black core surrounded by 2 layers of

clear ommatidia (in alcohol). Antenna 2 about as

long as head and pereon together, peduncle slender,

almost fully naked, flagellum slightly less than 4

times as long as peduncle, almost fully naked; antenna 1 scarcely more than half as long as antenna

2, reaching scarcely 30 percent along length of

flagellum on antenna 2, aesthetascs about 2 per

article on flagellum. Upper lip evenly rounded below, setules short. Mandibular incisors toothed,

right lacinia mobilis evenly bifid and with accessory

side tooth, spine row with 1 large, 1 small spine,

molar moderately triturative, right molar with short

plumose seta; left mandible with 3 spines in spine

row, long molarial seta. Maxillae as illustrated,

outer plate of maxilla 1 with 9 spines (some hidden

in figures), inner plate of maxilla 2 with 1 enlarged

inner seta. Each terminal segment of maxilliped

with 1 or more lateral spines, article 3 of palp

simple, apical spine of article 4 articulate, short,

with 1 accessory apical spine and several inner marginal spine-setae and 1 outer seta. Posterior acclivities of coxae 1-3 obsolescent and soft, setae on inner

faces vestigial, dorsoposterior excavation of coxa 4

scarcely bisinuate. Article 2 of gnathopod 1 with 5

posterior spinules, of gnathopod 2 with 6 or 7 small

spines, article 2 of gnathopod 1 broadly lobate distolaterally, article 4 with soft posterodistal protrusion, article 5 with several anterodistal spines

on protrusion set back from sleeve holding article 6,

posterior lobe of article 5 large and broadly

rounded, weakly scalloped and strongly spinose,

article 6 softly subrectangular but weakly tapering

distally, posterior margin with long row of setae

occupying nearly 40 percent of marginal length,

palm short, rounded and weakly oblique, defined

by 1 large lateral and 1 small medial spine, dactyl

fitting palm but slightly overlapping defining spines,

dactyl with medium development of minute ridges;

article 2 of gnathopod 2 grossly lobate and weakly

spinulose anteriorly, article 3 with nasiform lobe,

article 4 protrusively rounded apically, article 5 with

weak but cryptic lobe on medial surface, hand of

ordinary hyalid proportions, with 1 anteroproximal

spine on margin, palm oblique, slightly longer than

posterior margin of hand, latter almost naked, palm

defined by weak hump, spine and pocket for apex

of dactyl, palm lined with small stout spines, bearing weak quadrate protrusion near base of dactyl,

latter fitting palm, scarcely humped near inner base.

Pereopods ordinary (Figure 58) pair of locking

spines not larger than other marginal spines on

article 6, d¡stalmost locking spine slightly shorter

than proximalmost, distalmost weakly striate spirally, slightly bent, several setae between locking

spines, dactyls minutely striate, with obsolescent

inner notches, main inner seta very small, thin and

highly terminad. Uropod 1 with enlarged apicolateral spine, apicomedial spine not enlarged,

peduncles and rami of uropods 1-2 ordinarily

spinose; uropod 3 ordinary but peduncle slightly

elongate, ramus short, peduncle with 2 or 3 apicodorsal spines, ramus with about 6 apical spines,

none disjunct from others. Telson split only about

two-thirds towards base, lobes otherwise ordinary.

Epimeron 1 ordinary, epimeron 2 with anterofacial ridge, medium posteroventral sharp tooth,

epimeron 3 with tooth similar to that of epimeron 2.

Cuticle with split pits especially well developed on

coxae and article 2 of pereopods 3-5, pits with setule

and shaped like sagittal views of medusae with

tiny inserted setules on surface between pits, epimera rarely with pits but mainly with tiny inserted

setules, occasional pit on body segments dorsally

and appearing as craters from oblique views.

FEMALE.—Body about two-thirds length of male

(4.2 against 6.2 mm maximum sizes in material

available), antenna 2 reaching only two-thirds as

far as in male, peduncle even thinner than in male;

coxae 1-2 narrower relative to coxa 4 than in male;

article 2 of gnathopod 1 lacking posterior spines,

of gnathopod 2 with only 1 setule, dactyl with only

a few long ridges, hand narrower and more elongate

than in male and not tapering apically, posterior
NUMBER 271

105

FIGURE 57.—Hyale yaqui, new species, male holotype "a," 6.2 mm (n = female "n," 4.2 mm;

u — male "u," 4.2 mm).
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 58.—Hyale yaqui, new species, holotype, male "a," 6.2 n`m (n = female "n," 4.2 mm).
NUMBER 271

107

aE2

FIGURE 59.—Hyale yaqui, new species, holotype, male "a," 6.2 mm (n = female "n," 4.2 mm).

Below: Hyale zuaque, new species, holotype, male "b," 5.3 mm (a = female "a," 4.46 mm).

margin with setae divided into 3 groups of 1-2

each, sleeve of article 5 poorly developed, article 4

with long setae on apical protrusion; gnathopod 2

like gnathopod 1 but slightly larger, posterior setae

on article 6 (hand) more closely contiguous but

groups discernible.

REMARKS.—Specimens from KNO 1, Bahía Kino,

have the spines on the palm of male gnathopod 2

much longer than in the typical series and the locking spines on pereopods 1-2 have a stronger corkscrew pattern, but those on pereopods 3-5 are like

the typical series.

HOLOTYPE.—USNM 149451, male "a," 6.2 mm

(illus.).

TYPE-LOCALITY.—SCO 10, Gulf of California,

Puerto Peñasco, 23 February 1971, inter tidal, wash

of Sargassum sp.

VOUCHER MATERIAL.—Type-locality, female "n,"

4.2 mm (illus.); male "u," 4.2 mm (illus.).

RELATIONSHIP.—This species differs from its

nearby allopatriot and probable sibling, Hyale

frequens (Stout), in the wider setal distribution on

the posterior margin of article 6 on gnathopod 1,

on which H. frequens has several setae tightly compacted and occupying far less of the posterior margin; the division of these setae into groups on

female gnathopod 1 is another distinction because

males and females of H. frequens are alike in their

compact setal bundle. The protrusion on the palm

of male gnathopod 2, though weak, is better defined

in H. yaqui than it is in H. frequens.

Hyale yaqui has article 5 of antenna 2 on the

male relatively longer and article 4 relatively shorter

than in H. frequens. Pereopodal spines are generally

sharper in H. yaqui, the palmar protrusion on

gnathopod 2 is weakly larger and female gnathopod

2 has 3 sets of posterior setae on article 6 in contrast

to 2 sets in H. frequens.

Hyale nigra (Haswell) from Australia is another

sibling of this species group and differs from H.

yaqui in the simplicity and straightness of the distalmost locking spine on pereopods 1-5 and in the

smaller tooth on epimeron 2.

MATERIAL.—SCO 1, 5, 7, 8, 10, 11, 12, 13, 14,

15, 16, 17, 18, 19, 21, 22; KNO 1, 2; TOP 3; PAZ

2, 3, 10, 12, 13, 16, 17, 18, 20, 24; BRU 1, 2.
108

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Gulf of California: Puerto Peñasco, Bahía de Los Angeles, Bahía Kino, Bahía San

Carlos, Guaymas, Topolobampo, Bahía Concepción,

Bahía San Evaristo, Isla San Francisco, Isla Espiritu

Santo, La Paz, and Cabo San Lucas, intertidal to

7 m; outer Baja California, Bahía de San Quintín,

and Estero de Punta Banda, intertidal.

Hyale zuaque, new species

FIGURES 59 (part), 60, 61

DESCRIPTION OF MALE.—Anterior margin of

lateral cephalic lobe vertical. Eyes of medium size,

dark core surrounded by 1-2 layers of clear ommatidia (in alcohol). Antenna 2 about as long as head

and half of pereon together, peduncle of medium

stoutness, articles 4-5 with medioposterior setal

brushes, flagellum about twice as long as peduncle,

proximal articles with medial setal brushes; antenna 1 about two-thirds as long as antenna 2,

aesthetascs about 2 per article on flagellum. Upper

lip evenly rounded below, setules short. Mandibular

incisors toothed, right lacinia mobilis evenly bifid

and with blunt accessory tooth, spine row with 4-5

spines, molar moderately triturative, with long

plumose seta (broken off on left illustration and

usually broken on left side). Maxilla 1 as in Figure

6O:mXl, outer plate with 9 spines, inner plate of

maxilla 2 with 1 enlarged inner seta (like Figure

57:X2 of H. yaqui but outer plate slightly narrower).

Each terminal segment of maxilliped with 1 lateral

spine, article 3 of palp simple, apical spine of

article 4 articulate, short, with numerous inner

marginal spine-setae, inner plate with only 1 medial

seta and various setules. Posterior acclivities of

coxae 1-3 strong, blunt, margin below posterior

acclivity of coxa 4 sinuous, dorsoposterior excavation not bisinuate. Article 2 of gnathopod 1 with

2 posterior spines, of gnathopod 2 with 3 spines,

article 2 of gnathopod 1 thick but not lobate distolaterally, article 4 with subsharp posterodistal protrusion, article 5 with 1 anterior spine on distal

hump, posterior lobe ordinary, article 6 softly subrectangular, posterior margin with short row of setae

occupying about 30 percent of marginal length,

palm oblique, defined by 2 medium spines (lateral

and medial), dactyl scarcely overlapping palm, with

medium development of minute ridges; article 2 of

gnathopod 2 grossly lobate and naked, article 3 with

weakly nasiform lobe, article 4 protrusively rounded

apically, article 5 with weak naked lateral lobe and

subcryptic setose medial lobe, hand of ordinary

hyalid proportions, anteriorly naked, palm oblique,

much shorter than posterior margin of hand, latter

almost naked, palm defined by weak hump, 2 spines

and small pocket for apex of dactyl, palm lined

with cycles of long and short spines, bearing extremely weak protrusion near dactylar hinge (visible

only medially), dactyl fitting palm, not humped.

Pereopods ordinary (Figure 61) bearing only one

striate locking spine, this spine clavate and truncate

on pereopods 1-2, sharp on pereopods 3-5, smaller

than other spines on pereopods 3-5 but unknown

on pereopods 1-2 as latter without inflexible spines

on article 6, only flexible setae, dactyls minutely

striate, with weak inner castellations, main inner

seta long and of medium thickness. Uropod 1 with

enlarged apicolateral spine, ap¡comedial spine not

enlarged, outer rami of uropods 1-2 lacking dorsal

spines; uropod 3 ordinary, peduncle with 3 apicodorsal spines, apex of ramus with about 6 spines,

none disjunct from others. Telson split fully to base,

lobes triangular, lateral margins weakly concave,

apices thus subsharply extended. Epimeron 1 ordinary, epimeron 2 with subvertical anterofacial ridge,

broad and blunt posteroventral protrusion and

broad strongly extended anteroventral lobe, epimeron 3 with weakly convex posterior margin,

medium and blunt posteroventral tooth. Urosome

exceptionally tall and short relative to ordinary

species of Hyale. Cuticle with ordinary bulbar

setules and occasional minute punctation.

FEMALE.—Antenna 2 shorter and much thinner

than in male; setal brushes weaker; coxa 2 relatively

narrower than coxa 2 of male; gnathopod 1 like

that of male, dactylar ridges weak, gnathopod 2

similar to gnathopod 1 but article 2 with gross

distolateral lobe, article 5 with posterior lobe narrower than on gnathopod 1.

ILLUSTRATIONS.—Female antenna 2 (Figure 60:

hA2) drawn in same proportions as view of male

antenna 2 (Figure 60: C) attached to head; telson of

specimen "r" flattened, of holotype male unflattened.

HOLOTYPE.—USNM 142511, male "b," 5.3 mm

(illus.).

TYPE-LOCALITY.—PAZ-22, Baja California, 11 km

E of Cabo San Lucas, 4 December 1971, intertidal,

algal wash.

VOUCHER MATERIAL.—Type-locality, female "h,"

4.45 mm (illus.); male "m," 5.26 mm (illus.).
NUMBER 271 109

Glo

FIGURE 60.—Hyale zuaque, new species, holotype, male "b," 5.3 mm (h = female "h," 4.46 mm;

m =» male "m," 5.26 mm).
110

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 61.—Hyale zuaque, new species, holotype, male "b," 5.3 mm (h = female "h," 4.46 mm;

m = male "m," 5.26 mm).
NUMBER 271 111

RELATIONSHIP.—This species differs from Hynle

plumulosa Stimpson in the lateral position of the

enlarged spine on the peduncle of uropod 1 and

the absence of dense setular fringe in the male on

article 5 of the peduncle of antenna 2.

Hyale iole J. L. Barnard (1970:260) from the

Hawaiian Islands bears superficial resemblance to

H. zuaque because of setose antenna 2 and presence

of only 1 locking spine on the pereopods. Hyale iole

has the main enlarged spine of uropod 1 on the

mediodistal side, the main dactylar seta of the

pereopods is thick, and the lobes of the telson are

subquadrate.

Hyale ayeli J. L. Barnard (1970:254) is also in

this species complex but the locking spine of

pereopods I-2 is not apically truncate and the palm

of male gnathopod 2 has a conspicuous hump; the

absence of spines on the outer rami of uropods 1-2

and the enlargement of the distolateral spine on

the peduncle of uropod 1 suggest that H. ayeli and

H. zuaque are a sibling pair of species.

A few terminal males from the Galapagos have

the setae on the posterior margin of article 6 on

gnathopod 1 as widely spread as in H. yaqui, but

medium-sized males have the normal spread of setae

seen in Mexican specimens.

MATERIAL.—PAZ 14, 15, 21, 22; DAW 2, 10;

ECU 3; BRU 2.

DISTRIBUTION.—Gulf of California: Guaymas;

Cabo San Lucas; Galapagos Islands; Ecuador;

intertidal.

Hyale guasave, new species

FIGURES 62, 63

DESCRIP'TON OF MALE.—Anterior margin of

lateral cephalic lobe truncate, vertical. Eyes of

medium to large size, formed of dark core surrounded by 2 layers of clear ommatidia (in alcohol).

Antenna 2 scarcely as long as pereon alone, peduncle

slender, almost fully naked, flagellum less than

3 times as long as peduncle, with tufts of very short

setae on each article; antenna 1 about two-thirds

as long as antenna 2, reaching halfway along flagellum of antenna 2, aesthetascs 1-2 per article on

flagellum. Upper lip evenly rounded below but

unusually narrow for genus, setules long. Mandibular incisors toothed, right lacinia mobilis evenly

bifid and with accessory side tooth, spine row with

1 large, 1 small spine, molar triturative, with long

plumose seta; left mandible with 3 spines in spine

row, no molarial seta (unless rudimentary) (mandibles like those of H. yaqui as in Figure 57 with

exceptions noted). Following parts like Figures 57-59

for H. yaqui: lower lip, maxilla 2; maxilla 1 like

H. yaqui but spines of outer plate more densely

toothed (Figure 62.X1) and palp with 2-5 small

tubular truncate setules (?spouts) proximal to main

terminal spine. Maxilliped like H. yaqui but article

3 of palp broader (Figure 62:VS), and main spines

on inner plate pointed (Figure 62:Si). Posterior acclivities of coxae 1-3 strongly quadrate, only coxa

1 with 2 spines on inner face, dorsoposterior excavation of coxa 4 strongly bisinuate. Article 2 of

gnathopod 1 with 3 midposterior spines (2 sets), of

gnathopod 2 with 3 spines in 2 sets and 1 seta, article 2 of gnathopol 1 broadly lobate distolaterally,

article 4 with sharp posterodistal protrusion, article

5 ordinary, article 6 enlarged, broadened, with

hump defining expanded palm, hump bearing

stridulation ridges, posterior margin of article 6

with 2 weakly separated groups of several setae occupying about 25 percent of margin length, dactyl

failing to fit palm, middle of palm with 1 enlarged

submarginal spine, defining hump with second enlarged spine, dactyl with weakly defined inner

ridges; article 2 of gnathopod 2 broadly lobate and

bearing marginal spinules, article 3 with nasiform

lobe, article 4 protrusively rounded apically, article

5 cryptic, hand elongate, palm and posterior margin

of hand confluent and setose throughout, medial

margin with several rows of shorter setae, no spines

or spinules, dactyl nearly as long as palm and fitting

into pocket of hand near proximal end of palm,

latter weakly undulating but lacking defined protrusions. Pereopods ordinary (Figure 62), distalmost

locking spine small, next proximal spine large and

of size similar to other marginal spines of article 6,

spines all deeply striate in corkscrew fashion, numerous setae between locking spines but setae otherwise absent on article 6, dactyl minutely striate,

inner margins smooth, main inner seta very small,

thin. Uropod 1 with enlarged apicolateral spine,

apicomedial spine not enlarged, peduncles and rami

of uropods 1-2 ordinarily spinose; uropod 3 ordinary but ramus elongate, about three-fourths as

long as peduncle, bearing about 6 apical spines,

none disjunct from others. Telson split nearly to

base. Epimeron 1 ordinary, epimeron 2 with antero-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

vs

FIGURE 62.—Hyale guasave, new species, holotype, male "a," 4.8 mm (n = female "n," 4.0 mm;

u — male "u," 4.7 mm).
NUMBER 271 113

FIGURE 63.—Hyale guasave, new species, holotype, male "a," 4.8 mm (n

u = male "u," 4.7 mm).

female "n," 4.0 mm;

facial ridge tightly submarginal towards anterior

edge, posteroventral tooth obsolescent, epimeron 3

with medium sized posteroventral tooth. Cuticle

with very weak slit-pits, each bearing setule.

FEMALE.—Body about four-fifths as long as male,

antenna 2 scarcely shorter than in male, peduncle

thinner than in male; coxae 1-2 narrower relative

to coxae 4 than in male; article 2 of gnathopods 1-2

lacking posterior spines of male, gnathopods of

ordinary female dimensions, posterior margins of

hands on gnathopods with group of 4 or 5 setae, no

enlarged midpalmar spine, palm defined by large

spine, no hump, dactyl fitting palm, weakly striate.

ILLUSTRATION.—Female antenna 2 magnified to

similar dimension as antenna 2 of male shown on

head (Figure 62).

HOLOTYPE.—USNM 142502, male "a," 4.8 mm

(illus.).

TYPE-LOCALITY.—PAZ 22, Baja California, 11 km

E of Cabo San Lucas, 4 December 1971, intertidal,

algal wash.

VOUCHER MATERIAL.—Type-locality, female "n,"

4.0 mm (illus.); male "u," 4.7 mm (illus.).

RELATIONSHIP.—This species belongs with a

group formed of H. macrodactyla Stebbing, H.

chevreuxi K. H. Barnard, and H. honoluluensis

Schellenberg; these species are distributed in the

tropical Indo-Pacific region. Hyale guasave appears

to be closest to H. chevreuxi because of the distinct

hump defining the palm of male gnathopod 1, a

characteristic poorly developed on the other 2 species. Hyale guasave apparently differs from all other
114

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

species in the group in the presence of an enlarged

midpalmar spine on male gnathopod 1, similar to

that seen in species of Allorchestes and ParhyaJe.

This eastern Pacific species is, therefore, more distinctive than are H. chevreuxi and H. honoluluensis

between themselves, and indeed those may be

synonymous and actually be based on clines within

characters. Hyale chevreuxi, type area in the Seychelles, has a strong hump defining the palm of

gnathopod 1, whereas H. honoluluensis, type area

in Hawaii, has that hump reduced in prominence;

the two species are found sympatrically in Micronesia. Hyale macrodactyla is found in the Caribbean

Sea but appears less similar to nearby H. guasave

than does H. chevreuxi, and H. macrodactyla lacks

the palmar hump of male gnathopod 1 and has a

midhump on the palm of male gnathopod 2 and

apparently has but a single row of setae on the

palm of male gnathopod 2.

MATERIAL— PAZ 22; DAW ? 10, ? 17, 31.

DISTRIBUTION.—Baja California: Cabo San Lucas

intertidal; Galapagos Islands, 0-6 m.

Hyale frequens (Stout)

Allorchestes frequens Stout, 1913:650-651.

Hyale frequens.—Shoemaker, 1941b: 187; 1942:17.—Hewatt,

1946:199.—J. L. Barnard, l952a:23; l954a:23.

Hyale nigra.—]. L. Barnard, 1962c: 153-156, figs. 19, 20 [not

Haswell].

Hyale rubra frequens.—J. L. Barnard, 1969b: 139-141 [not

Thomson].

Not Hyale nigra, bay form.—J. L. Barnard, 1964a: 109, fig.

21A [not Haswell].

Not Hyale rubra frequens.—]. L. Barnard, l969a:2l2 [not

Stout].

RELATIONSHIP.—This species is neither identical

with, nor a subspecies of Hyale rubra (Thomson)

and H. nigra (Haswell). J. L. Barnard (1974:67) has

reexamined those species from Australia and found

them to be distinct from each other.

DISTRIBUTION.—Hyale frequens has not been

found in the collections of the present study and

may have its southern limit somewhere on the outer

coast of Baja California.

The rejected references in the synonymy are referred to Hyale yaqui, new species.

Hyale anceps (J. L. Barnard)

Allorchestes anceps J. L. Barnard, 1969b: 130-133, figs. 25, 26.

Hyale anceps.—]. L. Barnard, 1974:42.

DISTRIBUTION.—California: Carmel to Cayucos.

intertidal.

Hyale plumulosa (Stimpson)

Allorchestes plumulosus Stimpson, 1857:79.

Hyale plumulosus.—Thorsteinson, 1941:55-56, pi. 1: figs. 10-15.

Hyale plumulosa.—]. L. Barnard, 1969b: 138.—Bousfield, 1973:

155-156, pi. 44.2.

DISTRIBUTION.—Western Atlantic, Maine to North

Carolina; Eastern Pacific, southern Alaska to

southern California.

The Hyale grandicornis Complex

The grandicornis group of Hyale comprises

species with an immense inner setule or seta on the

dactyl of pereopods 1-5 close to the middle of the

margin. The eyes are very large, dark, and oblatovate. Uropod 1 has the apicolateral spine on the

peduncle either small or medium sized and the

apicomedial spine also middle sized, in neither case

is the spine enlarged. This eliminates H. longicornis

(Haswell) from the discussion.

This discussion concerns only those species of the

group in the Pacific Ocean, those having been described from Chile, New Zealand (2 species), Japan,

Hawaii, California, and, now, the Galapagos Islands.

The taxa are to be rectified herein nomenclaturally

so that each is established at full specific level. This

would appear to be the best way to treat these taxa

until much more extensive exploration and study of

morphs can be undertaken to determine whether

certain clusters are better reduced to a subspecific

level.

The first species described, Hyale grandicornis

(Krøyer, 1845) from Chile, was redescribed by

Stephensen (1949:33). It and Hyale thomsoni Hurley

(formerly known as Hyale grandicornis forma

thomsoni Hurley, 1957, here elevated to specific

level) from New Zealand differ from other Pacific

species in the presence of "fur" on pereopod 4, in

the absence of a disjunct subapical spine on the

ramus of uropod 3, and in the reduction or absence of the dorsal spination on the outer rami of

uropods 1-2. Hyale grandicornis also has a sharply

posteroventral lobe on pereopod 5 but this is

unknown in H. thomsoni.

All other Pacific species lack "fur" on pereopod

4, have a rounded lobe on article 2 of pereopod 5,

bear 2 or more dorsal spines on the outer rami of
NUMBER 271 115

uropods 1-2 and have at least 1 disjunct proximal

spine on the ramus of uropod 3.

The following species appear to form a group

characterized by normal spacing of dorsolateral

spines on the peduncle of uropod 1: Hyale novaezealandiae (Thomson, 1879, redescribed by Hurley,

1957, but formerly known as Hyale grandicornis

forma novaezealandiae) from New Zealand, and

Hyale bishopae J. L. Barnard, 1955a (formerly

known as H. grandicornis bishopae in J. L. Barnard,

1970) from Hawaii. These two species are very difficult to separate but apparently differ in the castellations on the pereopodal dactyls and the presence of

a supernumerary facial spine on the ramus of

uropod 3 in H. novaezealandiae. Hyale rupicola

(Haswell) (J. L. Barnard, 1974:54) is also indistinguishable from H. novaezealandiae as reflected

in the following key.

The final group is characterized by the extreme

discontiguity of dorsolateral spine 2 from spine 1

(apicalmost) on the peduncle of uropod 1: Hyale

californica J. L. Barnard, 1969b (formerly known as

H. grandicornis californica) from California, Hyale

species (formerly identified as H. novaezealandiae

by Iwasa, 1939) from Japan, and Hyale hurnboldti,

new species, from the Galapagos Islands. These species differ among themselves in castellations on dactyls of pereopods and a supernumerary facial spine

on the ramus of uropod 3 as shown in the key to

follow.

The 7 species form a Rassenkreis divisible into 3

parts, between Chile and New Zealand, between

New Zealand and Hawaii, and among Japan, California, and the Galapagos Islands. Study of collections from intermediate localities is now required to

demonstrate the validity of these species.

Key to the Males of the Species in the Hyale grandicornis Complex

from the Pacific Ocean

1. All spines on ramus of uropod 3 terminal 2

At least 1 spine on ramus of uropod 3 set subterminally and disjunctly from other spines

4

2. Outer rami of uropods 1-2 with 0 or 1 spine, articles 2 and 4 of pereopod 4 with posterior

"fur" 3

Outer rami of uropods 1-2 with 2+ spines, pereopod 4 lacking "fur" H. crassicornis

3. Uropods 1-2 lacking dorsal spines on outer rami H. grandicornis

Uropods 1-2 bearing dorsal spine on outer rami H. thomsoni

4. Gap between distalmost lateral spine on peduncle of uropod 1 and next proximal spine

about 1.6 times distance between spines 2 and 3 5

Gap between distalmost lateral spine on peduncle of uropod 1 and next proximal spine

about 2.3 times or greater distance between spines 2 and 3 6

5. Dactyls of pereopods 1-5 with minute castellations or pectinations

H. rupicola and H. novaezealandiae

Dactyls of pereopods 1-5 lacking pectinations H. bishopae

6. Dactyls of pereopods 1-5 castellate, hand of male gnathopod 2 strongly tapering distally

7

Dactyls of pereopods 1-5 not castellate, hand of male gnathopod 2 not tapering distally

H. californica, new status

7. Uropod 3 with supernumerary inner, midlevel facial spine on ramus

H. humboldti, new species

Uropod 3 lacking supernumerary inner facial spine besides ordinary subdistal spine

H. species

Hyale species

Hyale novaezealandiae.—Iwasa, 1939:276-278, fig. 16, pi. 16

[not Thomson].

This speciman is not H. novaezealandiae and according to notes of the late C. R. Shoemaker (USNM

files) is probably Hyale pugettensis (Dana), which is

probably synonymous with Allorchestes japonica

Stimpson. This taxon is included in the key to the

species of the Hyale grandicornis complex as Hyale

species.
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hyale californica J. L. Barnard, new status

Hyale grandicornis californica J. L. Barnard, 1969b: 133-138,

figs. 27, 28.

DESCRIPTION OF MALE.—Anterior margin of

lateral cephalic lobe truncate vertically. Eyes very

large, ommatidia all darkly pigmented. Antenna 2

only as long as first 4 pereonites together, peduncle

of medium stoutness, minimally setose, flagellum

about 1.35 times as long as peduncle, with tufts of

very short setae on each article; antenna 1 about

0.83 times as long as antenna 2, reaching about onefourth along flagellum of antenna 2, aesthetascs

about 2 per article on flagellum. Upper lip evenly

rounded below, broad, setules long. Mandibular

incisors toothed, right lacinia mobilis evenly bifid

and with accessory side tooth, spine row with 2

spines and 1 setal plume, these plumes filamentous,

molar triturative, with long plumose seta, left mandible with 4 raker spines and plume, seta on molar

present. Lower lip and maxilla 1 like H. yaqui but

mandibular lobes of lower lip broader, shape of

maxilla 2 as in H. humboldti, spines on outer plate

of maxilla 1 more densely toothed as in H. guasave,

palp apparently with 1 vestigial spout. Maxilliped

as in H. humboldti, palp especially stout. Posterior

acclivities of coxae 1^4 strongly extended and

pointed, no coxa with mediofacial spines. Article 2

of gnathopod 1 with 1 midposterior spine, of gnathopod 2 with 2 spines, article 2 of gnathopod 1

trapezoidal, not apically lobate, article 4 with protrusion, article 5 ordinary, article 6 scarcely broadened, palm defined only by pair of spines, no hump,

posterior margin of article 6 with 1 group of setae

occupying only 20 percent of marginal length,

dactyl fitting palm, palm ordinarily setose, dactyl

with small erect inner setules; article 2 of gnathopod

2 scarcely lobate, article 3 with medium anterior

lobe, article 4 pointed apically, article 5 cryptic,

hand stout, not tapering apically (in comparison to

H. humboldti), palm elongate, defined by 2 spines,

armed with rows of short spines and setae, convex

posterior margin of article 6 with bundle of setules,

dactyl fitting palm, apex inserting into pocket near

defining spines, inner margin of dactyl with erect

setules. Pereopods 1-2 illustrated in J. L. Barnard

(1969b, fig. 27), distalmost locking spine small, unridged, straight, next proximal spine short and immersed by setae, dactyl curved, with giant inner

setule, margin proximal to setule castellate, face

striate distally; pereopods 3-5 with distal locking

spine larger than on pereopods 1-2, larger than next

proximal spine, itself weakly enlarged, proportions

figured in J. L. Barnard (1969b, fig. 27), article 6

naked posteriorly. Uropod 1 with small apicolateral

spine on peduncle, next proximal spine separated

by especially long gap from apicalmost, apicomedial

spine of medium size, peduncles and rami of uropods 1-2 otherwise normally but sparsely spinose;

ramus of uropod 3 slightly shorter than peduncle,

bearing 3 apical spines and spinules, 1 subapical

spine. Telson almost split to base, apices rounded.

Epimeron 1 ordinary, epimeron 2 lacking facial

ridge, with medium posteroventral tooth, epimeron

3 with small to medium sized tooth. Cuticle with

short bulbar setules, no polygons (in contrast to

H. humboldti).

HOLOTYPE AND TYPE-LOCALITY.—See Barnard,

1969b:136.

REMARKS.—This subspecies is elevated to full

specific status owing to the reevaluation of the H.

grandicornis complex as discussed above. The description is based on reobservation of characters

from original materials of this species deposited in

the Smithsonian Institution.

The presence of this species in a sample from

Bahía Kino is exceptional. Barnard (1969b: 138) believed the species to be of cold-water provenance

and to be declining in density with southward

progression through California.

MATERIAL.—KNO 3.

DISTRIBUTION.—California: Carmel to La Jolla,

intertidal; Gulf of California, Bahía Kino, intertidal.

Hyale humboldti, new species

FIGURE 64

ETYMOLOGY.—Named for Alexander von Humboldt.

DESCRIPTION OF MALE.—Anterior margin of

lateral cephalic lobe truncate vertically. Eyes very

large, ommatidia all darkly pigmented. Antenna 2

only as long as first 4 pereonites together, peduncle

of medium stoutness, minimally setose, flagellum

about 1.5 times as long as peduncle, with tufts of

very short setae on each article; antenna 1 about

three-fourths as long as antenna 2, reaching about

one-third along flagellum of antenna 2, aesthetascs
NUMBER 271

117

`pG2o

FIGURE 64.—Hyale humboldti, new species, holotype, male "a," 9.75 mm (p = female *'p,'

7.61 mm).

w

about 2 per article on flagellum. Upper lip evenly

rounded below, broad, setules long. Mandibular

incisors toothed, right lacinia mobilis evenly bifid

and with accessory side tooth, spine row with 3

spines and 1 setal plume, these plumes thick and

leaf-like, molar triturative, with long plumose seta,

left mandible with 4 raker spines and leaf-plume,

seta on molar absent. Lower lip, maxilla 1, and

maxilla 2 like H. yaqui (Figure 57) but shape of

maxilla 2 as in Figure 64:X2, spines on outer plate

of maxilla 1 more densely toothed as in H. guasave

(Figure 62:XZ), palp apparently with 1 vestigial

spout. Maxill¡ped illustrated, palp especially stout.

Posterior acclivities of coxae 1-4 strongly extended

and pointed, no coxa with mediofacial spines. Article 2 of gnathopod 1 with 1 midposterior spine, of

gnathopod 2 with 3 spines, article 2 of gnathopod

1 trapezoidal, not apically lobate, article 4 with
118

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

protrusion, article 5 ordinary, article 6 slightly

broadened, palm defined only by pair of spines,

no hump, posterior margin of article 6 with 1 group

of setae occupying almost 30 percent of marginal

length, dactyl fitting palm, palm ordinarily setose,

dactyl with small erect inner setules; article 2 of

gnathopod 2 with small anterodistal mammilliform

lobe, article 3 with nasiform lobe laterally, article 4

pointed apically, article 5 cryptic, hand elongate,

pyriform, sharply tapering, palm elongate, defined

by 2 spines, armed with rows of short setae, weakly

sinuous posterior margin of article 6 with bundle of

setules, dactyl fitting palm, apex inserting into

pocket near defining spines, inner margin of dactyl

with erect setules. Pereopods 1-2 similar in proportions to those of H. zuaque, distalmost locking spine

small, scarcely ridged longitudinally, weakly curved,

next proximal spine thin and immersed by setae,

dactyl strongly curved, with giant inner setule, margin proximal to setule castellate, face striate; pereopods 3-5 with first spine proximal to locking spine

slightly larger than on pereopods 1-2, stouter, proportions of articles on pereopods 3-5 generally as in

H. zuaque but articles 4-6 thinner, article 6 naked

posteriorly (opposite to point of dactyl). Uropod 1

with very small apicolateral spine on peduncle, next

proximal spine separated by especially long gap

from apicalmost, apicomedial spine of medium size,

peduncles and rami of uropods 1-2 otherwise normally but sparsely spinose; ramus of uropod 3

longer than peduncle, bearing 2 apical spines, 1

subapical spine and 1 subfacial spine near middle

of ramus. Telson split to base, apices rounded;

epimeron 1 ordinary, epimeron 2 lacking facial

ridge, with strong posteroventral tooth, epimeron

3 with medium sized tooth. Cuticle with weak

polygons filled with tiny villi, occasional polygon

with disrupted margins, surface naked, bearing

short bulbar setule.

FEMALE.—Slightly smaller than male, antennae

slightly shorter, eyes fully enlarged, article 2 of

gnathopods 1-2 with posterior spines; gnathopod 1

overall like that of male but slightly smaller relative

to anterior coxae, palmar defining spines larger

than in male; gnathopod 2 forming enlarged version of gnathopod 1, stouter, posterior setae on

article 6 narrowly confined.

ILLUSTRATION.—Cuticle of Figure 64: QP5 with

small villi shown only in 1 polygon.

HOLOTYPE.—USNM 169026, male "a," 9.75 mm

(illus.).

TYPE-LOCALITY.—DAW 10, Galapagos Islands,

Hood Island, 3 February 1962, intertidal, wash of

algal turf.

VOUCHER MATERIAL.—Type-locality, male "b,"

7.68 mm; female "p," 7.61 mm (illus.).

RELATIONSHIP.—This species and the specimens

from Japan identified as H. novaezealandiae (Thomson) by Iwasa (1939:276), now named Hyale species,

are extremely similar but the Japanese species

has more apical spinules on uropod 3, the medial

spine is more disjunct from the apical spines and

the subfacial spine appears to be absent. Otherwise

the 2 species share the unusual gap between the

lateral spines on the peduncle of uropod 1 and

the castellate dactyls of the pereopods not reported

in combination for any other species of the Hyale

grandicornis group except H. humboldti. Article

4 of pereopods 3-5 is stouter in the Japanese species than in the Galapagan.

Hyale humboldti differs from H. californica J. L.

Barnard, 1969b (as H. grandicornis californica) in

the tapering hand on male gnathopod 2.

This species appears to have little affinity with

H. grandicornis, the basic species of the group from

Chile, in that uropod 3 bears not only the subdistal

spine on the ramus but an additional facial spine,

and in the spinal disjunction of uropod 1, the

absence of "fur" on pereopod 4 and the well spinose

outer rami of uropods 1-2. The closeness of relationship is to the Japanese species followed by the

Californian species.

MATERIAL.—GAL 113, 120; DAW 2, 5, 10.

DISTRIBUTION.—Galapagos Islands, Tower Island,

Isla Santa Cruz, Hood Island, intertidal.

Najna Derzhavin

Najna kitamati, new species

Najna ?consiliorum.—]. L. Barnard, 1962c: 157-160, figs. 21, 22

[not Derzhavin].

DIAGNOSIS.—Najna with molar composed of 1 or

2 long, wire-like spines. Dactyl of palp on maxilliped only 0.12 times as long as article 3. Article 4 of

pereopods 1-5 about half as long as wide. Posteroventral corner of epimeron 3 rounded. Ramus of

uropod 3 less than 0.15 as long as peduncle.

HOLOTYPE.—AHF 5745, male, 8.0 mm.
NUMBER 271 119

TYPE-LOCALITY.—AHF Velero IV station 4822,

southern California, 34°2l'l5" N, l2O°l4'45" W,

17 January 1957, 16.5 m bottom depth.

REMARKS.—This species is now seen to be clearly

distinct from Najna consiliorum Derzhavin on the

points in the diagnosis.

DISTRIBUTION.—California, 1-17 m.

Parallorchestes Shoemaker and Parhyale Stebbing

Parhyale Stebbing, 1897:27.

Parallorchestes Shoemaker, 1941a: 183.

REMARKS.—I do not intend to synonymize Parallorchestes with Parhyale but to discuss them together. Gurjanova (1951:813) and Bulycheva (1957:

78) believed that Parallorchestes is a synonym of

Parhyale. Parallorchestes formerly contained 2 species, the type-species, P. ochotensis (Brandt), and

P. zibellina (Derzhavin), the latter having been

originally described in Parhyale. Because the palp

of maxilla 1 is unknown, P. zibellina cannot be

confirmed as a member of Parallorchestes and in

some of its appearance better fits Parhyale.

The type-species of Parallorchestes, P. ochotensis,

differs from Parhyale only in the biarticulation of

the palp on maxilla 1 and in the unprotruding

posterior acclivities on coxae 1-3, the comparison

to be made especially on coxa 3 because coxae 1-2

are variable in Parhyale. Formerly, the 2 genera

also differed in the presence (Parallorchestes) or

absence of a posterior lobe on article 5 of male

gnathopod 2 fully separating articles 4 and 6; however, Krapp-Schickel (1974:326) has placed Hyale

plumicornis (Heller) in Parhyale and that species

also bears the parallorchestian lobe. J. L. Barnard

(1974:42) has dismissed that kind of character differentiation as having no generic value in two allied

genera, Hyale and A llorchestes.

Parallorchestes should be maintained as long as

possible to distinguish the primitive mark of a

2-articulate palp on maxilla 1. Because of the

extreme basal position of the articulation one might

suspect it to be a secondary development from a

uniarticulate condition but until that avenue can

be explored the retention of Parallorchestes is

desirable. Both Parhyale and Parallorchestes retain

an inner ramus on uropod 3, another mark of primitiveness in the Hyalidae. One might therefore suggest a scheme of evolutionary flow in this family

from Parallorchestes to Parhyale, then to Hyale,

and later to A llorchestes. Hyale and A llorchestes

lack the inner ramus of uropod 3 and Allorchestes

has a vestigial palp on maxilla 1 and what appears

to be a secondarily developed telson with mostly

amalgamated lobes. The geographic sequence would

proceed from cold temperate of North Pacific

(Parallorchestes) to tropical and Mediterranean

(Parhyale) to tropical and warm-temperate Hyale

and back to biboreal (Allorchestes), suggesting a

more logical sequence than if Allorchestes proceeded outward from Parallorchestes in a line of

development separate from Parhyale and Hyale.

If the biarticulate maxillary palp of Parallorchestes

is found to be a secondary development, then a

more logical geographic procession from Parhyale

to Hyale, Parhyale to Parallorchestes, and Hyale to

Allorchestes would ensue.

Apparently several of the species in Parhyale in

the warm-temperate fringe of the tropics show the

progressive loss of the inner ramus of uropod 3.

This is especially well seen in the 3 species KrappSchickel (1974:326-327) has discovered recently in

the Mediterranean Sea. This condition would appear to occur also in 1 or more species from southern Japan.

There remain some considerable problems regarding speciation to be studied in both Parhyale and

Parallorchestes. Herein, Parhyale fascigera penicillata is raised to full specific status, because its distribution is intercalated within or closely adjacent

to the distributions of P. fascigera and P. hawaicnsis and it probably, therefore, does not exist as an

allopatric subspecies.

Shoemaker (1956) studied P. hawaiensis and P.

fascigera intensively and concluded that P. hawaiensis occurred both in the Atlantic and Pacific

oceans and that P. inyacka from South Africa was

a junior synonym of P. hawaiensis. J. L. Barnard

(1977:298) points out that Atlantic and Pacific

specimens of P. hawaiensis have minor differences

in coxa 1 and that the name "P. inyacka" may have

to be restored to the Atlantic population to signify

at the least a degree of subspeciation. However, P.

inyacka from South Africa has not been compared

minutely to Atlantic tropical P. hawaiensis (unless

Shoemaker did this without referring to such action

in print) and might be distinctive.

Now that Parhyale has been found, by the actions
120

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of Krapp-Schickel and the following dissection of

the literature, to contain a wide array of species one

must examine various collections from tropical

waters with extreme care to be certain that more

species are not contained within the P. hawaiensis

and P. fascigera complex.

A key to the species of both genera combined together is presented below. The following possible

taxa have been omitted: Parhyale inyacka (K. H.

Barnard), Parhyale sp. of Sivaprakasam (see p. 126)

and the Parhyale inyacka of Ledoyer, 1972, (see

synonymy of P. hawaiensis, p. 122). The identification of Allorchestes plumicornis (Heller) byChilton

(1912:515) from St. Vincent Island in the Caribbean

Sea is removed to unknown status. It is an apparent

Hyale or Parhyale with tufted antennae.

Key to the Species of Parhyale and Parallorchestes

1. Outer ramus of uropod 3 with 1 or more spines located on dorsal margin disjunctly from

apical spines 2

All spines on outer ramus of uropod 3 apical and contiguous 5

2. Palp of maxilla 1 biarticulate, antenna 2 not exceeding antenna 1

Parallorchestes ochotensis*

Palp of maxilla 1 uniarticulate,-antenna 2 strongly exceeding antenna 1 3

3. Peduncle and flagellum of antenna 2 with numerous short tufts of marginal setae

; Parhyale plumicornis

Peduncle and flagellum of antenna 2 sparsely setose 4

4. Hand of male gnathopod 2 pyriform, posterior margin much shorter than palm

Parhyale? z¡bellina

Hand of male gnathopod 2 subrectangular, posterior margin as long as palm

Parhyalet iu·asai

5. Article 6 of pereopod 5 with posterier spines Parhyale hawaiensis

Article 6 of pereopod 5 naked posteriorly 6

6. Outer ramus of uropod 1 naked dorsally 7

Outer ramus of uropod 1 spinose dorsally 9

7. All rami of uropods 1-2 naked dorsally, inner ramus of uropod 3 fused to peduncle

Parhyale eburnea

Some rami of uropods 1-2 spinose dorsally, inner ramus of uropod 3 fully articulate 8

8. Male antenna 2 with dense elongate setal brushes on peduncular articles 4-5, male gnathopod 1 with sharp taper near defining corner of palm Parhyale penicillata, new status

Male antenna 2 lacking setal brushes, male gnathopod 1 with scarcely any or no taper near

denning corner of palm Parhyale fascigera

9. Article 2 of pereopod 5 with broadly rounded posteroventral lobe Parhyale aquilina

Article 2 of pereopod 5 with narrowly conical posteroventral lobe 10

10. Peduncle of uropod 1 with large apicolateral spike, palm and posterior margin of hand on

male gnathopod 2 confluent, palm with long setae, peduncle of antenna 1 with long

midventral setae on articles 1-3 ?Parhyale sp. of Bulycheva

Peduncle of uropod 1 with medium apicolateral spine, palm and posterior margin of hand

on male gnathopod 2 distinct, palm with short spines, peduncle of male antenna 1 lacking

long ventral setae Parallorchestes ochotensis

• P. ochotensis also keyed to couplet 10.

Parallorchestes ochotensis (Brandt)

Allorchestes ochotensis Brandt, 1851:143-144, pi. 6: fig. 27.—

Holmes, 1904:233-234, fig. 118.

Allorchestes Ochotensis.—Bate, 1862:36, pi. la: fig. 4.

Hyale ochotensis.—Stehbing. 1906:561-562.

Parhyale ochotensis.—Uschakov, 1948:182.—Gurjanova, 1951:

814-815, fig. 568.—Bulycheva, 1957:82-83, fig. 28.

Parhyale kurilensis Iwasa, 1934:1-7, fig. 1, pis. 1-2; 1939:284-

285.

Parallorchestes ochotensis.—Shoemaker, 1941a:184-185.—J. L·

Barnard, 1952a:23-24, pi. 5: fig. 1; l954a:24; 1962c: 160,

fig. 23; 1964a: 118; 1969b: 141.
NUMBER 271 121

DIAGNOSIS.—One or more spines on outer ramus

of uroped 3 located disjunctly from apex on dorsal

margin (or occasionally those extra spines absent),

inner ramus small and fully articulate. Palp of

maxilla 1 biarticulate, basal article very short. Hand

of male gnathopod 1 not expanded, thin and rectangular, bearing 2 spines side by side at defining

corner, palm setose; palm and posterior margin of

hand on male gnathopod 2 distinct, posterior margin much longer than palm, palm lined with thin

short to medium spines. Article 2 of pereopod 5

with narrow, conical posteroventral lobe. Peduncle

of uropod 1 with enlarged apicolateral spine, outer

ramus with 3 or more dorsal spines. Peduncle of

antennae 1-2 poorly setose. Coxa 3 with weak unprotuberant posterior acclivity. Epimera 2-3 moderately and sharply protuberant posteroventrally, or

epimeron 3 poorly produced.

VARIABLES.—Pleonites \-A carinate dorsally, or

weakly so, or nearly uncarinate. Occasional adults

from Bering Sea part of west Alaska (Smithsonian

collections) lacking supernumerary spines on uropod 3; southern "race" based on material from California (J. L. Barnard, 1962c: 160), with posteroventral corner of epimeron 3 much sharper and more

extended than in Alaskan specimens.

RELATIONSHIP.—This species differs from P.

zibellina Derzhavin in the thin, rectangular, and

elongate hand of male gnathopod 2.

DISTRIBUTION.—Japan Sea, Okhotsk Sea, Bering

Sea, eastern Pacific south to Bahía de San Quintín,

Baja California.

Parhyale plumicornis (Heller)

Nicea plumicornis Heller, 1867:5-6, pi. 1: figs. 8, 9.

Allorchestes plumicornis.—Stebbing, 1899:412-413, pi. 33c;

1906:583-584.—Walker, 1901:299, pi. 27: figs. 20, 21 —

Chevreux, 1911:241-242, pi. 17: figs. 1-3.—Chevreux and

Fage, 1925:291-292, fig. 302.—Ruffo, 1936:30-31; 1941:119.—

Soika, 1949:196.

Parhyale plumicornis.—Krapp-Schickel, 1974:326, pis. 3, 4.

Not Allorchestes plumicornis.—Iwasa, 1939:289-292, figs. 25,

26, pi. 22.—Nagala, 1965:308 [? = Hyale penicillata (Stimpson)].—?Stephensen, 1944:71-72, fig. 25.

Not Allorchestes ptilocerus Derzhavin, 1937:96-97, 111, pi. 6:

fig. 1.—Gurjanova, 1951:823-824, fig. 576 [probably allied

to Hyale plumulosa Stimpson].

DIAGNOSIS.—One or more spines on outer ramus

of uropod 3 shifted disjunctly from apex onto dorsal

margin, inner ramus strong but fused to peduncle.

Palp of maxilla 1 uniarticulate. Hand of male

gnathopod 1 unexpanded, rectangular, bearing 2

spines in tandem at defining corner of palm, none

facial, palm setose; palm and posterior margin of

male gnathopod 2 of approximately equal length,

distinct, palm lined with short spines. Article 2 of

pereopod 5 with broad, rounded but shallow posteroventral lobe. Peduncle of uropod 1 with large

apicolateral spine, outer ramus with 2 dorsal spines.

Peduncle of antenna 1 poorly setose ventrally; articles 4-5 of peduncle on antenna 2 and proximal

articles of flagellum with dense tufts of short setae;

coxa 3 with protruding, subsharp posterior tooth.

Epimera 1-3 weakly protuberant posteroventrally,

strongest on epimeron 2.

DISTRIBUTION.—Mediterranean Sea.

Parhyale? zibellina Derzhavin

Parhyale zibellina Derzhavin, 1937:92-93, 109-110, pi. 4:

fig. 1.—Gurjanova, 1951:815-816, fig. 569.—?Tzvetkova,

1967:182-183.

Not Parhyale zibellina.—Bulycheva, 1957:78-82, fig. 27 [see

Parhyale sp.].

DIAGNOSIS.—One or more spines on outer ramus

of uropod 3 shifted disjunctly from apex onto dorsal margin, inner ramus apparently strong and

articulate. Palp of maxilla 1 unknown. Hand of

male gnathopod 1 unexpanded, rectangular, palm

apparently with 1 small defining spine, no facial

spines, palm setose; palm and posterior margin of

hand on male gnathopod 2 distinct, posterior margin significantly shorter than palm, palm lined with

thin spines. Article 2 of pereopod 5 with shallow

but broadly rounded posteroventral lobe. Peduncle

of uropod 1 apparently lacking apicolateral spike,

apical spine apparently not enlarged, outer rami of

uropods 1-2 apparently with dorsal spines (highly

unclear). Peduncle of antenna 1 poorly setose ventrally. Antenna 2 poorly setose. Coxa 3 probably

with protruding subsharp posterior tooth (if similar

to coxa 2). Epimera 1-3 sharply protruding posteroventrally, epimeron 2 strongest.

DISTRIBUTION.—Japan Sea, 0-5 m.

Parhyale? iwasai Shoemaker

Hyale gracilis Iwasa, 1939:282-284, fig. 19, pi. 19 [homonym,

not Bate].

Parhyale iwasai Shoemaker, 1956:356 [new name].

Hyale iwasa [sic] Bulycheva, 1957:107-109, fig. 39 [new name].
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DIAGNOSIS.—One or more spines on outer ramus

of uropod 3 shifted disjunctly onto dorsal margin,

inner ramus unknown. Palp of maxilla 1 uniarticulate but constricted in middle. Hand of male

gnathopod 1 unexpanded, thin, rectangular, apparently bearing only 1 or 2 weakly enlarged spines

side by side at defining corner of palm, palm weakly

setose, setae elongate; palm and posterior margin

of hand on male gnathopod 2 of approximately

equal length, distinct, palm sparsely lined with

spines. Article 2 of pereopod 5 with broad, rounded

posteroventral lobe. Peduncle of uropod 1 with ordinary apicolateral spine, outer rami of uropods 1-2

with 2 and 1 dorsal spines, respectively. Peduncle

of antennae 1-2 poorly setose. Coxa 3 apparently

with protuberant subsharp posterior tooth. Epimera 2-3 with weak and sharp posteroventral tooth.

Article 6 of pereopod 5 with posterior spines.

RELATIONSHIP.—Shoemaker (1956:356) thought

this species to be close to P. hawaiensis. At present

it cannot be made a synonym of P. hawaiensis, because of the thin and normally armed hand of male

gnathopod I and the disjunct dorsal spine on the

outer ramus of uropod 3. The inner ramus of uropod 3 remains to be discovered. The posterior spines

on article 6 of pereopod 5 are known only for P.

iwasai and P. hawaiensis.

DISTRIBUTION.—Taiwan and southern Japan.

Parhyale hawaiensis (Dana)

Allorchestes hawaiensis Dana, 1853-1855:900, pi. 61: figs. 5a-h.

Allorchestes Hawaiensis.—Bate, 1862:47-48, pi. 8: fig. 1.

Hyale brevipes Chevreux, 1901:400-402, figs. 15-18.—Walker,

1909:337.—Chilton, 1921a:545-548, fig. 9; ?I921b.118, figs. 2,

2a; 1925:536.—Schellenberg, 1928:658-659.—K. H. Barnard,

1935:292-294, fig. 10.

Hyale nilssoni.—Walker, 1905:925, fig. 140.1 [not Rathke].

Hyale hawaiensis.—Stebbing, 1906:573.—Schellenberg, 1938a:

66-67, fig. 34.—Ruffo, 1950:57-60.—Bulycheva, 1957:109-

110, fig. 40.—Nayar, 1959:30-31, pi. 10: figs. 10-24.

Hyale prevostii.—Kunkel, 1910:66-69, fig. 25 [not Milne Edwards].—Shoemaker, 1920:378.

Hyale pontica.—Kunkel, 1910:69-72, fig. 26 [not Rathke].

Hyale trifoliadens Kunkel, 1910:72-74, fig. 27.

Hyale inyacka.—Chevreux, 1926:370-372, fig. 17.—?Stephensen, 1927:590; 1933:441-446, figs. 3, 4.—Bulycheva, 1957:88-

90, fig. 29 [not K. H. Barnard].

Parhyale fasciger — Fage and Monod, 1936:105-108, figs. 3-7a

[not Stebbing].

Hyaloides dartevillei Schellenberg, 1939:126-128, figs. 6-10.

Parhyale inyacka.—Stephensen, 1947:6-7.—J. L, Barnard,

l955a:23, fig. 12.—Mateus and Mateus, 1966:185—?Sivaprakasam, 1970:562-564, fig. 6.—?Ledoyer, 1972:275, pi. 80

[not K. H. Barnard].

Parhyale hawaiensis.—Shoemaker, 1956:351-356, figs. 3, 4.—

Ruffo, 1959:17-18—J. L. Barnard, 1965a:521-523, fig. 24.—

Nayar, 1967:156-157.—Ruffo, 1969:38.—Sivaprakasam, 1970:

560-562, fig. 5·—Bousfield, 1971:266-267.—J. L. Barnard,

1971b: 131.—Vader, 1972:15.—W¡lkens and Parzefall, 1974:

430.—J. L. Barnard, 1977:295-298.

?Hyale stolzmanni Wrzesniowski, 1879:201.

DIAGNOSIS.—All spines on outer ramus of uropod

3 apical and contiguous, inner ramus fully articulate. Palp of maxilla 1 uniarticulate. Hand of male

gnathopod 1 weakly expanded, bearing slightly enlarged spine at defining corner and 1 slightly enlarged spine on inner face along posterior margin

of hand, palm setose; palm and posterior margin of

hand on male gnathopod 2 of approximately equal

length, distinct, palm densely lined with medium

spines. Article 2 of pereopod 5 with broad, rounded

posteroventral lobe. Peduncle of uropod 1 with enlarged apicolateral spine, outer rami of uropods

1-2 with 2 and 1 dorsal spines, respectively. Peduncle of antennae 1-2 poorly setose. Coxa 3 with

weakly protuberant subsharp posterior tooth. Epimera 2-3 almost perfectly quadrate, scarcely but

very minutely protuberant posteroventrally. Article

6 of pereopod 5 with posterior spines.

REMARKS.—See "Remarks" under P. inyacka. The

synonymy and references for both species are

provisional.

Specimens reported by J. L. Barnard (l955a:23)

from Hawaii have been reexamined and found to

have sparse ordinary bulbar setules on the cuticle

lacking plaques. See P. penicillata and P. fascigera

for notes about this condition.

MATERIAL.—DAW 15.

DISTRIBUTION.—Apparently pantropical in shallow water, but not yet collected inside Gulf of

California.

Parhyale eburnea Krapp-Schickel

Parhyale eburnea Krapp-Schickel, 1974:327-328, pis. 8-11.

DIAGNOSIS.—All spines on outer ramus of uropod

3 apical and contiguous, inner ramus small and

fused to peduncle. Palp of maxilla 1 uniarticulate

but weakly constricted in middle. Hand of male

gnathopod 1 weakly expanded, bearing enlarged

spine at defining corner and enlarged facial spine

in middle, palm weakly setose; palm and posterior
NUMBER 271 123

margin of hand on male gnathopod 2 of approximately equal length, distinct, palm lined with elongate spines. Article 2 of pereopod 5 with broad,

rounded but shallow posteroventral lobe. Peduncle

of uropod 1 with tiny apicolateral spine, outer

ramus naked dorsally (all rami of uropods 1-2

naked dorsally). Peduncle of antenna 1 poorly setose

ventrally; antenna 2 poorly setose. Coxa 3 with protruding and subsharp posterior tooth. Epimera 1-3

rounded or unprotuberant posteroventrally.

DISTRIBUTION.—Italy, Sirolo (Ancona).

Parhyale penicillata Shoemaker, new status

FIGURES 65-67

Parhyale fascigera penicìllata Shoemaker, 1956:350-351, fig.

2g-i.

DIAGNOSIS.—All spines on outer ramus of uropod

3 apical and contiguous, though several spines somewhat facial and subterminal, inner ramus fully

articulate. Palp of maxilla 1 uniarticulate. Hand of

male gnathopod 1 expanded, bearing 2 enlarged

spines in tandem at defining corner, palm weakly

setose; palm and posterior margin of hand on male

gnathopod 2 of approximate equal length, distinct,

palm lined with short spines. Article 2 of pereopod

5 with broad, rounded posteroventral lobe. Peduncle of uropod 1 with enlarged apicolateral spine,

outer ramus of both uropods 1-2 naked dorsally.

Peduncle of antenna 1 poorly setose ventrally; articles 4-5 of peduncle on antenna 2 with dense but

short tufts of ventral setae. Coxa 3 with protruding

subsharp posterior lobe. Epimera 2-3 protruding

weakly posteroventrally, ep¡meron 3 strongest.

DESCRIPTION.—Male gnathopod 2 hand with up

to 9 setae in posterior acclivity; locking spines on

pereopods 3-5 accompanied by up to 4 setae in large

adults but setae absent elsewhere, occasionally

pereopod 5 with 3 locking spines and 3 setae;

gnathopods 1-2 each with 3 recurved posterior

spines on article 2; female coxae 2-A narrower anteroposteriorly than in male and posterior cusps

slightly sharper on coxae 2-3, cuticle with scattered

alate slit-pits each with a setule, surrounded by

simple setules (seen as holes on Figure 66:QE5) and

very finely punctate or frosted matrix on cuticular

surface, on dorsum of pleonites slit-pits with slightly

elevated rims; epimera with frosted surface but

bearing only simple bulbar setules (Shoemaker

found no setules on P. fascigera).

VOUCHER MATERIAL.—KNO 6: male "a," 7.8 mm

(illus.); female "n," 6.1 mm (illus.).

REMARKS.—Shoemaker distinguished P.f. penicillata also on the fact that most populations of circumtropical P.f. fascigera have 1 of the enlarged

spines on male gnathopod 1 moved anteriorly towards the middle of the palm but P.f. penicillata

differs little from specimens in the Virgin Islands

figured by Shoemaker in which that spine is scarcely

removed from the palmar defining corner.

Specimens from TOP-1 have the cuticular alae

poorly developed or absent and the posteroventral

tooth on epimeron 3 is poorly developed, almost

rounded.

This species differs from P. fascigera, to which it

formerly was assigned as a subspecies, in the characters of couplet 8 in the key to the species of

Parhyale and Parallorchestes (p. 120).

MATERIAL.—KNO 6; TOP 1.

DISTRIBUTION.—Gulf of California: Bahía Kino,

Topolobampo, La Paz, 0 m.

Parhyale fascigera Stebbing

Parhyale fasciger Stebbing, 1897:26-28, pi. 6.—?Schellenberg,

1925:162; ?l938b:2l5; ?I939:I28.

Parhyale fascigera.—Stebbing, 1906:556.—Shoemaker, 1935a:

244; 1956:346-350, figs. I, 2a-f.

Hyale brevipes.—Shoemaker, 1933a: 18, figs. 10, 11 [not Chevreux].

Hyale hawaiensis.—Shoemaker, 1942:18 [not Dana].

DIAGNOSIS.—All spines on outer ramus of uropod

3 apical and contiguous (fide Shoemaker, 1956, but

Stebbing, 1897, weakly disagreeing), inner ramus

fully articulate. Palp of maxilla 1 uniarticulate but

weakly constricted in middle. Hand of male gnathopod 1 expanded, bearing large spine in middle of

palm or divorced from defining corner plus enlarged facial spine, palm setose; palm and posterior

margin of hand on male gnathopod 2 of approximately equal length, distinct, palm densely lined

with medium spines. Article 2 of pereopod 5 with

broad, shallow, rounded posteroventral lobe. Peduncle of uropod 1 with enlarged apicolateral spine,

outer rami of uropods 1-2 lacking dorsal spines.

Peduncle of antennae 1-2 poorly setose. Coxa 3 with

protuberant subsharp posterior tooth. Epimera 2-3

very weakly and subsharply protuberant posteroventrally.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

V

XS``>

FIGURE 6.V—Parhyale penìcilluta Shoemaker, new status, male "a," 7.8 mm (n «= female "n,

6.1 mm).
NUMBER 271 125

QE5

FK.URE 6G.—Parhyale penicillata Shoemaker, new status, male "a," 7.8 mm (n — female "n,

6.1 mm; iv = female "w," 6.1 mm).
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PI El

FIGURE 67.—Parhyale pen¡cillata Shoemaker, new status, male "a," 7.8 mm.

G2

REMARKS.—Specimens from USNM 120194, Dominica, British West Indies, Soufriere, lower beach

in algae on rocks, 5 December 1966, coll. G. S.

Schultz and E. L. Bousfield, have been examined

and found to have sparse and ordinary bulbar

setules on the cuticle lacking plaques. See P. penicillata (p. 123) for importance of this observation.

DISTRIBUTION (based only on specimens identified by Shoemaker, 1956).—Atlantic; Florida, Gulf

of Mexico, Caribbean Sea; Pacific: west Mexico to

Peru and the Juan Fernandez Islands and the Galapagos Islands; not yet collected inside Gulf of

California.

Parhyale aquilina (Costa)

Amph¡thoe aquilina Costa, 1853:174; 1857:202-203, pi. 2: fig. 7.

Nicea fasciculata Heller, 1867:6-7, pi. 1: figs. 10,11.

Nicea nudicornis Heller, 1867:8-9, pi. 1: figs. 16-19.

M<r<ra rudis Heller, 1867:12, pi. 1: fig. 33.

Ållorchestes aquilina.—Walker, 1901:299, pi. 27: fig. 19.

Hyale aquilina.—Stebbing, 1906:565.

Ållorchestes aquilinus.—Chevreux, 1911:240-241, pi. 16: figs.

20-25.—Chevreux and Fage, 1925:289-291, figs. 300, 301.—

Chevreux, 1926:372; 1935:123.—Ruffo, 1941:119; 1947:126.—

Legrand, 1951:376.

Parhyale aquilina.—Krapp Schickel, 1974:326-327, 329, 334,

pis. 5-7.

DIAGNOSIS.—All spines on outer ramus of uropod

3 apical and contiguous, inner ramus vestigial, fully

fused to peduncle. Palp of maxilla 1 uniarticulate

but constricted in middle. Hand of male gnathopod

1 expanded, bearing 2 enlarged spines in tandem

near defining corner, none facial, palm setose; palm

and posterior margin of hand on male approximately equal in length, distinct, palm lined with

short stout spines. Article 2 of pereopod 5 with

broad rounded posteroventral lobe. Peduncle of

uropod 1 lacking spike, bearing medium apicolateral

spine, outer ramus with 1 dorsal spine. Peduncle of

antenna 1 poorly setose ventrally; antenna 2 poorly

setose. Coxa 3 with unprotruding posterior acclivity.

Epimera 2-3 only weakly protuberant posteroventrally, epimeron 2 strongest.

DISTRIBUTION.—Mediterranean Sea and Canary

Islands.

Parhyale species

Parhyale hawaiensis.—Sivaprakasam, 1970:560-562, fig. 5 [not

Dana].

REMARKS.—The shape of gnathopods 1-2 and the

presence of setules (possibly green algal tufts) on the

telson suggest that this may be a species distinct

from P. hawaiensis. The possible presence of posterior spines on article 6 of pereopod 5 and the pre-
NUMBER 271

127

cise spine distribution on the hand of gnathopod 1

must be determined.

DISTRIBUTION.—East coast of India.

Parhyale species of Bulycheva

Parhyale zibellina.—Bulycheva, 1957:78-82, fig. 27 [not

Derzhavin, 1937].

DIAGNOSIS.—All spines on outer ramus of uropod

3 apical and contiguous, inner ramus fully articulate. Palp of maxilla 1 uniarticulate. Hand of male

gnathopod 1 expanded, bearing 2 enlarged spines at

defining corner of palm, none facial, palm setose;

palm and posterior margin of hand on male gnathopod 2 confluent, palm long, posterior margin

short, palm lined with elongate thin setae. Article

2 of pereopod 5 with narrow, subsharp posteroventral lobe. Peduncle of uropod 1 with large apicodistal spike, outer ramus with more than 2 dorsal

spines besides apical spines. Peduncle of antenna 1

with long ventral setae in midmargins of articles

1-3, antenna 2 poorly setose. Coxa 3 with weak,

unprotuberant posterior acclivity. Epimeron 3 with

small but sharp posteroventral tooth (other epimera

unknown).

DISTRIBUTION.—Precisely constrained distribution

unknown owing to probable confusion with Parhyale zibellina; Bulycheva mentions Japan Sea,

Okhotsk Sea, and Bering Sea.

Parhyale inyacka (K. H. Barnard)

Hyale inyacka K. H. Barnard, 1916:233-234, pi. 28: fig. 4.

Parhyale inyacka.—K. H. Barnard, 1940:472-473.—Griffiths,

1973:229-300.

REMARKS.—Actual specimens from South Africa

have never been described in detail necessary to

identify them with Stephensen's (1933, figs. 3, 4)

and Shoemaker's (1956, figs. 3, 4) illustrations of

Atlantic P. hawaiensis, although Shoemaker considered the 2 species identical. J. L. Barnard (1977:

298) points out a difference in the posterior acclivity of coxa 1 between Atlantic and Pacific specimens

of P. hawaiensis and suggests the possibility that P.

inyacka might be a name available for recognizing

the Atlantic populations as distinctive subspecies or

species. However, it is also possible that South

African P. inyacka is itself distinctive from the tropical Atlantic form.

ISCHYROCERIDAE

Microjassa chinipa, new species

FIGURES 68, 69

DIAGNOSIS.—Rostrum and lateral cephalic lobe

projecting equally, lateral lobe subfalciform and

subacute. Eyes large. Mouthparts similar to those

of Parajassa angularis Shoemaker (1942) except for

expanded mandibular palp article 3. Inner plate of

maxilla 1 well developed as in P. angularis. Maxillipedal palp article 4 short as in P. angularis and

armed medially with 4 or 5 setae. Coxa 1 in male

about 50 percent as long as coxa 2, in female about

75 percent; coxae 3^i flat to excavate ventrally in

both sexes, very broad; coxa 5 in male about as

long as coxa 4 but shorter in female, posterior lobe

falcate, coxae 6-7 shorter than posterior lobe of

coxa 5. Gnathopod 1 small and similar in both

sexes, article 5 less than 60 percent as long as

article 6, latter moderately expanded, palm slightly

exceeding length of posterior margin, defined by 3

spines, gnathopod 2 of female similar to gnathopod

1 but article 5 shorter; gnathopod 2 of male with

elongate and enlarged article 6, palm and posterior

margin confluent, armed distally with 2 weak

humps, proximally with strong, slightly reverted

blunt process at joint with article 5, palm sparsely

setose, dactyl almost as long as palm, inner margin

evenly arcuate, article 5 very small, posterior lobe

narrow, short. Article 2 of pereopod 3 sexually

dimorphic, with posteroventral lobe in male, absent

in female, article 2 of pereopods 4-5 also lobed,

narrow on pereopod 4, broad on pereopod 5, but in

male somewhat narrower than in female. Pleonal

epimera almost straight, posteriorly notched. Outer

rami of uropods 1-2 shorter than inner, peduncle

of uropod 1 with sharp interramal tooth; uropod

3 elongate, rami about 25 percent as long as peduncle, outer ramus bearing 3 or 4 distolateral

cusps, apical cusp not an immersed or articulate

spine, margin proximal to cusps minutely scaly,

inner ramus simple, straight, bearing 1 basally immersed articulate spinule, lateral margin minutely

scaly. Telson short, subtriangular, apically blunt,

armed with 2 dorsal spines and several marginal

setules.

HOLOTYPE.—USNM 168025, male "q," 2.58 mm

(illus.).

TYPE-LOCALITY.—DAW 26, Galapagos Islands,
128

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 68.—Microjassa chinipa, new species, holotype, male "q," 2.58 mm (w

2.11 mm).

female "w.

Isla Santa Cruz, Academy Bay, 1962 (date otherwise

unknown), intertidal, rock wash.

VOUCHER MATERIAL.—Type-locality, female "w,"

2.11 mm (illus.), DAW 40, female "x," 2.08 mm

(illus.).

RELATIONSHIP.—This species is very close to Aí.

claustris J. L. Barnard, 1969b, from California, and

may be the fully developed stage of that species

which might, itself, represent the retarded northernmost fringe of the species. Both males and females

of Aí. ch¡nipa have broadened coxae 3^1 which are

flat or excavate ventrally, whereas Aí. claustris has

much narrower coxae with convex ventral margins.

Even young males and females of Aí. chinipa have

these typical coxae so that the somewhat youthfulappearing Aí. claustris differs in that character. The

posterior lobe of coxa 5 is sharper and more elongate in Aí. chinipa than in Af. claustris. Male gnathopod 2 of Aí. chinipa passes through the stages

shown for Aí. claustris.

MATERIAL.—GAL 114, 115, 116, 119, 120; DAW

1, 3, 4, 9, 13, 15, 16, 19, 20, 23, 26, 27, 31, 33, 35,

37, 40; PAN 14.

DISTRIBUTION.—Galapagos Islands, 0-6 m; Pacific

Panama, 0 m.

Microjassa macrocoxa Shoemaker

Microjassa macrocoxa Shoemaker, 1942:44—47, figs. 16, 17.—

J. L. Barnard, 1964a:111, fig. 2ID.

MATERIAL.—TOP 3 (15).

DISTRIBUTION.—Gulf of California, Topolobambo, 1 m; outer Baja California, Bahía Magdalena to Bahía de San Quintín, 0-27 m.

LEUCOTHOIDAE

Leucothoe alata J. L. Barnard

Leucothoe alata J. L. Barnard, 1959:19-20, pi. 1; l%2c:l32,

figs. 7DJ·,F; l964b:227; I966a:22; l%9a:2l4; 1969b: 164.—

Nagata, 1965b: 158-159, figs. 9, 10.
NUMBER 271 129

FIGURE 69.—Microjassa chinipa, new species, holotype, male "q," 2.58 mm (a;

2.11 mm; x - female "x," 2.08 mm).

female "w,"

MATERIAL.—SCO 1, 8, 14, 15, 16, 18; KNO 1;

TOP 3; PAZ 15, 24; GAL 108, 114, 115, 118; DAW

3, 8, 17, 27, 28, 31, 36; BRU 1, 2. Presumed to be

associated with sponges and tunicates.

DISTRIBUTION.—Gulf of California, Galapagos

Islands, outer Baja California and California northward to Monterey Bay, 0-24 m; Japan, 2-4 m.

Leucothoe spinicarpa (Abildgaard)

Gatnmarus sp¡n¡carpus Abildgaard, 1789:66-67, pi. 119: figs.

1-4.

Leucothoe sp¡n¡carpa (Abildgaard).—Sars, 1895:283, pis. 100,

101: fig. 1.—Stebbing, 1906:165-166.—J. L. Barnard, l954a:6;

1962c: 1S2, fig. 7A,B,C; l964b:227; l966a:22.

REMARKS.—Another cosmopolitan species requiring closer study; referred to scores of times in the

literature, references above limited to early definitive studies and those of eastern Pacific Ocean. Associated with tunicates and sponges.

MATERIAL.—TOP 3; PAZ 24; DAW 31.

DISTRIBUTION.—Cosmopolitan, 0-1505 m; found

here in the Gulf of California at Topolobampo and

Bahía Conœpción, 0-1 m, and in Galapagos Islands,

6 m. Apparently not as tolerant of intertidal condi-
130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tions as L. alata; also not recorded yet from intertidal of California, though abundant in very shallow sublittoral depths.

MATERIAL.—PAZ 3.

DISTRIBUTION.—Gulf of California, Isla Espiritu

Santo, 1 m; Australia.

Leucothoid.es pacifica J. L. Barnard

Leucothoides pacifica J. L. Barnard, 1955c:26-28, figs. 1,

2e,h,n; 1959:21; 1964a: 114; 1969b: 165, fig. 24n-r.

MATERIAL.—SCO 14; PAZ 12. Probably associated with tunicates and sponges.

DISTRIBUTION.—Gulf of California: Puerto Peñasco and Bahía San Evaristo, 0 m; California from

Carmel to Newport Bay, 0-8 m.

Leucothoides pottsi Shoemaker

Leucothoides pottsi Shoemaker, 1933b [only the holotype, not

the description and figures].—J. L. Barnard, 1974:103.

REMARKS.—The single specimen fits the diagnosis

of Barnard (1974:103) and differs from L. yarrega

in the presence of serrations on apposing margins of

articles 5-6 on gnathopod 1 and uneven distribution of spines on article 6 of pereopods 1-2.

MATERIAL.—DAW 31.

DISTRIBUTION.—Galapagos Islands, 6-9 m; Caribbean Sea, occurrence in Mediterranean Sea and

India doubtful.

Leucothoides ?yarrega J. L. Barnard

[}]Leucothoides yarrega J. L. Barnard, 1974:103-104, figs.

62: f, 62: fj, 63 f.

MATERIAL.—PAZ 3; DAW 31.

REMARKS.—Despite the nearness of Baja California to the Caribbean Sea, the material at hand

differs from Caribbean species of the genus and resembles the Australian species very closely. The

absence of serrations on the apposing margins of

articles 5 and 6 of gnathopod 1 is a crucial feature

in this resemblance. Coxa 1 of the material at hand

is identical to that of L. yarrega, the head is even

more broadly rounded than in L. yarrega but article

2 of pereopod 5 is slightly less beveled than that of

L. yarrega. Gnathopod 2 is of the purely tropical

form in contrast to gnathopod 2 of L. pacifica, the

subtropical species.

LYSIANASSIDAE

Lysianassa dissimilis (Stout)

Nannonyx dissimilis Stout, 1913:638-639.

Aruga dissimilis.—Shoemaker, 1942:7.—J. L. Barnard, 1955b:

100-103, pi. 29g,i; l946b:23O.

Lys¡anopsis? dissimilis,—Hurley, 1963:76-77, fig. 2ld.

Lysianassa dissimilis—J. L. Barnard, l969a:2l8; l969b:l86-

187, fig. 47a-f.

MATERIAL.—SCO 14, 16; KNO 1; TOP 3; PAZ

24; GAL 115.

DISTRIBUTION.—Gulf of California, at Puerto

Peñasco, Bahía Kino, Bahía de Los Angeles, Topololambo, Bahía Concepción, 0-1 m; Galapagos

Islands, 0-18 m; generally from Tomales Bay, California to Isla Isabel, Mexico, 0-73 m.

Lysianassa holmesi (J. L. Barnard)

Aruga holmesi J. L. Barnard, 1955b: 100, pis. 27, 28; 1959:18;

1964a: 79.

Lysianopsis(?) holmesi.—Hurley, 1963:74-75, fig. 21b.

Lysianassa holmesi.—J. L. Barnard, l966a:25; l966b:69.

MATERIAL.—PAZ 1, 3, 6, 7, 8, 9, 12, 13.

DISTRIBUTION.—Gulf of California at La Paz, Isla

Espiritu Santo, Isla Partida and Bahía San Evaristo,

0 m; generally Monterey Bay, California to Ecuador, 0-183 m.

Lysianassa macromera (Shoemaker)

Aruga macromerus Shoemaker, 1916:157-158.

Lysianopsis(}) macromerus.—Hurley, 1963:77.

Lysianassa macromerus.—J. L. Barnard, l969a:2l8; 1969b:187-

189, figs. 48, 49.

MATERIAL.—PAZ 10, 12, 20.

DISTRIBUTION.—Gulf of California at Cabo San

Lucas, Bahía San Evaristo, Isla San Francisco, Bahía

de Los Angeles, 0 m; California, from Cayucos to

La Jolla, 0 m.
NUMBER 271

131

PHLIANTIDAE

Heterophlias seclusus escabrosa J. L. Barnard

FIGURE 40 (part)

Heterophlias seclusus escabrosa J. L. Barnard, 1962b:79-80,

fig. 5; l969a:2l9; 1969b: 195-196.

VOUCHER MATERIAL.—SCO 7, sex unknown "p,"

3.12 m (illus).

REMARKS.—Three figures of this subspecies from

California and Mexico are given for comparison to

the new species from the Galapagos Islands.

Epimeron 3 is like that shown for H. galapagoanus; Shoemaker's (1933b, fig. 4c) view of this epimeron for the typical subspecies from the Caribbean Sea is distinctive. The pereopodal dactyls

and locking spines of H. s. escabrosa also fit those

of H. galapagoanus (Figures 71, 72). The telson of

H. s. escabrosa is slightly elliptical side to side.

Juveniles from Puerto Peñasco have elongate

processes on antenna 1 and the long inner ramus

on uropod 2 typical of H. s. escabrosa, but adults

have a short inner ramus on uropod 2 typical of the

Caribbean stem subspecies, H. s. seclusus.

MATERIAL.^SCO 1, 7, 8, 12, 14, 18, 19; KNO 1;

PAZ 13, 18, 22.

DISTRIBUTION.—California from Cayucos to La

Jolla, 0-16 m; Gulf of California, Puerto Peñasco,

Bahía de Los Angeles, Bahía Kino, Bahía San

Evaristo and Cabo San Lucas, intertidal.

Heterophlias galapagoanus, new species

FIGURES 70-72

DIAGNOSIS.—Rostrum 1.1 times as long as broad,

half as broad as head; dorsal outline of pleonite 1

broadly deltoid, posterolateral margins nearly

straight; medial protrusions on peduncles of pleopods reaching approximately to vertical tangents

from medial edges of inner rami.

DESCRIPTION.—Right mandible with tiny tripartite comb proximal to lacina mobilis, this comb

presumably remnant of spine row, left mandible

with conical projection bearing tiny denticles, perhaps combination of lacinia mobilis and comb row;

maxilla 1 with 5 spines on outer plate; cuticle

minutely punctate, bearing stout setules and various-sized tubes opening externally, remaining parts

as illustrated.

Lateral view with coxae depressed and flattened

unnaturally, coxae normally splayed; lower lip

shown as outer view in situ; detached pereopods

3-5 reduced in size in comparison to pereopods 1-2

and gnathopods 1-2; lateral view of epimera and

urosome of specimen "b" joined by view of epimeron 1 from specimen "a" in magnification similar

to relative body sizes.

HOLOTYPE.—USNM 142293, sex unknown "a,"

2.28 mm (illus.).

TYPE-LOCALITY.—DAW 31, Galapagos Islands,

Isla Santa Cruz, halfway between Tortuga Bay and

Academy Bay, 22 February 1962, 6-9 m, algal rock

wash.

VOUCHER MATERIAL.—Type-locality, sex? "y,"

2.12 mm (illus.); sex? "c," 1.28 mm (illus.).

REMARKS.—Lengths of these specimens are taken

from the anterior edge of the rostrum to the posterior edge of pleonite 1 from dorsal view. Sex of the

specimens is unknown and presumably they are

juveniles as no brood plates nor penial protrusions

are present.

RELATIONSHIP: The diagnosis distinguishes this

species from the only other species of the genus and

its holotype. In H. seclusus Shoemaker (1933b) pleonite 1 is weakly trifoliate, the rostrum is broader and

shorter and the medial protrusions on the pleopodal

peduncles are much longer than in H. galapagoanus.

The one tiny juvenile, 1.28 mm, of H. galapagoanus at hand is very similar to H. seclusus in the

characters mentioned and is, therefore, unidentifiable. Presumably hatched juveniles resemble the

parent species H. seclusus but the rostrum, pleonite

1, and the pleopods transform during maturation.

The Pacific subspecies, H. s. escabrosa, resembles

the typical subspecies from the Caribbean Sea in

the characters being discussed but differs from the

typical subspecies in stronger protrusions on antenna 1 and in the longer inner ramus of uropod 2.

These characters on H. galapagoanus resemble the

situation in the Pacific subspecies of H. seclusus.

Pariphinotus Kunkel (1910) is presumably a senior synonym of Heterophlias Shoemaker (1933b),

but Kunkel reported the mandibular molar as

absent; until this character is confirmed the two

genera should be held separate. The type-species,

P. tuckeri Kunkel, from Bermuda, is highly similar

to H. seclusus in the strong protrusion on the pleopodal peduncles, but the head resembles that of H.

galapagoanus; Kunkel (1910) did not describe pleonite 1.
132

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

M

FIGURE 70.—Heterophlias galapagnanus, new species, holotype, sex unknown "a," 2.28 mm

(/o — opposite sides of apical nipple on maxilla 1; Id = dorsal view of complete specimen).
NUMBER 271 133

X2

FIGURE 71.—Heterophlias galapagoanus, new species, holotype, sex unknown "a," 2.82 mm.

MATERIAL.—GAL 106; DAW 17, SI.

DISTRIBUTION.—Galapagos Islands, Isla Santa

Cruz, intertidal.

PHOXOCEPHALIDAE

Metaphoxus frequens J. L. Barnard

Metaphoxus frequens J. L. Barnard, 1960:304-306, pis. 51, 52;

l964b:242; l966a:28; l966b:88; 1969b: 196; l97la:68.

MATERIAL.—PAZ 3.

DISTRIBUTION.—Gulf of California at Isla Espiritu Santo, 0 m; generally Oregon to Isla Isabel,

Mexico, 0-458 m, rarely intertidal.

"Paraphoxus" spinosus Holmes

Parapho×us spinosus Holmes, 1905:477-478, unnumbered

fig.—Kunkel, 1918:76-78, fig. 13.—Shoemaker, 1925:26-27.—

J. L. Barnard, 1959:18; 1960:243-249, pis. 29-31; 1961:178;

1964a: 105; l966b:89; l%9a:224; 1969b: 197-198.—Reish and

Barnard, 1967:18.—Bousfield, 1973:12, pi. 34.1.

REMARKS.—All members of so-called "Paraphoxus" in America will be assigned to other genera;

see also Barnard and Drummond (1976).

MATERIAL.—SCO 1, 17; PAZ 3, 5, 11.

DISTRIBUTION.—Gulf of California, widely distributed, 0-24 m; ranging northward to Puget

Sound, Washington, 0-73 m; western Atlantic

Ocean.
134

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 72.—Heterophlias galapagoanus, new species, holotype, sex unknown

(c = sex unknown "c," 1.28 mm, y = sex unknown "y," 2.12 mm).

'a," 2.28 mm

PODOCERIDAE

Podocerus fulanus J. L. Barnard

FIGURES 73, 74

Podocerus sp.—J. L. Barnard, 1959:40, pi. 14.

Podocerus fulanus J. L. Barnard, l962a:69; l%9b:224.

VOUCHER MATERIAL (all illustrated).—SCO 1:

male "p," 4.2 mm; male "g," 4.0 mm; male "h," 3.4

mm; male "m," 3.6 mm. SCO 10: male "a," 5.4 mm;

male "x," 5.3 mm; female "w," 5.1 mm.

REMARKS.—The size and shape of posterodorsal

segmental teeth vary widely among specimens of a

single collection; females have low, evenly developed teeth, whereas males vary from low and evenly

developed teeth to strongly projecting teeth. Certain males have the tooth of pereonite 7 largest,

others have the teeth of pereonite 7 and pleonite

1 equally dominant.

The eye forms a bulge on the side of the head in

many specimens but in others, and especially in the

largest adults, the eye forms no distinct bulge.

The ordinary specimen, 3-4 mm long, has 2 telsonic spines, but a few specimens larger than 4.0

mm have 4 spines. The 5 mm holotype male from

Newport Bay has 5 spines.

This species is widely distributed in the opensea of Mexico but in southern California at the

periphery of its northern limits the species occurs

only in warm, ponded embayments.

Weak lateral cusps occur on 1 or more pereonites

6—7 and pleonites 1-2 especially on males with

largest teeth; giant specimens, near 5 mm in length,

have a cusp on pereonite 7 just above coxa 7.
NUMBER 271 135

FIGURE 73.—Podocerus julanus J. L. Barnard, male "a," 5.4 mm (g = male "g," 4.0 mm;

p •= male "p," 4.2 mm; x = male "x," 5.3 mm).
136

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

wG2o

FIGURE 74.—Podocerus fulanus J. L. Barnard, male "a," 5.4 mm (g — male "g," 4.0 mm;

h -» male "h," 3.4 mm; m = male "m," 3.6 mm; p = male "p," 4.2 mm; w = female "w,"

5.1 mm; × = male "x," 5.3 mm).
NUMBER 271 137

MATERIAL.—SCO 1, 10, 14, 15, 18, 21; KNO 1;

TOP 3; PAZ 3, 13.

DISTRIBUTION.—Gulf of California at Puerto Peñasco, Bahía de Los Angeles, Bahía Kino, Topolobampo, Bahía San Evaristo and Isla Espiritu Santo,

0-42 m; California, Newport Bay, 0-2 m.

Podocerus spp.

Podocerus brasiliensis.—J. L. Barnard, l962a:67.

P. cristatus.—J. L. Barnard, l962a:67.

REMARKS.—The true P. brasiliensis from Brazil

and true P. cristatus from Australia have not been

clarified recently, although they have been identified from widely spread localities throughout the

world. Until they have been analyzed satisfactorily

there is little purpose in continuing to identify

these species from places outside their type-areas.

The material at hand will require clarification. A

variety of cuspidation patterns occurs on male

gnathopod 2, mixing both brasiliensis and cristatus

forms and gradations between. All but a few specimens bear the brasiliensis body form.

MATERIAL.—The brasiliensis form: SCO 14 (3),

SCO 15 <3), SCO 18 (3); TOP 2 (2), TOP 3 (16);

PAZ 24 (26); GAL 120 (2); DAW 19 (1); cristatus

form: SCO 18 (1), PAZ 12 <l).
Appendix

Station Data

SCO, KNO, TOP, PAZ, BRU, and SNY samples

collected by J. L. Barnard and associates from the

Gulf of California at Puerto Peñasco, Bahía Kino,

Topolobampo, and region of La Paz on Baja

California.

SCO 1, Puerto Peñasco, 17 October 1970, intertidal, general

algal wash, 12 rocks, 2 liters algae.

SCO 2, Puerto Peñasco, 18 October 1970, intertidal, thin

layer of turf algae on platform. No amphipods.

SCO 3, Puerto Peñasco, 18 October 1970, intertidal, one rock

on moist substrate {shell sand) in platform zone. No

amphipods.

SCO 4, Puerto Peñasco, 18 October 1970, intertidal, 0.02 m2

of sandy algal turf. No amphipods, tanaids only.

SCO 5, Puerto Peñasco, 18 October 1970, intertidal, 1 piece

of algae.

SCO 6, Puerto Peñasco, 18 October 1970, intertidal, shale

rock 51 × 33 cm, 0.02 m^ of lower surface covered with

sponges and Amaroucium.

SCO 7, Puerto Peñasco, 18 October 1970, intertidal, 0.02 m2

of brown alga in wet pool.

SCO 8, Puerto Peñasco, 18 October 1970, intertidal, 0.02 m2

of sandy Padina in wet pool.

SCO 9, Puerto Peñasco, 14 November 1970, 29-38 m, shrimp

trawl, coll. J. R. Hendrickson and Fidel Mendoza Lopez

on trawler, squid embryos.

SCO 10, Puerto Peñasco, 23 February 1971, intertidal, Sargassum sp. (thin form), 0.5 liter.

SCO 11, Puerto Peñasco, 23 February 1971, intertidal, one

red alga, cf. C·igartina johnstonü, hanging on rock.

SCO 12, Puerto Peñasco, 23 February 1971, intertidal, wash

cf. Coìpomenia sp.

SCO 13, Puerto Peñasco, 23 February 1971, intertidal, 0.005 m2

of zoantharians.

SCO 14, Puerto Peñasco, 24 February 1971, intertidal, rocks

at low water, washed.

SCO 15, Puerto Peñasco, 24 February 1971, intertidal, sponge,

cf. Leucetta losangelensis.

SCO 16, Puerto Peñasco, 24 February 1971, intertidal, bryozoan turf and sponges on underhang.

SCO 17, Puerto Peñasco, 24 February 1971, intertidal, 0.02 m2

sandy algal turf free of Coìpomenia near lowest tide level.

SCO 18, Puerto Peñasco, 24 February 1971, intertidal mix of

Padina and brown Viva on cataract and sandy broken rock.

SCO 19, Puerto Peñasco, 25 February 1971, intertidal, general

wash of Padina sp. plus Sargassum canouii (recent winter

bloom).

SCO 20, Puerto Peñasco, 25 February 1971, intertidal, 0.02 m*

algal turf, same place as SCO-2, tanaids only.

SCO 21, Puerto Peñasco, 25 February 1971, intertidal, sprigs

of Sargassum johnstonii (about 1 year old), with conspicuously attached sandy agglutinations.

SCO 22, Puerto Peñasco, 25 February 1971, intertidal, sprigs

of Sargassum canouii (new bloom), sandy agglutinations

present only on axils.

KNO 1, Bahía Kino, 31 October 1970, 0-2 m, mass wash,

brown algae, corallines, sponge, Amaroucium, gorgonians,

scuba diving, coll. J. R. Hendrickson, J. Wood, F. Wood.

KNO 2, Bahfa Kino, 1 November 1970, intertidal, 0.02 m2

small brown algae, barnacles on platform 0.67 m above

water. (Alga turns green in formaldehyde.)

KNO 3, Bahía Kino, 1 November 1970, intertidal, 0.02 m2

Enteromorpha on crustose nobs 0.67 m above low water.

KNO 5, Bahía Kino, 1 November 1970, intertidal, 0.02 m2

Viva tufts on knobbly surface.

KNO 6, Bahía Kino, 1 November 1970, intertidal, rock wash

at splash zone, rocks bare above, with holothurians and

tube worms underneath.

KNO 7, Bahía Kino, 1 November 1970, intertidal, algae in

hermit crab zone, 2 specimens of Elasmopus sp., 3 specimens of Hyale sp.

KNO 8, Bahía Kino, 1 November 1970, intertidal, brine pool

at upper level, 1 amphipod, 5 worms, vinegaroons, flying

insects.

KNO 9, Bahía Kino, 1 November 1970, intertidal, combined

scrapings of Viva and Enteromorpha on knobbly surface,

coll. J. R. Hendrickson. No amphipods.

KNO 10, Bahía Kino, 1 November 1970, intertidal, general

gravel wash including clumps of Enteromorpha, coll. J. R.

Hendrickson.

TOP I, Topolobampo, 25 November 1971, 1.6 km seaward of

town, upper intertidal, shale and flint rocks washed.

TOP 2, Topolobampo, 25 November 1971, 1.6 km seaward of

town, 1 m, tunicates and sponges on rocks.

TOP 3, Topolobampo, 25 November 1971, 1.6 km seaward of

town, 1 m, tunicates and sponges on rocks, whole rocks

washed.

PAZ 1, Bahía Pichilinque, La Paz, 27 November 1971, intertidal, masses of dead red algae adjacent to mangrove

swamp. Mostly nebalians and snails.

PAZ 2, Bahía Pichilinque, La Paz, 27 November 1971, intertidal, open pebble reef, 0.5 liter red algae washed.

PAZ 3, Isla Espiritu Santo, Bahía San Gabriel, 28 November

1971, 0.3-1.0 m, rocks covered with finely anastomosed

brownish red alga in band of phyciferous rocks 3-7 m wide.

138
NUMBER 271 139

PAZ 4, Isla Espiritu Santo, Bahía San Gabriel, 28 November

1971, old oyster ponds packed with Caulerpa sertulartoides,

wash. No amphipods.

PAZ 5, Isla Espiritu Santo, Bahía Ballenas, 28 November 1971,

1 m, wash of 6 rocks covered with Caulerpa sertularioides.

PAZ 6, Isla Espiritu Santo, Isla Partida rift, 29 November

1971, intertidal, wash of 4 rocks covered with Caulerpa

sertularioides and brownish red alga.

PAZ 7, Isla Espiritu Santo, Isla Partida rift, 29 November

1971, intertidal, wash of 30 pebbles and substrate on shingle

beach.

PAZ 8, Isla Partida, Bahía Cardonale, 29 November 1971,

intertidal, old oyster weirs on N side, wash of brownish

red algae on sand bottom.

PAZ 9, Isla Partida, Bahía Cardonale, 29 November 1971,

intertidal wash of old coral rubble cobbles covered with

Caulerpa sertularioides.

PAZ 10, Isla San Francisco, east bay, 30 November 1971, wash

of intertidal cobbles covered with green felt turf.

PAZ 11, Isla San Francisco, east bay, 30 November 1971, 2 m,

wash of 4 liters of Caulerpa sertularioides on sand bottom.

PAZ 12, Bahía San Evaristo, 1 December 1971, intertidal, SE

point, wash of 4 large muddy rocks covered with short

brownish felt turf and green patches of algal scum.

PAZ 13, Bahía San Evaristo, 1 December 1971, intert¡dal, NE

point, wash of nonmuddy rocks covered with short tufted

brownish red algae.

PAZ 14, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 200 cc closepacked green alga.

PAZ 15, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 200 cc of Codium.

PAZ 16, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 200 cc wiry brown alga.

PAZ 17, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 200 cc of small Padina.

PAZ 18, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, wash of 1 liter golden Pelvetiops¡s(Y) with golden shrimp.

PAZ 19, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 200 cc of small polychaete

tubes in sand at low water.

PAZ 20, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 1 liter "pinetree" red alga.

PAZ 21, 11 km E of Cabo San Lucas at Bajo Colorado Hotel.

4 December 1971, intertidal, 1 liter heavy green subcalcareous algae.

PAZ 22, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intert¡dal, 1 liter curl-stripe golden

bronze alga in tidepool of barnacle zone.

PAZ 23, 11 km E of Cabo San Lucas at Bajo Colorado Hotel,

4 December 1971, intertidal, 300 cc green scum algae at

high tide level, probably Enteromorpha and Viva.

PAZ 24, Bahía Concepción, 24 km on east side from entrance,

6 December 1971, 1 m, wash of 6 rocks covered with short

cropped brownish red alga in zone 2 m wide.

PAZ 25, Bahía Concepción, 16 km on east side from entrance,

6 December 1971, 1 m, wash of almost dead ?Pelvetiopsis on

rocks.

PAZ 26, Bahía Concepción, 16 km on east side from entrance,

6 December 1971, beach wrack of ?Pelvetiopsis, wash of 3

liters. No amphipods.

BRU 1: Bahía San Carlos, 24 January 1971, rocky intertidal,

collected by Dr. R. C. Brusca.

BRU 2: Guaymas, 25 January 1971, rocky intertidal, collected

by Dr. R. C. Brusca.

SNY 1: Puerto Peñasco, 27 January 1975, in hermit crab shells,

collected by Elaine Snyder.

GAL, ECU(ador), COCOS, and PAN, samples collected by J. L. Barnard and associates from the

Galapagos Islands, Ecuador, and Cocos Island,

especially Isla Santa Cruz and Tower Island (Isla

Genovesa); note that 36 samples (GAL 122-157)

from Isla San Salvador, Isla Baltra, and northern

Isla Santa Cruz were not analyzed.

GAL 101, Isla Santa Cruz, Academy Bay, 23 January 1964,

area east of lower bodega (warehouse), alga Cystophyllum?',

4 liters washed, strongly wave-dashed, strained through 32

Tyler mesh, 15:06 hours, 0.0 tide.

GAL 102, same data as GAL 101, 20 rocks in tidepool

washed, 30° C water temperature in tidepool partially

sea-connected.

GAL 103, same data as GAL 101, mangrove t¡depool 305 m

from sea near lower bodega, Hyale, and palaeomonid

shrimps, tidepool fluctuates with tide but not sea-connected at surface.

GAL 105, same data as GAL 101, wash of 25 Eucidaris collected subtidally by Durham and Abbott, many amphipods on spines.

GAL 106, Isla Santa Cruz, 24 January 1964, Academy Bay,

offshore of lower bodega in skiff, underwater rocks, 0.3-

1.0 m deep, inside reef, wash of 8 liters of Padina in

plankton net for tanaids; sort in 120 mesh screen.

GAL 107, same as 106 but screen sort of 32 mesh of remainder of sample.

GAL 108, same as 106, wash of 3 underwater rocks covered with short tufted brownish algae, some sponge, numerous gastropods, hermits.

GAL 109, Isla Santa Cruz, Academy Bay, 25 January 1964,

offshore of lower bodega in skiff, underwater rocks, 1.0-

1.3 m, 32 mesh screen.

GAL 110, same as GAL 109, dredge, sorted in 32 mesh,

coral and shell sand, a few cumaceans.

GAL 111, same as GAL 109, coral sand bottom dredged, 32

mesh, ghost shrimps and enteropneust bottom.

GAL 112, Tower Island (Genovesa), Darwin Bay, 29 January 1964, intertidal, scrapings of zoanthids, 5 juvenile

amphipods.

GAL 113, same as GAL 112, wash of about 30 small rocks

and fragments of large slabs of intertidal lava overturned

by crowbar.

GAL 114, same as GAL 112, rock 46 × 20 × 30 cm covered with 3-5 cm of red algal tufts, rock from lateral

channel on side of sandbed in quiet water 1 meter deep.

GAL 115, same as GAL 112, wash of Pocillopora-Porites

heads recovered bv divers.
140

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

GAL 116, Tower Island, Darwin Bay, 30 January 1964, intertidal, wash of 5 rocks and 5 pieces coral head on outer

coast.

GAL 117, same as GAL 116, half liter of algae from trapped

pool hole behind volcanic dike connected to sea by

tunnels.

GAL 118, Isla Santa Cruz, Academy Bay, 4 February 1964,

intertidal adjacent to Nelson cottage, wash of ?Cystophyl·

him.

GAL 119, same as GAL 118, 3 small rocks in 0.3 m water

depth.

GAL 120, same as GAL 118, 1 liter of red algae in about

0.66 m

GAL 121, Isla Santa Cruz, Academy Bay, 5 February 1964,

southwest end, in Braza type lagoon, wash of one liter of

Caulerpa, Tyler 32 mesh.

ECU 2, 3, Ecuador, Punta Centinela, 5 March 1964, N of

Libertad, half low tide on sandstone reef point, unscreened samples of miscellaneous algae.

COCOS 1, Cocos Island, Chatham Bay, 8 March 1964, wash

of Pocillopora coral and attached algae, 1 m.

COCOS 2, same as COCOS 1, but dried moss above high tide

on cliff. No amphipods.

COCOS 3, same as COCOS 1, wash of rocks covered with

tubes and sand, mud, and encrusted coral heads. Largely

shrimp recovered.

COCOS 9, same as COCOS 1, 5 March 1964, wash of intertidal rocks with minute turf and tiny sponges.

PAN 14, Panama, Pacific side of Canal Zone, Diablo Swimming Club, 17 April 1955, wharf, wash of sponges and

fouling matter from inner brackish tub.

DAW samples collected by the late Dr. E. Yale

Dawson from the Galapagos Islands

DAW 1, Isla Santa Cruz, Academy Bay, 20 February 1962,

wash of Caulerpa sp.

DAW 2, Isla Espagnola (Hood Island), 3 February 1962,

algal turf wash in extreme surf.

DAW 3, Isla Santa Cruz, Academy Bay, 17 February 1962,

near town landing, wash of intertidal rocks with thin

turf cover of articulated corallines (Jenia amphiroa).

DAW 4, Isla Santa Cruz, Academy Bay, 16 February 1962,

intertidal, Hypnea wash, 2 sets.

DAW 5, Isla Santa Cruz, 11 February 1962, intertidal wash

of Padina durvilliae.

DAW 6, Isla Santa Cruz, Punta Nunez, 21 February 1962,

intertidal, wash of Cystophora galapagaea.

DAW 7, same as DAW 6, wash of Ochtodes crockeri.

DAW 8, Isla Santa Cruz, off Academy Bay, Isla Coamaño

(Jensen Island), 13 February 1962, wash of Cystophora.

DAW 9, same as DAW 4, intertidal rock wash.

DAW 10, same as DAW 2, second sample.

DAW 11, Isla Santa Cruz, Academy Bay, filter tide pool

back of beach next to Distichlis grass, blue green algal

wash. No amphipods.

DAW 12, same as DAW 1, wash of Gigartina.

DAW 13, same as DAW 1, wash of rocks, Ulva and Ecto·

carpus. `

DAW 14, Isla Santa Cruz, no date, wash of Cystophora. No

amphipods.

DAW 15, same as DAW 5, wash of Cladophoropsis from

under mangroves.

DAW 16, Isla Santa Cruz, Bahía Tortuga, 9 March 1962,

algal wash from warm quiet lagoon.

DAW 17, same as DAW 8, wash of Sargassum and Spathoglossum.

DAW 18, same as DAW 8, intertidal, wash of Plocamium.

DAW 19, same as DAW 3, wash of small coral head.

DAW 20, same as DAW 1, wash of Ochtodes.

DAW 21, same as DAW 5, wash of Ulva.

DAW 22, same as DAW 4, wash of Centroceras turf.

DAW 23, same as DAW 4, wash of Ectocarpus on upper

rocks.

DAW 24, Isla Santa Cruz, Academy Bay, 18 February 1962,

wash of rocks covered with Ulva, in mangrove cover,

intertidal.

DAW 25, same as DAW 8, wash of Cladophoropsis in mangrove ground cover; 1 juvenile of Hyale sp.

DAW 26, same as DAW 14, wash of intertidal rocks covered

with short "Ulva."

DAW 27, same as DAW 4, rock wash in upper intertidal.

DAW 28, same as DAW 5, wash of Blossevillea galapagensis.

DAW 29, same as DAW 6, wash of Glossophora Knuthii.

DAW 30, same as DAW 1, wash of Laurencia Hancockii.

DAW 31, Isla Santa Cruz, halfway between Tortuga Bay

and Academy Bay, 22 February 1962, 6-9 m, wash of

algae and rocks.

DAW 32, same as DAW 7, wash of Caulerpa racemosa.

DAW 33, same as DAW 5, wash of Laurencia sp. turf.

DAW 34, same as DAW 1, wash of Ga¡exora.

DAW 35, same as DAW 1, wash of coral head.

DAW 36, Isla Santa Cruz, bay southwest of Academy Bay,

22 February 1962, 6 m, wash of Padina.

DAW 37, same as DAW 3, wash of Hypnea.

DAW 38, same as DAW 1, wash of Glossophora.

DAW 39, same as DAW 24, wash of Bostrychia, beach hoppers only.

DAW 40, same as DAW 4, intertidal rock wash.
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144

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1879. New Zealand Crustacea, with Descriptions of New

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1941. New or Noteworthy Amphipods from the North

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1901. Contributions to the Malcostracan Fauna of the

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146 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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Index

(Page numbers of principal accounts in italics)

achire, Lembos, 25, 26, 27

Acuminodeutopus, 36, 38

periculosus, 23

alata, Leucothoe, 128

Allorchestes, 90, 114, 119

anceps, 114

angusta, 90, 91

angustus, 91, 94

aquilina, 126

aquilinus, 126

bellabella, 90, 91, 94

carinata, 90, 91, 96

carinatus, 96

compressa, 90

frequens, 114

hawaiensis, 122

japonica, 115

malleolus, 91

malleolus carinatus, 96

novizealandiae, 90

ochotensis, 120

oculatus, 91

plumicornis, 120 121

plumulosus, 114

ptilocerus, 121

vladimiri, 91

Aloiloi, 34

Ampelisca lobata, 13,

schellenbergi, 14

Amphideutopus, 36

Amphilochus neapolitanus, 14

picadurus, 14

Amphithoe aquilina, 126

inermis, 42

Ampithoe guaspare, 16, 20

kulafi, 21

lindbergi, 20

plumulosa, 18

plumulosa tepahue, 18

pollex, 18, 20, 21

ramondi 20

tahue, 18, 20, 21

vacoregue, 18, 21

Anamixis linsleyi, 21

anceps, Allorchestes, 114

Hyale, 99, 114

Anchialella, 53, 54

vulcanella, 53, 54

angusta, Allorchestes, 90, 91

angustus, Allorchestes, 91, 94

antarctica, Polycheria, 38

antennatus, Elasmopus, 60, 61, 67, 79

Neogammaropsis, 61

aquilina, Allorchestes, 126

aquilina, Allorchestes, 126

Amphithoe, 126

Hyale, 126

Parhyale, 120, 126

armatus, Microdeutopus, 35

Aruga dissimilis, 130

holmesi, 130

macromerus, 130

Aspidophoreia, 90

audbettius, Lembos, 25

Audulla chelifera, 35

aviculae, Chevalia, 24

ayeli, Hyale, 111

baciroa, Citanopsis, 14

baconi, Corophium, 24

bampo, Elasmopus, 57, 59,60, 61, 64, 68

bartschi, Pontogeneia, 41, 49

Batea rectangulata, 21

susurrator, 21

transversa, 23

transversa coyoa, 23

bellabella, Allorchestes, 90, 91, 94

Bemblos macromanus, 25

bishopae, Hyale, 115

brasiliensis, Ericthonius, 24, 98

Podocerus, 137

Pyctilus, 24

brevipes, Hyale, 122, 123

californica, Hyale, 98, 99, 115 116, 118

Caliniphargus sulcus, 88

calliactis, Elasmopus, 59, 60

canalina, Hyale, 98, 101,102, 103

carinata, Allorchestes, 90, 91, 96

carinatus, Allorchestes, 96

Cedrophotis, 31

Ceradocus paucidentatus, 54

Cheiriphotis megacheles, 23

cheli£er, Microdeutopus, 35

chelifera, Audulla, 35

Chevalia aviculae, 24

chevreuxi, Hyale, 113, 114

Chevreuxius, 38

chinarra, Maera, 83, 86, 87

chinipa, Microjassa, 727

cinaloanus, Zoedeutopus, 35, 38

claustris, Microjassa, 128

Colomastix pusilla, 23

species, 23

compressa, Allorchestes, 90

concavus, Lembos, 25, 30

consil¡orum, Najna, 118, 119

Corophium baconi, 24

crassicornis, Hyale, 115

cristatus, Podocerus, 137

dartevillei, Hyaloides, 122

darwini, Hyale, 98, 99, 101, 102, 103

diffidentia, Meximaera, 88

dissimilis, Aruga, 130

Lysianassa, 130

Lysianopsis, 130

Nannonyx, 130

Dulzura gal, 54

hamakua, 57

eburnea, Parhyale, 120, 122

ecuadorensis, Elasmopus, 60, 64, 67

Elasmopus, 57, 58, 59, 60, 98

antennatus, 60, 61, 67, 79

bampo, 57, 59, 60, 61, 64, 68

calliactis, 59, 60

ecuadorensis, 60, 64, 67

hawaiensis, 67

holgurus, 60, 82

hooheno, 59

mayo, 60, 67, 68, 82

mutatus, 60, 68

ocoroni, 60, 68

piikoi, 59

pocillimanus, 57, 59, 79

pseudaffinis, 59

rapax, 57, 58, 59, 60, 61, 64, 68, 69, 72,

73, 77, 79, 82

serricatus 60, 73

temori, 60, 75

tiburoni, 57, 60, 68, 77

tubar, 60, 79

zoanthidea, 60, 77, 79, 82

elephant¡s, Photis, 30

Ericthonius brasiliensis, 24, 98

Eriopisa, 53, 54

Eurystheus tonichi, 25

fasciculata, Nicea, 126

fasciger, Parhyale, 122, 123

147
148

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fascigera, Parhyale, 119, 120, 122, 123

fascigera penicillata, Parhyale, 119, 123

frequens, Allorchestes, 114

Hyale, 98. 99, 101, 102, 104, 107, 114

Metaphoxus, 133

fulanus, Podocerus, 134

gal, Dulzura, 54

galapagoanus, Heterophlias, 131

Galapsiellus, 58

leleuporum, S3

Gammaropsis, 81

lina, 85

photosimilis, 31

semichelatus, 35

tonichi, 25

Gammarus spinicarpus, 129

G¡tanopsis badroa, 14

vilordes, 16

Gondogeneia, 41

gracilis, Hyale, 121

grandicornis, Hyale, 98, 114, 115, 116,

118

grandicornis bishopae, Hyale, 115

Grandidierella, 38

guasave, Hyale, 99, 111

guaspare, Ampithoe, 16, 20

hamakua, Dulzura, 57

hancocki, Microdeutopus, 30

h?waiensis, Allorchestes, 122

Elasmopus, 67

Hvale, 122, 123

Parhyale, 119, 120, 722, 126, 127

Heterophlias galapagoanus, 131

seclusus escabrosa, 131

holgurus, Elasmopus, 60, 82

holmesi, Aruga, 130

Lysianassa. 130

Lysianopsis, 130

honoluluensis, Hyale, 113, 114

hooheno, Elasmopus, 59

humboldti, Hyale, 99, 115, 116,118

Hyale, 90, 98, 119, 120

anceps, 99, 114

aquilina, 126

ayeli, 111

bishopae, 115

brevipes, 122, 123

californica, 98, 99, 115, 116, 118

canalina, 98, 101,702,103

chevreuxi, 113, 114

crassicorn¡s, 115

darwini, 98, 99, 101, 102, 103

frequens, 98, 99, 101, 102, 104, 107,114

gracilis, 121

grandicornis, 98, 114, 115, 116,118

grandicornis bishopae, 115

guasave, 99, 77¿

hawaiensis 122, 123

honoluluensis, 113, 114

humboldti, 99, 115, 116, 118

inyacka, 122, 127

iole, 111

iwasa, 121

iwasai, 121

longicornis, 114

macrodactyla, 113, 114

nigra, 98, 104, 107, 114

nilssoni, 122

novaezealandiae, 98, 115, 118

ochotensis, 120

penicillata, 121

plumicornis, 119

plumulosa, 99 111, 114, 121

plumulosus, 114

pontica, 122

pugettensis, 115

rubra, 98, 707, 102 103, 104, 114

rupicola, 98, 115

schmidt¡, 101

species, 98, 101,775

stolzmanni, 122

thomsoni, 114, 115

trifoliadens, 122

yaqui, 98, 101,104, 107, 111, 114

zuaque, 98, 108

Hyaloides dartevillei, 122

inaequipes, Maera, 83, 86, 88

inermis, Amphithoe, 42

Pontogeneia, 38, 41, 42

intermedia, Pontogeneia, 41, 42

intermedius, Lembos, 27, 30

inyacka, Hyale, 122, 127

Parhyale, 119, 120, 122, 727

iole, Hyale, 111

iwasa, Hyale, 121

iwasai, Hyale, 121

Parhyale, 120, 121, 122

japonica, Allorchestes, 115

kitamati, Najna, 775

Konatopus, 36, 38

kulañ, Ampithoe, 27

kurilensis, Parhyale, 120

leleuporum, Galapsiellus, 83

Paraniphargus, 83

Lembos achire, 25, 26, 27

audbettius, 25

concavus, 25, 30

intermedius, 27, 30

macromanus, 25, 30

tehuecos, 25, 26, 27, 30

Leucothoe alata, 128

spinicarpa, 128

Leucothoides pacifica, 130

pottsi, 130

yarrega, 750

lina, Gammaropsis, 31

lindbergi. Ampithoe, 20

Linguimaera, 88

linsleyi, Anamixis, 27

lobata, Ampelisca, 13

longicornis, Hyale, 114

longleyi, Pontogeneia, 41, 46

Lysianassa dissimilis, 130

holmesi, 130

macromera, 130

Lysianopsis dissimilis, 130

holmesi, 130

macromera, 130

macrocoxa, Microjassa, 128

macrodactyla, Hyale, 113, 114

macromanus, Bemlos, 25

Lembos, 25, 30

macromera, Lysianassa, 130

Lysianopsis, 130

macromerus, Aruga, 130

Maera chinarra, 83, 86, 87

inaequipes, 83, 86, 88

pacifica, 86

quadrimana, 86

rathbunae, 85, 87

reishi, 83, 85, 86, 87

simile, 88

Mallacoota, 58

malleolus, Allorchestes, 91

malleolus carinatus, Allorchestes, 96

mayo, Elasmopus, 60, 67, 68, 82

megacheles, Cheriphotis, 23

Megamphopus, 23

Megamphopus, 35

megacheles, 23

Melita palmata, 88

sulca, 88

Metaphoxus frequens, 133

Meximaera diffidentia, 88

Microdeutopus, 35, 36, 38

armatus, 35

chelifer, 35

hancocki, 30

schmitti, 30

Microjassa chinipa, 727

claustris, 128

macrocoxa, 128

minuta, Pontogeneia, 42, 46

mutatus, Elasmopus, 60, 68

Najna consiliorum, 118, 119

kitamati, 118

Nannonyx dissimilis, 130

nasa, Pontogeneia, 49, 53

Tethygeneia, 42 49, 53

neapolitanus, Amph¡lochus, 14
NUMBER 271 149

Neogammaropsis antennatus, 61

Neomegamphopus, 34, 35, 36, 38

Neomicrodeutopus, 38

Nicea fasciculata, 126

nudicornis, 126

plumicornis, 121

rubra, 101

rudis, 126

n¡gra, Hyale, 98, 104, 107, 114

nilssoni, Hyale, 122

novaezealandiae, Hyale, 98, 115, 118

novizealandiae, Allorchestes, 90

nudicornis, Nicea, 126

ochotensis, Allorchestes, 120

Hyale, 120

Parallorchestes, 119, 120

Parhyale, 120

ocoroni, Elasmopus, 60, 68

oculatus, Allorchestes, 91

opata, Pontogeneia, 41, 42, 46

osborni, Polycheria, 38

pacifíca, Leucothoides, 130

Maera, 86

palmata, Melita, 88

Paradexamine, 58

Parallorchestes, 119, 120

ochotensis, 119, 120

Paraniphargus, 54

leleuporum, 83

Paraphoxus spinosus, 133

Parhyale, 114, 119, 120

aquilina, 120, 126

eburnea, 120, 122

fasciger, 122, 123

fascigera, 119, 120, 122,123

fascigera penicillata, 119, 123

hawaiensis, 119,120,122,126,127

inyacka, 119, 120, 122,127

iwasai, 120,121, 122

kurilensis, 120

ochotensis, 120

pencillata, 120, 122, 123, 126

plumicornis, 119, 120, 121

species, 120, 121, 126

species of Bulycheva, 120,127

zibellina, 119, 120,121, 127

Pariphinotus, 131

tuckeri, 131

paucidentatus, Ceradocus, 54

pencillata, Hyale, 121

Parhyale, 120, 122,123, 126

periculosus, Acuminodeutopus, 23

Photis, 31

elephantis, 30

photosimilis, Gammaropsis, 31

picadurus, Amphilochus, 14

piikoi, Elasmopus, 59

Pleonexes species, 21

plumicornis, Allorchestes, 120, 121

Hyale, 119

Nicea, 121

Parhyale, 119, 120,72/

phimulosa, Ampithoe, 18

Hyale, 99, 111, 114, 121

plumulosa tepahue, Ampithoe, 18

plumulosus, Allorchestes, 114

Hyale, 114

pocillimanus, Elasmopus, 57, 59, 79

Podoceropsis, 35

Podocerus brasiliensis, 137

cristatus, 137

fulanus, 134

species, 134, 137

pollex, Ampithoe, 18, 20, 21

Polycheria antarctica, 38

osborni, 38

pontica, Hyale, 122

Pontogeneia, 38, 41

bartschi, 41, 49

inermis, 38, 41, 42

intermedia, 41, 42

longleyi, 41, 46

minuta, 42 46

nasa, 49, 53

opata, 41, 42, 46

quinsana, 41, 42, 53

rostrata, 41, 49

Posophotis, 30

seri, 31

pottsi, Leucothoides, 130

Psammoniphargus, 54

pseudaffinis, Elasmopus, 59

Pseudeurystheus, 31

Pseudomegamphopus, 34

ptilocerus, Allorchestes, 121

pugettensis, Hyale, 115

pusilla, Colomastix, 23

Pyctilus brasiliensis, 24

quadrimana, Maera, 86

quinsana, Pontogeneia, 41, 42, 53

Tethygeneia, 42, 53

ramondi, Ampithoe, 20

rapax, Elasmopus, 57, 58, 59, 60, 61, 64,

68, 69, 72, 73„ 77, 79, 82

rathbunae, Maera, 85, 87

rectangulata, Batea, 21

reishi, Maera, 83, 85, 86, 87

rostrata, Pontogeneia, 41, 49

rubra, Hyale, 98,101, 102 103, 104,114

Nicea, 101

Rudilemboides stenopropodus, 34

rudis, Nicea, 126

rupicola, Hyale, 98, 115

schellenbergi, Ampelisca, 14

sfhmidti, Hyale, 101

schmitti, Microdeutopus, 30

seclusus escabrosa, Heterophlias, 131

semichelatus, Gammaropsis, 35

seri, Posophotis, 31

serricatus, Elasmopus, 60, 73

simile, Maera, 88

species, Colomastix, 23

Hyale, 98, 101, 115

Parhyale, 120, 121, 126

Pleonexes, 21

Podocerus, 134, 137

species of Bulycheva, Parhyale, 120,127

spinicarpa, Leucothoe, 128

spinicarpus, Gammarus, 129

spinosus, Paraphoxus, 133

stenopropodus, Rudilemboides, 34

stoìzmanni, Hyale, 122

sulca, Melita, 88

sulcus, Caliniphargus, 88

susurrator, Batea, 21

t;ihue, Ampithoe, 18, 20, 21

tehuecos, Lembos, 25, 26, 27, 30

temori, Elasmopus, 60, 75

tepahue, Ampithoe plumulosa, 18

Tethygeneia, 41

nasa, 42, 49, 53

quinsana, 42, 53

thomsoni, Hyale, 114, 115

tiburoni, Elasmopus, 57, 60, 68, 77

tonichi, Eurystheus, 25

Gammaropsis, 25

topianus, Varohios, 34, 35

transversa, Batea, 23

coyoa, Batea, 23

trifoliadens, Hyale, 122

tubar, Elasmopus, 60, 79

tuckeri, Pariphinotus, 131

vacoregue, Ampithoe, 18, 21

Varohios, 34, 35

topianus, 34, 35

vilordes, Gitanopsis, 16

vladimiri, Allorchestes, 91

vulcanella, Anchialella, 53, 54

yaqui, Hyale, 98, 101,104, 107 111, 114

yarrega, Leucothoides, 130

zibellina, Parhyale, 119, 120,121, 127

zoanthidea, Elasmopus, 60, 77, 79, 82

Zoedeutopus, 35, 36, 38

cinaloanus 35, 38

zuaque, Hyale, 98,108

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